PT) A

PAGE

ACACIA, Peculiar foliage in . 727

Beddard’s Manii QII

Allen, J. A

— N rth American deer 755

—— Upland game birds reti

Angiosperms, Embryo sac of . . 777

Animals in the drift. line . 855

VESPA Buddin . 295

Anodon cialis of mido ..395

Mie 'M. H. GETIN

of Sisyra

Ants, Aiidh Oar ee eee Os

—— Guests of . 365

——— Key to a ee et Oy

Aster, Variations in = 242i. IRI

BavLEY, W. S. Tillman's Miner-

— Weinschenk’s Rockforning

mine cea

Bibio fuere tuc LAM S IBI

Bigelow, M. Jordan and

Heath's Animal forms .

Pratt's Invertebrate tenar 502

Bird migration and wind . 735

tc rae ; x *

Birds, Notes on - 503

Blood flow in Lumbricus < ips

Botanical notes . 16

Brues, C. T. Gunis of Tenia

legionary ants. . eek a OG

Bufo, Gastrulation in . . . . . 527

CAMPBELL, D. H. Affinities of

certain dicotyledons ^ 7

——— Anew work on Pinoa 72

—— Embryo sac of angiosperms . 777 |

Cancer magister, Breeding o poi

Carboniferous fishes, New .

. 849

PAGE

Cats' paws, Abnormality in ; Ss

Chamberlin, R. V. Geophiida of

Utah .

Chilopoda of Ut ih - 473

Cockerell, T. D. A. eae sad

insects in New Mexico

Hymenoptera of Hawaii . . 591

—— The cee gga! bape

Depressaria. . plv

—— Ridgway's Finches . ue uA c:

Twenty new pocket mice . . 587

Coe, W. R. A new genus of Ne-

s rimas

- 758

Nem i die. of iube 43I

Collateral sheets in annelids .

— of pigeon flowers . . 203

Co ck, J. H., and Chujiro, K

era of head of insects . . 13

Corymorpha palma . D

Cushman, J. A.

alized stages

of growth in New England plants 865

DavENPORT, C. B. Gardiner's Mal-

dives and Laccadives . 819

Dean, B. Biometric evidence in

e problem of the ori

limbs of vertebrates . . 837

—— Origin of vae fade of

vertebrates s; 9267

De Bary's Babtesfs - 340

Dicotyledons, Affinities j

Diemyctylus, Sexual feature | in 643

Dinoflagellata in California . . . 187

D e OL Y (5.9. OUI

owning, E. R. Variation in posi-

tion of muscles in Anodonta . 395

Duerden, J. E. Aggregated

rals. . 46t

PAGE

EASTMAN, C. R. New carbonifer-

ous fishes . . 849

Position of ibis iva ple - 505

— Unpublished observations on

Paleozoic fishes EA Orestes St.

ohn .

Eckel, E. c. quis variation in

garter snake . . 481

Embryo sac of angiosperms . e riri

Emery, C. Key to' genera of For-

micidz a dup d ru MM

FisHES, Colors of . : . 803

New carboniferous . . 849

Pioners and insects in New in

ios of inim. 20

Formica cinerea, Occurrence of, in

erica

Formica rufibarbis Fy oma al of,

in America

. 947

ifie Key t to genera . 797

CEU ur. 653, 849

pad W.F. The habits of tor-

Gasteropoda, S Studies a.

Gastrulation ja

Gibbs, R. E " Phos as a

beach-buil I

- . 917

Gifts of the id ien à : T

Grabau, A. W. Studies of Casters:

poda .

» O17

HAFTED flint knives . .

Hargitt, C. W. P a N isang

of Woods Hole . - 549

Hazen, A. P. Regeneration. in

Hydractinia and errs : gr

Heath, H. Breeding habits

Cancer magister . . Ta

Henshaw, S. Habits of i Sects . 407

Hilton, W. A. A structure con-

nected with mating of Dente

lus

terous | *

Homoplasy as a i - homology 2

Howe, F. usc etaed pen

: Leavitt's Outlin

— A :

INDEX.

Hrdlička, A. Instances of divi-

sion of the malar bone

Hydra, regeneration and hiersiogh

cal changes in :

Hydroids, Faisean in

INsECTS and flowers in New

Mexico

and iir distribution yw wid

skeleton of head

Iron and living matter

JENNINGS, H. S. Structure and

metamorphosis of Actinotrocha

Johnson, H. P. Collateral bud-

ding in Annelids .

Johnston, J. B. and ceo s.

Course of blood flow in

Lumbricus . .

Jordan, D. S. Boulenger's clas

cation of trachinoid fishes

Fishes of jira i

—— — Hawaiian a :

—— New Pse

—— Notes on pene 6 164 336

—— Oarfish in Californ

—— Paired limbs of bissl A

—— Synonymy of the fish skele-

- 159,

TON 35 ag X ee

— — The colors of fishes.

KANGAROO rats, Habits

Kellogg, V. L. visse and

homologies of mouth pa:

insects

King, H. D. Ga isi la Bufo

Knives, Prehistoric hafte:

Kofoid, i A. etm of tho

Azor

— bab a the Pacific

Law of adaptive Hdiation ;

. 68

527

PAGE

INDEX. v

PAGE

Lovell, t H. Reply to Professor

Rober » B32

ee Cutie of blood iow’ in 317

MADREPORE corals, Colonies in . 461

Malar bone, Division of . . . . 273

Marsh mice . . 661

McClendon, = F. Life history of

Ulula hyal

McDougall’s Tus: "sd ; pe

Meek, S. E. Contribution to

museum technique . . . . . 53

Mechanism and vitalism . . . . 154

Mouth parts of insects . 683

Museum technique. . . . . + 53

Myriapods of Utah 473

NEEDHAM, J. G. A course in

insect anatomy oo

——— Remarkable occurrence of tié

fly Bibio fraternus . 2 Se

Nemertean sisi of SM ep Ri!

Notes and literature:

Biograph V I É

Biology . 3» 154, 329, 665

Botany, 72, 165, 247, 340, 408, 592,

671, 761, 822, 981

Petrography 255, 344, 766

Zoology, > I st 243, 330 401, 497,

5, 667, 755, 819, 911-

gea A. E. pem

region

d s Faai of ore

land -e 5 V s. o 499

Osborn, H. F. Homoplasy as a

law of latent or potential homol-

ogy 259

— bus of A SMS canes -353

Osborn, H. L. Anatomy of a

double calf i515. . 001

Osmia, New speci of 816

PAMMEL, L. H. Garden beans . 593

Parker, G. H. Circulation in the

nervous s : pe 65

—— Dispersal UL Gigustin lucis : pe

Gegenbaurs Comparative

Ce ae 4 x os os

, W. The Tremataspide . 379

derit P as a beach-builder . 101

Pierce, G. J. Pfeffer's Plant physi-

ology - 594

Planaria, Regeneration in - 633

Plant growth, Stag

Pratt, H. S. resin of digsa-

etic Trematodes . 887, 953

Prest C Peculiar tolsa

In AÁcarin 0; 1223

Pteropus haldemani, aang of 201

Publications received, 80, 178, 350, 508,

QUARTERLY record, 172, 415, 678, 915

RADIATION, Law of adaptive .

Rand, H. W. Hair in Equidz

—— Osteology of flamingoes . . 588

E VA in Hydra.

Vascular system ~ € - 338

Secesodin and ological

changes in Hydra 561

Regeneration in Sisi + 193

Regeneration in Plana » 633

ehn, J. A. G. Standing of Piwo

pus haldemani . . 201

Reviews :

Agardh’s genera et ordines Al-

garum -413

Mahood Pussitic. Hynes:

optera of Haw - 59

grins s Tpit of

Pm Cyclopedia E horti-

culture . 71

Becker on Phoridz iua Hd vee

Beddard’s Mammalia . . . QII

osos onflatworms . . . d

Biometri

cendi s Cleudücatibe. ot

trachinoid fishes

Britton’s Manual of de dora

of the northern states . . 247

Busck on Depressaria . .` . 670

€ Mechanism ar

gre

Canpbets University taxi.

of botany . 671

vi INDEX.

PAGE PAGE

Reviews : Reviews:

e Methods in Leonard on basic rocks in

plant histology . 165 Matyland . . . . sana eee

Comstock a Kellogg's Ja Lühe’s Sporozoan studies . . 69

sect anatomy . - 500 MacDougall’s Plant physi-

Coulter and iodain” s GONE. X tee) a 94 GE

orphology of spermato- Marshall on hair in the

phytes : 72 Equide - 339

De Parys Bacteria e - 340 Meier's Hadeudion ate plant

Doelter’s Densities of d description . ee

magmas . 348 Merriam on eine mice... 1° 5Rg

Dresser’s Leccodite of Shefford Nott’ s Notophylla of Califor-

Mountain. . . 348 cwn

Driesch on organic mie k p Packie Life of ‘Camere’ - 495

Fabre’s Insect life . . 4 Pammel’s Grasses of Iowa . 249

Falkenberg’s Rhodome cin id Parke's Regeneration in Hydra 589

Ganong's Plant physiology . 76 Pfeffer's Plant physiology . . 595

Gardiner's Maldive and Lacca- Pratt's Invertebrate zoólogy . 502

dive archipelagoes. . . . 819 Ridgway's Birds of North

oólogy of the Maldive PEMEX 4s o s 6-333

and Laccadive archipelagoes 67 Roosevelt's Deer family . . 755

Gasser's Circulation in the Sacharoff’s Iron and living

nervous system. . . 65 matter

Gattinger’s Flora of atin 249 Sandys ind Vx Diii Up

Gegenbaur’s — land game birds . A iri

Anatomy . 4 567 Saunder's Alaskan Alp oy" gas

Griffon on Cuoi. € » 240 Schwartz on forest trees . . 250

acude Outlines of biology. 666 Seale’s Fishes of Guam . . 585

n electric lobes of tor- Seeliger's Deep sea animals . 668

edd Selous's Bird watching ; 54

Esatit s Tises i in winter. ded Shipleyand McBride's Zoblogy 243

Ikeda on Actinotrocha . . . 499 gore Osteology of -fla-

Irish’s Garden beans . - 593

Jackson on the blood aid Mit. Classtilentiin of the

of Bdellostoma 8 Enteropneusta . - $90

Jacobi's s Biopnghical regions . 157 Spurr on rock atfuctdte. £o MF

Jenkins and Kellogg's Nature Studer's Corals of the Pacific. 669

study . . 63 ——— Alcyonaria of the Azores 669

Jordan's Distribution of fishes 162 Thompson on Zygeupolia . . 758

m and Heath’s Animal Tillman’s Text-book of impor-

. 821 tant minerals and rocks 344

Kerr on the piirsi Tabe of 2 Turner on perknite . . . . 255

ertebrates . 163 Wallengren's Proboscis of

Kidd's Direction of the hair . 669 Pob e o. oro ces

Se in Hydra 589 Ward's Disease in plants . . 248

Kolbe's Econo: entomol- Washington's Differentiation

ogy - 64 | at Magnet Cove . . . .

MEOS ety CHR NE o0 BEC o0 0o

INDEX.

2 PAGE

Reviews:

Weinschenk’s Rock-forming

Wimmera TU UM. vx

popeye Nestlings of for-

t and marsh S757

Need vi the Ponana

of Agricu

Zschokke’s ay of dui

land ... . s uns s

Rhoads, S. N. Marsh or rice-

field mice ot eastern states . . 661

Rice-field m . 661

Ritter, W. E. Ravina’ ae

tion of the Enteropneusta

. Colors of nido

polypetalous flowers .

Rowley, H. T. Histological hens

in Hydra during regeneration . 579

SAGARTIA, Dispersal of . . . . 491

St. John, Orestes on foin fishes .

Sargent, F. L. rbariu

and plant description . . . . 592

Sense hairs in Lepidoptera . . . 561

Variation in Aster . III

Shull, G. H.

Sar Metamorphosis of . . . 615

land, M. V. New economic

Entomology

Snow, L. M. de of the

drift line. . ows BSS

Sponge inven: Origin «. v a v UE

Squid, Vascular system of . . .787

Stages in plant growth. . 865

Synopsis of invertebrates. XIL Di-

genetic trematodes . . 887, 953

. 601

TERATOLOGICAL calf . .

Dinoflagellata. . . - 187

Trelease, W. Bailey's Cyclopedia

of horticulture. . . . . . . 671

PAG

Trelease, W. Campbell’s Text-

ook of botany 671

Leavitt's buiton. of liotiny 342

—— Schwartz on forest trees .

— Trees in winter SUM,

—— — Ward's Disease in Sdn ie B

Tremataspide . . : 379

Position of . . eee

Trematodes, Synopsis dis 887, 953

Trowbridge, C. C. Relation of

wind to bird migration . « 735

True, R. H. Chamberlain's Meth-.

ods in plant histology . . 165

——— Ganong’s Plant Pháis. (c 5

—— MacDougall’s Plant igi

ology . 74

Tubularia Md Corymorpha esos 02 ONE

Identity of . = > 2 E

Tubularia parasitica . . . . . 987

ULULA, Life history of. . . . . 421

VARIATIONS in Aster... . «. . WII

—— in garter snake . . . . . 481

—— odonta - 395

Maprani system of squid. . . . 787

Vertebrates, Origin of limbs of, i 837

pw H. B. Benham on flat-

worm - 497

—— elon sporozoan investiga

Wate F. M. Winds sid im

and the distribution of insects .

Wheeler, W. M. New cnc

tural ant from Texas . . 85

Occurrence of Formica cin-

erea and F. rufibarbis in

i - 947

795

qu Temperature of insects . . 401

Williams, L. W. Vascular system

of the squid . 2 4*4 989

Willoughby, C. c. Prehistoric

ed flint knives . . .

"VOL. XXXVI, No. Ps

; A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

lud e

pile ses ON

aspe SUE

The American Naturalist.

ea ASSOCIATE EDITORS:

JA. I PH.D., American Museum of Natural ns New York.

: = H. COMST TOCK, S.B., Cornell University, Ithaca.

AVIS, M.E., Harvard — Cambridge.

eid

THE

AMERICAN NATURALIST

VoL. XXXVI. January, 1902. No. 421.

PREHISTORIC HAFTED FLINT KNIVES.

CHARLES C. WILLOUGHBY.

Tue question of the function of the stone implements

commonly called arrow and spear points has been a vexed

one. There are few references by early writers to the use

of chipped flint by the American Indians for other purposes

than as points for projectiles. It is very probable, however,

that a majority of these implements were used as knives or

cutting tools and were attached to short handles of wood or

antler.

Major Powell found such knives in use among the Pai Utes.

Colonel Ray collected similar implements from the Hooper

valley Indians of California, and there is in the Peabody Museum

at Cambridge a finé collection of leaf-shaped jasper blades fas-

` tened with pitch and cord wrappings to short handles of wood. -

These were obtained from the Klamath Indians of southern

Oregon.

The finding of a few similar tools with wooden hafts still

attached in prehistoric burial caves, cliff houses, and graves,

shows that such implements were in use in prehistoric times

over a large portion of North — Prof. F. W. Putnam

2 THE AMERICAN NATURALIST. [VoL. XXXVI.

has described and illustrated hafted stone knives from graves in

the Santa Barbara Islands and from prehistoric burial caves in

the state of Coahuila, Mexico; also one knife with a handle of

antler from a mound in Ohio.!

The blades of the eight knives from the mummy packs of the

Mexican burial caves referred to above are chipped from light-

colored chalcedony, the largest one being seven and one-fourth

inches in length, and its greatest width being three and one-

fourth inches. The smallest blade is three and one-fourth inches

long and two and one-fourth inches wide near its base. The

blades vary in form, some being proportionally short with round-

ing points, others being comparatively narrow and sharply

Fic. 1. — Prehistoric knives from the cliff houses, 14.

pointed. The hafts of these knives are of wood, about six inches

in length, the majority of them being made of a section of a

limb with the bark removed. One is fashioned from the dis-

carded hearth of a fire-making set, and still retains the burnt

depression in which the fire drill revolved. The thin base of

the blades is inserted into a notch extending across the end of

the haft, and is fastened with gum.

The knives from the Santa Barbara graves have small blades

of flint of the leaf-shaped type. The hafts are of split wood,

the blades being inserted into a notch at one end and secured

with asphaltum.

1 Bulletin of the Hee Institute, vol. xv, 1883 ; Wheeler’s made West A the

basin quide vol. vii; ee Museum Reports, vol. i p- 4)

ee ee So ee AN C PEOR MET CEREREM IINE S

No. 421.] PREHISTORIC HAFTED FLINT KNIVES. 3

The knife from the Marriott mound, Ohio, has a blade of

black flint, nearly triangular in outline, inserted into a notch

cut in the larger end of a curved antler prong. This knife

and those from the Mexican caves, together with one from

the Santa Barbara Islands, are on exhibition in the Peabody

Museum. 2d

While studying the remarkable collections from the cliff

houses of the Southwest collected a few years since by the state

of Colorado and by private individuals, the writer had unusual

opportunities for examining a number of prehistoric flint knives

Fic. 2. — Prehistoric knives from the cliff houses, 44.

hafted in wooden handles. He is indebted to those in charge of

the collections for permission to make the drawings accompany-

ing this paper. Several of these knives are now in the Free

Museum of Science and Art in Philadelphia.

The blades are chipped from different varieties of flint and

chalcedony, and are of the common typical forms (leaf-shaped,

triangular, stemmed, and notched) usually found in a collection

of chipped implements. The handles are of wood, in most

instances symmetrically fashioned and well finished, the blades

being inserted into a deep notch and cemented with gum, prob-

ably of Larrea mexicana. In a few examples, windings of sinew

or of cord made from yucca fiber furnish additional security.

THE AMERICAN NATURALIST. [VoL. XXXVI.

The blades of the knives shown in Fig. 1 are attached to their

handles with gum only. Their bases fit the notches snugly. The

M Fie. 3

— Prehistoric

cement with which they are fastened projects

slightly beyond the end of the haft and is pressed

against the blade at either side. The blade of a

is of black flint; 4 is also of dark flint, and c

is chipped from gray chalcedony. A suspending

cord of twisted cotton passes through the handle

of c. The handle of a is also perforated for the

passage of a cord.

The leaf-shaped blade of the knife illustrated

in Fig. 2, a, is of light-colored chalcedony and is

inserted for about half its length into the wooden

handle, being firmly secured with gum. The

drawing shows both front and side.

The beautiful knife shown in Fig. 2, 4, has

a blade of the stemmed variety, chipped from

pink chalcedony. It is attached to the wooden

handle with gum only.

The broken blade of Fig. 2, c, is fastened with

gum. As an additional security it is wrapped

with a cord of twisted yucca fiber. The handle is

perforated for the passage of a suspending cord.

The unique example illustrated at æ has a blade

of dark flint secured at an angle to the well-made

wooden haft. A cord of twisted yucca fiber

passes through a perforation near the end.

The implement represented in Fig. 2, e, would

perhaps be most correctly classed as a saw. The

blade is of a type widely distributed but nowhere

common. It is chipped from dark flint. The

teeth are thin and sharp, and in the hands of a

skilled prehistoric workman it has doubtless done

good execution in wood, bone, and antler. The

handle is a section of a limb or shoot, and the

double-aded knife blade is secured with

adi house, - The double-bladed knife E igatested in Fig. 3 has

for a handle a section of a small sapling or limb, with the bark

No. 421.] PREAISTORIC HAFTED FLINT KNIVES. 5

still adhering. Its length, including the blades, is eleven and

one-half inches. The notched blades are fastened with sinew.

A similar double-bladed knife is shown in the hand of a god

issuing from the mouth of a serpent sculptured upon one of the

lintels of a ruined temple at Yaxchilan, southern Mexico.

It is very probable that the implements shown in Fig. 4 were

primarily intended as foreshafts for light spears projected with

a spear thrower, a few examples of this ingenious device having

been found in the cliff houses of the Southwest. These spear

throwers have double finger loops, and are in other respects very

similar to the ancient Mexican atlatl used by both Mexicans and

Mayas, and represented in their sculptures. The foreshafts of

the spears accompanying the atlatl in the carvings resemble

Fic. 4. — Prehistoric knives or foreshafts, 14.

those illustrated in Fig. 4. A foreshaft similar to Fig. 4, 4,

has the end opposite the point beveled for inserting into a socket

at the end of the spear shaft. The lower end of Fig. 4, c, is

also slightly beveled. The ends of æ and 2 show no beveling,

and these implements may have been intended for knives only ;

but if originally constructed for foreshafts to spears, it is prob-

able, as they are detachable, that they were also used as cutting

implements. The blades are secured to the hafts with sinew,

no cement being visible.

In one of the collections was a spear-like implement tipped

with a point of black flint closely resembling in form the knife

blade illustrated in Fig. r, 6. This was secured in the notch

with cement and cord wrappings. The shaft, forty inches in

length, is worked smooth and polished, its lower end terminating

. ina rounding point.

6 THE AMERICAN NATURALIST.

Besides the tools whose functions are indicated by their forms,

as perforators, scrapers, etc., chipped flint was used in America

for cutting implements and as points for various kinds of pro-

jectiles, including arrows, light spears thrown with a throwing

stick, harpoons with detachable heads, hand lances, small jave-

lins, and thrusting weapons; but the greater number of flint

implements of the common types, of lengths varying from

about two inches to seven inches, were probably used as knife

blades. Nor are we safe in assuming that the stemmed and

notched forms were all projectile points, as an examination of

Fig. 2, 6, and Fig. 3 will show. It is of course impossible with

our present knowledge to classify correctly all forms of chipped

implements, but a study of the few prehistoric hafted examples

known will materially aid us in the work. .

PEABODY MUSEUM OF AMERICAN ARCH/EOLOGY AND ETHNOLOGY,

CAMBRIDGE, Mass.

a Lic e e MM Eder

ON THE AFFINITIES OF CERTAIN ANOMALOUS

DICOTYLEDONS.

DOUGLAS HOUGHTON CAMPBELL.

THERE is still a difference of opinion among botanists as to

the relationship between the two great divisions of angio-

spermous plants, monocotyledons and dicotyledons. The two

groups have evidently been derived from a common stock, but

which branch is the older is still somewhat doubtful.

Among the genera commonly referred to the dicotyledons

are several which differ from the typical forms both in the

structure of the flowers and in the character of the tissues,

especially the structure and distribution of the vascular bundles.

Among these anomalous dicotyledons may be mentioned the

Nymphaacez, certain Ranunculacez (Actaa, Thalictrum),

and among the Berberidacez, Podophyllum, Diphylleia, and

Caulophyllum.

A recent paper! on the embryo of Nelumbo has called atten-

tion to the importance of a thorough study of these anomalous

genera, whose embryogeny is almost completely unknown.

The result of Lyon's examination of Nelumbo was the dis-

covery that the apparently dicotyledonous embryo has really

but a single cotyledón and resembles that of the aquatic Alis-

males, an order of monocotyledons that shows numerous analo-

gies with the Nymphzacez.

This interesting discovery suggested to the writer the advis-

ability of looking up what had been already done with the

. study of the embryos of some of the other forms referred to,

and although the results of these inquiries have been very

incomplete, they have revealed a number of “extremely sugges-

tive facts, which are here given, and which emphasize the

desirability of more thorough work in the same direction.

1 Lyon, H. L. Minnesota Botanical Studies, Ser. 2, pt. v, 1901.

; :

8 THE AMERICAN NATURALIST. [VoL. XXXVI,

NYMPH/EACE.

In several respects the Nymphzaceze suggest the Alismales,

rather than the dicotyledons, with which they are usually asso-

ciated. Some of the earlier botanists, among them Jussieu,

considered the Nymphzeacez as true monocotyledons; but all

later botanists have agreed in placing them with the Poly-

carpice, or Ranales, among the most primitive of the dicoty-

ledons. The discovery that Nelumbo has a monocotyledonous

embryo at once raises the ques-

tion whether the other genera

are also monocotyledonous.

The structure of the flowers,

especially in the simpler genera,

Cabomba and Brasenia, is very

much like that in some of the

Alismales, and the character of

the vascular bundles, as well

as their arrangement in all the

Nymphezeacez, is similar to that

in the typical monocotyledon-

ous stem. The form of the

leaves, also, is often very sug-

gestive of the sagittate leaves

of Alisma or Sagittaria. This

is seen in Nuphar, especially

Fic. 1. — Seedling of Victoria regia,showing When growing in shallow water,

" Pages iie pee (From and the early leaves of other

Nymphzacez (Fig. 1) are re-

markably similar to those of Sagittaria. Whether or not this

resemblance of the young leaves of these Nymphzacez to

those of the Alismales is really an indication of relationship,

it is worth noting in connection with the other points of

resemblance.

From Lyon's studies upon the embryo of Nelumbo it appears

that in this genus the apparent two cotyledons are the result

of a bifurcation of a single cotyledon. The stem apex arises

laterally as in typical monocotyledons, and, as in these, the

No. 421.] CERTAIN ANOMALOUS DICOTYLEDONS. 9

cotyledon enwraps the plumule, which becomes largely devel-

oped. In Aponogeton, one of the Alismales, the sheathing

base of the single cotyledon develops a pair of stipule-like

lobes, which remotely suggest the two large lobes described

for Nelumbo.

Whether the other Nymphzacez will show a monocotyled-

onous embryo remains to be seen, and the removal of the

family to the Alismales is, perhaps, premature.

RANUNCULACE.

A number of Ranunculacez resemble superficially the Alis-

males, especially as regards the structure of the flowers. The

genus Echinodorus, of the Alismacez, for instance, has flowers

closely resembling those of Ranunculus. Other Ranunculacez

—e.g., Actzea, Thalictrum— have the vascular bundles arranged

much as in the typical monocotyledons instead of in the single

ring characteristic of the typical dicotyledons.

Ranunculus ficaria has but a single cotyledon, which it is

usually supposed is the result of the abortion of one of two

cotyledons; but this has not been demonstrated, and it is-

barely possible it may prove to be truly monocotyledonous.

Certain peculiarities in the embryos of other genera, which

will be referred to later, point to the possibility of the embryo

in these forms also having but a single cotyledon.

One important difference, however, must be noted between

the Ranunculacez and the Alismales or Nymphæaceæ. In

the two latter groups the embryo in the ripe seed is large,

with very little endosperm about it. In the Ranunculacez the

embryo in the ripe seed is always minute and is surrounded by

abundant endosperm.

ANOMALOUS BERBERIDACE®.

Under the Berberidacez are generally included several pecul-

iar genera of somewhat doubtful affinities. Of these, Jeffer-

sonia, Podophyllum, and Diphylleia are especially interesting, -

as they are all small genera of peculiar distribution, being

IO THE AMERICAN NATURALIST. [VoL. XXXVI.

represented in eastern Asia and Atlantic North America, but

not occurring elsewhere. This points to their being old types

which have persisted in these two widely separated regions,

and adds to the interest in their history.

The development of the embryo is quite unknown in all of

them, but the germination has been observed in Podophyllum,

both in our native P.

peltatum and in the Asiatic

P; emodt, which agrees

closely with P. peltatum.

The latter species has

been carefully studied by

Holm,! and there are certain

peculiarities in the germi-

nation, which, in view of

the recent discovery in

Nelumbo, are extremely

significant. According to

Holm (and the same is

shown by Lubbock for P.

emodt) there are apparently

two cotyledons, with com-

pletely united, much elon-

gated petioles, which form a

hollow tube, at the base of

which the plumule is placed

d p (Fig. 2, 2). Thelatter finally

grea rinse : bn. wangen emai breaks through the base of

mule st) at the base of the coli adodaty hic: the coty ledonary tube.

as seis erar Prec M There would be nothing

E i especially significant about

this were it not found that the second leaf, as shown by

Holm's studies, is deeply bilobed (Fig. 2, C ) and resembles to

an extraordinary degree the supposed fair of cotyledons. The

thought was at once suggested, — Is not the supposed pair of

cotyledons in Podophyllum really a single one, as in Nelumbo,

the two apparent cotyledons being merely lobes of a single

on ive mg Botas i IG " a 1899. a : : RU WE #

No. 421.] CERTAIN ANOMALOUS DICOTYLEDONS. II

leaf? This view is much strengthened by the persistence of

the bilobed form in the leaves of the young plant for several

years, and the permanently binate leaves of the allied genera

Jeffersonia and Diphylleia. The position of the plumule at

the base of the cotyledonary tube suggests a possible lateral

origin for this, as in typical monocotyledons and in Nelumbo.

The character of the flowers and the tissues in Podophyl.

lum and Diphylleia, as well as in Caulophyllum, are in accord

with a possible monocotyledonous affinity for these forms. It

is, at any rate, highly desirable that a careful embryological

study should be made of these extremely peculiar plants.

The formation of a cotyledonary tube apparently similar to

that in Podophyllum has been described for a number of widely

separated forms, e.g., Anemone, Delphinium, various Umbel-

liferze, Megarrhiza californica, Dodecatheon meadia. Whether

in all of these there is really a coherence of two cotyledons

must be determined by a study of the embryogeny.

It is evident that the last word has not yet been spoken as

to the interrelationships of the angiosperms. The numerous

studies upon the development of the embryo sac, which have

appeared during the past few years, have shown that there is

much more variation in the structures of the embryo sac than

was supposed to be the case. The most marked departure from

the angiospermous type is the genus Peperomia. The embryo

sac of this genus has regularly sixteen nuclei instead of the

eight usually present, and in this respect shows an approach to

the condition obtaining among the gymnosperms and higher

pteridophytes.

Peperomia, which appears to be genuinely dicotyledanpus,

nevertheless in the structure and distribution of the vascular

bundles, as well as in the character and arrangement of the

flowers, approaches some of the simpler monocotyledons, espe-

cially the Aracez. It seems possible, as the writer has else-

where suggested, that there may be two points where the two,

great divisions of angiosperms come together.

From the evidence at hand it would appear that the two

"uM Se and dicotyledons — are of about equal

1 Holm, doc cit., p. 422.

12 THE AMERICAN NATURALIST.

antiquity; but as a whole, the former have remained simpler

than the dicotyledons. The scattered, closed vascular bundles

probably represent a more primitive type of structure than the

ring of open bundles characteristic of the more highly devel-

oped dicotyledons.

It is exceedingly important for a clear understanding of the

affinities of such anomalous types as Podophyllum that a com-

plete study should be made of the development of the embryo

sac and embryo. Results of importance ought to reward the

student who will make a critical study of the development of

these puzzling forms.

LELAND STANFORD JR. UNIVERSITY.

THE SKELETON OF THE HEAD OF INSECTS.

JOHN HENRY COMSTOCK anp CHUJIRO KOCHI.

Tue skeleton of the head of an insect is composed of several

sclerites more or less closely united, forming a capsule, which

includes a portion of the viscera, and to which are articulated

certain appendages.

The early entomologists, among whom were Fabricius (1775),

Illiger (1800, 1806), Kirby (1802, 1826), Savigny (1816), Straus-

Durckheim (1828), Burmeister (1832), Newman (1834), and

Newport (1839), laid the foundation of our knowledge of the

structure of this skeleton; and it is remarkable, considering

the extent of entomological literature, how little has been

added in this particular field since the publication of the article

* [Insecta " by the last-named writer.

Although comparatively little progress has been made in the

study of the sclerites of the head during the last sixty years,

very much has been learned by the workers of to-day regarding .

the development of this region of the body ; and the time has

come when, using the known facts of embryology as a starting

point, one can hope, by comparative anatomical studies, to gain

a clearer idea of the structure of the skeleton of the head than

has been set forth as yet. To do this has been the aim of the

writers of this paper.

THE AREAS OF THE EXTERNAL SKELETON,

In descriptions of insects it is necessary to refer to the dif-

ferent regions of the surface of the head. This has resulted in

the establishment of a nomenclature, which, although based on

the work of the early insect anatomists, is really of compara-

tively little morphological value ; for but few of the primitive

sclerites of the head have remained distinct, and some of them

greatly overshadow others in their development. The result

+3...

14 THE AMERICAN NATURALIST. [Vor. XXXVI.

is that in some cases a named area includes several sclerites,

while in other cases only a portion of a sclerite is included.

This nomenclature, however, is sufficient for the needs of

describers of species, and will doubtless continue in use. It

is worth while, therefore, to review it briefly and to attempt

_where necessary to make it more definite.

The best landmark from which to start

for this purpose is the epicrantal suture, the

inverted Y-shaped suture on the dorsal

part of the head, in the more generalized

insects (Fig. 1, ¢.sz.). Behind the arms of

this Y there is a series of paired sclerites,

which meet on the middle line of the dor-

sal wall of the head, the line of union being

the stem of the Y; and between the arms

of the Y and the mouth there are typically three single sclerites

(Fig. t, A C, L). It is with these unpaired sclerites that we

will begin our definitions of the areas of the head.

The Front (frons, Kirby; clypeus posterior, Newport). —

The unpaired sclerite between the arms of the epicranial

suture (Fig. 1, F).

In the more generalized insects at least, if not in all, the

front bears the median ocellus; and in the a the

paired ocelli also. Frequently the suture

between the front and the following

sclerite, the clypeus, is obsolete; but as

it ends on each side in the invagination

which forms the anterior arms of the

tentorium (Fig. 2, a7), its former position

can be inferred, at least in the more gen-

eralized insects, even when no other trace

of it remains. In Fig. 2 this is indicated

by a dotted line.

The Clypeus (clypeus, Fabricius; chaperon, Straus-Durckheim ;

clypeus anterior, Newport). — The intermediate of the three

unpaired sclerites between the epicranial suture and the mouth

— (Fig. 1, C). To this part one gente (the d of the

andis articulates.

Fic. 1. — Head of a cricket.

No. 421.] SKELETON OF THE HEAD OF INSECTS. I5

Although the clypeus almost always appears to be a single

sclerite, except when divided transversely as indicated below,

it really consists of a transverse row of three sclerites, — one

on the median line and one on each side articulating with the

mandible. The median sclerite may be designated the clypeus

proper, and each lateral sclerite, the antecoral piece of the man-

dible. Usually there are no indications of the sutures sepa-

rating the clypeus proper from the antecoxal pieces; but in

some insects they are distinct. In the larva of Corydalis

the antecoxal pieces are not only distinct

but are quite large (Fig. 3, ac, ac). TT

In some insects the clypeus is com-

pletely or partly divided by a transverse |

suture into two parts (Fig. 1). These

may be designated as the first clypeus

and the second clypeus, respectively ; the

first clypeus being the part next the front

(Fig. 1, Cr) and the second clypeus being l

that next the labrum (Fig. 1, C2). ic

There is a great lack of uniformity in L

the application of the term cZyfeus, aris- es Ha dde ra

ing from the fact that many writers apply

it to the entire area between the epicranial suture and the

labrum; either overlooking the fact that the part here

designated as the front is a distinct sclerite, or, following

Newport, terming it the cZypeus posterior. But as the front

and the clypeus (in the more restricted sense) pertain to

different segments of the head, it is desirable to use dis-

tinct names for them; and as the names proposed by New.

port are morphologically incorrect, the so-called clypeus

posterior being in front of the so-called clypeus anterior, as

will be shown later, it is doubtless better to use the older

term frons, or front, for the sclerite next the epicranial

suture, and to restrict the term clypeus to the part termed

clypeus anterior by Newport.

The Labrum (labrum, Vlliger). — A movable flap which con-

stitutes the upper lip of the mouth (Fig. 1, Z). The labrum.

is the last of the series of unpaired sclerites between the

16 THE AMERICAN NATURALIST. | [Vor. XXXVI.

epicranial suture and the mouth. It has the appearance of

an appendage but is really a portion of one of the head

segments.

The Epicranium (épicráne, Straus-Durckheim). — Under this

term are included all of the paired sclerites of the skull, and

sometimes also the front. The paired sclerites constitute the

sides of the head and that portion of the dorsal surface that is

behind the arms of the epicranial suture. The sclerites con-

stituting this region are so closely united that they were

regarded as a single piece by Straus-Durckheim, who also :

included the front in this region, the epicranial suture being

obsolete in the May beetle, which he used as a type.

The Vertex (vertex, Kirby). — The dorsal portion of the epi-

cranium ; or, more specifically, that portion which is next the

front and between the compound eyes (Fig. 1, V, V). In many

insects the vertex bears the paired ocelli. It is not a definite

sclerite; but the term vertex is a very useful one and will

doubtless be retained.

The Qeciput (occiput, Kirby). — The hind part of the dor-

sal surface of the head. When a distinct sclerite, it is

formed from the tergal portion of the united postgenae

described below (Fig. 3, O, O).

The Gene (gene, Kirby). — The lateral

portions of the epicranium. Each gena, |

in the sense in which the word was used

by the older writers, includes a portion of

several sclerites. Like vertex, however,

the term is a useful one.

The Postgene. — In many insects each

gena is divided by a well-marked suture.

This led Comstock! to restrict the term

gena to the part in front of the suture

: (Fig. 4, G), and to propose the term

postgene for the part behind the suture (Fig. 4, Pg).

The Gula ( gula, Kirby ; piece basilaire, Straus-Durckheim), —

A sclerite forming the ventral wall of the hind part of the head

dn certain orders of insects, and bearing the labium or second

` ¥Comstock and Kellogg. d MK 1895.

— ud aai aah of

Fic. 4.

a cockroach,

No. 421.) SKELETON OF THE HEAD OF INSECTS. 17

maxillz (Fig. 5, Gz). In the more generalized orders, the sclerite

corresponding to the gula does not form a part of the skull.

The Cervical Sclerites (cervical sclerites, Huxley). — Small

sclerites found in the neck of many insects. Of these there

are dorsal, lateral, and ventral sclerites ; the lateral cervical

sclerites have been termed the jugular sclerites (pièces jugu-

laires, Straus-Durckheim) (Fig. 4, es, em).

Other Sclerites. — 1n addition to the areas and sclerites

named above, the following sclerites will be described in later

pages of this essay: the ocular sclerite,

the antennal sclerite, the trochantin of the

mandible, and the maxillary pleurites.

These terms should be added to the list

of those available for the purposes of

systematic entomology.

THE SEGMENTS OF THE HEAD.

The determination of the number of X P. UN

segments in the head of an insect is a

problem that has been much discussed

since the early days of entomology. The

first important step towards its solution was made by Savigny

(1816), who suggested that the movable appendages of the head

were homodynamous with legs. This conclusion has been

accepted by all; and as each segment in the body of an insect

bears only a single pair of appendages, there are at least four

segments in the head; z.e., the antennal, the mandibular, the

maxillary, and the second maxillary or labial.

As the compound eyes are borne on movable stalks in cer-

tain Crustacea, it was held by Milne-Edwards that they represent

another pair of appendages; but this view has not been gen-

erally accepted. It is not necessary, however, to discuss in

this place whether the eyes represent appendages or not ; the

existence of an ocular segment has been demonstrated in

another way, to be discussed later.

This is the point to which the solution of the problem was

carried by the methods of comparative anatomy. The existence

Fic. 5. — Head of Corydalis,

adult, ventral aspect.

18 THE AMERICAN NATURALIST. [VoL. XXXVI.

of four segments was demonstrated, and the presence of a fifth

em ocular) suggested. From this point the work has been

carried on chiefly by the embryologists.

The existence in the embryo of distinct

segments, each corresponding to a pair of

mouth parts, was early shown. Among the

better of the older figures for this purpose

are those of Brandt (69, Fig. 12) and of

Packard (71). Fig. 6, which is copied from

Brandt, represents an early stage in the devel-

opment of the embryo of a damsel fly (Calop-

teryx) In this stage the labial and maxillary

| segments are quite distinct, appearing to be

body segments rather than cephalic. This

Mag poco doubtless represents a phylogenetic stage, in

Mors el fy (Calopter, which the head proper consisted of fewer seg-

ments than it does in existing insects.

It was also found that the subcesophageal ganglion, which

innervates the mandibles, maxilla, and labium, is formed by

the union of at least three pairs of primitive ganglia. Fig. 7,

from Heider (89), represents a stage in the development of

Hydrophilus, in which these ganglia are still distinct, each pair

of ganglia corresponding to a pair of mouth parts.

So far the results of embryology merely con-

firm the conclusions of comparative anatomy.

But the embryologists have also demonstrated

the existence of vestiges of segments, which had

not been recognized as such by the early writers.

In his work on the embryology of the honey-

bee, Bütschli ('70) described a pair of evanescent

appendages situated between the antennz and

the mandibles. Later these were observed by

= others, and writers began to refer to a “pre-

mandibular," or * intercalary,' ' segment in the

head of insects. More recently the appendages !

of this vestigial segment, which is properly ^fe" Heider.

. termed the second antennal segment, were observed in the

oe rhon M n

Fic. 7. — Embryo

of H 1

No. 421.) SKELETON OF THE HEAD OF INSECTS. I9

They were also observed in Campodea by Uzel (97) The

last writer states that these appendages persist in the adult

Campodea, and Folsom ('99b) says, * I may add that rudimen-

tary chitinized intercalary appendages persist in adults of

Tomocerus, Orchesella, and other Collembola.”

Equally important evidence as to the existence of a second

antennal segment in insects has been furnished by studies of

the nervous system. It was shown by Viallanes (87a) in his

study of the structure of the brain (supra-cesophageal ganglion)

of Vespa that there are three principal divisions in the brain

of insects. These he named the fprotocerebrum, the deutocere-

brum, and the ¢ritocerebrum. Almost immediately after Patten

(88) demonstrated that the brain is formed from three pairs of

primary ganglia; and the same fact was shown by Wheeler ('89).

Viallanes also showed that the protocerebrum innervates the

compound eyes and ocelli; the deutocerebrum, the antennae ;

and the tritocerebrum, the labrum. This demonstrates the

existence of three premandibular segments: first, an ocular

or protocerebral segment, without appendages, unless the com-

pound eyes represent them (the supposed discovery of other

appendages on the ocular segment by Carriere ('90) has not

been confirmed); second, an antennal or deutocerebral seg-

ment, bearing antennz ; and third, a second. antennal, or trito-

cerebral segment, of which the labrum is a part, and to which

the so-called intercalary appendages doubtless belong. As

Viallanes has shown that the tritocerebrum of Crustacea inner-

vates the second antennz, we are warranted in considering the

tritocerebral segment of insects to be the second antennal

segment.

The evidence thus far brought forward demonstrates the exist-

ence of six cephalic segments, — three innervated by the brain

and three by the subcesophageal ganglion. We have now to

refer to the evidence indicating the existence of a seventh

cephalic segment. .

The hypopharynx of insects is usually, in the Pterygota, a

tongue-like organ lying below and projecting in front of the

beginning of the alimentary canal In the Apterygota it

consists of three distinct parts, — a median organ termed th:

20 THE AMERICAN NATURALIST. [Vor. XXXVI.

*lingua" and a pair of organs termed “ paraglossz " by writers

on the Thysanura and Collembola. As the term “ paraglossae "

has long been used for a part of the labium or second maxilla,

Folsom I deed maintains that it should not be applied to a

part of the hypopharynx, and

proposes for these paired organs

the term superlingue.

In his work on the develop-

ment of apterygote insects, Uzel

(98) showed that the lingua arises

between the rudiments of the

maxilla; hence it may be regarded

it LIT ___} as pertaining to the sternum of

Pio. 8.— Head of embryo of Anurida, — the maxillary segment. Uzel also

showed that the superlinguz arise

as a pair of appendages between the mandibles and the maxil-

lae. , This indicates the existence of a segment between the

mandibular and the maxillary segments.

Similar results were obtained by Folsom ('00) in his work

on the development of the mouth parts of Anurida. Fig. 8

is copied from Folsom and illustrates the relative positions of

the rudiments of the mouth parts. To Folsom also belongs

the credit of completing the :

evidence of the existence of a

superlingual segment, by demon-

strating the existence of a pair

of primary ganglia between those

of the mandibular and of the

maxillary segments (Fig. 9, 5).

And in a preliminary paper (99b),

he set forth the first complete

account of what promises to be

our final view regarding the seg- —

mentation of the head. ———

In confismation of the view

bees the: sut 'sophageal ganglion consists of four pairs of

| y ganglia, it should be mentioned that wis ago Patten

a ‘these ganglia. It has been believed , however, that

No.421.] SKELETON OF THE HEAD OF INSECTS. 21

the pair immediately in front of the maxillary ganglia were

the mandibular ganglia (see Korschelt and Heider, '99b, p. 326);

but it is more probable that the first of these four pairs of gan-

glia (Fig. 10, 7) pertains to the man-

dibular segment and that the second

pair are the homologues of the super-

lingual ganglia figured by Folsom.

The seven segments of the head

are designated as follows:

First, ocular, or protocerebral.

Second, antennal, or deutocerebral.

Third, second antennal, or tritocerebral.

Fourth, mandibular.

Fifth, superlingual.

Sixth, maxillary. Fic. ro. — Head of embryo of

Seventh, labial, or second maxillary. Acilius, after Patten.

DIAGRAMS OF THE ELEMENTS OF THE HEAD.

The head of an insect consists of seven segments closely

consolidated, greatly reduced in length, and, in the case of

some, bent out of the original line. The morphological rela-

tions of these segments can be shown by representing them

as distinct, of uniform size, and in a direct line. This is done

in the accompanying diagram (Fig. 11, A).

Let us trace the steps by which this diagram was made.

First, the outlines of the seven segments were drawn as if no

Fic. 11.— Diagrams of the elements of the head: A, lateral aspect; B, ventral aspect.

reduction or consolidation of any of them had occurred; at

this stage the diagram might represent the cephalic end of

an earthworm, except that the prostomium is not represented.

22 THE AMERICAN NATURALIST. [Vor. XXXVI.

Second, a longitudinal line was drawn representing the line

of separation of the sternal from the pleural elements of the

segments ; it is along this line that the appendages are borne.

Third, the chain of ganglia were added, a pair of ganglia in

each segment; -as the nervous system is developed from the

ectoderm on the ventral side of the body between the append-

ages, it is represented in the sternal portion of the preoral

segments as well as in the postoral segments. The diagram

being of a side view, only a single member of each pair of

ganglia and longitudinal commissures are shown. Fig. 11, B,

is a diagrammatic representation of a ventral view of the

nervous system. Fourth, the compound eyes and the ocelli

were represented in the first segment (Fig. 11, A), because

they are innervated by the protocerebrum. The reasons for

the exact positions within the segment assigned to these

organs will be discussed later. Fifth, the position of the

appendages was indicated, a pair to each segment except the

first. It will be remembered that the antennz are innervated

by the deutocerebrum; hence they pertain to the second seg-

ment. Sixth, the mouth was represented as opening in the

ventral wall of the third segment; this is in accordance with

the results of the studies of Viallanes (87a), who has shown

that, although the third pair of ganglia enter into the com-

position of the supra-cesophageal ganglion, the commissure

- Which connects them passes behind the cesophagus. This is

shown in the second diagram. Although the position assigned

to the mouth in these diagrams was suggested by the results

of the studies of Viallanes, it is not in accordance with his

conclusions; for he evidently believes that the mouth opens

between the third and fourth segments. Our view is based

on the well-known fact that the mouth of the embryo is

formed zz the labrdl rudiment and not behind it.

The determination of the position of the mouth is one of the

most striking of the results of the later studies of the head.

Naturally the older entomologists believed that the mouth-

opening was at the cephalic end of the body, and this effectually

onites; trois sont prebuccaux et trois

Peet = whe oS im : £.

| post-buccaux." — Villanes (87b, p- 117).

SET ee

UAE ey ENEE e SEC PEE CUPS

TS TET SETS S

Very E A E gem ED

No. 421.) .SKELETON OF THE HEAD OF INSECTS. 23

prevented a correct homologizing of the sclerites of the head.

It is evident that the old belief is still held by many ; thus Hey-

mons ('95) designates what he believes to be the first segment

the primäres Kopfsegment oder Oralstück.

The earliest suggestion of a different position of the mouth

that we have met was by E. Ray Lancaster (73), who refers

to an *adaptational shifting of the oral aperture." The later

writings contain many references bearing upon this, although

the full force of them is evidently not appreciated by the writers.

Thus it has been said by many that although the antenna were

doubtless originally postoral they have become preoral. The

facts would be more accurately stated by saying that although

the mouth was doubtless originally preantennal it has become

postantennal. This, however, would only partially indicate the

change that has taken place; for, as will be shown later, the

antennz have moved cephalad at the same time that the mouth

has moved caudad.

THE STRUCTURE OF A TYPICAL SEGMENT.

In order to determine the homologies of the sclerites of the

head, it is necessary to decide what sclerites were probably

present before the consolidation and reduction of its segments

took place. We have decided that the head is composed of

seven segments; let us now determine the elements in the

skeleton of a single segment. This necessitates a study of

segments in other regions of the body.

In the abdomen it is evident that a reduction of certain parts

has taken place, correlated with the loss of the abdominal

appendages; it is to the thorax, therefore, that we must look

for the more typical insectean segment.

The parts of a thoracic segment that are commonly recog-

nized are those described by Audouin (24): a ventral part,

sternite; two lateral parts, p/eurites; and a dorsal part, tergite.

- These are most easily seen in the wing-bearing segments ;

but they can be recognized also in the prothorax of certain

generalized insects. This is especialy the case in many

Orthoptera, as cockroaches and walking-sticks, where the

24 THE AMERICAN NATURALIST. [Vor. XXXVI.

pleurites of the prothorax are distinct from the sternum on

the one hand and from the tergum on the other; more often,

however, the tergite of the prothorax is not separated from

the pleurites. This is also the case in the segments of the

head: sometimes the tergite is distinct from the pleurites;

|. but more often the tergite is merely a

continuation of the pleurites over the

dorsal side of the segment. In such

cases the combined lateral and dorsal

. parts are designated as the pleurites ; for

we find that they bear the most charac-

teristic feature of the pleurites, the

lateral apodemes, to be described later.

owes Equally important for the purposes

Fic. r2.— Ventral aspect of the of this study is the fact that each

metathorax of a nymph of K n

PATRONS, thoracic segment is composed of two

subsegments. It is not necessary for

us to decide in this place whether or not this indicates a fusion

between two primary metameres, as has been suggested by

_ various writers!; the essential fact which must be taken into

account is the transverse division of each segment. The line

separating the subsegments. passes, on the pleural aspect,

between the episternum and the epimeron ; and, on the tergal

aspect, between the scutum and the scutellum. On the sternal

aspect, in most insects, the

division is not easily recog-

nized; but in certain Ple-

coptera and Orthoptera it

can be readily seen.

Fig. 12 represents the

sternite of the metathorax

of a nymph of Pteronarcys, pia. r3.—Ventral

2 aspect of the metathorax of Steno.

and Fig. 13 the same part pelmatus. The position of the furca within the

body i ted by a dotted li

of Stenopelmatus. In each is represented by a dotted line.

case it can be seen that the sternite is composed of two

distinct sclerites, — one lying between the episterna and one

: 1 The reasons for believing that each soient e ipon dC two, primy

netame cuir apu ee

Lowe

No. 421.) SKELETON OF THE HEAD OF INSECTS. 25

between the coxa; the former pertains to the first subsegment,

the latter, to the second.

In the furrow or suture between these two sclerites are the

invaginations forming the furca of that segment. The position

of the furca is, therefore, a good landmark for determining the

line of union of the two scle-

rites forming a sternite, or, in

other words, the division be-

tween the subsegments.

The second of the sclerites

forming a sternite is smaller

than the first, even in those in-

sects where it is best developed;

and in most insects it is greatly —

reduced or obsolete, so that the vote ita Viris ug alamo

furca appears to arise from the pne — Hie body sre amined

caudal margin of the segment

(Fig. 14). It is for this reason that the sternite of the

second subsegment has been overlooked heretofore. The ster-

nite of the first subsegment is retained in all insects, and is

the sclerite to which the term sternum has been universally

applied. The smaller sternite of the second subsegment may

be termed the sternellum (Fig. 12, S2; Fig. 13, S2).

Sometimes, as in Pteronarcys, the cephalic portion of the

sternum is more or less separate from the main part; this

detached portion may be termed the presternum (Fig. 12, je

A poststernellum, corresponding to the

postscutellum, has not been observed.

It has been shown by Haase (89) and

= | Heymons (95) that in a comparatively

| | early embryonic stage each segment of

M iei the body is composed of three parts, —

Frc. 15.— Diagram ofa segment a median field and two lateral fields

aan embryo, air Heymons- (Fig. r5). and that the appendages

are developed as evaginations of the lateral fields. Each ster-

nite is therefore composed of three parts, — the portion derived

from the median field of the segment, and, on each side, a

portion derived from that part of. the lateral field which lies

26 THE AMERICAN NATURALIST. [VoL. XXXVI.

between the appendage and the median field. This portion

may be designated the /ateral element of the sternite. Such a

division is well shown in the abdominal sternites of the adult

Gryllus (Fig. 14).

More frequently, however, when a sternum in an adult

insect is divided longitudinally it is by a single median suture,

which perhaps represents the neural groove of the embryo.

A sternite of a subsegment may be composed, therefore, of

either two or three elements: in the one case the sutures

between the median field and the lateral fields are preserved ;

in the other, a trace of the neural groove is indicated. But

as a rule, each sternite is an un-

divided sclerite.

In the same way that the posi-

tion of the furca determines the

line of union of the subsegments

on the ventral aspect of a thoracic

segment, the line of union of the

a subsegments on the pleural aspects

Ka: 16.— Ental surface of the pleurites is determined by the position of

ee ts the lateral apodemes. Each of

these is an invagination of the

body-wall between the episternum and the epimeron. Fi ig. 16

represents the inner surface of the pleurites of the meso- and

metathorax of Melanoplus and shows the form of the lateral

apodemes (af).

For the purposes of this paper, it is not necessary to discuss

the structure of the tergal aspect of the typical segment beyond

a reference to the median suture, which represents the line

of the closure of the embryo. This suture has been well pre-

served in the head and thorax, as it is the chief line of rupture

of the cuticle at the time of molting.

The relations of the appendages to a typical segment are

illustrated by the accompanying figure (Fig. 17) of the base of

aleg of a cockroach. Near the point marked x the coxa artic-

ulates with the ventral end of the foot of the lateral apodeme

of the segment, Ze. with the ventral end of the episternum

and the epimeron. This may be termed zhe pleural articulation

EX S i

E i ECCE

No. 421.] SKELETON OF THE HEAD OF INSECTS. 27

of the coxa. In front of the coxa there is a triangular /7o-

chantin (tr), with its apex pointing towards the middle line of

the body. Between the trochantin and the ventral arm of the

episternum there are two sclerites, — one next the trochantin,

the antecoxal piece ; and one next the episternum, not yet named,

which may be termed the second antecoxa! piece. The antecoxal

piece at its mesal extremity ( y)articulates with

the coxa. This articulation may be termed

the ventral articulation of the coxa.

THE SCLERITES OF THE HEAD AND NECK.

The segments of the body of an insect

do not fall into three well-marked groups, —

head, thorax, and abdomen, — as commonly pi

defined in the text-books. Between the head Fis. :7.— The base of a

and the thorax there is a more or less distinct — 4^ ^ "ere

neck, which in some insects contains several sclerites. As we

believe that these cervical sclerites form a part of the second

maxillary or labial segment, it is necessary to include them in

a discussion of the sclerites of the head. We shall return to

them later.

If an embryo insect be examined after the appearance of

the mouth and the appendages, the cephalic end of the body

will be found to consist of a central swelling, the procephalon

(Fig. 7, fr), and a large lobe on each side, the cephalic lobes

(Fig. Joel).

The procephalon has been commonly described as the rudi-

ment of the clypeus and thelabrum. This is doubtless correct

if the term c/ypeus be taken in the broader sense by which it

includes the clypeus anterior and the clypeus posterior of

Newport; and this is evidently the sense in which it has

been used by nearly all writers on the embryology of insects.

Hence, according to the nomenclature adopted in this paper, .

the procephalon is the rudiment of the front, clypeus, and

labrum. he

This conclusion makes evident the significance of the epi-

cranial suture. The sclerites lying in front of the arms of ©

28 THE AMERICAN NATURALIST [Vor. XXXVI.

this inverted Y-shaped suture are those developed from the

procephalon, while the stem of the Y represents the line of

union of the cephalic lobes.

When we take into account the position of the mouth (see

Fig. 11), it is evident that the parts developed from the pro-

cephalon pertain to the ventral aspect of the body. In the

course of development there is a dorsal flexure of the cephalic

region by which the preoral sterna are bent up towards the

tergal aspect (Huxley, '78, p. 343); this has been described by

several observers (see Korschelt and Heider, '99b, p. 302).

As a result of this dorsal flexure, the former most anterior

part of the procephalon assumes a more backward position,

which led to the part derived from it being termed by Newport

the clypeus posterior. As this term is morphologically incor-

rect, we have adopted the name front for this part, and restrict

the term c/ypeus to the clypeus anterior of Newport.

It seems obvious that the three sclerites derived from the

procephalon, — the front, the clypeus, and the labrum, — rep-

resent the sternites of the three preoral segments..

As to the front, it bears the median ocellus, and, in the

Plecoptera, the paired ocelli also ; and as the ocelli are inner-

vated by the protocerebrum, it is evident that the dere is a

part of the protocerebral segment.

In this connection reference should be made to a migration

of the paired ocelli. The condition in the Plecoptera, where

the front bears all the ocelli, is probably the most generalized ;

for in this order, as will be shown later, the most nearly primi-

tive position of the antennz is found; and, too, in this order

the most generalized condition of the tracheation of the wings

exists (Comstock and Needham, '98, p. 237). In the Orthoptera

(Blattidae and Gryllidae) the paired ocelli are in the suture

between the front and the vertex. In certain Ephemerida the

paired ocelli are in this suture, while in others they have passed

on into the vertex. In the more specialized orders, wherever

we have been able to distinguish between the front and the

vertex, we have found the paired ocelli in the vertex.

The labrum is innervated by the tritocerebrum ; for this

-~ reason we — s it as the sternite of the tritocerebral segment,

No. 421.] SKELETON OF THE HEAD OF INSECTS. 29

or rather as a part of this sternite; for, as the invagination of

the stomodzeum is surrounded by the labral rudiment, the labrum

represents only that part of this sternite that lies cephalad of

the mouth.

The clypeus (clypeus anterior) isa sclerite between the front

and the labrum ; for this reason, we believe it to be the sternite

of the intermediate of the three preoral segments, the deuto-

cerebral.

We have described the sclerites derived from the procephalon

as representing the sternites of the preoral segments. But

strictly speaking, we believe that each represents only the

median field of a sternite (Fig. 15, wf), and that the lateral

elements of the sternites have not been separated from the

pleural portions of the lateral fields of the segments; in other

words, that the early embryonic divisions of the segments have

been retained, and that those parts derived from the lateral

fields of the segments form a single sclerite on each side of

each segment.

In the ocular segment each lateral sclerite constitutes one-half

of the vertex and the corresponding gena, the line of union of the

lateral sclerites being the stem of the Y-shaped epicranial suture.

Each lateral sclerite of this segment bears a compound eye,

except in cases where they have been lost and except in the

larvae of metabolous insects, in which the development of these

organs is retarded; this is obviously a secondary condition,

like the internal development of the wings in the same forms.

The position of the compound eye, in the lateral sclerite

slightly removed from the middle field of the sternite (the

front), is that in which one would expect to find an appendage,

and it seems to us that the question whether or not the com-

pound eyes represent the appendages of the ocular segment is

still an open one.

Heretofore the chief reason for regarding the compound eyes

as representatives of appendages has been the stalked condition

of them in certain Crustacea ; but later writers are inclined

to regard the eye-stalks **as secondarily abstricted lateral por-

tions of the head which have.become independently nuosable a

gosc and Heider, '99a, p. 165).

30 THE AMERICAN NATURALIST. [VoL. XXXVI.

We are inclined, however, to return to the old view ; for we

find that in many insects each compound eye is situated in the

axis of an annular sclerite, which may be the basal segment of

the ocular appendage. Certainly if the eyes were merely spe-

cialized portions of the lateral sclerites, we should not expect:

them to be surrounded by a ring-like

suture, which in some cases is com-

paratively remote from the specialized

portion of the body-wall that forms the

eye.

These sclerites bearing the compound

eyes may be termed ¢he ocular sclerites;

they are represented in Fig. 18, os.

Passing to the second segment, we

uM find at the base of each antenna an

. 1%. — Head of a cricket, annular sclerite (Fig. 18, as), which is

T mM EM linc iu comparatively few insects and

which has not been described. It is

most clearly shown in the Plecoptera (Fig. 19, as). This we

believe represents the lateral field of the antennal segment,

7.£., the lateral element of the sternite and what is left of the

pleural element of the segment, which is greatly reduced. This

sclerite may be termed she antennal sclerite.

The position of the antennal sclerites should be discussed.

If the clypeus represents the median field of the sternite of

the antennal segment, as we believe, the primitive position of

the antennal sclerites was laterad of the clypeus, and we should

Fic

expect to find the rudiments of them in this position, t.e., lat-

erad of the procephalon, in the early stages of the embryo.

Many observers, however, have described the antenna as aris-

ing in a postoral position ; how can this be explained? A

study of the figures given by these authors! shows that while

a line connecting the two antennz would pass in some cases

behind the mouth, it is by no means so clear that the basal

part of the rudiment of the antennal sclerite does not abut

against the procephalon. In fact, the very figures given to

~ ? Weismann, '63, Fig. 22; Graber, '88; Figs. 1 and 2; Heider, '89, Fig. 102

a (Fig. 7 above); Patten, '88 (Fig. 10 above). | oe

Ei a Sce. E,

ra seine

TUS

No. 421.] SKELETON OF THE HEAD OF INSECTS. 31

support the view that the antenna are postoral in the early

embryo support the opposite view.

A migration cephalad of the antennz has been noted by many

observers ; and it is obvious that the position of the antennz in

adult insects is more or less remote from the primitive position.

In the Plecoptera is to be found the most generalized condi-

tion of the antennal sclerites yet observed in postembryonic

stages of insects (Fig. 19) ; here they are distinct sclerites,

and are only slightly removed from the clypeus, compared with

their position in the more specialized insects. Even here,

however, they are opposite the front, having been pushed out

of place by a migration of the mandibles and the antecoxal

pieces, to be described later.

In most insects each antenna has migrated along the suture

between the front and the gena, and occupies a position on the

lateral border of the front remote from the clypeus. Even in

so generalized an insect as a cockroach (Fig. 2), the antenna

has reached a point opposite the cephalo-lateral angle of the

vertex. Here it is remote from the anterior arm of the ten-

torium ; while in Pteronarcys it is very close to it.

In the case of the second antennal segment, the reduction

has been so great that we have been able to find in the Ptery-

gota no trace of the parts derived from the lateral fields of the

segment ; the labrum is the only well-

marked remnant of this segment rep-

resented in the skeleton ; it is possible

that the lateral elements are fused with

the gena. It is probable that a study

of those Apterygota in which the second

antennze are retained will reveal the pres-

ence of distinct lateral sclerites pertain-

ing to this segment. Fic. r9. — Head of a nymph

In taking up the study of the postoral E rim

segments of the head, the subject can be treated most easily

by beginning with the labial segment and proceeding forward

to the point reached in the above discussion.

In the early embryonic stages the labial segment is obviously

a body segment (Fig. 6, 2°mx); but in the course of the

32 THE AMERICAN NATURALIST. [Vor. XXXVI.

development of the embryo it moves forward, and in adult

insects it constitutes the dividing line between the head and

the thorax, forming the neck. Its appendages, the second

maxillae or labium, however, have moved forward so that they

are either loosely attached to the ventral wall of the head

(Plecoptera, Orthoptera, e/ al.) or, in the more specialized

orders, they contribute to the formation of the fixed parts

of the head.

While the appendages of this segment have been retained

and play an important róle as a part of the mouth organs, the

segment itself is greatly reduced, being represented by small

and more or less detached sclerites, the cervical sclerites.

Straus-Durckheim (28) suggested that the cervical sclerites

represent the remains of two segments situated originally

between the head and prothorax. Newport (39) regarded

i them as detached portions of the pro-

thorax; and Huxley (78) wrote: “I

think it probable that these cervical

sclerites represent the hindmost of the

cephalic somites.” But we find no

account of these sclerites that contains

5 | _| more than a suggestion regarding their

Fic. 20.— Lateral cervical scle- homologies. No evidence has been

rites of Melanoplus. :

brought forward to support any of the

conclusions, beyond the position between the head and thorax

occupied by these sclerites. We have been led to adopt the

view put forth by Huxley for reasons that seem to us conclusive, |

and which we will now state.

The cervical sclerites are best preserved in the Orthoptera.

In this order a variable number occur in the ventral wall of the

neck ; two in each lateral wall ; and in some forms, two in the

dorsal wall.

Between the two lateral cervical sclerites there is in certain

forms, as Melanoplus and Stenopelmatus, a prominent apodeme

_ (Fig. 20, ap). This apodeme we regard as homodynamous with

the lateral apodemes of the thoracic segments. Each of these

apodemes is an invagination between an episternum and an

epimeron ; we, therefore, conclude that the anterior lateral

ers

No. 421.] SKELETON. OF THE HEAD OF INSECTS. 33

cervical sclerite is the episternum of the labial segment, and

the posterior one the epimeron. This conclusion is confirmed

by the fact that the posterior of the two lateral cervical scle-

rites articulates with the episternum of

the prothorax, and the anterior one with

what we believe to be the epimeron of

the maxillary segment.

The ventral cervical sclerites in this

order are either two in number, Peri- br

planeta (Fig. 21), or constitute two trans- 5... — Vental und asc

verse series, Stenopelmatus (Fig. 22), cervical sclerites of Peri-

Gryllus (Fig. 23). We regard these as He

constituting the sternites of the two subsegments of the labial

segment. If this view be correct, the anterior sclerite or series

: of sclerites represents the sternum of

the labial segment, and the posterior

the sternellum. The division of a

sternite into a transverse series of

either two or three sclerites is com-

parable with what frequently takes

place in the sternites of the thorax

and of the abdomen, already discussed

on an earlier page.

The appendages of this segment, the

ime E —-j] second maxillae, project forwards from

Fic, 22. — Head of Stenopelma- the front margin of the segment as a

5n i single organ, the labium. This anoma-

lous condition is the result of a migration of the appendages

forwards and towards each other and of a coalescence, which

has been figured by many writers and

often described, lately in detail by

Deegener (00). As a result of this

coalescence the united cardines be-

come the submentum, and the stipites ; |

the mentum. Fic. 23. — Ventral cervical sclerites

2 of Gryllus.

In the more generalized insects a

gula is not developed as such. We have devoted much study

to the question of the homology of the gula and conclude that -

34 THE AMERICAN NATURALIST. [Vor. XXXVI.

it is the sternum of the cervical segment which has migrated

cephalad and become a part of the skull. In Corydalis the

sternellum of the cervical segment is retained back of the gula

(Fig. 5, Sz).

Although the maxilla are well-developed organs, the maxil-

lary segment itself is greatly reduced. The most conspicuous

element of it is the lingua, the unpaired portion of the hypo-

pharynx. This, as has been shown (Fig. 8, 7), arises between

the rudiments of the maxilla and evidently pertains to the

sternite of this segment.

The opening of the salivary glands is in thelingua. Carriére

has shown that these glands arise as the spiracular invagi-

nations of the prothorax, and that their openings migrate

cephalad, and towards each other, finally forming a single open-

ing in the lingua (Carriére and Bürger, 97). This explains the

absence of spiracles in the prothorax, and is one of the most

remarkable instances of the migration of organs and change

in function yet described.

Regarding the maxillary pleurites of the completed head,

almost nothing has been published. Huxley, in his descrip-

tion of the structure of the cockroach (78), stated that the

cardo of the maxilla is articulated ** with a thin skeletal band

which runs round the posterior margin of the epicranium." He

made no suggestion regarding the homology of this sclerite ;

and subsequent writers do not appear to have done so.

This lateral band (Fig. 4, m.em.) is one of two sclerites,

between which is the invagination which forms the posterior

arm of the tentorium ; the other of these two sclerites, the

anterior one, is much more reduced than this one, still it can

be seen in Periplaneta and in Gryllus. The articulation of

the maxilla is at the ventral end of these sclerites just ventrad

of the invagination between them, the open mouth of which is

very conspicuous in the more generalized insects.

The relation of these parts corresponds exactly with what

exists in a thoracic segment, where each leg is articulated

just ventrad of the lateral apodeme, which is an invagination

between the episternum and the epimeron. Evidently the band

described by Huxley is the epimeron of the maxillary segment,

x d. c

$ f ek ics iy Y ]

ROSE EN AU E ao ME TENTE TS,

PR; CORRIERE OU AN Mr EAA dE UNE C QUSS Ej ES. RT ERE MESE e LE ay eot TRE ir go) oe UNT

No. 421.] SKELETON OF THE HEAD OF INSECTS. 35

the thinner band in front of the invagination is the epister-

num, and the invagination itself is the lateral apodeme of this

segment.

From this it will be seen that the posterior arms of the

tentorium are to be homologized with lateral apodemes instead

of with spiracles, as is often done. It should be borne in mind,

however, that the spiracles are lateral invaginations between

segments, and that the lateral apodemes are invaginations in

a similar position between subsegments (Fig. 16). If each

segment consists of two consolidated metameres, the lateral

apodemes and spiracles are homodynamous structures. The

solution of this question must wait, however, the solution of

the larger question, the structure of the segment in air-

breathing arthropods ; we need not dwell upon it here.

The superlingual segment is so greatly reduced that we are

able to find no trace of the lateral elements of it in the skele-

n; if they exist, they are inseparably united with the mandib-

ular pleurites. The sternal elements are represented by that

part of the floor of the mouth cavity that bears the superlinguz ;

and the appendages of the

segment, by the superlinguz

themselves. In the Ptery-

gota these parts are greatly

reduced and have received

but little attention. Fig. 24

represents the hypopharynx

of Melanoplus, in which the

lingua is very prominent, and what we regard as the superlinguze

are reduced to a pair of small sclerites. As the superlinguz

have been carried into the mouth cavity by its invagination,

they lie behind the lingua, although they originated in front

of it. The superlinguz are connected with the skull by a

membranous portion of the body-wall, which, on each side,

extends between the attachments of the maxilla and of the

mandible.

In the mandibular segment the pleurites are represented by

the postgenz, which, excepting the vertex and genz, are the

most prominent sclerites of the head in many Orthoptera.

Fic. 24. — Hypopharynx of Melanoplus.

36 THE AMERICAN NATURALIST. [Vor. XXXVI.

The suture separating the postgenz from the genz is well-

marked on the lateral aspect of the head in the Orthoptera.

In most forms it is obsolete on the dorsal aspect but in a large

South American cockroach that we have studied the postgenae

are separated from the genæ and vertex throughout their entire

extent. Upon the presence or absence of this suture on the

dorsal wall of the head depends the presence or absence of

the so-called occiput ; the occiput being the tergal portions of

the postgenz (Fig. 3, O, O).

In the ventral end of each postgena there is an acetabulum

into which a condyle of the mandible fits (Fig. 25). Beginning

à ——— in this acetabulum and extend-

ing dorsad there is a suture which

divides the postgena into two parts ;

this suture is the more or less open

mouth of an apodeme which extends

into the cavity of the head.

Here again the same relation of

parts exists that is found in a tho-

e racic segment. The mandible is the

Fic. a basal segment (coxa) of an append-

age, which articulates with the ven-

tral ends of two sclerites (episternum and epimeron), between

which there is a lateral apodeme.

Thus we see that three of the head segments — the labial,

the maxillary, and the mandibular — closely resemble a thoracic

segment, in having on each side two sclerites, with an apodeme

between, and an appendage below except in the case of the

labial segment, where there has been a cephalization of the

appendages.

In the floor of the mouth cavity of Melanoplus there is on

each side just behind the superlinguz a sclerite (Fig. 24, ps)

which may represent a sternal element of the mandibular seg-

ment. The position of this sclerite farther in the mouth cavity

than the superlinguz is that which would be occupied by a

mandibular sternite, as such a sternite must precede the super-

linguæ in the course of the invagination of the mouth; equally

suggestive is the fact that this sclerite is closely connected

No. 421.] SKELETON OF THE HEAD OF INSECTS. 37

with the mandible, joining it near the insertion of the flexor

muscle. But in the present state of our knowledge little stress

can be laid on supposed homologies of the parts of the pharyn-

geal skeleton; for it is evident that in this region sclerites are

frequently developed secondarily.

The articulation of the mandible with the postgenz is its

pleural articulation ; the ventral articulation appears to be

with the clypeus in Periplaneta (Fig. 2), Ze, with the sternite

of the second segment in front of the mandibular segment.

It is difficult to imagine the steps by which, in the course of

the phylogenetic development of cockroaches, this condyle of

the mandible could pass from one segment to another with-

out interfering with the usefulness of the mandible during

the transition period. This was for us a perplexing problem

for a long time.

Later it was found that in Gryllus (Fig. 1) the mandible

articulates with a small sclerite which is sometimes distinct

from the clypeus. The same thing was found in the larva of

Corydalis, except that here (Fig. 3, ac) the sclerite is large

and clearly distinct.

As the ventral articulation of a coxa is with an antecoxal

piece (see page 27), we regard this sclerite as the antecoxal

piece of the mandible. In the course of the consolidation of

the segments of the head and of the dorsal flexure of the

sternites of the first three segments (the procephalon), the

antecoxal piece of the mandible has been pushed out of its

own segment past the labrum (the sternite of the second

antennal segment) and, in Gryllus, nearly past the clypeus.

Such a migration of the antecoxal piece could take place

without interfering with the action of the mandible.

Doubtless a factor in bringing about these changes is the

fact that while there has been a marked reduction in the length

of the head segments (the entire head composed of seven seg-

ments being approximately of the same length as a single seg-

ment elsewhere), there has been no reduction in the width of

the base of the mandible in biting insects. It is not strange,

therefore, that the least firmly fixed point of articulation, the

antecoxal piece, should be pushed out of its primitive position.

38 THE AMERICAN NATURALIST. | [Vor. XXXVI.

In most insects the antecoxal piece of the mandible coalesces

with the clypeus, so that the mandible appears to articulate with

this sclerite.

In some insects, as Sioned there is a distinct sclerite

between the mandible and the gena (Fig. 1, 77. This from

its position on the cephalic side of the base of the mandible

between the pleural and ventral articulations must be regarded

as the ¢rochantin of the mandible.

This completes our account of the sclerites found in the

external skeleton of the head of the more generalized insects.

A résumé of the conclusions reached is indicated by the fol-

lowing table.

SEGMENTS, SCLERITES, AND APPENDAGES OF THE HEAD.!

SEGMENTS. ScLERITES. APPENDAGES.

Vertex and gene.

L Oéular (Protecerebral).. © |. =... 4. ess 4 Ocular sclerites.

Front.

Antennal sclerites.

II. Antennal (Deutocerebral). «ce OO cQ Sts v.c tenen.

Clypeus proper.

III. 2d Antennal (Tritocerebral).| . . . . . . . . . | 2d Antenne of

Labrum (Mouth). Campodea etal.

Postgenæ.

IV. Mandibular. ` Costes. 0 20 7s s T ] MaundIbs.

Antecoxal pieces. Trochantin.

Pharyngeal sclerites.

V. Superlingual. ‘is wi. c € x » | Operos

: Maxillary pleurites.

VI. Maxillary. vta re wo se deN | Mana.

Lingua.

Dorsal cervical sclerites.

VII. Labial. Lateral cervical sclerites. | Labium.

Ventral cervical scl. (Gsl), |

pe Š In each section of the middle column the dotted line indicates the division

EO EE cd iM c E DEE DD t ME

PE RDUM TIMENS

No. 421.] SKELETON OF THE HEAD OF INSECTS. 39

THE ENDOSKELETON OF THE HEAD.

The endoskeleton of the head, like that of the thorax, con.

sists of invaginations of the body-wall and of chitinized tendons.

Some of these invaginations may be homodynamous with tho-

racic trachez, but others are obviously apodemes.

In many insects the mouths of some of the invaginations of

the wall of the head remain open in the adult ; in Corydalis, for

example, they are very conspicuous.

Some of the apodemes remain separate, and are comparatively

simple ; but in the case of two or three pairs of invaginations,

they meet and coalesce. In this way there is formed in the

interior of the head a complicated structure which is known as

the Zentorium (Burmeister, '82, Vol. I, p. 25).

The three pairs of invaginations which may enter into the for-

mation of the tentorium are known as the anterior, the posterior,

and the dorsal arms of the tentorium respectively. The coalesced

and more or less expanded tips of these invaginations constitute

the central portion of the tentorium, and may be designated as

the body of the tentorium. From the body of the tentorium there

extends a variable number of processes or chitinized tendons.

The Posterior Arms. — The posterior arms of the tentorium

(Figs. 26, 28, 29, pr) are the lateral apodemes of the maxillary

segment. In many Orthoptera the open mouth of the apodeme

can be seen on the lateral aspect of the head, just above the

articulation of the maxilla (Fig. 4). In the Acrididz (Fig. 28)

these apodemes bear a striking resemblance to the lateral apo-

demes of the thorax (Fig. 16), except that the ventral process

of the maxillary apodeme is much more prominent, and the two

from the opposite sides of the head meet and coalesce, thus

forming the caudal part of the body of the tentorium.

The Anterior Arms. — Each anterior arm of the tentorium

(Figs. 26, 27, 29, a£) is an invagination of the body-wall which

opens on the cephalic margin of the antecoxal piece of the

mandible (Fig. 2, a¢),! or, when the antecoxal piece is not distinct,

1 Note that owing to the flexure of the head in those insects, like Corydalis, in

which the mouth is at the anterior end of the body, the opening of the anterior

usi apte t OU O e e T T

40 THE AMERICAN NATURALIST. (VoL. XXXVI.

usually on the cephalic side of the cephalo-lateral angle of

the clypeus (Fig. 2, af). (It should be borne in mind that the

cephalic margin of the clypeus is that margin which joins the

front ; that morphologically the labrum is caudad of the clypeus.)

It has been shown by Carriére and Bürger (97) that the

position of this invagination in the young embryo indicates

that it is the spiracle of the mandibular

segment. It is easy to see that the

migration cephalad of the antecoxal

piece of the mandible, already described,

would push this invagination into the

position which it occupies in the adult

insect.

Bearing on this point is the fact that

| in Smynthurus, according to Lubbock

(73), the spiracles “open on the inner

d asi E dh cork” side of the bases of the mandibles.” It

; remains to be determined whether in

this case the anterior arms of the ten-

torium are wanting or not. Folsom

(00) found that, although the three

pairs of arms are present in the collem-

bolan Orchesella, the anterior arms are

wanting in Anurida. In Orchesella

where the anterior arms are present

they are described by Folsom ('99a) as

being joined to the paraglossz (super-

a | linguæ). Butas the invaginations form-

par EP nre ing the anterior arms arise cephalad of

the mandibles (Carriére and Bürger,

'97), they cannot pertain to the superlingual segment.

Usually the invagination forming the anterior arms is ex-

tended to a greater or less degree along the sutures that con-

verge upon it. This is well shown in Gryllus, where it is

furnished with three buttress-like extensions : one along the

suture between the front and the clypeus ; another between

. the front and the gena ; and a third between the gena and the

trochantin of the mandible.

No. 421.] SKELETON OF THE HEAD OF INSECTS. AI

In the cockroach (Fig. 26) the buttress of the anterior arm

that extends along the suture between the front and the gena

is twisted in a way that suggests that it has been pulled out of

place by the migration of the antenna. If we imagine the

antennal sclerite pushed into a position which would untwist

this buttress, it would bring this sclerite opposite the clypeus,

that is, in its assumed primitive position.

In the forms upon which the study is

chiefly based, — the Plecoptera, the Orthop-

tera, and the Neuroptera (Sialidz), — the

antecoxal piece of the mandible is joined to

the clypeus near its cephalo-lateral angle as

described above. But in the Hymenoptera

that we have studied it is joined to the

.caudo-lateral angle of the clypeus very near

the labrum. A result of this in some forms

is that the point of invagination of the

anterior arm is remote from the front, as in

the ants, although even here the buttress-

like extension follows the margin of the

clypeus and extends along the suture be-

tween the clypeus and the front. In other

Hymenoptera the opening of the invagina-

tion has migrated to the cephalo-lateral angle

of the clypeus and is thus remote from the |

articulation of the mandible ; this is the case Frc. 29.— Teniorium of

Fic. 28. — Head of Mela-

noplus, caudal aspect.

. Melanoplus, cephalic

in Bombus. cei

The Dorsal Arms. — Each dorsal arm of yay dorsal arms

the tentorium arises from the side of the

body of the tentorium between the anterior and posterior arms

and extends either to the front or to the margin of the antennal

:sclerite (Figs. 26, 27, 29, dz).

In the Plecoptera it appears to be merely a chitinized tendon,

the peripheral end of which is less chitinized than the base and

is only loosely attached to the skull. Here the point of attach-

ment is on the front, some distance from the antennal sclerite,

beneath a spot in the cuticle, which is marked by numerous

polygonal areas. This spot is very distinct and bears some

42 THE AMERICAN NATURALIST. [VoL. XXXVI.

resemblance to an ocellus (Fig. 19, dt). In the Orthoptera the

peripheral end is more strongly chitinized than in the Plecoptera

and firmly attached to the skull between the front and the

antennal sclerite. It is small in the cockroaches, but is larger

and easily seen in Gryllus (Fig. 27). In this case it bears some

resemblance to an apodeme.

The resemblance to an apodeme is much more marked in

some other insects. Thus in the Hymenoptera each dorsal

arm is firmly attached to the skull near the antennal sclerite,

and, in Cimbex for example, it is hollow and has an open

mouth, appearing exactly like an apodeme.

It remains to be determined whether or not the dorsal arms

in the Plecoptera are homologous with the apodeme-like dor-

sal arms in other insects, and, if so, which type is the more

generalized.

The Body of the Tentorium. — This is the part to which the

name ¢entorium was originally applied. It is the prominent

bridge which divides the occipital foramen into two parts. The

alimentary canal in entering the head passes above it, and the

nervous system below it. It is formed of the coalesced and

expanded tips of the arms of the tentorium (Fig. 28, 2).

The Frontal Plate of the Tentorium. — In the cockroaches

the anterior arms of the tentorium meet and fuse, forming a

broad plate situated between the crura cerebri and the mouth;

this plate may be termed the frontal plate of the tentorium

(Fig. 26, ff). On each side, an extension of this plate con-

nects it with the body of the tentorium; these enclose a cir-

cular opening through which pass the crura cerebri.

The Tendons of the Esophageal Muscles. — Muscles extend

from the body of the tentorium to the cesophagus, passing

between the crura cerebri. The tendons connecting these

muscles with the body of the tentorium are frequently chiti-

nized. In Pteronarcys there is a single chitinized tendon; in

Periplaneta there are two (Fig. 26, toe).

_ The Tendons of the Flexors of the Head. — These arise from

"e bed lower surface of the body of the tentorium and extend

Sometimes, as in dtt tron eel are e and

chitinized.

No. 421.] SKELETON OF THE HEAD OF INSECTS. 43

The Tendons of the Extensors of the Head.— These are

strongly developed in Gryllus; they project into the occipital

foramen (Fig. 25, ex.A.).

The Dorsal Apodeme of the Head. — This is also well devel-

oped in Gryllus (Fig. 25, d.ap.).

The Lateral Cervical Apodemes.— These have been described

above (see Fig. 20, af).

The Mandibular Apodemes. — These also have been described

above (see Fig. 25, maf).

In this study of the skeleton of the head our attention has

been confined almost entirely to representatives of the more

generalized orders of insects. We have felt that to do so was

the surest way to gain an idea of the fundamental plan of

structure. The working out of the ways in which this plan

has been modified in the more specialized groups of insects

must be left for the future and perhaps for other workers.

LIST OF ABBREVIATIONS.

ac. Antecoxal piece. dg. Leg.

ant. Antenna. md. Mandible.

ap. Apodeme. m. em. Maxillary epimeron.

as. Antennal sclerite. mf. ce field.

at. Anterior arm of the tentorium. Mx. axilla.

bt. Body of the tentorium. O. "usen

C. Clypeus. os. Occular sclerite.

C: First clypeus. Pg.

C2 Second clypeus. pl. Pleurite

cl Cephalic lobes. pr Procephalon

d. ap. Dorsal a e prs. Præstemum.

dt. Dorsal arm d i. tentorium. ps. Pharyngeal sclerites.

em Epimeron. pt. Pesterior arm of the tentorium.

es. Episternum. S.

e. su. Epicranial suture. Sp. Sternellum

ex. h. Extensors of the head. x Superlingua.

PF. Front. sp. Spiracle.

Jb. Frontal plate of the tentorium. z. oe. Tendons of the «esophageal

G. Gena. muscles.

Gu. Gula. tr. Ticchantin.

£. Labrum. : E. Vertex.

Z. i 2d mx. Second maxillæ or labium.

Lingua.

Jf. Lateral field.

44 THE AMERICAN NATURALIST. [VoL. XXXVI.

LIST OF REFERENCES.

1775. FABRICIUS, J.C. Systema Entomologiz.

1800. ILLIGER, KARL. Versuche einer systematischen Terminologie für

das Thierreich und Pflanzenreich.

1802. Kinsv, W. Monographia Apum Angliz.

1806. ILLIGER, K. Zusätze zu der Terminologie der Insekten. Llliger’s

Magazin. Bad. v, pp. 1-27.

1816. Savicny, J. C. Mémoires sur les Animaux sans vertèbres.

1824. AUDOUIN, J. V. Recherches anatomiques sur le thorax des ani-

maux articulés. Ann. Sci. Nat. Tome i, Pp: 97-135, 416-432.

1826. KinBv and SPENCE. Introduction to Entomology. Vol. iii.

1828. Srraus-DuRCKHEIM, H. Considérations générales sur l'anatomie

comparée des animaux articulés.

1832. BURMEISTER, H. Handbuch der Entomologie. Bd. i.

1834. NEWMAN, E. Osteology, or External Anatomy of Insects. Ænto-

mological Magazine. Vol. ii, pp. 60-93.

1839. NEwPoRT, G. Insecta. Todd’s Cyclopedia of Anatomy and

Physiology.

1863. WEISMANN, A. Die Entwicklung der Dipteren. Zeit. wiss. Zool.

Bd. xiii, pp. 107-220.

1869. BRANDT, A. Beiträge zur Entwicklungsgeschichte der Libelluliden

und Hemipteren. Mém. /' Acad. St. Petersb. Tome xiii.

1870. BürscHLI O. Zur Entwicklungsgeschichte der Biene. Zeit. wiss.

Zool. Bd. xx, pp. 519-564.

1871. PACKARD, A. S. Embryological Studies. Mem. Peabody Acad.

Science, Salem. Vol. i.

1873. LANCASTER, E. R. On the Primitive Cell-Layers of the Embryo.

Ann. and Mag. Nat. Hist. Vol. xi, pp. 321-339.

1873. LuBBOCK, J. Monograph of the Collembola and Thysanura.

1878. HuxLEv, T. H. Manual of the Anatomy of Invertebrated Animals.

1887a. VIALLANES, H. Etudes histologiques et organologiques sur les

centres nerveux et les organes des sens des animaux articulés.

4° Mémoire. Ann. Sci. Nat. Tome ii, pp. 5-100.

1887b. VIALLANES, H. 5e Mémoire. Ann. Sci. Nat. Tome iv, pp. 1-120.

1888. GRABER, V. Ueber die Polypodie bei Insekten-Embryonen.

Morph. Jahrbuch. Bd. xiii, pp. 586—615.

1888. PATTEN, W. Studies on the eyes of Arthropods. Journ. Morph.

| Vol. ii, pp. 97-190. |

1889. Haase, E. Die Abdominalanhange der Insekten. Morph. Jahr-

No. 421.] SKELETON OF THE HEAD OF INSECTS. 45

1890.

CARRIERE, J. Die Entwicklung der Mauerbiene. Archiv für

Mik. Anat. Bd. xxxv, pp. 141-

PATTEN, W. On the Origin of Seen from Arachnids.

Quart. Journ. Micr. Sci. Vol. xxxi, pp. 317-378.

WHEELER, W. M. A Contribution to Insect Embryology. Journ.

Morph. Vol. viii, pp. 1-160.

. HEvwows, R. Die Segmentirung des Insektenkórpers.

. COMSTOCK and KELLOGG. Elements of Insect Anatomy.

. CARRIÈRE and BURGER. Die Entwicklungsgeschichte der Mauer-

biene. 4A. d. k. Leop.-Carol. Deut. Ak. Bd. xlix.

UzeL, H. Vorläufige Mittheilungen über die — der

Thysanuren. Zool. Anz. Bd. xx, pp. 125, 129, 2

CLAYPOLE, A. M. The Embryology and Pe eee of Anurida

maritima. Journ. Morph. Vol. xiv, pp. 219-

Comstock and NEEDHAM. The Wings of Insects. American

Naturalist. Vols. xxxii and xxxiii.

UZEL, H. Studien über die Entwicklung der Apterygoten Insecten.

. FoLsow, J. W. (4) The Anatomy and Physiology of the Mouth-

Parts of the Collembolan Orchesella cincta. Bull. Mus. Comp.

Zool. Vol xxxv, No. 2. (6) The Segmentation of the Insect

Head. Psyche. Vol. viii, p. 391.

KORSCHELT and HEIDER. Text-Book of the Embryology of Inver- ;

tebrates. (a) Vol. ii; (4) Vol. iii.

Forsow, J. W. The Development of the Mouth-Parts of Anurida

maritima. Bull. Mus. Comp. Zool. Vol. xxxvi, pp. 87-157.

ON THE HABITS OF THE KANGAROO RATS

IN: CAPTIVITY.

DR. R. W. SHUFELDT.

SOME time during the early part of the month of June or

the latter part of May, 1901, Mr. Edward S. Schmid, a dealer

in pets and animals, with an establishment at 712 Twelfth

Street, Washington, D.C., received from one of his collectors

in Kansas some two dozen specimens of “kangaroo rats." At

first glance I did not recognize the species; but Mr. Schmid,

with his usual generosity, presented me with three of the finest

specimens in the lot, — two males and a female.

Upon taking these to my study I consigned them to a roomy

cage with an inch or more of soil on the bottom of it, and I

soon found that these very interesting little mammals fed with

great avidity upon hemp and canary seed mixed up with a

supply of wheat grains. They also drank freely of water

placed for them in little china vessels.

After they had been in my possession a day or two, I found

they had become sufficiently accustomed to my presence and

handling to allow me to make the attempt to obtain photo-

graphs of them. This I undertook on two separate occasions,

selecting for the purpose the darker and better marked male

animal of the trio. Both times I succeeded in obtaining life-

size pictures, and the reproductions of my results, reduced rather

more than one-half, illustrate the present article. Fig. 1 rep-

resents the animal as he appears when asleep during the day-

time, and Fig. 2 shows him when wide awake and engaged in

busily nibbling upon a piece of root at the entrance of a shallow

burrow he had dug for himself. As my methods of obtaining

such photographs as these have been fully set forth by me

during the past year or two in the technical journals devoted

to scientific photography in this country and abroad, it will not

be necessary to touch upon that question here.

48 THE AMERICAN NATURALIST. (VoL. XXXVI.

Upon exhibiting my photographs to Mr. Gerret S. Miller, Jr.,

of the Mammal Department of the United States National

Museum at Washington, D.C., he at once pronounced them to

be specimens of the kangaroo rat described by Dr. J. A. Allen

as Perodipus richardsoni, a species, so far as at present known,

confined to Indian Territory, Kansas, and Oklahoma.

During the last sixteen or seventeen years the number of

new North American species of jumping mice and kangaroo rats

described by our mammalogists has been something phenomenal.

Fic. 1.—P. ip ithardsoni(Allen). ĝ (asleep). Less than 14 nat. size.

Photographed from life by the author.

Other genera have been similarly increased. When the United

States National Museum published its provisional list of mam-

mals of North America in 1884 (Proc. U. S. Nat. Mus., Vol. VII,

Appendix, p. 585), there were but two subspecies of Dipodomys

recorded, and but one species of Zapus. If we turn now to

the excellent volume published by Mr. D. G. Elliot, curator of

the Department of Mammals of the Field Columbian Museum

of Chicago (Zoól. Ser, Vol. II, Chicago, 1901), entitled A

Synopsis of the Mammals of North America and the Adjacent

Seas, we find that there was recorded at the time of the

issuance of that book no less than 23 kangaroo rats and 20

Oe SUE RE :

No. 421.] KANGAROO RATS IN CAPTIVITY. 49

jumping mice (Zapus) ; 13 of the former are of the genus Dipo-

domys and 10 of Perodipus, the genus to which our present

subject belongs. (See pp. 236, 237; Family (VI) Heteromy-

idee; and for three views of the skull of Perodipus agilis,

p- 2 Other genera of this extensive group of our smaller

rodents have likewise been largely added to, and a great many

new species described of recent years. Mr. Elliot has since

published a supplement to the volume here referred to, and

this contains the descriptions of still other species.

I found my specimens of P. richardsoni extremely gentle in

disposition, and any one of them would allow me to catch hold

of it with my hand, and no amount of handling or even gentle

squeezing could induce the little animal to bite or scratch. On

one occasion I let them all out in my study, when it was very

amusing to see their various antics and capers. They were as

full of fun and play as could be, and soon seemed to take a

genuine delight in my attempts to recapture them. They

hopped about with great agility on their hind pair of kangaroo-

like legs, while the little short pair of anterior limbs were

curled inwards on the chest. At these times the animal’s tail

is simply dragged behind it, being in contact with the ground

for its entire length. Its body is held obliquely, its axis making

rather a small angle with the surface over which it is passing,

but if occasion occurs for it to use its fore legs, they are

brought into play at once, either for feeding, climbing, or

burrowing. They are able climbers, and the rapidity with

which they can dig a burrow in ordinary ground is astonishing.

They use the fore feet to perform the digging part and the

long and strong hind legs to kick the loosened soil out of their

way behind, as it accumulates every moment or so. In soft

soil one of these little mammals can put itself out of sight in

less than a minute by digging, and it really seems to enjoy the

operation. Among themselves they are somewhat inclined to

be quarrelsome, biting and scratching one another sometimes

without any apparent cause, while at other times they huddle

together in a corner and sleep as peacefully as so many snails.

- The borders of the ears of one of these males, however, exhibit

a few small healed-up nicks and notches that have very much

Mo. Bot. t Gardou

50 THE AMERICAN NATURALIST. (Vor. XXXVI.

the appearance of having been acquired during combats with

its fellows. During most of the day they pass the time in

sleep, but they become very active towards dusk and probably

are active all night. So it is that they rarely feed during the

daytime, while after dark they are hearty consumers of any-

thing that takes their fancy, especially of such grain and seeds

as I have mentioned above. When sleeping they sometimes

curl their long tails about them in a circle on the ground, while

the nose and face is poked well down in between the fore legs,

the body thus looking like a round puffball of hair surrounded

Fic. 2. — P. vichardsont. Same specimen as shown in Fig. 1. Feeding. Less than % nat. size.

Photographed from life by the author.

by a single coil of the bicolored and longitudinally striped tail,

the latter terminating in a brush at its end. Again they may

sleep in the attitude shown in Fig. r, where the animal had

mounted a little log for the purpose and had been in sound

repose for fully ten minutes, without moving, before I exposed

my plate upon him. At other times they sleep upon tbeir

backs or sides, stretched out like little kittens or other small

animals that assume such attitudes when enjoying a blissful

doze in the warm sunshine.

This kangaroo rat is a very neat and cleanly little creature,

frequently dressing it$ soft fur much after the fashion of the

No. 421. KANGAROO RATS IN CAPTIVITY. I

common house mouse. Sitting up like a kangaroo, it will

vigorously, and with both fore paws, dust its nose and face for

a few seconds, when, with equal alacrity, it will pass to a

general scratch of its sides and back, terminating the operation

by starting at the root of its long and stout tail and rapidly pass- `

ing the entire appendage through its fore paws to the very tip,

-while it is, as it were, at the same time briskly titillated with

the lips and teeth at the front of the mouth. It is very amus-

ing to observe the evident satisfaction it has in performing

this act.

In addition to preening thus the face, body, and tail, it will,

to dress the hair of the belly and back, push itself along in

the dust or earth by means of its hind legs, and afterwards

vigorously brush out the fur with its fore paws.

Another peculiar habit I observed in Perodipus was the way

it had of using its fore paws for quickly pulling out the cheek

pouches. By this means the pouches were completely turned

inside out ; they were then stretched, scratched, and dusted

for a second or two, after which the animal tucked them back

into position with equal rapidity.

As in the case with the majority of the Muridæ, when

startled by sharp and sudden noises these nervous little ani-

mals will involuntarily spring from the ground for a centimeter

or more ; when coming down again they will stand and gently

grit their teeth together, while their eyes appear as though

they were about to pop out of their orbits. At other times,

particularly when feeding, the eyes are often kept not more

than two-thirds open, and the animal then has an especially

gentle look (Fig. 2).

Perodipus occasionally, at long intervals, gives vent to a

peculiar note not altogether unlike the low sound of a cat when

calling her kittens together, though not nearly so loud or distinct.

If kept in a roomy cage with five or six inches of soil on the

bottom, where it is perfectly quiet and sunny, I see no reason

why this species would not breed in captivity, and in any event

they make very interesting and gentle little pets in confine-

ment, quite equaling any of the squirrels, or even the famous

dormouse of Europe, in this regard.

A CONTRIBUTION TO MUSEUM TECHNIQUE.

S. E. MEEK.

To exhibit fishes properly in a museum has been no easy

task. Many methods have been devised, but none have as

yet given universal satisfaction. Land animals, such as mam-

mals, birds, reptiles, etc., are mounted and arranged in cases

according to their natural order, to show relationship, or they

are mounted in groups illustrating some of their habits and

natural surroundings. These methods have received universal

approval. With fishes the case is quite different. To mount

them is difficult, and in most cases unsatisfactory, while many of

the smaller soft-rayed fishes cannot be mounted. Painted plas-

ter casts, or casts made of other material and painted, are used

in some museums. In case nothing better can be had, these

serve a good purpose and are especially desirable when a

collection is to be exhibited at different times in various places.

They are simply representations, and do not meet the desire

of the visitor as do the real fishes themselves. A mounted

fish is “a fish out of water," a fish robbed of his natural

» surroundings.

Three years ago we began some experiments in this museum

to devise a metal case or vessel with a plate-glass front in which

we could exhibit fishes in alcohol in a horizontal or natural

position. A joint between the metal vessel and the plate glass,

which would hold alcohol, and which would compensate at any

temperature for the unequal expansion of the plate glass and

metal, was devised by Mr. Wines, our building superintendent.

Under his direction a vessel was made in December, 1898, with

a plate-glass front of 18 by 36 inches. Nine species of sunfishes

were placed in this vessel, and it was filled with 7o per cent

alcohol, and put on the wall of one of our exhibition rooms. A

few weeks later another case of 14 by 32 inches was made, and

in it was placed a large blue-black trout from Lake Crescent,

54 THE AMERICAN NATURALIST. [VoL. XXXVI.

Washington. These two cases have so far held satisfactorily,

having been subjected to temperatures from 20° F. to 90° F.

Two smaller cases put up at the same time have also proved

very satisfactory.

In 1899 Dr. A. B. Meyer, director of the Natural History

Museum of Dresden, made a tour of this country to study our

museums and scientific institutions. It was my pleasure to escort

him through the department of zoólogy in this museum. I called

his attention to one of these cases and he remarked, * Very

attention to an article, * Ein Beitrag zur Museumtechnik," by

Dr. Max v. Brün, published in the Abhandlungen aus dem Gebiete

der Naturwissenschaften (Bd. XIII), herausgegeben vom Natur-

wissenschaftlichen Verein in Hamburg, 1895. As this article

is not easily accessible to all who are interested in the exhibition

of fishes, I give here an abridged translation of it.!

The zoólogical museums are constantly progressing in their

efforts to acquaint the public with the animal world by means of life-

like representations. Those animals which are robbed. of their

natural appearance, in a greater or less degree, by insufficient

1 Published with the permission of Dr. Brün.

No. 421.] MUSEUM TECHNIQUE. 55

methods of preservation and preparation, are gradually being

replaced by properly prepared material, so that museums are con-

stantly asking less of the fancy or imagination.

The problem of preserving natural colors in alcoholic specimens is

still unsolved, and yet the impressions which animal forms make on

the eye, and through it on the understanding, depend much on color.

It is not so much the task of our museums to preserve the natural

color of an animal as to give it a lifelike appearance after the

color has faded or has been lost. If for exhibition purposes we

do this properly, our methods need not be questioned. To repre-

sent absolutely true to nature an individual specimen, is in most

cases as impossible as it is unnecessary; for even in nature the

individuals of any species are not all made according to the same

mould, but they differ from each other within certain limits, not less

in color than in other characteristics. The problem of preserving

natural colors is reserved for preserving liquids, as Wiese-liquid,

which by means of its chemical action seems well adapted for this

purpose.

Preceding the opening of the Natural History Museum of Ham-

burg, it was my wish so to exhibit fishes that they should appear as life-

like as possible. The methods known to me up to that time did not

appear favorable, so I began to think of something new. I did not

believe fishes should be exhibited in a dry state, — vis., embalmed or

stuffed, — nor in tall bottles, standing on head or tail, but that they

should be in a horizontal position, seemingly swimming in a medium

representing water.

This was attained by means of oblong glass boxes filled with

alcohol. It was then observed that the colors which gave the fish a

true natural appearance were absent; this was to be helped only by

painting, and accordingly an attempt was made with water colors

which led to satisfactory results. It has so met the favorable judg-

ment of visitors and professional men that I venture to believe

myself in a position to commend this method as worthy of imitation.

. In Bergen, Norway, the entire fish fauna has been set up in this way,

and is very satisfactory. This method is also in use in the British Museum,

as well as in German museums.!

The method consists in painting the fish, preserved in alcohol, on

one side with water colors, as nearly natural as possible; and then

1The paragraphs in smaller type are from letters written to the author by

Dr. Brün.

56 THE AMERICAN NATURALIST. [VoL. XXXVI.

fastening it in a horizontal position by means of gelatin to the wall

of the glass box, which is later filled with alcohol. A fish thus suc-

cessfully prepared appears, to the visitors at least, as a living fish in

water. Its lifelike appearance is much improved in the proper

painting of the eye, which is not at all difficult. These water colors

are durable in alcohol only when they are carefully selected, as has

been proved by four (now ten) years’ experience. A fish thus com-

pleted needs no repairing for some time. While in a general way

this method can be recommended, yet in a few instances it may not

work, especially when we are dealing with forms that have a bright,

sparkling appearance, such as goldfishes. In such cases the Wiese-

liquid seems to promise good results. A large goldfish preserved

in this liquid for a period of two (now eight) years, though kept in

the dark, has lost none of its brilliancy and beauty of color.

' The using of oblong glass boxes is more expensive than glass

bottles, but when we are dealing with a native fish fauna with com-

paratively few specimens of considerable size, this evil need not be

permitted to play a large part. It is a fact that the mechanical pos-

sibilities of producing sufficiently large glass boxes for an available

price are limited. The largest glass boxes prepared for this museum

are 7o cm. long, and cost 24 marks each. Because of this, only the

smaller specimens of the larger species, such as salmon and catfishes,

can be easily exhibited. Very large specimens of these can be

shown along with the smaller ones, either as mounted specimens or

plaster casts, or they may be exhibited in glass boxes, made of glass

plates cemented together.

The German marine exhibit in 1896 afforded me an opportunity to dem-

onstrate the fact that these glass boxes with their contents could be easily

transported from one place to another. Besides some small ones, the

large glass boxes containing the salmon and the lamprey were taken from

Hamburg to Berlin and returned in fine condition. The transportation was

by boat on the Elbe, but, being taken to and from the boat by wagon, they

were subject to considerable jarring.

The task of putting up one of the above-mentioned fishes may be

divided into three parts: (1) preparation and preservation ; (2) paint-

ing ; and (3) setting up in the glass boxes.

(1) As the fish must be presented in a swimming position in the

glass box with parallel walls, it must be preserved as perfectly as

possible. In selecting a specimen one must pay special attention to

form, color, proper size. Most of the fishes found in the markets

are iex about the munde fins, or scales. Pretty badly injured

WES ee sts R EE Ie WORE SY hey Goble NEAR te du o X I ERE nU UP eee TE PENES PEN e T {

Pale ge cae

No. 421.] MUSEUM TECHNIQUE. 57

fins can be repaired. It is always best to select fishes to be mounted

at the place of capture, and thus avoid the injuries occasioned by

fish dealers in frequently throwing fishes from one vessel to another.

As only one side of the body is exhibited, it should be the better

one, though as far as possible, for sake of uniformity, the same side

of each should be presented to the visitor, as in my opinion it does

not make as good an impression when some fishes are turned to the

right and others to the left.

It is advisable to keep the live fish for some time in an aquarium,

and make a study of its coloration. Then kill the fish in a 10 per

cent solution of alcohol.

In using strong alcohol the mucous layer on the fish becomes too much

hardened and cannot be removed.

After one half hour or so the mucous layer can be removed by

means of a brush. Next lay the fish on its side in a shallow vessel,

the bottom of which is lined with wax, and put on it some stronger

alcohol. The fins are then spread, and held in position with insect

needles. The torn fins are mended with fine silver wire, which is

later removed. After the fish is in its proper position it is hardened

by the gradual strengthening of the alcohol; the time required for

this depends on the size of the fish, one of about two pounds’ weight

requiring about six days to bring the alcohol to 75 per cent.

In successful cases the form of the body is scarcely changed at

all, and the eye fills its cavity as when the fish was alive. It is

well, in case of larger fishes, to inject some alcohol into the vent

after killing and to close it with cotton. The intestine should first

be pierced, also the inner muscles of the body, with a fine scalpel,

that the injected alcohol may fill the body cavity and more easily

enter the flesh.

Externally the fish should not be injured in order to make the

alcohol penetrate more rapidly, since where injured the fish would

not get dry enough to paint, and injury to the farther side would

‘interfere with fastening the fish to the gelatin. Should the belly still

sink to an interfering extent, the proper roundness can finally be

secured by stuffing it with cotton.

Later I have used formaldehyde 1:20, which I believe is to be com-

mended to keep the body shape well. The Sad is first killed in weak Mx

then hardened in formalin, and later transferred to alcohol f.

preservation. With the formalin treatment t ‘ales of some e species come

off too easily.

58 THE AMERICAN NATURALIST. (VoL. XXXVI.

(2) The painting of the fish faded by use of alcohol. For this pur-

pose I have used water ground color and occasionally some marine

blue. A few kinds of these colors are, however, to be avoided;

in the first place, those containing lead, such as white lead and

chrome yellow. The alcohol in which a fish has been for some time

finally becomes somewhat acid, and at the same time the process of

decomposition forming H,S turns the colors containing lead dark or

black. That is the way I was served in several cases ; the pretty

white belly of the fish finally appeared quite blackened. Chemical

examination of the alcohol showed that there was present .034 per

cent of free acid, supposed to be acetic acid (.348 gr. to one liter of

alcohol). Under these conditions silver and brass bronze, usually

appearing so durable, turned dark. Experiments with H,S + H,O

resulted in great durability of the same, but changed immediately

after any trace of acid was added.

Aluminium bronze remained unchanged, even after the addition of

considerable acid. Therefore one should use permanent white fine

aluminium bronze and clear gold bronze. For use these are worked

with a little liquid gum arabic.

Dr. W. G. Ridgwood has carried out in the British Museum very thor-

ough tests for the most varied colors, as to their permanency in alcohol and

sunlight, and has found decided differences.

The method of painting is as follows. The hardened fish is laid

on a suitable surface. It is best to color the trunk and head first.

In the mean time the fins must not be allowed to dry. The eye

should be kept moist before and after painting, else these parts will

dry out too quickly and shrink. If after a few minutes the body

seems sufficiently dry, then one may begin with the painting, which

after a little practice may be done without any special artistic

ability. The paint cannot be applied as perfectly as it can be on

paper, yet still in a sufficient degree to obtain the desired effect.

This is most easily done in case of small scaled fishes, for example

the tench; in other cases patience and practice lead to the goal.

The process of painting is delayed very much because of one's being

occasionally compelled to moisten the parts with alcohol, in order

_ to observe the action of the colors, and eventually to change the

tone of the same. |

Because of the escape of fat or similar substances, the opercle,

and many scales, and fin rays occasionally do not seem to take on the

Eod EMT

No. 421.] MUSEUM TECHNIQUE. 59

would perhaps be best to give them a coat of ox gall, which is used

in painting on glass. The pupil should be colored a bluish black

. and surrounded with a golden circle verging into the iris; to the iris

one can reproduce the niceties of its natural appearance, namely, its

silver or gold glint, red circles and spots, black stripes, dots, etc.’

As soon as the paint is dry the eye should be moistened; otherwise

it would sink, though a little sinking is hardly noticeable when the

fish is put in alcohol.

Injured fins must be repaired before painting, which may be done

by pasting on underside small pieces of silk paper with a thin solu-

tion of gelatin. The repaired places must be carefully dried before

painting.

(3) The fastening of the fish in the glass.

_ The glasses used in our museum are blown oblong, with parallel

walls. There are three special sizes produced by the glass factories

of Gundlach & Miiller of Altona-Ottensen. Their comparative sizes

in centimeters and their prices, with 5 per cent discount, are as

follows :

LENGTH. HEIGHT. DEPTH. PRicE.

35 20 9 8 marks

50 25 II ER n

70 30 14 "M. ”

These glasses are unground: those with polished face are more expen-

sive, but appear also much more elegant. The firm already mentioned has

filled many orders for such glasses, and has of course much experience.

Transportation of the same, even of the largest glass boxes, has been suc-

cessfully made to America without material loss.

These glass boxes are closed with sheets of glass about 3 mm.

thick and the edges ground to correspond to the thickness of the

walls of the box, to which they are securely fastened with gelatin.

The closing of glass boxes, especially the larger ones, is not easy.

When the collection is to stand some time undisturbed it is well to close the

larger ones by simply covering with the glass lid. As the alcohol slowly

evaporates it may be replaced. To open the glass boxes which have been

closed with gelatin, it is better to cover the lid with layers of blotting paper

and these with a wet cloth. After some time, say over night, the gelatin is

softened, and the cover can be released with a little care.

In order to fasten the fish in a horizontal position in the glass

case, it should first be laid on its broad side in the manner in which -

it is to be fastened in the glass. The tail should be supported with

a piece of cork, or something of the kind, which is covered with

60 THE AMERICAN NATURALIST. [VoL. XXXVI.

gelatin. Remove the fish and place where the head is to lay a layer

of pretty stiff gelatin liquid, also place some on the cork ; then lay

the fish, previously well dried, on its unpainted side in its proper

position in the glass. Soon the gelatin will be sufficiently hardened

to permit the filling of the glass box with alcohol. If necessary, a

hot knife may first be applied to the gelatin to fasten the fish

securely to the glass. An eight-pound salmon fastened in this way

has remained unchanged. The visitor sees nothing of this manner

of fastening.

As I have in the foregoing only spoken of the painting of fishes, I

should here like to say that other objects to be kept in alcohol may

be treated in the same manner, — reptiles, amphibians, invertebrates,

anatomical preparations, plants, etc.

In our museum at Hamburg there are numerous painted objects of this

kind from both fresh and salt water, and though very imperfectly painted,

they have a lifelike appearance. Especially is this true of water plants.

Such objects as sponges, which should have a uniform coloring,

and which will stand being in water for a little while, may be put

into a watery solution of the color, where they quickly become

impregnated with it.

Moulded glass vessels are not as perfect as they should be ; the

surface is not exactly smooth; besides, it is made more or less

dim by being in contact with the mould. The glass is thicker

at the center and gradually thins out toward the edges and cor-

ners. The side of an oblong moulded glass vessel is quite inferior

to a similar surface of polished plate glass. It is not necessary

that a vessel be made entirely of glass; the top, bottom, two

ends, and farther side may as well be made of any other suitable

materia. The side through which the observer must see the

specimens cannot be too perfect.

A few years ago Mr. J. E. Benedict, of the United States

National Museum, appreciating the value of a glass vessel whose

sides (front one at least) are planes, experimented considerably,

constructing vessels by cementing together sheets of plate-glass.

To hát extent he now regards this method as a success I am

I never did believe his method received the recog-

t deserved. It seems to me, however, that by using

tho 2 the HM 1 surface must necessarily be limited,

No. 421.] MUSEUM TECHNIQUE. 61

just as in the case of the cast-glass boxes. The idea of repre-

senting a fish in a museum in a fluid so that it “appears, to the

visitor at least, as a living fish in water” is an excellent one.

Following out this idea, our exhibition vessel partakes somewhat

of the nature of an aquarium. A well-constructed aquarium

needs but one glass side, and this side should be polished plate

glass. It is necessary to make a joint between the material

used and the plate glass which will not be effected by changes

of temperature, and this presents no serious difficulties.

We have not tried the painting of fishes or fastening them in

the vessel by means of gelatin. The use of round or oval glass

bottles has been, within the past few years, quite universal.

But both of these bottles are objectionable, because in them the

fishes must stand on head or tail, and besides suffer more or less

distortion. Among fishes are many peculiar and interesting

forms, and this class ‘of animals are as deserving of suitable

methods of exhibition as are the other classes.

The time has certainly come when some method which will

permit fishes to be placed in our museums as lifelike as possi-

ble, and one which will be economical and durable, will be much

appreciated. If those who are interested in this question, and

who have given it some attention, will publish the results of

their experiments, it is possible that enough good features can

be selected to discover the proper method.

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Jenkins and Kellogg on Nature Study. — Under the title of

Lessons in Nature Study Professors Jenkins and Kellogg of Stanford

University have issued a volume (Whitaker & Ray, San Francisco)

which should be of great service to teachers. It contains simple

directions for the study of natural objects within reach of children.

Among these are the dandelion, the mosquito, the toadstool, pond

insects, spiders, crystals, ferns, birds, etc. These lessons are based

on actual experience in dealing with children. The book is well

illustrated and fairly printed. Bis,

ZOOLOGY.

« Bird Watching " ! is the modest title of a new volume in the

Haddon Hall Library. There are a dozen chapters, dealing with a

variety of British birds, and constituting a record of patient * watch-

ing" of great plover on open, sandy wastes ; wheatears on warren

lands; great skuas and shags on far northern islands ; winter birds

at a straw stack, etc. "The account of the actions of these and other

birds often takes the form of long extracts from the author's note-

book, but his personality and feeling for style give his observations

an unusually bright and readable character. Occasionally he pushes

his fancies to the verge of extravagance, and frequent digressions

increase the burden of legitimate detail which the reader willingly

bears. While there is constant evidence of painstaking accuracy of

statement, the author's interest in the problems connected with the

evolution of habit and plumage tempt him often into rather fruitless

speculation. :

The book must be very welcome to the lovers of British birds and

will be read with interest not only by American ornithologists but

' Selous, Edmund. Bird Watching. The Haddon Hall Library. London,

J. M. Dent & Co., 1901. 8vo, x + 347 pp., 14 illustrations.

64 THE AMERICAN NATURALIST. [Vou. XXXVI.

also by many of the growing class of amateur naturalists. Detailed

observation of familiar birds (such as Mr. Frank Bolles’s paper on

* Sap Suckers ") will constitute an ever-increasing proportion of the

literature of our popular ornithology, which has hitherto been marked

by the not unnatural tendency to magnify the importance of a new or

unfamiliar species.

The paper, print, and binding are extremely attractive. There are

fourteen full-page drawings by J. Smit. These drawings, if neces-

sarily less accurate than the now omnipresent photograph from nature,

possess a welcome softness and repose. R. H.

A New Economic Entomology.!' — In a neat little volume of about

three hundred pages, Prof. H. Kolbe has compiled an admirable

treatise on Gartenfeinde und Gartenfreunde; it constitutes Vols.

XXXIV-XXXVI of the great German *Garten-Bibliothek." This

little venture of the author into the economic field will be of interest

to American entomologists, as we are familiar with his great work,

Einführung in die Kenntnis der Insekten. Notwithstanding the wealth

of German literature on injurious insects and plant diseases, Garten-

Jeinde und Gartenfreunde fills an unoccupied niche, for it deals with

both the animal (principally insects) and fungous enemies in a way

suited to the needs of every German gardener.

After a brief discussion of the structure, the classification, and the

various groups of insects, a dozen pages are devoted to remedial and

preventive measures. Here the author introduces some of our Ameri-

can methods, but it is evidently unfamiliar ground. Sirrine's poisoned-

resin mixture is recommended for garden caterpillars, and the Bor-

deaux mixture for nearly all fungous diseases; but our kerosene

emulsion he calls * petroleumbrühe," and under “kerosene emulsion ”

he gives the formula for our potash whale-oil Soap. Among reme-

dies for plant-lice and thrips he mentions Paris green used dry. It

is also an interesting fact that nowhere else in the book are any of

our arsenical or other poisons recommended for killing insects; cur-

rant worms are to be shaken off onto a sheet instead of poisoned, and

the universal poison spray of American orchardists for the codling-

moth and other caterpillars is not mentioned. Most of the remedial

measures involve hand work. The discussion of preventive meas-

ures is excellent, and the chapters on state, communal, and social

! Kolbe, H. Gartenfeinde und Ga de. Die für den Gartenbau schüd-

lichen und niitzlichen Lebewesen. Garten-Bibliothek, Bd. xxxiv-xxxvi. Berlin,

Karl Sigismund, 1901. 8vo, iii-- 518 pp., 76 figs.

No. 421.] NOTES AND LITERATURE. 65

institutions and regulations relating to plant protection in different

countries will interest American readers.

The admirable method is followed of discussing the enemies under

their respective food-plants, and over sixty pages are devoted to fruit-

tree enemies. The San José scale receives more space than any

other enemy except the grape phylloxera, and curiously enough the

Aspidiotus ostraformis scale is called the “ yellow European pseudo-

San José scale." Injurious mammals and birds are briefly discussed,

and then fifty pages are devoted to “Gartenfreunde,” mostly the

parasitic and predaceous insects.

I know of no similar work in any language covering so much

ground so briefly and yet so well adapted for its field.

M. V. S.

The Circulation in the Nervous System. — In a pamphlet of some

hundred and fifty pages bearing the above title, Dr. Gasser! develops

what to his mind *is the only true conception that is in entire

harmony with the established order of facts in the world of thought "

concerning the action of the nervous system. Precisely what this

conception is the author nowhere makes very clear, but so far as can

be gathered his experience as a physician has profoundly impressed

him with the belief that the nervous system works in a circular

fashion. What circulates, whether matter or energy, and through

what particular channels the circulation is accomplished, is left mostly

to the imagination of the reader, though we are told that the evidence

for this movement is as substantial as that for the circulation of the

blood. As a figure of speech the circulation in the nervous system

may be innocent enough; as a description of what actually takes

place it is utterly without foundation.

The Oarfish, Regalecus, on the Coast of Southern California. —

On the 25th of February a huge “sea serpent," with bright colors

and the conventional mane, was reported in the newspapers as hav-

ing come ashore near Santa Ana in southern California. No part

of the animal was preserved, so far as known, but a good photograph

was taken by Mr. G. T. Peabody of Santa Ana. It evidently repre-

sents an oarfish or herring king, some species of Regalecus. The

fish was fifteen feet long and something over a foot in depth, weigh-

ing about five hundred pounds. The dorsal is considerably injured,

1 Gasser, H. The Circulation in the Nervous System. oe

Publishing ee 1901. 156 pp.

66 THE AMERICAN NATURALIST. (Vor. XXXVI.

but as nearly as can be counted the number of rays is about four

hundred. The long spines on the head do not show in the picture.

The species of Regalecus have never been defined, and the present

one can hardly be certainly identified. It may be the New Zealand-

Japanese-Indian species, Regalecus russelli or argenteus. 5.8. T.

Notes on Recent Fish Literature. — In the Annotationes Zoologice

Japonenses (Vol. IV, Pt. I), Dr. S. Hatta has a review of the Japa-

nese lampreys, with notes on their local distribution and their ana-

tomical structure. The new species, Lampetra mitsukurii Hatta,

already characterized by Dr. Jordan, is here described. In another

note Dr. Hatta discusses the metameric segmentation of the meso-

blast in the lamprey and the lancelet. In the earlier stages the

segmentation in the lamprey agrees with that of the lancelet ; in its

later stages with that of the higher craniota.

In the Atti de la Società di Naturalisti, etc., of Modena, Dr. Luigi

Facciolà continues his valuable series of studies of the development

of the larvz of different forms of eels. These larva, known as Lep-

tocephali, are ribbon-shaped, translucent, and soft, with very small

heads. In their further development they undergo a shrinkage in

size, attaining greater compactness of substance.

In the Proceedings of the United States National Museum (Vol.

XXIV, pp. 33-132), Jordan and Snyder continue their series of

monographic reviews of the fishes of Japan. Of the gobies, 57 spe-

cies are described, representing 33 genera. Of these species, 21 are

new and are here represented in the excellent drawings of Mrs.

Chloe Lesley Starks.

We may here note that the generic name Chasmias, proposed by

Jordan and Snyder for a genus of gobies in 19oo (page 76 1), is ante-

dated by Chasmias Ashmead, a genus of Ichneumon flies published

on page 17 in the same volume of the Proceedings. The genus of

fishes, type Chasmias misakius, may receive the new name of Chas-

michthys Jordan and Snyder. I am indebted to Mr. W. H. Ash-

mead for calling attention to the prior establishment of his genus.

Of sea horses, pipefishes, and their relatives, nineteen species are

described from Japan. Of thet seven are described and figured as

new. The sea horses exist in espe cially large number and pur

. along the headlands | washed by « the warm “ Black cc o

; “ Kuro Shiwo.” 2

ss s eoa

No. 421.] NOTES AND LITERATURE. 67

Professor Gilbert L. Houser, of the University of Iowa, contributes

to the Journal of Comparative Neurology (Vol. XI, No. 2) a valuable

monograph on the ** Neurones of a Selachian," Mustelus canis, The

anatomy of the nerve structures is given in great detail. Professor

Houser shows commendable caution in refraining from “the tend-

ency to elevate the results of specific methods into exclusive dogma.”

He closes his paper with these remarks suggestive of Golgi: * The

knowledge which we possess, either anatomical or physiological, is

not yet such as to permit us to interpret with certainty the greater

number of the facts discovered, much less to attempt doctrinal con-

structions of a high order on the functional mechanism of the ner-

vous elements."

In the Bulletin of the United States Fish Commission Dr. Eigenmann

gives a useful account of the development of certain eggs supposed

to be those of the Conger.

In the same Buletin Dr. Hugh M. Smith gives a list of species of

fishes the young of which are borne to Woods Hole in the Gulf

Stream. Four of these, Exonautes rondeleti, Ocyurus chrysurus,

Scarus croicensis, and Sparisoma flavescens, had not been previously

recorded to the north of Florida. Among the other tropical forms

are Sardinia pseudohispanica, Mycteroperca bonaci, Mycteroperca vene-

nosa, Lutianus apodus, Lutianus analis, Chetodon ocellatus, Chetodon

bricet, Teuthis ceruleus, Teuthis hepatus, Teuthis bahianus, Lactophrys

tricornis, and Scorpena plumieri.

In the Annotationes Zoologica Japonenses (Vol. III, Pt. IV), Dr. C.

Ishihawa describes and figures two new gudgeons, Leucogobio giin-

theri and L. jordani. Both species belong to the rich fauna of Lake

Biwa, the largest lake of Japan.

In the Proceedings of the United States National Museum (Vol.

XXIV) Jordan and Starks give an account of the anatomy of Apha-

reus, a Polynesian genus of “red snappers," rare in collections.

In the Proceedings of the United States National Museum (Vol.

XXIV) Jordan and Starks give an account of the Atherinidz, or

silver-sides, found in the waters of Japan. Five species are enu-

merated, four of them being new. Two new genera, Atherion and

Iso, are chamoterised, : D. s. L

Zoitogy | of the Maldive and Laccadive Archipelagoes. — The

epu b ecu one Su Gardiner in the Maldive and

68 THE AMERICAN NATURALIST. (VoL. XXXVI.

Laccadive archipelagoes of the Indian Ocean’ are appearing in a

series of parts that closely resembles the series of Willey’s “ Results.”

The first part contains an eleven-page “ Narrative and Route of the

Expedition,” with two maps. This shows that one who goes on such

a journey may expect exasperating delays of weeks at atime. Next

follows * An Account of the Coral Formations of the Indian Ocean,”

by Gardiner, which is not completed, but shows that atolls have been

formed in regions of e/evation (and not always of depression, as called

for by Darwin’s theory). The accompanying papers are by Borradaile

(who was with Gardiner in the early part of his trip) on the ** Land

Crustaceans”; by Cameron on “ Hymenoptera”’; and by Punnett on

“ Nemerteans.” In the last paper the first mesonemertean from the

Indian Ocean is recorded, — a Cephalothrix. The prevailing genera

are Eupolia, Cerebratulus, and Drepanophorus, — genera widely dis-

tributed. As to the Hymenoptera, Cameron says the known species

are Indian forms of wide distribution in the Oriental zodlogical

region, and all the genera are of universal distribution in temperate

and tropical countries, — such familiar genera as Crabro, Bembex,

Polistes, and numerous Apide. The most general interest attaches,

however, to the paper on “Land Crustaceans.” ‘Their numbers,

their ubiquity, their activity, combine to give them a prominence

which is all the more marked from the absence of so many other land

animals of continental areas. They are the chief scavengers of the

island, play a great part in the destruction or disintegration of fruits,

and probably aid in the distribution of seeds. The work done by

them in burrowing along the sandy lagoon. shore has a possible

importance not hitherto noticed." The paper gives an exhaustive

account of the anatomy of Ccenobita, the land hermit crab. The gills

seem to have undergone little modification, and the gill chamber is

not at all a lung, but provision is made for keeping the gills moist by

means of salt water apparently retained from possible rare visits

to the sea. If this salt water is removed, a sticky fluid is exuded over

thegills. If thegills be cut off, the crab lives by virtue of its abdomi-

nal respiration. Continuous submersion is fatal after a longer or

shorter time, — one to six days. The habitation of the land hermit

crab is "X varied, — usually any kind of land or sea gastropod shell;

i: 1 gemens J. Stanley (editor). The Fauna and Geography of the Maldive and

ac rk on and of the collec-

tions made by an expedition during he years 1809 and 1900. Vol. i, pt. i, Pls.

I-V, bue ire A = = University P Serene C uà —-

1905. be. m

No. 421.] NOTES AND LITERATURE. 69

but also a serpulid worm tube, a half cocoanut shell, and a broken

glass tube were appropriated. Ocypoda occurs here in two species,

one of which lives and burrows in the sand, the other inland. Three

grapsids are found ; two Palæmonidæ, a Ligia, four Oniscidz, and

one of the Armadillidiide.

Recent Sporozoan Investigations.' — This pamphlet, which is a

revision and expansion of the articles published in the Centra/blatt

für Bakteriologie (Bde. XXVII and XXVIII), gives not only the

most extensive, but also the fullest and most reliable, discussion of

present knowledge on this group, which has been almost neglected

until the appearance of recent studies on its structure and develop-

ment. The chapters of Lühe's work take up the life history (1) of

the Coccidia, (2) of the Hamosporidia (malarial parasites), (3) of

the Gregarinida, Myxosporidia, and the little-known groups of Micro-

sporidia, Sarcosporidia, and Haplosporidia. The first two sections

are particularly complete and satisfactory, and it is hard to find

omissions, while the discussion of the various authors cited is admi-

rably fair. The comparative table of terms used by different writers

in describing the development of Coccidia will prove very useful in

view of the entire lack of uniformity as to terms employed, — a defect

so universal that even successive publications of the same investigator

differ in terminology. Much would be gained by the adoption of a

uniform set of terms as advocated by Lühe, but unfortunately the

papers which have appeared since his have added to the confusion

by making further changes.

The third chapter is the least satisfactory, probably, since the field

covered by it is the least well known and is consequently most diffi-

cult to bring into relation with the other groups. Furthermore, the

introduction of numerous additions to the original articles, in the

form of lengthy footnotes and appendices, makes the treatise difficult

to use at some points; and yet the gain in accuracy compensates

for the slight lack of clearness.

In one point the work must be strongly criticised: the biblio-

graphic methods employed are antiquated and cumbersome to an

extent that interferes greatly with the clearness of the text. One

may well wonder how the author could have done so well with such

a confused system of reference, the same papers, 6g., Labbé, ** Sporo-

zoa,” being referred to in three different literature lists by as many

. different numbers. |

. $ Lühe, M. Ergebnisse der neueren Sporozoenforschung. Jena, 1900.

70 THE AMERICAN NATURALIST. [Vor. XXXVI.

Lühe is nevertheless to be congratulated on having produced the

first manual on the entire group, and in having made it a well-

balanced and useful treatise. H. B. W

Notes. — Under the name of Causeries Scientifigues, the Zoölogical

Society of France began last year the publication of a series of lec-

tures on various topics of scientific importance. Among the subjects

treated thus far are “ Vibratile Cilia,” by P. Vignon; the “ Tecti-

branch Mollusks,” by J. Guiart ; and “Coccidia and their Patho-

genic Rôle,” by R. Blanchard. While the method of treatment

shows all the advantages given by the freedom of a lecture and the

articles are very readable, they are not lacking in scientific accuracy.

Stiles and Hassall (Annual Report of the Bureau of Animal Indus-

try for 1899) have described carefully the mammalian lung fluke

recorded previously from dog and cat in the United States and

believed to be identical with the Asiatic lung fluke of man. The

parasite is reported as frequent in the lungs of hogs slaughtered in

Cincinnati, and the authors are inclined to regard it as identical

with the Asiatic form, and hence as a real menace to the health

of man in this country as it is in Asia.

Braun has added to his previous studies on the trematodes two

further articles. The first, on the trematodes of the chelonians

(Mittheil. Zool. Mus. Berlin, Bd. II, 1901), deals with twenty-two

species in detail, the large majority of which are new or poorly

known forms. The second paper, on the trematodes of mammals

(Zool. Jahrb., Syst., Bd. XIV, 1901), discusses thirteen forms, chiefly

also from the Berlin collection. Noteworthy is the demonstration of

the specific difference between Ofisthorchis tenuicollis Rud. and

O. felineus Riv., which have recently been thrown together by a

number of authors. The careful comparisons between related spe-

cies instituted by Braun in these studies furnish the only hope for

the clearing up of this much-confused territory.

Among the parasites from Lake Nyassa recently described by von |

Linstow (Jenaische Zeitschr., Bd. LIII, p. 408), Ziznia africana n. sp.,

from man, and Moniezia amphibia n. sp., from the | PIppopotamus, are

noteworthy.

The embryonic development of Anoplocephala has been worked

out in detail by Saint Remy (Arch. Parasitol., Vol. ITI, p. mica who

has observed for the first time in this group the formation of polar

corpuscles, —

pss i Sate enews

CIR WEE S q

E EE dps

DE c RE CS Lp AA Ir

No. 421.] NOTES AND LITERATURE. 71

The life cycle of the gregarines has been studied by Cuénot

(Arch. Biol, Vol. XVII, p. 581). He finds that the conjugation of

the sporoblasts is a regular feature of the encysted condition.

Shipley has given (Quart. Journ. Mier. Sci., Vol. XLIV, p. 281)

an account of the structure of Syndesmus echinorum, an imperfectly

known turbellarian which parasitizes in the sea urchin.

Loxosoma davenporti, which is described by Nickerson (Journ. of

Morph., Vol. XVII, p. 351), is the first representative of this genus

to be recorded from this continent.

Klunzinger has published (Stuttgart, 1901) a valuable discussion

on the physical, chemical, and biological causes of color in bodies

of fresh water.

Elrod has published (Zrans. Amer. Micr. Soc., Vol. XXII, 1901)

an extended account of the work done at the Montana Biological

Station, Flathead Lake, during 1899. The paper gives a full

description of the region, and lists of some collections made at

different points, with a few striking features in the distribution of

such forms as have been studied. The nine plates are good illus-

trations of the general appearance of the region.

Fordyce has worked up (Trans. Amer. Micr. Soc., Vol. XXII,

19o1) the Cladocera of Nebraska. The paper gives a valuable sum-

mary of our knowledge of the distribution of these forms in the

United States, and a discussion of the conditions in the plains

region, and then records important notes on the vertical distribution

of species in small lakes. In all, twenty-six species were found, of

which five — Macrothrix tenuicornis, Dunhevedia setiger, Bosmina

obtusirostris, Daphnia curvirostris, and Chydorus rugulosus — are

rare in the United States; Pleuroxus truncatus has not yet been

recorded elsewhere from this country; and three — Daphnia par-

vula, Bosmina ornamenta, and Leydigia fimbriata — are new to sci-

ence; descriptions and figures are given for these and the rarer

forms. S i

There has been received through the courtesy of Dr. Mitsukuri a

copy of a work entitled * Classified Catalogue of the Specimens of

Vertebrates in the Collections of Kiyoiku Hakubutsukuan (Educa-

tional Museum)," published in Tokyo in 1881 by Motokichi Namiye,

| now curator in the Imperial University of Tokyo. The work is espe-

"cially interesting as the earliest contribution of the Japanese (known

to me) to the systematic zoólogy of their country. It gives a list of

72 THE AMERICAN NATURALIST. [Vor. XXXVI.

1180 species of vertebrates, systematically arranged, with the vernac-

ular names Of each species. The identification is made from stand-

ard authorities and is generally correct, reflecting much credit on

„the patience and conscientiousness of its author, Mr. Namiye.

R. Lauterborn (Zool. Anzeiger, Bd. XXIV (1901), pp. 50-55) has

found that shallow fresh-water ponds covered with such floatihg plants

as duckweed, and with bottoms of black vegetable ooze often gener-

ating hydrogen sulphide, contain a very uniform and characteristic

life. The flora is mostly sulphur bacteria, the almost complete

absence of desmids and diatoms being very striking. The fauna

contains, in addition to certain rhizopods and flagellates, no less than

eighteen exclusively characteristic species of Infusoria. The author

proposes to call this well-defined fauna and flora sapropelic, because

of its association with decaying mud.

BOTANY.

‘A New Work on Gymnosperms.' — This is a very welcome addi-

tion to the list of books dealing with the special morphology of the

higher plants.

The last five years have been notable for the number of impor-

tant contributions to our knowledge of the Gymnosperms, and these

recent discoveries have very materially changed the older views as

to the affinities of some of them, notably the Ginkgoales. The book

before us summarizes, very successfully, the results of these investi-

gations, and should be very much appreciated by all botanical

students.

While the book is largely a compilation, it includes some original

MUS especially in the illustrations, many of which are vessel

The book deals principally with the reproductive parts, which are

treated very much in detail While the writer agrees with the

authors that an elaborate discussion of the vegetative organs was,

impracticable in a book of this character, still the value of the book

would have been much enhanced bya proper treatment of the more

1 Coulter, J. M., and Chamberlain, C.J. Morphology of nate mci New

oo TAE pd 8vo, prit. NA.

No. 421.] NOTES AND LITERATURE. 73

important anatomical characters of the vegetative organs of the prin-

cipal groups. This is especially the case in regard to the apical

meristems. A single borrowed figure (Fig. 48) is hardly sufficient

to make clear the characters of the apical meristems of the whole

group.

Four coordinate series of living Gymnosperms are recognized, vis. :

Cycadales, Ginkgoales, Coniferales, Gnetales. These are considered

successively, the details of the gametophyte and embryo being treated

very completely. On the whole, the treatment of the Cycads and

Ginkgoales is most satisfactory, although including less original

matter than the chapter on the Conifers. The important work of

Webber, Lang, Ikeno, and Hirase are given full attention, and the

chapters are fully illustrated by numerous well selected and well

executed figures.

The chapter on the Conifers is to a great extent a study of Pinus

laricio, presumably the work of the junior author. This is very com-

plete and admirably executed. It is to be regretted, however, that

the author did not include some of the numerous important Ameri-

can types as well as the highly specialized genus Pinus. The work

hitherto done upon the genera Taxodium, Sequoia, Torreya, Thuja,

etc., is very incomplete, and the inclusion of these in the book would

have been a real contribution to the literature of the subject. As it

is, the chapter on the Conifers must be considered much less satis-

factory than those upon the other orders.

The later chapters treat very satisfactorily the fossil Gymno-

sperms, comparative morphology of Gymnosperms, phylogeny and

geographical distribution. A valuable feature in the chapter upon

the fossil forms is a series of photographs illustrating some of the

Mesozoic Cycads of the United States. 3

The conclusions as to the origin and affinities of the Gymnosperms

are essentially the same as those given by Professor D. H. Scott in

his recent work on fossil plants, £.e., all of the existing Gymnosperms

are referable to a common ancestral stock, of filicineous origin.

This conclusion, we surmise, will not be admitted, without reserva-

tion, by all botanists.

The illustrations are, for the most part, extremely good. Some of

the half-tone figures, however, such as the sections of leaves shown

in Figs. 5 and 47, are too blurred to have much value, and might

well be replaced by good line drawings.

An admirable feature of the book is the carefully prepared bibliog-

. raphies at the end of the chapters.

74 THE AMERICAN NATURALIST. [Vor. XXXVI.

The book has a distinct place, and ought to be very welcome,

both as a work of reference and as a stimulus to further work in

this important group of plants, which are so well represented in our

own flora. D. HC.

MacDougal’s Plant Physiology.'— Those who have had occa-

sion to feel the lack of a clear, concise, and up-to-date text in plant

physiology will welcome the appearance of any promising book on

this subject. Dr. D. T. MacDougal, of the New York Botanical

Garden, is the author of a work of this character.

Dr. MacDougal points out in his preface the increased recognition

of irritability in its various manifestations, due chiefly to the work of

Pfeffer, and makes this the cardinal point in his arrangement of the

subject-matter. The first seven chapters are devoted to a considera-

tion of this subject in its different phases. The study of these

functions and properties, or the organism and its interaction with

environmental conditions, is thus the first task of the student. The

composition of the plant body, exchange and movements of liquids,

metabolism, growth and reproduction, are taken up in the order

named. To some this order of consideration may seem open to

question on the ground that the student is plunged immediately into

the thick of the most obscure of plant activities, where he can find

little by way of physical or chemical foundation to lend him

support.

In general, the treatment of the different phases of the subject of

irritability is very satisfactory. One notes with pleasure that recent

work has been incorporated. A sense of being in touch with the

movement of the science is worth not a little as a stimulus to the

student. This prompt acceptance, however, in an elementary text,

of work that has not been thoroughly verified, unfortunately has its

possible disadvantages. Occasionally, without criticising a given

piece of work, one hesitates to accept the author's statement of

results without a grain of salt. Explanations, even though plausible,

may rightly fail to win unreserved adherence until more evidence is

at hand. The work of Nemec on transmission of stimuli, Loew's

explanation of the action of various classes of poisons, and (in

‘another part of the book) Nathansohn's amitotic nuclear division

under the circumstances reported, illustrate the principle just enun-

ciated. In some cases, through oversight, minor errors have crept

Tiea D. T. Practical Text-Book of Plant dom s London,

— GM CR ee Sage 352 PP. >-

PAESE e cr.

UM y rE E EEE TS

Wu :

Lunge ENS SB ip qae LL og Ay Mon en

Bie ee ee

[gere ee! ETUR cs en

IRR ROS ice kee

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No. 421.] NOTES AND LITERATURE. 75

in. The strong mineral acids in normal solutions are by no means

completely dissociated in all cases (p. 53). A “normal solution ”

as defined in botanical literature should conform to the chemical

usage (p. 51). Unfortunately, in a number of instances botanists

have failed to be thus exact in the use of terms, and confusion is

likely to follow. These matters will doubtless correct themselves in

later editions of the book under discussion.

The chapter dealing with the composition of the plant body, writ-

ten by Mr. J. E. Kirkwood and Dr. W. J. Gies, introduces the

beginner to the methods of detecting and estimating many of the

more important substances that may be extracted from plants. This

chapter will probably prove very useful, since it furnishes in clear,

brief terms much information to which the student will be glad to

refer often. The chapters dealing with the chemical changes taking

place in the plant, with growth and with reproduction, call forth

much by way of commendation.

In view of the fact that the book is intended to contain “ details

of experimental methods suitable for the exact analyses requisite

in research work,” it demands notice as a laboratory guide. The

experiments are closely interwoven throughout the book with the

discussion of the various phases of the subject which they illustrate.

Generally speaking, the experiments are abundant and well chosen,

and the directions for work are couched in clear terms. In the man-

ner of arranging this matter, one might perhaps see some grounds

for question. The juxtaposition of experiment to principle illustrated

aids a possible shrewd guess on the part of the student occasionally,

but the author has generally avoided undesirable suggestion in

describing the experiments. Of course, this intermingling of discus-

sion and experiment makes the book more difficult of use as a work

of reference.

As a whole the illustrations are good, whether original or bor-

rowed. ‘An occasional exception may be noted (Fig. 134).

An appendix contains numerical data of use to the laboratory

student. Tables for converting units of various kinds into those of

other systems, densities of gases, expansion of air at different tem-

peratures, and an abridgment of Pfeffer’s table of osmotic values

may be mentioned. :

The book has so many excellencies that it is entitled to a wide

use, and the occasional errors will be remedied as later editions call

the author's attention more critically to his text. RET

76 THE AMERICAN NATURALIST. [Vor. XXXVI.

Ganong’s Plant Physiology.’

importance of plant physiology as a source of illuminating informa-

tion on the conduct of living things, as well as a valuable means of

discipline, the very scanty assortment of English texts available for

classes taking up this study has been a source of inconvenience to

many teachers. Laboratory manuals and concise, up-to-date treatises

on the subject have both been conspicuously lacking.

Fortunately, several additions to our texts on this subject have

recently been made; among these 4 Laboratory Course in Plant

Physiology, by Prof. W. F. Ganong of Smith College (New York,

Henry Holt & Co., 1901), outlines a series of experiments covering

a school year in which eight hours are given weekly for laboratory

work. Dr. Ganong has chosen experiments for their teaching value,

making use of those, so far as possible, which require simple appara-

tus, purchasable at small expense. Inspection of the figures and

descriptions given discovers much ingenuity in this selection. Since

Dr. Ganong believes that in elementary courses “it is mainly quali-

tative results that are of value,” although “the exact quantitative

methods and spirit are scientifically and educationally the best,” the

correct use of simple apparatus is emphasized. Comment upon the

experiments is chiefly by the way of suggesting to the student lines

of thought and reading. When experiment is out of the question

proportion is secured by references for outside reading. Much use

of synoptical essays is recommended.

The suggestions: are, in most cases, stimulating and within the

range of the possible. When, however, the student is directed to

form a clear mental picture of the molecular processes and energy

involved in osmosis, one cannot refrain from asking whether the

picture is to be that in the mind of Graham or that in the mind of

van’t Hoff. When physicists are still divided on the subject the

undergraduate deceives himself who fancies that he has the data

necessary for the formation of a clear mental picture.

The book is clearly the result of careful work and ample experi-

ence. It cannot but be of the greatest assistance to the teachers of

plant physiology, and it deserves to be most cordially received.

Rou. T.

Notes. — Part I of a botany of the Faeróes, based upon Danish

Investiganeayy has been published, by aid of the Carlsberg fund, by

x Ganong; W.F. A Laboratory Course in Plant Physiology, cedi as a Basis

7 Ecology. New York, eame Holt & Co., 19o1. w vi-146 pp., 35 fi

No. 421.] NOTES AND LITERATURE. 77

the Nordiske Forlag of Copenhagen. Dr. Warning contributes the

historical introduction, and systematic and ecological discussions are

given of all but the marine algz, which, with a comparison of land

and sea vegetation and certain economic discussions, is reserved for

a concluding part.

The Congo Museum of Brussels has commenced the publication,

as a series of its Annales, of a systematic enumeration of the Congo

plants collected in 1895-96 by Dewévre. The descriptions are by

De Wildeman and Durand.

Some botanical matter of interest to students of our northern

vegetation is contained in No. 21 of North American Fauna, referring

to the natural history of the Queen Charlotte Islands and the Cook

Inlet region.

The willows of Alaska are discussed by Coville in the Proceedings

of the Washington Academy of Sciences, under date of August 23.

Part VI of Dr. Rydberg’s “ Studies on the Rocky Mountain Flora,”

in the September Bulletin of the Torrey Botanical Club, contains a

considerable number of new species, chiefly gamopetalous.

A second part of Barber's “ Flora der Oberlausitz ” is contained in

Vol. XXIII of the Abhandlungen der Naturforschenden Gesellschaft zu

Görlitz.

Professor Arechavaleta’s flora of Uruguay, in course of publication

in the Anales del Museo Nacional de Montevideo, has reached the

group Mimosez.

A number of papers on the oe flora of Java, by

Koorders, are contained in Vol. LX of the Natuurkundig Tijdschrift

voor Nederlandsch- Indie.

The holly-leaved barberries, constituting the genus Mahonia, are

revised by Fedde in the opening number of Vol. XXXI of Engler's

Botanische Jahrbücher, which also contains a monograph of the orchid

group Disez, by Schlechter.

The signatures of Vol. IV of Pittonia, issued September 30, con-

tain descriptions of a number of violets and crucifers, by Professor

Greene.

The origin of Liriodendron stipules is discussed by E. W. Berry in

the Buletin of the Torrey Botanical Club for September. _

Hybrid currants are discussed by Janczewski in the July Bulletin

International de D Académie des Sciences de Cracovie.

78 THE AMERICAN NATURALIST. [VoL. XXXVI.

A monograph of the genus Sorbus, by Hedlund, is reprinted from

Vol. XXXV of the A. Svenska Vetenskaps-Akademiens Handlingar.

The chemistry of the bark of Rodinia pseudacacia, by Power, and

the anatomy of the bark of the same species, by Perrédés, form the

subject of Nos. 20 and 21 of the Publications of the Wellcome Chemical

Research Laboratories of London. A paper by von Schrenk, on

the decay of the wood of the same tree, induced by Polyporus

rimosus, has been separately printed from the Zwe//tà Report of the

Missouri Botanical Club.

Dr. Heyl, of Darmstadt, has distributed a paper, “Ueber das

Vorkommen von Alkaloiden und Saponinen in Cacteen,” from the

June Heft of the Archiv der Pharmazie. Pilocereus sargentianus,

Cereus pecten-aboriginum, and C. gummosus were studied.

Professor Hume publishes an interesting account of Citrus decu-

mana as Bulletin 58 of the Florida Experiment Station.

Aligera patelliformis and Collinsia breviflora, from California, are

described in Zhe West American Scientist for August, by Suksdorf.

Agave langlasset is the name applied by André, in the Revue

Horticole for August, to a species of Manfreda from the Pacific slope

of Mexico.

Professor Nelson publishes an economic treatise on the species of

Bromus occurring in Wyoming, in Buletin No. 46 of the Wyoming

Experiment Station.

In a paper published as No. 3 of the current volume of Proceedings

of the California Academy of Sciences Professor Peirce discusses the

curious etiolated suckers which are sometimes produced by Seguoia

sempervirens,

Separates of a paper on hybrid conifers, by Dr. Masters, have

been distributed from the Journal of the Royal Horticultural Society.

Dr. E. F. Smith publishes an extended account of the cultural

characters of four species of Pseudomonas of economic importance,

in Bulletin No. 28 of the Division of Vegetable Ph ystology and

Pathology of the United States. Department of Agriculture.

In a paper reprinted from the Proceedings of the Indiana Academy

of Science for 1900, Professor Arthur reaches the conclusion that the

l "eedanapple " fungi, usually known by the generic name Gymno-

Paes

ROI ee ue

unc

gium, should really be called Tremella, and he consequently

No. 421.] NOTES AND LITERATURE. 79

renames them under this genus, — without, however, considering the

nomenclature of the many fungi usually treated as constituting the

latter.

A revision of North American puffballs of the group Tylosto-

macez, by V. S. White, appears in the Buletin of the Torrey pet

Club for August.

A revision of the genus Tilletia, by Massee, is contained in the

Kew Bulletin of Miscellaneous Information for 1899.

Professor Bailey contributes to Ze World’s Work for September

an illustrated account of Luther Burbank’s work as a plant breeder.

Current numbers of the Revue Générale de Botanique contain a

treatise by Jumelle on the rubber plants of northeastern Madagascar.

Raphia bast, much used by florists, is considered at length in a

paper by Sadebeck, reprinted from Vol. XVIII of the Jahrbuch der

Hamburgischen Wissenschaftlichen Anstalten.

An illustrated paper on the anatomy of the cocoanut, by Winton,

is published in the October number of The American Journal of

Science.

The weeds of Montana are treated by Professor Blankinship in

Bulletin No. 30 of the Experiment Station of that state.

Dr. Willis, director of the gardens, has begun the publication of

an irregular-period journal under the title Annals of the Royal

Botanic Gardens, Peradeniya. he first number, issued in June last,

contains a brief history of the gardens and an account of the facilities

offered for research.

Volume VIII of the Proceedings of the Towa Academy of Sciences

contains the following botanical articles : Graw, “ Preliminary List

of the Flowering Plants of Adair County” ; Fitzpatrick, “ Juglanda-

cee, Betulacew, and Fagacez of Iowa”; Mueller, ** Shrubs and:

Trees of Madison County”; Pammel, “Thistles of Iowa, with

notes on a few other species"; Faurot, “Notes on the Early

Development of Astragalus caryocarpus.”

The initial numbers of the Bulletin du Jardin Impérial Botanique.

de St.Pétersbourg are largely concerned with fungi and lichens. The

Russian text is accompanied by abstracts in French.

An account of the Geneva meeting of the newly organized Associ-

ation I t js printed in the current volume :

of the Bulletin de r eder Boissier. |

80 THE AMERICAN NATURALIST.

The World’s Work for September contains an illustrated account

of the Arnold Arboretum, by Sylvester Baxter.

A little handbook of nature-teaching based upon the general

principles of agriculture, by Francis Watts, has been prepared at

the wish of Commissioner Morris, of the Imperial Department of

Agriculture for the British West Indies, and is printed by Messrs.

Dulau & Co., of London.

Some phases of the conflicting interests of people who are trying

to teach city pupils about nature, and those who are trying to pre-

serve natural objects, are well presented in an article by Mrs. Britton

in Zorreya for August.

A suggestive essay entitled * How shall a Young Person study

Botany?" by Professor Beal, is reprinted from the Proceedings of

last winter's conference of the New York State Science Teachers'

Association.

Another of the interesting and well illustrated memoirs on plant

ecology that the Botanical Gazette is bringing out is by Professor

Bray, and deals with western Texas.

Phytogeographic nomenclature, discussed by F lahault in the July

Bulletin of the Torrey Botanical Club, was the subject of an extended

paper by Clements at the recent Denver meetings.

À paper on some changes effected in plants by frost is published

by Lapeyrére in the current volume of the Buletin de la Société de

Borda, of Dax.

An essay on old herbaria, by Matouschek, is printed in Vol.

XXXII of the Mittheilungen aus dem Vereine der Naturfreunde in

Reichenberg for 1901.

PUBLICATIONS RECEIVED.

(Regular exchanges are not included.)

BACHMETZERD, P. Experimentelle entomologische Studien vom physikalisch-

chemischen Standpunkt aus. Erster Band. Temperaturverhiltnisse bei Mini

Leipzig, Engelmann, 1901. Svo, ix, 160 pp., 7 figs. 4 Marks. — BENH . B.

: Treatise on Zoólogy. Edited by E. Ray Lankester. Pt. iv, The Phybeliia.

zoa, and Nemertini. London, Adam and Charles Black, 1901. 8vo, vi, 204 pp.

nnd $5.25. — Brirron, N. L. Manual of the Flora of the Northern

States and Canada. New York, Henry Holt & Co., 19or. 8vo, x, 1080 pp.

BürscHLI O. Mechanismus und Vitalismus. Leipzig, Engelmann, 19or. biu

107 pp. 1.60 Marks. — CHESTER, F. D. A Manual of Determinative Bacteriology.

New York, The Macmillan Company, 1901. 8vo, vi, 401 pp- 13 figs. — DNIESCH, H.

Die organischen Regulationen. Vorbereitungen zu einer Theorie des Lebens.

ipzig, Engelmann, 1901. Svo, xvi, 228 pp., 1 fig. 3.40 Marks. — HOERNES, M.

Primitive Man. Translated from the German by J. H. Loewe. Temple Primer

Series. London, J. M. Dent & Co., 1901. 8vo, 135 pp. 48 figs. — Kipp, W

Use-Inheritance illustrated by the Direction of Hair on the Bodies of Animals.

London, Adam and Charles Black, 1901. 8vo,47 pp., 16figs. 9o cents. — KOLBE, H.

Gartenfeinde und Gartenfreunde. Die für den Gartenbau schädlichen und nütz-

li en. Gartenbau-Bibliothek, Bd. xxxiv-xxxvi. Berlin, Karl Siegis-

mund, Igor. $vo,ii,318 pp, 76 figs. 3. 60 Marks. — PENFIELD, S. L., and PIRSSON,

L. V. (Editors). Contributions to Mineralogy and Petrography from the Labora-

tories of the Sheffield Scientific School of Yale University. Yale Bicentennial Pub-

lications. New York, Charles Scribner's Sons, 1901. 8vo, xii, 482 pp. Illustrated.

— SHIPLEY, A. E., and MacBripg, E. W. Zoólogy, an Elementary Text-

New York, The Macias Company, 1901. 8vo, xxi, 632 pp. 349 figs.

Banxs, N. Some Spiders and Other Arachnida from Porto Rico. Proc. U.S.

Nat. Mus. Vol. xxiv, pp. 217-227, Pl. XV. — BEAL, F. E. L. Some Common

Birds in their Relation to Agriculture. U.S. Dept. Agr., Farmers’ Bull. No. 54.

40 pp., 22 = ind e J.E. The Anomenan Collections made by the Fish-

hawk n to Porto Rico. Bull. U.S. Fish Com. for 1900. Vol. ii, pp. 129-

148, Pls. EVI — BicELow, M. A. The Cirripedia collected near Porto Rico

by the Fishhawk Expedition. Bul. U.S. Fish Com. for 1900. Vol. ii, pp. 177-

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THE

AMERICAN NATURALIST

Vor. XXXVI. February, 1902. No. 422.

A NEW AGRICULTURAL ANT FROM TEXAS, WITH

REMARKS ON THE KNOWN NORTH-

AMERICAN SPECIES?

WILLIAM MORTON WHEELER.

THE genus Pogonomyrmex, comprising the true “ agricul-

tural ants," is one of several formicid genera peculiar to the

American fauna. It comprises more than a dozen species

which range from Montana to Argentina, often over wide

areas, though apparently absent from considerable portions of

this vast region. As the species are mostly large and conspic-

uous and inhabit exposed situations, they have attracted more

attention than many of our American ants. Notwithstanding

this fact, however, we are still very far from possessing an

adequate knowledge of the habits and —— relationships

of the various members of the genus. :

The species described in the following pages seems to have

escaped attention hitherto on account of its idiosyncrasies. It

is small and inconspicuous, of a timid disposition, and lives

under stones, instead of in exposed grassy regions like the

other North-American species. It is, moreover, rather rare

1 Contributions from the Zological Laboratory of the University of Texas, No. 24.

85 oo

86 THE AMERICAN NATURALIST. | [Vor. XXXVI.

and local. Upto the present time I have seen it in only one

locality, on the flat limestone terraces which form the southern

slope of Mt. Barker, a short distance from the Colorado River,

near Austin, Texas. Though but a few acres in extent and on

warm days fully exposed to the rays of a pitiless sun, these

terraces are, nevertheless, a rich collecting ground for the

myrmecologist. All about the place there is something of the

local color of the dry Mexican plateau, and this peculiarity

extends also to the ant-life of the region. Here, under the

flat, detached pieces of lime-

stone scattered among a

sparse but interesting vege-

tation,! occur at least four

species of grain-storing

ants: the new Pogonomyr-

mex described below, a

golden yellow variety of the

ubiquitous subtropical and

tropical “fire ant” (Sode-

nopsis geminata Fab.), and

, two species of Pheidole (a

diminutive new form and

Ph. kingii André, var. insta-

ilis Emery). This is also

one of the few localities in

which I have seen the little

mushroom-growing ant

Cyphomyrmex wheeleri Forel, the first of its genus to be taken

in the United States.? Here, too, occur Odontomachus clarus

Roger, Pheidole hyatti Emery, X; iphomyrmex spinosus Pergande,

and, of course, Dorymyrmex pyramicus Rog., Forelius fatidus

Buckley, and Camponotus fumidus Rog., var. festinatus Buckley.

While many of these species abound in this locality, I have

failed to find more than a dozen nests of the new Pogono-

myrmex, and these were so close together — within an area of

.1 A brief account of the flora of this region is given by Oberwetter (86).

: we oasis discovered a dark variety of C. rimosus Spinola at New Braun-

No. 422.] NEW AGRICULTURAL ANT FROM TEXAS. 87

a few square rods — as to suggest that they may have been

merely parts of a single colony. These nests were all under

rather small flat stones, which were often located by following

up the foraging workers as they trudged home slowly over the

hot soil in the intense glare of the sun. The nest is a simple

structure consisting of a few broad and very shallow surface

chambers (134-3 inches in diameter) connected by one or two

vertical or oblique galleries with a few chambers situated at

lower levels in the soil. The superficial chambers always con-

tained from about 34 to 24 of a teaspoonful of seeds, mostly,

but not exclusively, from the grasses of the neighborhood.

These seeds were all dry and unhusked, and hence of a very

different appearance from those found in neighboring nests of

Solenopsis geminata. This ant carefully shells its seeds and

treats them in some singular manner, so that they all have a

glistening yellow color like the ants themselves. Although I

collected the Pogonomyrmex at different times of the year and

excavated their entire nests, it was impossible to discover

either the queens or the males. Even the larvae and pupe,

found in great numbers in the chambers ot the nests June

1-10 were all of the worker type. The specific description

which follows is drawn therefore exclusively from the worker.

This, however, can scarcely be confounded with the workers of

any of the other North-American species of the genus.

Pogonomyrmex imberbiculus n. sp.

Worker: Length 4-4.8 mm. Color rich ferruginous red, legs somewhat

paler, eyes and edges of mandibles black; hairs covering the body yel-

lowish. Head quadrangular, scarcely longer than broad, its posterior

margin hardly incised. Mandibles sexdentate, the two apical teeth largest,

blades traversed nearly their entire length by coarse longitudinal ridges.

Clypeus subopaque, with longitudinal ruge separated by series of faint

stria and provided with long, anteriorly projecting hairs. Antennal scape

covered with faint longitudinal ridges. the hairs on its anterior surface sub-

erect, on the posterior surface more appressed. Dorsal and lateral surfaces

of head covered with coarse rugas, which are scarcely divergent behind and

connected with one another by irregular transverse ridges; the areola thus

enclosed are subglabrous, coarsely and confluently punctate. Hairs on the

upper and lateral surfaces of the head short, erect, subobtuse. Lower sur-

face of head more delicately longitudinally rugose, with somewhat longer

88 THE AMERICAN NATURALIST. (VoL. XXXVI.

and more tapering hairs, which, however, do not form a conspicuous beard

as in the other North-American species of Pogonomyrmex. Thoracic dor-

sum and pleurz covered with coarse reticulate rugze, enclosing more finely

reticulate rugose and confluently punctate, polygonal areole. In some

specimens the rugze have a transverse trend on the pronotum and a slightly

longitudinal trend on the meso- and metanotum ; promesonotal suture usu-

ally indistinct. Epinotum armed with two pairs of rather blunt spines,

scarcely longer than the breadth of their bases; anterior pair connected

with each other at the base by a transverse ridge and with the spines of the

posterior pair on either side by a longitudinal ridge; the space thus

enclosed is subglabrous and traversed by a few longitudinal ruge. Hairs

covering the thorax short,

subobtuse, and perfectly

erect. Stem of petiole

laterally compressed,

slender, provided below

near its insertion with a

small but distinct tooth ;

node scarcely longer than

the stem, its apex obtuse

in profile, its dorsal sur-

face subelliptical, covered

with coarse reticulate rugæ like those of the thorax, but bearing somewhat

longer and more pointed hairs. Postpetiole campanulate, subdepressed

dorsally, with a prominent rounded projection below near its base ; sculpture

decidedly fainter than that of the petiole and consisting of rather indistinct

rugz interspersed with punctate spaces. Gaster small, smooth, and shining

throughout, without basal stria and punctures, and covered with prominent,

suberect hairs. Legs glabrous, clothed with suberect hairs.

Fic. 2. — Pogonomyrmex imberbiculus n. sp. Worker.

ri iew.

While P. tmberbiculus is very sharply distinguished from any

of the other North-American species of Pogonomyrmex by its

small size, peculiar sculpture, and the lack of the beard of long

hairs which suggested the generic name to Mayr (68, p. 11), it

is, singularly enough, very closely related both in these and

other particulars to a Brazilian species, P. negelit Forel ('86

pp. 4, 5). Through the kindness of Professors Forel and

Emery, who have sent me specimens of the Brazilian form, I

have been able to compare the two species, which at first sight.

_ would almost certainly be confounded. More careful examina-

_ tion, however, reveals the following differences: In negelit the

~ gaster is of a distinctly darker color than the head and thorax,

= and its extreme basal portion is longitudinally striated and finely

No. 422.] MEW AGRICULTURAL ANT FROM TEXAS. 89

punctate. The head, thorax, and petiole are somewhat more

coarsely rugose than in zmberbiculus, and the epinotal spines

more acuminate at their tips. The most striking difference,

however, is in the sculpture of the postpetiole, which in xegelii

is but little finer than that of the petiole, whereas in the Texan

species this segment is nearly smooth.

Recently Forel (99, pp. 61, 62) has discovered in Columbia

still another beardless and otherwise aberrant Pogonomyrmex

(P. mayri), which he assigns to a new subgenus, Janetia. This

is based very largely on the predaceous, non-granivorous habits

of the species and on the neuration of the male fore wings,

which exhibit only a single cubital cell. He expresses doubt

as to whether 7.

negelit should be in-

cluded in his new sub-

genus, but leaves the

matter undecided, as

he supposes the male

of this species to be Fic. 3.— Pogonomyrmex negelii Forel. Worker.

unknown. This, how- ;

ever, appears to be an oversight, since Mayr (87, p.612) describes

both the male and female of P. nægelii. He clearly states that

the female has two cubital cells and that the wings of the male

are the same as those of the female. Now from the very close

affinity of P. zmberbiculus with P. negelit it is safe to predict

that the latter is also a grain-eating species. Weare compelled,

therefore, to regard the small group of Pogonomyrmex —

ing the beardless Texan and Brazilian species as transitional

between Pogonomyrmex sensu stricto and the subgenus J —

rather than as belonging to the latter. It may be advisable

ultimately to erect a special subgenus for the two small grain-

storing species, but a careful study of the males and females

of all the known species of the genus should be previously

undertaken. HE :

The workers from two nests of P. zmberbiculus, with their

numerous pupæ, nearly mature larvae, and their store of vp

were put together in the same artificial nest. : The ants from

different nests fraternized without the slightest signs of hostility,

90 THE AMERICAN NATURALIST. [VoL. XXXVI.

thereby indicating that they were perhaps members of the

same colony. They soon distributed their progeny and provi-

sions in three separate piles — one for the larvae, one for the

pupæ, and one for the seeds. During the first few days of

their captivity the ants were fed on house flies. These were

not only eaten with avidity by the adult Pogonomyrmex, but cut

into pieces and fed to the larvae in the same manner as I have

described for the Ponerinz and some Myrmicinz ('00 and '00a).

On one occasion nearly every larva in the nest could be seen

munching a small piece of house fly. But a still more remark-

able method of feeding was adopted after a few days, when the

supply of insect food was exhausted. Then the ants were seen

to bring seeds from their granary, crack them open with their

strong mandibles, and, after consuming some of the softer por-

tions themselves, to distribute the remainder among their larva.

The latter could be seen under the lens cutting away with their

mandibles and devouring the softer starchy portions of the seeds.

The hard and useless hulls were afterwards carried away by the

ants and placed on the refuse heap. These observations show

that the Jarve of certain ants are not only able to subsist on solid

food, but even on food of a vegetable nature. The adaptation of

what were probably once exclusively carnivorous ants to a

vegetable diet, although not yet complete, is, nevertheless, so far

advanced that the larva already participates in the peculiar feed-

ing habits of the adult insect. The P. zmberbiculus seem not

to possess the power of feeding one another or their larvae by

regurgitation. At any rate they were not seen to make use of

this method in the artificial nests.

These observations are quite in line with some which I

made on artificial nests of the large “agricultural ant of Texas ”

(P. barbatus Smith, var. molifaciens Buckley). In this case the

workers carried the seeds, a few at a time, into the chamber

containing the queen and her attendants. Here the ants,

including the queen, gnawed away the soft portions of the

seeds till they had satisfied their hunger. Thereupon the

empty hulls were carried out. Even when the nest was sup-

~ plied with honey or syrup, each ant helped herself from the

v food supply, and neither fed other ants nor ed herself

No. 422.] NEW AGRICULTURAL ANT FROM TEXAS. 91

to be fed by regurgitation. I deem it probable, therefore, that

the larvz of moltfactens are also fed like those of zmberbiculus

by what we may call the direct method, to distinguish it from

the indirect method adopted by the Camponotinæ. In the

Ponerinz and many Myrmicinz, including Pogonomyrmex, the

direct appears to be the prevailing, if not the exclusive, method.

In the Camponotinz, on the other hand, the indirect method

prevails, since at a given time only a comparatively small num-

ber of ants function as caterers for the whole colony and dis-

tribute the food by

regurgitation to the

larvae and the other

ants.

It may not be

altogether out of

place in this paper

to record a few other

observations on P. molifaciens, inasmuch as this form has

been singled out among all the known members of the genus

as presenting certain remarkable instincts. Lincecum is

responsible for the myth that this Pogonomyrmex sows a

certain species of grass, the “ant rice” (Aristida oligantha),

protects it from harm and frees it from weeds while it is grow-

ing, for the purpose of reaping the grain. This notion, which

even the Texan schoolboy has come to regard as a joke, has

been widely cited, largely because the great Darwin stood spon-

sor for its publication in the Journal of the Linnean Society

(62). McCook, after spending a few weeks in Texas observing

P. molifaciens and recording his observations in a book of 310

pages ('79), failed to obtain any evidence either for or against

the Lincecum myth. He merely succeeded in extending its

vogue by admitting its plausibility.

1 Not only have able myrmecologists like Forel (99, p. 63) pem deceived by

the accounts of Lincecum and McCook into assuming the existence of a kind of

symbiotic relation between the Pogonomyrmex and the “ant rice," but this —

now in its fortieth year, still flourishes in the newspapers. n M grows d

CH. pies of May 19, 1901: “ Many species of ants € pn

apparently cultivate many varieties of foodstuffs. The trimmer ants

Fic. 4. — Pogonomyrmex barbatus F. Smith (typical).

Worker.

92 THE AMERICAN NATURALIST. [Vor. XXXVI.

Two years of nearly continuous observation of P. molifactens

and its nests enable me to suggest the probable source of Lince-

cum's and McCook's misconceptions. In either case the

observer has started with a few facts and has then stopped

short to draw inferences before gathering more facts. If the

nests of moltfaciens be studied during the cool winter months,

— and this is the only time to study the nests leisurely and

comfortably, since the cold subdues the fiery stings of their

inhabitants, — the seeds which the ants have garnered in many

of their chambers will

often be found to have

sprouted.! On sunny

days the ants may

often be seen remov-

ing these seeds when

they have sprouted

too far to be fit for

food and carrying

them to the refuse heap, which is always at the periphery of the

cleared earthen disk or mound. In this place the seeds thus

cast away as inedible often take root and somewhat later form

an arc of tall grass more or less closely approximating a com-

plete circle around the nest. Since the Pogonomyrmex feeds

. largely, though by no means exclusively, on grass seeds, and

since, moreover, the seeds of the Aristida are a very common

and favorite article of food, it is easy to see how this grass

should often predominate in the circle. In reality, however,

Fia. 5. — Pogonomyrmex occidentalis E.T. Cresson. Worker

harvesting ants of Texas are both of this kind. The trimmers prune a sort of

weed which is to their taste so that it shall grow strong and sturdy, and the har-

s ants go even further than this. They clear disks several yards across

around about their nests of all manner of vegetation. Then they plant these

farms with ant rice, which they watch d tend until it ripens, keeping the crop

carefully free of weeds and insects. The ants’ dogs keep the ant cows out of the

growing grain, and the farmer ants probably sit around themselves at night with

shotguns to shoot colored ants suspected of pilfering."

1 The same is true of the seed stores of Pheidole kingii, var. instabilis. It is

. : this presen seems to be of no avail.

No. 422.] NEW. AGRICULTURAL ANT FROM TEXAS. 93

only a small percentage of the Pogonomyrmex nests, and only

those situated in certain localities, present such circles. Now

to state that the mo/ifaciens, like a provident farmer, sows this

cereal and guards and weeds it for the sake of garnering its

grain is as absurd as to say that the family cook is planting

and maintaining an orchard when some of the peach stones

which she has carelessly thrown into the back yard with the

other kitchen refuse chance to grow into peach trees.!

There are several other facts which show that the special

ring of grass about the mo/ifaciens nest is an unintentional

and inconstant by-product of the activities of the ant colony.

First, the Aristida often grows in flourishing patches far from

the nests of molifactens. Second, one often finds very flourish-

ing ant colonies that have existed for years in the midst of

much-traveled roads or in stone sidewalks often a hundred

or more feet from any vegetation whatsoever. In these cases

the ants simply resort for their supply of seeds to the nearest

field or lawn, or pilfer the oat bin of the nearest stable. Third,

it is very evident that even a complete circle of grass like those

described by Lincecum and McCook would be entirely inade-

quate to supply more than a very small fraction of the grain

necessary for the support of a flourishing colony of these ants.

Hence, they are always obliged to make long trips into the

surrounding vegetation, and thereby wear out regular paths

which radiate in different directions, often to a distance of

forty to sixty feet from the entrance of the nest. These

paths in the case of the Mexican agricultural ant (P. barbatus

sens. str.) remind one of human footpaths, as they may be as

much as four to six inches wide in places. The existence of these

paths, which are often found in connection with grass-encircled

nests, is alone sufficient to disprove Lincecum's statements.

McCook's conceptions of the external architecture of the

molifaciens nest are hopelessly confused, notwithstanding the

1 Lincecum was fond of attributing agricultural and horticultural propensities

to ants. Thus he states ('67, pp. 28, 29) that the leaf-cutting ant pen v giis

plants trees and vines on its nest! At the same time of course, like McCook, he

failed to observe the marvelous mushroom-gardening habits of these ants, —

another instance in which truth is stranger than fiction.

94 THE AMERICAN NATURALIST. [Vor. XXXVI.

fact that he seems to have been much interested in ant archi-

tecture, and has devoted no less than thirty-five pages to a

presentation of this feature. It does not seem to have occurred

to him that the character of the architecture of molifactens

must be profoundly affected by two factors, — the nature of

the soil and the age of the ant colony. Gravel-cone nests can,

of course, be built only in soil that abounds in small pebbles,

whereas nests dug in a uniform soft, loamy soil, like that of

northern Texas along the Red River, must be simple disks or

very low mound nests, as the soil brought up by the ants is

spread out by the rains and the movements of the ants them-

selves. On the other hand, a small, incipient colony of ants

is unable to clear away much of the vegetation about the

entrance to the nest. At least the tougher plants, like the

grasses, whose hard siliceous stems offer considerable resistance

to the mandibles of the ants, cannot be cut away till the colony

waxes strong both in the size and number of its individuals.

Then the work proceeds rapidly, the circular area coéxtensive

with the subterranean galleries is completely cleared and

opened up to the sun's light and warmth. This clearing is

evidently an adaptation for insuring the greatest possible dry-

ness in the granaries

of the nest. The

circular denuded disk

or mound enlarges

slowly year after year,

and it should be noted

that during this pro-

Fic. 6. — Pogonomyrmex subdentatus Mayr. Worker. S ressive enlarg ement

even the peripheral

circle of ant rice is quite as ruthlessly cut down and cleared

away along its inner edge as any other plants that may cast a

shadow on the disk, and thereby enable the soil to retain the

moisture.

While we possess the observations of Buckley (61), Lince-

cum ('62, '66, '74), and McCook (79), on the habits of the Texan

P. molifaciens, of Mrs. Mary Treat (77) and McCook (79) on

the * Florida harvester” (P. badius Latr.), and of McCook (82)

No. 422.] NEW AGRICULTURAL ANT FROM TEXAS. 95

on the “occident ant" (P. occidentalis Cresson), no observa-

tions seem to have been published on the two distinctively

Californian species (P. californicus Buckley and P. subdentatus

Mayr) My friend Dr. Harold Heath, who has made strenuous

effort to fill this gap in our knowledge, kindly supplies me with

the following notes on the latter species :

The red agricultural ant (P. suédentatus) is one of the most abundant

ants in the neighborhood of Pacific Grove, Cal. Here one is constantly

coming upon them and their nests along the roadsides and in the sandy

soil of the woods and fields. The nests, so far as I have been able to

observe, are never placed under stones or logs, but in exposed regions, —

that is, away from the shadow of vegetation. Little attempt is made to

clear away the short grass in their vicinity. The earth carried out from

their burrows is usually deposited several inches from the opening, espe-

cially along their runways, which extend out in various directions into the

surrounding region. Large quantities of chaff and the hulls of seeds are

also scattered about, usually in fairly definite dumping grounds, but neither

these materials nor the earth are ever fashioned into a mound. Some of

the ants entering the nest carry pods, others bits of leaves and grass, all

well dried, while an equal number of the insects leaving the nest carry away

similar materials, but the pods are emptied of their seeds and the leaves are

evidently thrown away as non-nutritious and useless. Within the nest there

are several little granaries, or accumulations of seeds, each sometimes

amounting to as much as a teaspoonful, though usually considerably less.

The foodstuffs seem to be carried to one spot within the nest and there

hulled and assorted. The seeds are then carried to the storehouses, while

the chaff is at once carried out, although it may accumulate and almost

completely fill a burrow for a distance of several inches. On comparing

the seeds taken from the nest with those of the surrounding plants, I find

them to be chiefly those of a species of grass and of two species of Com-

posite. At the present writing these seeds are fully ripe, but as soon as

those of other plants mature they appear to be equally acceptable. : I may

add that these ants defend their homes with extraordinary pugnacity and

inflict stings more painful than those of the honey-bee. As I write I feel

the dull ache of several stings inflicted more than a day ago.

These observations, by a thoroughly competent zoólogist,

show that at least one of the Californian species of Pogono-

myrmex conforms rather closely to what is known of the other

species of the genus.

Some interesting

problems center about the geographical

distribution of the species of Pogonomyrmex.

These ants

96 THE AMERICAN NATURALIST: [VoL. XXXVI.

evidently represent an extreme adaptation to the open, dry and

sunny, and more or less grass-covered regions of the New

World. Such regions are, perhaps, most typically represented

by the deserts of Wyoming, the plateau of central and northern

Mexico, and the pampas of La Plata. The area occupied by

the genus and extending, as above stated, from Montana to

Argentina, presents in North America an eastern offshoot to

Florida (P. badius)

and a peculiar west-

ern offshoot to the

Sandwich Islands

(P. occidentalis). In

South America there

is between Colombia

and Argentina a con-

siderable area from

which species of

Pogonomyrmex are unknown, but the continuity of the distri-

bution, though broken at this point, is at least in part preserved

further to the east by the Brazilian P. negelii.

This distribution over two continents naturally suggests an

inquiry as to whether the species arose in North America and

migrated thence along the Andes into South America, or had

their origin in the pampas of Argentina and migrated into

North America over the same lofty road. Two authors,

v. Ihering (94, p. 416) and Emery (94, p. 354), who have seri-

ously studied the interesting problems suggested by the dis-

tribution of the American ants, agree in regarding North

America as the primeval home of the species of Pogonomyr-

mex. Concerning this genus and the genera Dorymyrmex

and Forelius, which have a very similar distribution, Emery

says:

Fic. 7. — Pogonomyrmex californicus Buckley. Worker.

Their migration probably proceeded along the Andes at a time when

the climate was cooler and the vegetation therefore different from the

present. Later, on the supervention of new floral conditions, they were

crowded out of a portion of their former domain by the tropical ant fauna.

For the reason that the southern species of Pogonomyrmex and Dorymyr-

mex are more numerous than the northern, we might, perhaps, assume that

these animals had migrated from the south to the north. But it is not in

No.422.] NEW AGRICULTURAL ANT FROM TEXAS. 97

the least improbable that these ants, like the South-American species of

Didelphys, deer, camelids, and mastodons, are of North-American origin.

Without being able to adduce stringent proof in favor of my opinion, I

nevertheless incline to accept this latter view.

The migration between the continents is supposed to have

taken place during the Pliocene. This view of the North-

American origin of Pogonomyrmex is supported to some extent

by the flourishing condition of the closely allied holarctic

genera Myrmica and Stenamma (including the subgenera

Aphzenogaster and Messor) in the United States and Canada.

A problem of more subordinate interest is suggested by the

close morphological relationship of the Brazilian P. negelii

and the Texan P. imberbiculus, without known forms of a simi-

lar aberrant character in the intervening geographical region.

It is possible, how-

ever, that a more

searching investiga-

tion of the Mexican

and West-Indian

fauna may bring to

light still other beard-

less forms of Pogono-

myrmex and thereby fill this gap. It should be mentioned,

nevertheless, that the new Texan species has all the appearance

of being a geological “ relict.”

In conclusion I subjoin a dichotomic table to aid in the

identification of the workers of the North-American species of

Pogonomyrmex : |

badius Latreille. Worker.

Fic. 8. — P E Y

I. Small species, less than 5 mm. long ; under surface of head without a

beard of long cürved hairs; epinotum armed with four spines; head,

thorax, and petiole coarsely reticulate rugose, base of gaster not

striated. Formicary under stones. (Central Texas.)

P. imberbiculus n. sp.

2. Larger species, more than 5 mm. long; under surface of head with a

beard of long, curved hairs; epinotum with only two spines or none;

head and thorax finely rugose, the ruge being more or less parallel

with one another, not reticulate. Formicary not under stones,

exposed... ea oA ee LE NISL T 3

3. Epinotum with a single pair of spines .

* - *

ec] $

98 THE AMERICAN NATURALIST. [Vor. XXXVI.

4. Epinotum unarmed . . ORA P

5. Head finely and densely rugose, ruga but little divergent posteriorly,

without or with very indistinct interrugal sculpture . . 7 a—e

6. Head less densely rugose, rugæ very distinctly divergent debtor,

interrugal sculpture distinct, consisting of dense foveolate punctures 8

7 a. Head, thorax, and legs black; d postpetiole, and gaster red.

(Mexico.) . . P. barbatus F. Smith ('58, p. 130)

. Cephalic ruge der "id denser, dus ferruginous red throughout.

(Mex, Test, Ind; Ter, Ark,

rs harhat, var. pee Ae Buckley ('61, p. 445)

7c. Head and thorax brownish red, gaster in part or entirely brown.

Rugosity as in 7 è or somewhat stronger. (Tex., Col.)

P. barbatus, var. fuscatus Emery (94, p. 309)

7d. Rugosity a little coarser than in 7 æ; head, thorax, and legs black,

petiole and postpetiole brown, abdomen red, node of petiole longitu-

dinally rugose. (Marfa, Tex.) . P. barbatus, var. marfensis n. var.

. Head and thorax much more coarsely rugose than in 7 a-d. Ruge

irregular in direction on the pro- and mesonotum, on the other regions

transverse. Petiole rather strongly and irregularly rugose; its ante-

rior stem-like portion shorter than in P. barbatus ; postpetiole rugose-

punctate. (Cal) . P. barbatus, subsp. rugosus Emery ('94, p. 309)

8. Head in. iront rugose, the rugæ distinctly divergent posteriorly,

interrugal spaces densely fov: eolate punctate 9

9. Lower surface of petiole without a distinct tooth ; intraspinal TERE

of epinotum rugose, scarcely shining. . Io a-ó

104. Head opaque, interrugal punctures distiuct. (Col, New Mex.,

Utah, Ariz, Nev., Wyo., Mont, Kans., Neb., Honolulu.)

P. occidentalis Cresson ('65, pp. 426, 427)

10 6. Head more shining, interrugal punctures more indistinct; petiole less

opaque than in toa. (S. Cal.

P. occidentalis, vat. subnitidus Emery ('94, p. 310)

11. Petiole with a distinct tooth below; infraspinal concavity of epinotum

shining, without ruga. (Cal) . P. subdentatus Mayr. (O, p. 971)

12. Interrugal spaces of head rather indistinctly and oe punctate.

Workers monomorphic . . . inåt

13 a. Color yellowish red, stem of RS about ‘Ge same birth as its

nodal portion ; postpetiole as high as long. (Cal, Lower Cal.)

. californicus Buckley ('66, p. 236)

13 À. sow red than 13 æ; apical third or more of gaster more or less

black; petiole and postpetiole often brown, the former slender, its

digit and less erect, with rounder or but slightly pointed apex.

ow Cal.)

“I

P. californicus, var. apens Pergande (93, p. 33)

"ne Dee red, gaster brown except at the base; stem of petiole

—— key very long nodal portion, which oe above ;

No. 422.] NEW AGRICULTURAL ANT FROM TEXAS. 99

postpetiole not as high as long. Sculpture fainter than in 1 $ es

petiole and postpetiole punctate, without ruga. (Cal.

P. californicus, subsp. /onginodis Emery ('94, p. 311)

14. Interrugal spaces regularly foveolate punctate. Color ferruginous

red. Workers Eee ié., with size of head greatly varying.

(N.C., Ga, Fla.) Co . - P. badius Latreille ('02, p. 238)

BIBLIOGRAPHY.

'61 BuckLEY, S. B. Myrmica (Atta) molifaciens, “ Stinging Ant," or

* Mound-making Ant,” of Texas. Proc. Acad. Nat. Sci. Phila.

(1860), 1861. Pp. 445-44

'66-67 Descriptions of New aan of North-American Formicida.

Proc. Ent. Soc. Phila. Vol. vi (1866-67), pp. 152-172, 335-350.

'65 CnEssoN, E. T. Catalogue of Hymenoptera in the Collection of the

Entomological Society of Philadelphia from Colorado Territory.

Proc. Ent. Soc. Phila. (1865). Pp. 242-313, 42

'93—94 Emery, CARLO. Beiträge zur Kenntniss der nordamerikanischen

. Ameisenfauna. Zool. Jahrb. Abth. f. System. Bd. vii (1893),

pp. 633-682, Taf. XXII; Bd. viii (1894), pp. 257-360, Taf. VIII.

'86 FOREL, AUGUSTE. Espèces Nouvelles de Fourmis Américaines.

Compt. Rend. Soc. Ent. de Belg. (6 fev., 1886), pp. 1-12.

Formicide. Biologia Centrali-Americana, Hymenoptera. Vol.

iii (1899), pp. 1-169. Pls. I-IV.

'94 IHERING, H. von. Die Ameisen von Rio Grande do Sul. Berl. Ent.

Zeitschr. Bd. xxxix, Heft 3 (1894), pp. 321-446. 1 plate and

text-figures.

02 LATREILLE, P. A. Histoire Naturelle des Fourmis. 1802.

'62 Lincecum, GIDEON. Notice on the Habits of the “ Agricultural

Ant" of Texas [“ Stinging Ant" or * Mound-making Ant,” Myr-

mica malefaciens Buckley]. Communicated by Charles Darwin.

Journ. Proc. Linn. Soc., Zoology. Vol. vi (1862). pp. 29-31.

'66 — — On the Agricultural Ant of Texas. Proc. Acad. Nat. Sci. Phila.

Vol. xviii (1866), pp. 323-331-

‘67 The Cutting Ant of Texas, (Ecodoma Texana Buckley. Proc.

Acad. Nat. Sci. Phila. (1867). Pp. 24-31-

74 The Agricultural Ant. Am. Nat. Vol. viii, No. 9 (Sept., 1874),

» 613—517

"79 et Sed n. C. The Natural History of the Agricultural Ant of

| Texas. Author's edition. Acad. Nat. Sci. Phila. (1879).

'82 —— The Honey Ants of the Garden of the Gods and the Occident

Ants of the American Plains. Philadelphia, Lippincott & Co., 1882.

100 THE AMERICAN NATURALIST.

'00a

Mayr, Gustav. Formicidae Nove Americana Collectz a Prof. P. de

Strobel. Annuario della Soc. d. Naturalisti. Ann. III (Modena,

1868), p. 1-20.

Neue Formiciden. Verh. k. k. zool. bot. Ges. 1870.

Siidamerikanische Formiciden. Verh. k. k. zool. bot. Ges.

Bd. xxxvii (1887), pp. 511-632.

MocGRIDGE, J. T. Harvesting Ants and Trapdoor Spiders. London,

L. Reeve & Co., 1873.

OBERWETTER, P.H. The Wild Plants of Texas. Gardiner’s Monthly

and Horticulturist. Vol. xxviii, Feb., 1886.

PERGANDE, THEO. Ona Collection of Formicidae from Lower Califor-

nia and Sonora, Mexico. Proc. Cal. Acad. Sci. (Ser. 2). Vol. iv

(1893), pp. 26-36.

SMITH, FREDERICK. Catalogue of Hymenopterous Insects in the

Collection of the British Museum. Pt. vi, Formicidae. London,

1858.

TREAT, Mary. Chapters on Ants, in Harper’s Half Hour Series.

1877.

WHEELER, W. M. A Study of Some Texan Ponerine. Biol. Bull.

Vol. ii, No. 1 (Oct., 1900), pp. 1-31, figs. 1-10.

The Habits of Ponera and Stigmatomma. Biol. Buli. Vol. ii,

No. 2 (Nov., 1900), pp. 43-69, figs. 1-8.

CoLEBROOK, CONN., September 9, 1901.

PHYLLOSPADIX AS A BEACH-BUILDER.

RALPH ERWIN GIBBS.

In December, 1898, while overhauling a heap of seaweeds

recently collected by Prof. W. A. Setchell and myself at

Bodega Bay, California, I discovered, clinging to a branch

of Amphiroa, a small brown object which bore, at first sight,

a rude resemblance to a beetle's head with rigid, bristle-fringed

antenne. On examination, however, it was evident that this

nondescript, clasping the Amphiroa in such a way that the

reflexed bristles, or “barbs,” of its arms resisted attempts

to detach it, was the seed or fruit of some flowering plant.

Further, it was obvious that this contrivance of arms and

bristles was a unique and most interesting example of dis-

semination mechanism.

As there are extensive beds of eelgrass (Phyllospadix)

growing in shallow water along the rocky shores of Bodega

Bay, the possibility suggested itself that this was the ripe

fruit of Phyllospadix, and upon comparison with the figures

of Ruprecht and others it seemed that such was the case, —

although our find was, in some respects, very different from

what Ruprecht supposed to be the “ripe fruit.” ?

A few weeks later, at Monterey Bay, the matter was put

beyond doubt, and given new complexity and interest, when

we collected not only an abundance of the fruits, but also

young plants in several stages of development. Following up

these discoveries, and with the help of Professor Setchell and

Dr. W. L. Jepson, I made an investigation, the results of

which are here set forth. These results, to anticipate a sum-

mary, are, first, some data as to the life history of Phyllospadix,

and, second, some speculation as to the significance of the

plant from a geological point of view.

1 Thesis for the degree of M.S., University of California, May, 1900. _

Stillen Oceans.

es F. J Neue... Pflanzen aus dem... 1852.

lor

102 THE AMERICAN NATURALIST. [Vor. XXXVI.

First we consider the fruit itself. Upon comparison of

Figs. 2 and 3 it is seen that our wave-beaten specimen

from Bodega, with its long stiff arms, lined with brushes of

inflexed bristles, is a decided departure from Fig. 2, which

represents the most mature

fruit I have been able to

find still attached to the

spadix. Fig. 2 is also an

approximate reproduction of

. Ruprecht’s figure. Fig. 1

shows a younger stage.

The feature which catches

the attention in Fig. 3 is, of

course, the arrangements of the two arms, with the bristles

or barbs. This is what I have called the ‘dissemination

mechanism"; but that is rather a misnomer, for it is a device,

not for scattering the seeds, but for anchoring them after ied

have been drifted away from the parent plant.

But the first question is, What is the origin of these OM

and how are they developed from the object we see in Fig. 2?

Curiously enough, the key to this puzzle was unwittingly

stumbled upon by Ruprecht in 1852. He noticed upon the

fin-like expansion which extends between

the body and the arms of the green fruit

the parallel darker streaks marked f in

Fig. 2; for he remarks that, upon dissec-

tion, he found imbedded in the softer,

semitranslucent tissue, bundles of braune

Fasern, and it is these brown fibers which,

as we shall see, play an important part in

one chapter of Phyllospadix life history. / Z

In Fig. 4 is shown a diagrammatic sec- “ ia

tion of the stage of fruit shown in Fig. 2, F'S- 3-— Fruit as found attached

cut in the plane of the two arms. The m

pericarp is here so differentiated that we may distinguish, for

convenience, two parts, — what we may call the exocarp, the

| Soft, spongy outer portion; and the endocarp. The latter is

- eue or absent at the lower end of the seed, but it | Caeps

FiG. 1.

Fic. r.— Early stage

of fruit.

Fic. 2.— Later stage

of same.

No. 422.] PHYLLOSPADIX AS A BEACH-BUILDER. 103

the body of it as in a sling of tough, compact tissue, com-

posed of elongated lignified cells. Further, this endocarp,

prolonged downward, forms the axes of the arms; and, finally,

it produces a great number of very long and thick-walled

cells, which lie loosely imbedded in the softer substance of

the “fin” (f). These “fibers” may be easily separated from

one another, and from the enclosing exocarp, except at their

lower ends, where they are conjoined with the endocarp of the

arms. These slender lignified cells are the Fasern of Ruprecht,

and now it is to be inquired how i

they are metamorphosed into the a

barbs of Fig. 3.

2. -tesla

= ps "i -endocarp

When the ripe fruit is ready to [ uw"

break loose from the spadix, its

arms have grown longer and Dex

: ; . grammatic

stiffer, as in Fig. 3, but are still, section of

of course, enclosed in the exo-

carp. Now the fruit begins

to drift about, — flung against the rocks, washed up on the

beach, and sucked back again, bruised and scoured, in the

swirling sand and pebbles of the undertow; and after a little

of this rough-and-tumble existence the spongy exocarp begins

to wear away, leaving exposed, in the arms and about the upper

part of the seed, the hard endocarp, while about the base (7.¢.,

the lower middle part of Fig. 3) there is laid bare the smooth

surface of the shell-like testa. But the important point is that

the exocarp in which the barbs are imbedded is also got rid of.

It easily flakes off and is washed away, and then, with a little

more sand-scouring, the freed barbs, springing apart a little,

stand out clean, like so many whalebones.

Now our “anchoring mechanism” is uncovered and ready for

use. By this time, too, the seed may be ready to pe

and, for the safe putting forth of leaves, a fixed abiding place

is necessary. The seed must be anchored, and, the e

being ready, the next requisite is a suitable anchoring groun

‘Along our coast there are various species of coralline hend

weeds (Corallina, Amphiroa) which abound wherever t

are rocks between tidelines. These alge have slender,

104 THE AMERICAN NATURALIST. [VoL. XXXVI.

lime-impregnated stems, made flexible by being broken into

short joints, and constricted between the joints like a string of

coral beads ; and they form an intricate, often turf-like, growth

over the wave-washed rock.

We could not imagine a better opportunity for Phyllospadix

in search of a lodging place; and it seems that Phyllospadix is,

in fact, not slow to catch on, for experience showed that the more

profitable way to collect Phyllospadix seed was to search at low

tide, not in the eelgrass itself, but rather among the corallines

covering some flat rock just inshore from the eelgrass beds.

When the fruit, hurried landward by a wave, blunders

against a tip of Amphiroa, there is a fair chance that the

stem will slip into the crotch of one of the arms. When

once this has happened the wanderings of Phyllospadix are

over, The many barbs unite to hold whatever is caught, and

further knocking about by the waves only serves to wedge the

alga more tightly into the grasp of the seed. The segmented

structure of the Amphiroa prevents it from slipping through

the grip of the arm, and so the seed is safely planted, as it

were, in a tree-top. Frequently it even catches two branches

of its host, one in either armpit, and so

swings secure on two anchors.

Time and seed are now ripe for ger-

mination. What next takes place will

be better understood upon referring to

Fig. 5. Here is a longitudinal, dorsi-

Fic. s- — Diagram of dorsiventral ventral, median section of the immature

median section of Fig.2. c,coty- 2

edon; i, insertion of pistil; fruit. The cavity of the seed is filled

tie by the embryo, af which the bulk cor-

sists in the hypocotyl (%, ^). The coty-

ledon (c) is short, straight, and tubular, and serves merely as a

sheath for the plumule (^). It lies upon the ventral side —

as regards the pistil — of the hypocotyl, and points toward the

base, t.e., in the direction of the arms. The testa (7) is formed

of a single layer of cells with thick, lignified cell walls, and

thus, while it serves, like an arch of bricks, to protect the

embryo from outside pressure, it is readily burst open by the

-~ swelling within of the hypocotyl. —

No. 422.] PHYLLOSPADIX AS A BEACH-BUILDER 105

Perhaps the exposure, and consequent partial drying of the

seed, at low tide helps to crack the testa. At any rate, when

germination begins the basal end (2) of the hypocotyl bursts

through the testa and presses the broken edge of it back, leav-

ing a clear passage for the cotyledon. The

hypocotyl grows no farther than this, and

eventually, when its store of substance has

served to give the young plant a start, it

dies away.

The cotyledon grows out little beyond

the hypocotyl, and is quickly outstripped by

the ensheathed plumule, which turns upward

and shortly unfolds to the waves several

grass-like leaves (Fig. 6). Henceforth it is

these leaves only which grow upward ; for,

except for the slender peduncle which bears

the inflorescence, the only stem of the

mature Phyllospadix is a creeping root-

stock, or rhizome.

Our seedling (Fig. 6) has now quite a start in life, and yet

it has no root. Its leaves, meanwhile, whipped back and forth

by the waves, are gradually beaten to shreds, and replaced by

others. In this way several of the first leaves may

have lived and died before any root appears. At last,

when roots are put forth, there are two of them pro-

duced, not from the hypocotyl, as we might expect, but

from the stem above the cotyledon (7, Fig. 7).

That is to say, these are, in the technical

sense, adventitious roots. When full grown

. the roots are about two centimeters long,

stout, and unbranched. . This first pair hang

Fic. 7. — Young seedling down, one on either side of the stem, and

with coats removed. c, in- d :

sertion and of are soon followed by several more which

Fic, 6. — Seedling with first

leaves.

cotyledon ; Z, second node i

with remnant of leaf; », grow out from the second internode.

oe When the roots are several millimeters

long they begin to produce, near their tips, a dense, woolly

covering of root-hairs, and when a root comes in contact with

any object, — be it the stem of the Amphiroa (Fig. 8), the rock

106 THE AMERICAN NATURALIST. . [Vor. XXXVI.

beneath, or another root, — these rhizoidal hairs spread, as a

closely adhering film of whitish fibers, over the surface touched,

binding the root to it. The root-hairs assume fantastic shapes

in order to conform with the irregularities of the rock. If a

root has, by chance, entered the loose sand, it becomes enclosed

in a compact cylinder of sand, bound

together by the myriad of branch-

ing, interwoven hairs.

The plant has now taken a firm

hold on life. Supposing that the

start has been made aloft in the

branches of the Amphiroa, the elon-

gating stem now dips downward till

it strikes the rock. Thenceforth it

creeps along, taking, as it goes, a

firm grip upon every inch. At each

node it bears a leaf, and each of the

short internodes produces, on one

side a supra-axilary bud, and on

the other what Professor Dudley!

has aptly termed an *'epaulette "

of six or eight roots; alternating,

so that, if one internode has its

roots on the right side, the next

will have roots on the left. Thus

the stem, though in itself weak and

brittle, keeps a close, broad grasp

upon the rock, while the wiry

leaves, buoyed by intercellular air-

Spaces, stream upward sometimes

Fic. 8.—Seedling (older) with rou, COT @ length of two meters. Before

these leaves have been whipped to

tatters by the waves, their place is supplied with new ones

from the lateral buds.

When the rhizome has reached a length of, often, only one

. or two decimeters, it begins to die away behind. At the same

time the lateral buds begin to push out for themselves, so that

n ! Cf. Dudley, R. W., in Zee, vol. iv, p. 381.

No. 422.] PHYLLOSPADIX AS A BEACH-BUILDER 107

as the parent stem creeps on it leaves on either side of its trail

a series of new rhizomes, starting out at right angles to the old

one. In like manner this new generation of rhizomes branch

and rebranch, growing over and upon one another, till, in a few

years, the bowlders, as well as the bed-rock, are covered bya

patch of eelgrass, — a thick mattress, which, as it lies in sinu-

ous tangles at low tide, quite dissipates the force of a hammer,

though swung with all the energy of an enthusiastic botanist,

and so must, one would think, considerably lessen the effect of

the forces which are ordinarily at work reducing the bowlders

to pebbles and grinding down the rock itself.

This brings us to the second phase of our subject, namely,

the significance of such a plant as Phyllospadix to the geolo-

gist. What we wish to show is that under some conditions

the effects of the plant in modifying the results of wave and

current action are worthy of consideration.

Various geological authorities have commented upon the

protection afforded to shore rocks by some marine plants, and

upon the accumulation of detritus by others. Certainly no

plant is better adapted to either of these functions than Phyllo-

spadix. In the first place its manner of growth is exceptional,

resulting, as it does, in the formation of a broad, continuous

patch, instead of a scattering of individuals. Again, unlike the

soft or rubbery fronds of the algz, its leaves are strengthened

by extremely tough bands of collenchyma fibers. :

A moment ago we noticed how the eelgrass bed, the growth

of which from a seedling we have sketched, covers the bowlders

with a thick mattress which must prevent attrition between

the loose stones and pebbles. At Bodega Bay may be seen

such a bed, where, at low tide, one may stumble for rods along

a bowlder-strewn shore without seeing the rock under foot, all

being overlaid by the tangled mat of eelgrass. Now, it might

be objected that the corrosive action of the acids excreted by

root-hairs would offset the protection the plant might afford

to the rock. But it is at once obvious that the root-hairs of

Phyllospadix are, in this respect, unlike those of most plants.

Their essential function is that of attachment, not patrition,

and, of course, if they should cut the gr ound from under their

108 THE AMERICAN NATURALIST. [VoL. XXXVI.

own feet, so to speak, by dissolving away the rock to which

they ought to hold fast, they would be useless.

Suppose, now, that a wave-cut terrace, which is being

widened by the inroads of the sea, and which is swept by a

littoral current bearing the shore-drift along with it, becomes

overspread by a growth of coralline algz. Geike, in his Tert-

Book of Geology, speaks of the preservation of shore rocks by

the overgrowth of these corallines (“calcareous nullipores "').

These plants cover the substratum with a brittle, calcareous

crust, which, though a considerable protection against the

cutting of water-borne sand, is shattered by the blow of a

pebble. This calcareous enameling retards, to some extent,

the lowering of the terrace by the sand-bearing current. That

is, while the terrace widens, the water above it may remain

comparatively shallow. Moreover, many of these corallines, as

Amphiroa and Corallina, produce, in addition to the calcareous

crust, numerous erect, jointed fronds ; and the latter, as we have

seen, offer the best possible lodging place for Phyllospadix. Here,

then, we have the conditions most favorable to Phyllospadix, —

shallow, but not quiet, water, and corallines to anchor the seeds.

Gradually, if there is eelgrass within drifting distance, the

terrace becomes dotted with green tufts, the tufts spread

into patches, and ultimately the higher border of the terrace

presents, at low tide, the appearance of a wind-laid hay-field,

the “ grass,” one to two meters long, lying in prostrate tangles.

Now begins the accumulation of debris. Stones and pebbles

being carried alongshore by the current, odds and ends of

seaweeds, and all the multifarious small drift of the shore,

are caught in the network of rhizomes and wiry leaves. The

larger stones may themselves serve as footholds upon which

the rhizomes climb higher and wave their leaves higher in the

water. Every stone entangled serves to stop more pebbles

and sand, and, as the mass continues to pile up, the rhizomes

are at last buried deep under it; but as long as the tips of

the leaves wave free the plant thrives. On almost any of our

beaches there may be found between tide-lines tufts of slender

leaves apparently growing in the sand, but in reality anchored

to the rock a meter, perhaps, below.

No. 422.] PHYLLOSPADIX AS A BEACH-BUILDER IO9

The result of the accumulations, then, is that the water is

made shallower, so that, though the pounding of the heavy

seas upon the shore is lessened, the waves still race in over

the shallows and carry up the smaller particles to deposit them

on the beach. At the same time, even though the littoral

drift be not held permanently by the eelgrass, yet the time

required for it to pass the place, and hence its chance of

contributing to the beach, is increased.

Shoaler and shoaler grows the water, the shore line advan-

cing as a low beach, and finally, — the littoral current being

deflected seaward and the wave deposition continuing, — the

terrace that was is overlaid by a sand-flat.

That the long, hemp-like fibers of eelgrass lend coherence to

the mass of sand and stones in which they are imbedded is

attested by the fact that where, as above, we find a clump

of eelgrass half-buried in the sand, the level of sand within the

clump is often several inches above that of the surrounding

beach.

In closing, and to recapitulate, it seems probable that the

spreading over a terrace of such a plant as Phyllospadix must

tend, first, to protect the rocks from erosion and attrition ;

second, to help, by trapping the shore-drift, to raise the terrace

so as to form a beach, or a sublittoral sand-flat ; and last, by

binding together its materials, to render the foundation of the

beach, once formed, more coherent and stable.

UNIVERSITY OF CALIFORNIA, BERKELEY, (CAL,

A QUANTITATIVE STUDY OF VARIATION IN

THE BRACTS, RAYS, AND DISK FLORETS OF

ASTER SHORTII HOOK., A. NOVZE-ANGLLE L,

A. PUNICEUS L, AND A. PRENANTHOIDES

MUHL., FROM YELLOW SPRINGS, OHIO.

GEORGE HARRISON SHULL.

CONTENTS. PAGE

I. Introduction III

IL Matora ios wc 112

III. Methods and Precautions 114

IV. Results 115

Aster shortii Hook. I15

Aster nove-anglie L. 118

Aster puniceus L. IIQ

Aster prenanthoides Muhl

Result of Successive Collections

(1) Bracts

(2) Rays tie ta

(3) Disk Florets eee a i

(4) Correlations 145

V. Discussion of Results 146

VI. Summary 15!

Bibliography 552

I, INTRODUCTION.

THE work upon which this paper is based was done in the

Biological Laboratory of Antioch College, during the academic

year 1900—1901, under the direction of Prof. W. L. Tower.

My primary object was to study the variations of several

species of Aster by means of the statistical methods, to deter-

mine whether the results of Ludwig (95, '96, '98) upon Chrysan-

themum leucanthemum L. were also true for other nearly allied

forms, and to find out, if possible, how much correlation there

1 Contributions from the Biological Laboratory of Antioch College, No. 5.

III

Ti THE AMERICAN NATURALIST. [VOL XXXVI.

is between the observed variation and the environment, or

between parts of the same plant. Secondarily, it was hoped

that this research might show something of the applicability

of these methods to taxonomic or monographic work in a genus

so difficult from the systematic standpoint as Aster.

II. MATERIAL.

This consisted of the blooming capitula of Aster shortzi

Hook., A. nove-anglie 1., A. puniceus L., and A. prenanthoides

Muhl.

The capitula were cut off without any conscious selection,

folded in papers, labeled, and preserved in alcohol. With the

exception of those plants where series of pickings were made,

the stems from which the capitula had been gathered were

preserved for future reference.

All of the species of Aster studied are perennial, but they

differ in the manner in which the annual stems are produced. In

the following description of the material used in this study I

shall use the term “ individual" in its broader sense, including

all stems which have been derived from a single seed.

Aster shortii Hook. has from one to three or four slender

annual stems arising from a small perennial root. The material

which forms the basis for the study of this species was obtained

Sept. 26, 1900, and consisted of 226 capitula from three isolated

individuals and from a group of ten stems growing near to each

other. The stems of this group were probably mostly distinct

individuals, though they may have been of close genetic rela-

tionship. These plants grew in the thin limestone soil at the

foot of the Niagara limestone cliffs bordering the northern end

of Sheldon's Glen, one-half mile southeast of Yellow Springs,

Ohio.

Aster nove-anglie L. has a heavy mass of perennial roots,

and from the base of the annual stems of one year's growth

may arise a considerable number of heavy stems of the next

year's growth, forming a clump. The 199 capitula used in the

study of this species were collected Sept. 30, 1900, from five

individuals growing in the flood plain of a tributary to the Glen

No: 422.] QUANTITATIVE STUDY OF VARIATION [13

Stream, one-half mile east of Yellow Springs, and from four

individuals similarly located in the valley of the Glen Stream

about forty rods above the point at which it empties into the

Little Miami River, one mile southeast of Yellow Springs,

Ohio. |

The manner of production of annual stems in Aster puniceus L.

resembles closely that of Aster nove-anglie L. The material

of Aster puniceus L. used in this study was collected Sept. 25,

1900. It consisted of 798 capitula from thirteen stems arising

from three perennial roots growing at the margin of a bog five

miles west of Yellow Springs, Ohio. These three individuals

were identieally located, being separated by a space of but a

few yards. Environmental conditions had, therefore, no known

influence in oe the differences in the heads from the

three clumps.

Aster prenanthoides Muhl. differs from all of the other species

studied, in the manner in which its annual stems are produced.

It sends out slender rootstocks, which give rise to new stems

at a little distance from the old ones, thus forming patches

with the stems growing singly.

Eighty-three capitula of Aster prenanthoides Muhl. were col-

lected Sept. 27, 1900, from seven stems, apparently belonging

to two individuals. The remaining material of this species,

making a total of 658 capitula, was all collected from a single

small plot. Four successive collections were made on Sep-

tember 27, September 30, October 4, and October 8, 1900.

All the capitula which were blooming at the time of each col-

lection were taken, amounting respectively to 117, 143, 139,

and 176 heads, and comprising all the heads produced by the

selected plot during the season. The object in collecting in

this way was to test, at least within narrow limits, the con-

stancy of the garri « constants " throughout the flowering

season.

The plants from which all the material of 4. prenanthoides

Muhl. was collected grew at the bottom of a small ravine near

Clifton, Ohio, at the margin of a permanent stream, so that

heat, light, soil, and moisture conditions were nearly constant

throughout the growing period.

114 THE AMERICAN NATURALIST. [VoL. XXXVI.

III. METHODS AND PRECAUTIONS.

In order that the personal equation should modify results as

little as possible, all of the heads of each individual were col-

lected, and none were discarded on the ground of abmodality.

The only material rejected was such as was eaten by insects,

or had been blighted to such extent as to make a correct count

impossible. It is evident that by the discarding of healthy

material because of its great departure from the usual condition,

or, as Strong (1901, p. 295) says, the choice of ‘individuals

which appear on inspection to be typical the statistical

method may be made to give any result for which the investi-

gator may be looking.

There is one element of error in the choice of material which

must be mentioned. In those cases in which it seemed desir-

able to get curves and * constants," representing the conditions

in the capitula of single individuals, there was necessarily a

choice of those individuals which had the largest number of

blooming heads. In this respect, therefore, the determinations

would represent the conditions in the more robust specimens

rather than in the general population. To counteract this

tendency, there was an occasional collection of material from

a sufficiently large group of smaller individuals to give equally

valid results. As these collections from a number of smaller

individuals did not show a marked difference from those of

single larger individuals, this selection has probably not greatly

modified the results.

In making counts the liability of error is not so great as in

the taking of measurements. However, even in counting there

are sources of error, which I found it necessary to eliminate as .

far as possible. To free my work as far as possible from these

sources of error, the capitula were carefully dissected and the

parts kept separate until the count was completed. When

there was the least doubt as to the correctness of the result

they were recounted. Also in cases of great abmodality the

. results were verified by a recount.

In calculating the various constants I have used the formula

given by Davenport (99). All the mathematical processes

No. 422.] QUANTITATIVE STUDY OF VARIATION 115

were carried out to the tenth decimal place, and wherever

practicable they were checked. It is probable, therefore, that

all calculations are correct to the fifth decimal place.

In plotting the variations I have used the “method of rec-

tangles" as best representing the actual conditions found.

In those cases in which material was insufficient compared

with the range of variation, I have doubled the classes, but

in no case have I grouped more than two classes together.

In this way the essential features of the curves have remained

unchanged, while the lesser irregularities have been eliminated.

It must be remarked, however, in regard to the grouping of

classes, that the method should be used with much caution.

By combining a sufficient number of classes every multimodal

curve may be made monomodal.

It is also essential, in doubling, that a definite plan be

adopted in order that the results may be comparable. The

writer followed the plan of throwing together the two classes

nearest the mean. As the mean is a constant, this makes the

resulting double classes of the various curves strictly compar-

able. If I had begun at the lower limit of range instead of at

the mean, the count of one more variate might have lowered

the range by one, and thus have changed the combinations,

with the result that the character of the curve might be much

modified.

IV. RESULTS.

Aster shortii Hook. — Fig. 1 represents the frequency poly-

gon of the bracts. The range, 28 to 49, was so great that the

226 counts were insufficient, and it was deemed best to double

the classes. In this way the curve becomes monomodal, with

the mode on 36-37, and the mean on 36.800884. The coeffi-

cient of variability was 10.727157.

The frequency curve of the rays (Fig. 2) shows a remarkably

strong mode on 13, 38 per cent of the variates falling into that

class, with the mean on 14. This strong mode on 13 suggests

at once Ludwig's (95,'96,'98) interesting results on Chrysan-

themum leucanthemum L., etc.; but, as nowhere else in the

Asters studied has there been any apparent tendency of modes

116 THE AMERICAN NATURALIST. [Vor. XXXVI.

B 5

Lp SR

-e ——

E a summ UR 80

SN RS a ot BE

a A Bi =

AE E A 3 70

= 6) 65

uo e i)

55 »

59 50

30 wT b:

b 40)

30 30

25 25

15| 15

0 10

5 5

E 50

3l 5l E 5 »

Fic. x. Fic: 2. Fic. 3.

Fic. 1. — Aster shortit Hook. Bract 6h Cl loubled. Mean = 36.800384 +

7121; ierat: 36-37; 09 — 3. ceo +

Fic. 2. S shortii Hook. Ray curve Pa ar bedda Mean = 14 + .068448; mode = 13;

= 1.525592 + .048400.

Fic. m Aster storti Hook. Disk f 226 head

= 1.513025 + .o48001.

Mean = 22,053097 + .067885 ; mode = 22 ;

No. 422.] QUANTITATIVE STUDY OF VARIATION 117

to fall into the series 8, 13, 21, 34, etc., this condition in Aster

sortii Hook. cannot be considered as having any special sig-

LIJIIIPIIITIIIFITITITITLIE

1-8 77-6 2,4 3 2-1 i| 2

28/29/30 |31 |32]33 |34|35| 36|37| 38] 39/40/47 [42/43 [t4 | 5] 6147 148 [49

eae T Pit Ya) tte Pare ey gd

LI 12 Lii213|8|3[u| 3| 1 LH 1:1 Tr

ee AR AF 18]10| 8/9 ES

ERRA 14 1j (24 5]12|89|5161/0|3]2| 11 |

PTT. ae 1|[112/[58111215|316] I RILIS) |2

ion 16 31) 2 affi

TA S I7 nn fit | 3

LE] dH gla eA EI OE III T

aama BEEOTHENO

- Ra H H ae

HAT Pia be

Fic. 4. — Aster shortii Hook. Correlation surface for 226 heads. Rays subject and bracts relative.

= .549555 + .025157

EISIIBDILLILIA

X4—3|-4 -1| I 3 y

19|20|21|22[2324|25|26 [2728]?

| [28 fil 1|

L [^ Hs 117/6] 71

D 13 3/8, 2 | 1

L| po fw 1318/5

| | B [es] 12141216] 5/2]5)2

|] B [i6 44 1121213) 1

| | Bl 21 1213/2/1

ga 18 112 i

GERE ]

20 AES

LI 9 M |

Fig. 5. — Aster aded Hook. Correlation surface for 226 heads. Rays subject

d disk florets relative. p= .44664: 9861

nificance in this connection. The coefficient of variability for

the rays was found to be 10.897091, or a little greater than

that of the bracts.

118 THE AMERICAN NATURALIST. [Vor. XXXVI.

The curve of the disk florets (Fig. 3) in this species shows a

remarkable lack of variation in parts which, owing to their

indeterminate character, might be expected to be the most

variable. The mode is on 22, the mean on 22.053097, and the

coefficient of variability is only 6.860830. It should be noted

that in all these curves there is negative skewness,! though it

is slight in the bracts and disk florets. The variability con-

stants for Aster shortii Hook. are shown in the following

table.

TABLE A.— CONSTANTS OF ASTER SHORTII Hook.

Bracts. Rays. Disk FLORETS.

Noc auc 226 226 226

Meu inco oS 36.800884 14 22.053097

Moden c uve ue us 36-37 13 22

MD serit 3.029211 1.176991 1.115200

K aS E ry 3.947688 1.525592 1.513025

FEA Dea) 5... + 2.662716 + 1.029012 + 1.020535

FEM 3:7 + 77121 + .068448 + .067885

PRO. | x .125243 + .048400 | + .o48oor

e. v. 10.727157 10.897091 6.860830

In Fig. 4 is represented a “correlation surface” with rays

subject and bracts relative. The coefficient of correlation was

found ‘to be .549555 (P. E. p= .025157). The correlation

between rays and disk florets is shown in Fig. 5, in which the

coefficient of correlation is -446645 (P. E. p = + .029861).

Aster nove-angliz L. — In this species the range of varia-

tion in all the parts was great, the least range, 30 to 60,

occurring in the bracts. "

All the frequency polygons, Figs. 6, 7, and 8, show a multi-

modal condition, but this may be due to too scanty material.

Although no dependence can be put upon the multimodal

condition of these curves, the “varia

perhaps not differ widely from results w

* 1 The degree of Skewness has not been computed for any o£ the polygons of

distribution because the range of material was too small to make this index of |

any value. - | ura

No.422.]] QUANTITATIVE STUDY OF VARIATION I19

by counting larger quantities of the same material These

appear in the following table.

TABLE B.— CONSTANTS OF ASTER NOV/E-ANGLLE L.!

Bnacrs. | Rays, Disk FLORETS.

PG ee eee Ce 199 199 | 199

NUR E 44.030150 42.874371 | 62.452261

Mode PS Be 3549 a ae 1 35 aan st | 51, 60, 63, 70

ad ou oe * 4-111310 5.048407 7.688088

c PATRE NOKIA: 5.212961 6.308112 | 9-314270

POR AI i so: + 3.516142 + 4.254822 | + 6.282475

PREM ee + .249252 + .301616 + -445352

PES Y 00 0 + .176248 + .213267 + -314912

C. V 11.839527 14.710682 14.91422

There is a high degree of correlation between rays and

bracts (Fig. 9), which is the more apparent because of the

close agreement in the number of rays and bracts; 10.54 per

cent of all the capitula had the number of rays and bracts

equal. The coefficient of correlation between these was

.802388 + .012685. Between rays and disk florets the corre-

lation (Fig. 10) was much lower, the coefficient being .594798

= .024859.

Aster puniceus L.— The polygons of distributign of the

bracts (Fig. 11), rays (Fig. 12), and disk florets (Fig. 13) of

Aster puniceus L. are all multimodal ; but it would not be fair to

assume that this condition is a specific one, for, although the

number of variates was 798, they represent only three individ-

uals, and these individuals had an exceedingly wide range of

variation and also differed widely in the values of their “ con-

stants.” These facts will be best appreciated by a study of

the table on the following page.

1 The table of constants for Aster nove-anglie L. is presented because the multi-

modal condition of the polygons of distribution of this species may be due to too

limited material. If more material shows this same condition, then all the con-

stants except the mean and mode must esca suco olus The same is

ate mn a t Ae Sie alt At ch

TOY ali OF Te sp B5*

120 THE AMERICAN NATURALIST.

[Vor. XXXVI.

TABLE C.— CONSTANTS OF ASTER PUNICEUS L.

INDiviDUAL | IwDbiviDUAL

OV. | Now 2.

337 heads. | 246 heads.

Mean. . . 42.501483 | 45.378048

| Modes :

|A-D.. . .| 3965747 | | 3.140095

BRACTS Jig... 4.760197 | 3.959072

||P. E. A. D. | + 3210752 | + 2.670394

[IP E. M... |] E 04901 x .170258

FE i .123673 | + .120390

CV. 11.200072 8.724641

Mean... -| 35.020771 36.321138

Modes :

NE gXEK ui 3.039368 3.421607

Bi N ul 3.904214 4-297721

Poe A. D. + 2.633392 | + 2.898813

EM- . | x -143450 | + -184821

P.E.c . .| + .101434 | + .130688

EIC Vie s 11.148282 11.832838.

f|Mean. . .| 60.513353 | 76.609756

Modes ‘

Disk 4 A. D. 5.805756 | 8.068

m> 7.338405 | 10.083765

P. E. A. D. | + 4.949754 | + 6.801500

P.E.M + .26963o | + .433647

P.E. v + .190657 | + .306628

tic: v. 12.126919 | 13.162508

INDIVIDUAL SUMMATION,

No. 3. OS. 1, 2, and 3.

215 heads.

46.795348 | 44.546365

35 39% 49,

46, 48

2.377 501 3.571510

3-035446 4-497254

X 2.047408 | + 3.033397

+ .139632 | + .107381

+ .098734 | + .075929

6.486640 10.09 5669

39-660465 36.671679

7h 3» 37

3-175164 3-577188

4.018685 4.480251

+ 2.710603 | + 3.021929

+ .18486r | + .106975

+ .130716 | + .075642

10.1 32723 12.217198

74.241860 69.174185

57-58, 63-64,

67-68, 73-74,

5.644478 8.947764

7.115330 11.116989

+ 4.799290 | + 7.498400

+ .327308 | + .265440

+ -231442 | + .187694

9.583987 | 16.071008

This work on Aster puniceus L. shows what a wide difference

may exist, even in such constants as the means, standard devi-

ations, and coefficients of variability, in individuals growing

under apparently identical conditions.

In Fig. 14 is shown the correlation surface for rays and -

_ bracts of all the heads counted. The coefficient of correlation

was .705100 + .009194, while that for rays and disk (Fig. 15)

was 674928 + 010045. t

No. 422.] QUANTITATIVE STUDY OF VARIATION 12!

Aster prenanthoides Muhl. — The frequency polygons for all

the heads counted in this species are shown in Figs. 16, 17,

and 18, All are multimodal, but by doubling the classes in

Fig. 17, which represents the variation in the rays, the curve

becomes monomodal with negative skewness, the mode being

on 26-27 and the mean on 28.037993.

Doubling the classes in Fig. 16 still leaves three modes in

the frequency polygon of the bracts. These modes occur on

3 ELI |_|

ILITITIITIEII

Saas

oS 2S ee a es

Lt iti m) -

3) ul 5i e

IG. 6. — Aster nove-anglie L. Bract curve of 199 heads. Classes doubled. Mean = 44.030150 =

jui sue eae x aee 37, 43, 47, 575 € = 5.212961 + 176248.

40-41, 44-45, and 48-49. Fig. 18 shows the curve of the disk

florets to have two nearly equal modes on 48-49 and 52-5 3-

The correlation of rays and bracts in the 658 or a

counted (Fig. 19) is expressed by the coefficient 77

+ .007820, and that of the rays and disk florets (Fig. 20) by

770316 + .008042.

122 THE AMERICAN NATURALIST. [Vor. XXXVI.

TABLE D.— CONSTANTS OF ASTER PRENANTHOIDES Munur.

BRAcTs. | Rays. | Disk FLorRETS.

a ace nee aaan E NESE S EEE MS A EANA PASESER SS UM DN

No. : | 658 658 658

Mean 44.044072 28.037993 50.297872

| ( 40-41 ^ ( 48-49.

Mode 1 44-45 26-27 1

| | 48-49 | L 52-53

A.D, | 4-497002 31313229 4.884951

Sio 5.716510 4.070071 6.310315

PRAD + 3-855786 + 2.745263 + 4.256308

PLE, M. | zx .150314 + .10702I + .165889

REg. + .106288 + .075675 £ 417301

CV. | 12.979068 14.516272 12.545890

Result of the Successive Collections. — The remaining figures

represent the conditions found in four successive pickings

made from a single group of individuals of Aster prenanthoides

Muhl

I. Bracts. Figs. 21, 22, 23, and 24 show the frequency

polygons of the bracts in the successive collections. In the

first collection (Fig. 21) there was a single mode on 49-50,

with the mean on 47.410256 and a strong positive skewness.

In the second collection (Fig. 22) the curve broke up into

three modes on 40-41, 44-45, and 48-51, while the mean fell

to 44:342657. In the third collection (Fig. 23) the material

exhibited two modes on 44-45 and 50-51, which correspond

closely with the upper two modes of the second picking, the

mean having fallen to 43.834532. At the last collection the

curve, given in Fig. 24, showed a strong mode on 41-42,

while the upper mode, which at this time occurred on.49—50,

had become much less prominent, and a small mode had

appeared on 33-34. The mean had continued to fall, and in

this last collection was only 41.92045. ic :

. . The table on page 124 will facilitate a comparison of the

bracts from the four collections. e Ne

No. 422.] QUANTITATIVE STUDY OF VARIATION 123

30 B i

25 d

E ]

x & ]

20| | | E] |

am |

ae |

FE 3

eL EEI

[i |

1T |

B q

10 11 IL TT Li

i] IL 11 L1

HT I

HT |

5 ITI gi

1T [i

IH {|

IT] l!

30 m 50 G0

31 ly 5! 6i

Fic. 7. — Aster nove-anglie L. Ray curve of 199 heads. Premi — Mean = 42.874371 =

.301616 ; modes = 37, 43, 47, 51

20 LT PTT titi

E Bate S

EI E

Li E

‘5 2

Ed IH:

10 ae |

T TT HT

5| T !

ail

EN i

y t) 3) 70

gl rt 61 7 BI

Fic. 8. — Aster nove-anglie L. Disk curve of 199 heads. a (€ Mean = 62.452261 +

.445352 ; modes = 51, 60, 63, 79; € = 9-314270 + -3149

124 THE AMERICAN NATURALIST. [VoL. XXXVI.

TABLE E. — CONSTANTS OF Bracts oF Four PICKINGS OF

ASTER PRENANTHOIDES Muur.

117 CAPITULA, 143 CAPITULA, | 139 CAPITULA, 176 CAPITULA,

COLLECTED COLLECTED COLLECTED COLLECTED

SEPTEMBER 27. SEPTEMBER 30. OCTOBER 4. OCTOBER 8.

Min . RS 47.410256 44. 342657 43.834532 41.92045

40-41 44-45 33-94

Mode . . 3 49- 50 , 44-45 50-51 41-42

48-51 49-50

K 1h o2 9 3 4.350646 4.256345 4.308211 ' 3.855552

DÀ E 5.524237 5.152370 5.275976 | 4.889626

Ek E. hes D. . .[ x 3.726098 + 3.475273 | + 3-558646 | + 3.298052

FEM |. | + .344540 + .290617 | + .301840 | + .248600

PRA. -4 243626 + .205497 | + .213433 | + .175786

CV | | 11.664053

al Steet. 11.619444 | 12.036119

2. Rays. The frequency polygon for rays from the first

collection (Fig. 25) shows a strong mode on 32-35 and a lesser

mode on 26-27. The mean of this collection was 30.769230.

In the second collection (Fig. 26) the principal mode had

fallen to 30-33, filling up the sinus and forming a monomodal

curve. The mean had also fallen to 28.706293. In the third-

collection the rays again broke up into a multimodal condition,

as shown in Fig. 27. The modes of this curve are on 24-25,

28-29, and 32-33, and the mean on 28.251798. In the fourth

collection the rays again exhibited a strongly monomodal con-

dition (Fig. 28), with the mode on 26-27, while the mean had

fallen to 26.335227.

3. Disk Florets. In Fig. 29 is shown the multimodal fre-

quency polygon for the disk florets of the first collection, with

modes on 56-57 and 60-61, and the mean on 56.427350. In

the second collection (Fig. 30) modes occurred on 45-46 and

51-52, and the mean had fallen to 51.713286. In Fig. 31 is

represented the variation of the disk florets of the third collec-

tion. The curve is monomodal and nearly normal, the mode

occurring on 49-50 and the mean on 49.158273. The disk

curve of the fourth collection (Fig. 32) is also strongly mono-

modal and nearly normal, but the mode has fallen to 45-46

and the mean to 45.778409.

No. 422.]

QUANTITATIVE STUDY OF VARIATION 125

] FA 71-5 J- | Ej

[3031 |32]33|3]35 36 [3738 39 |o ur [23 |a usne| 7 48 1[52/53 |54[55|56 a

| [38 2 |!

| | al [30 I i

{all DI l i Ili

i I l

33 i l ar

[ [35 34 j 11a i

| |-1/ |35 Ur ee

E. 36 [ l 43|2|2

A Eag 2[3] 12 uy| |l

Li 111/21]

Lal 39 12H igs l

[2 M0 113|3] |

m l Loy 22a iit [

kd 11412]113 l i

| 0 tr] [EINI] ta

| | i tat tt 22 I

EEES 11212121113

E THil2litia3i2lult] [0] ti

E. 1|2 1 [ 2|i|i]r] |i

E] 1 Hi iH LH

ae 1|1|2. I I

Ba a. I il2/i2]1| [14

Ej 51 i ia] tt

A 52 l l

|| I8 53 i 2

L| JM Se RB

| I B B l

E i I i

E! 1 Ü

E] i

E 1i I

Fic. 9. — Aster nove-angliz L. Correlation surface for 199 heads.

relative, p= -802388 + .012685.

Rays subject and bracts

126 IHE AMERICAN NATURALIST. (Vor. XXXVI.

en |

em MI. Dip

= =

—-

= Lad -- bL.

- -|-— =a

S Sa

—T—3 aan ————À —1—41

= ai

-leQ =) -

I—-—1———7TL-—. SS oe O

= pan n —| |- -

= - iat —

— | = je

— —]— D - — — -p

—-—-— ——L—]1—1 “se

—- -

~ -— - cr ~~ =

I——]1——24——— —1—

—- "| - -

am ns i ee oe es O =

<j -1— - -

a os on oo ——— ——

- N —j—| ~AN —

ES51-—1——1——- ee ee Gd GONS ad SN S OO O I] Se ee

—|— ~ - —-

= banal

— Sed fend -|-—

on aaan OS S Da

-i|-— N N —_

- e ~ |=

[ N - -l-— A

: F—L—3——3-——-— —1 LL.

Me =) |=] - —| —

L—3— —T— a ae a —] E—À————4——3 I-—4

4 —— | = — | =] — —

oon oe oe SED GS Gees eee ee ee ee ee eee ee cae Cente E = ee ae

met - - —| i—-le

I——4—3— eo eee mas ee on oe oe ee

Es - -| -—

- ——41 e e —ÍL— Tm uM tt HSH HH SH

tis —- - - — N pae

Si it tte

Te re N -

: Lad

[i -| ——

pt L1

—77]1—3—34-—4-— — oe CN =a

— —Do— | jew, —]|—

us ae oe See See = a ae

Nj == - -|-

= —]—31-—-———À—L-1.1.

—]——L—1—1L—L-]

—- -| i

4 en SE SNNN

— — — i

I - a

T —d4———34À

rz) HN ||

~ ~ —-

$ B aanas pani eee ee SONS GNN

| —

a — ie —=

— i —

—LÉ—-— L—IL—L—

> |

m i

— + —LL——L

i =

258 E]

" | | p UH am ^

: I i; LII

Fic. 10. — Aster nova-angiiz 1. Correlation surface for 199 heads. Rays subject and disk florets

relative, p= .594798 = .024559.

127

ARIA TION

STUDY QF

ANTITATIVE

QU.

No. 422.]

rH [ BER TERRE I rT

HH : LET pies) ] H3

; ] Lr CeCe | H

= | cc irri] + H

} l E LI ] | crt

] LL LLLI [ rri

| FIT LILITI | Ld

i C CCEE ] Tit

i ZI rre : oe

pr c

ema esent jet I

EAR m - Hl z [

EERI Li r1

EEI Li rt i f |

| | Li Tt t t ^

l mi PEETI |

| Li t 1 i

: H | | [

| 1 |

i | i

l | |

| | | i

| | i |

| | | i

: | | i

l | | | :

: l | | E |

: - | | Li l

& R 3 z

44.546365 + .107381

; q = 4497254 + 075929

of 798 heads. Mean

modes = 32, 35, 39

Fic. 11. — Aster puniceus L. Bract curve

THE. AMERICAN NATURALIST. [VoL. XXXVI.

LLLI

LELE

LII

Fic. 12. — Aster puniceus L. Ray curve of 798 heads. Mean — 36.671679 « -075642 ;

modes = 27, 35, 37; T= 4-480251 + .075642,

No. 422.] QUANTITATIVE STUDY OF VARIATION

e Eee wl wE

HET

]

i 1

[]

3 n

Imi i

T LE

40 50 g » 30 100 d

FIG. 13. — Scl ee L. Disk curve of 79

; modes =

57-38, 63-64, 67-68, 73-74: 79-

80; d — 11.

116989 =

129

8 heads. Classes doubled. Mean = 69.174185 +

187694.

[Vor. XXXVI.

THE AMERICAN NATURALIST.

—

Se SESS LEL

eS ee 22-3

29)30/31[32]33]34)35]36]37| 38|39 [90 | [42] 3 |] us |46|u7 48 |u9|50 [5152 [53 [9^ [55

BEEN

Amm

irme edo est» 2

Fic. x4. — Aster puniceus L. Correlation surface for 798 heads. Rays subject and bracts relative.

P= .705100 + +009 194,

131

— fm mre

—-[—L- Pa m

— "n Tol z

Gi pepa | EA 1

— EI ELLE Ed

Niem -— uw]

—IE--1- £

— | mei be

[

bi m F

T -N -leos X i

— = e — El

p ET - -|= mu mm i

— — es oe at i 2

E sa at je.

^ I2

mi f

LTD ele Rut um Is

— II I3.

TET EM

= I.

aum ga VEN pe

p om — | A m] x

A _

= we wea js

pn del S T

e; — -

— -

— | pom ees pt" re

— - pg fa

— i a | mae [ee y

— = fe]

=a me pm

— — -— —

eei D

— "m

| - -

— iss AE

-— =

SS OOS A ee aS —-——-——

Sa E S ae S ee a A ———À——4À ———52: EB

—

L| E

S mol

| and disk

Fic. 15. — Aster puniceus L. eee surface for 798 heads. Rays subject

florets relative. p= .674928 = 010045.

132 THE AMERICAN NATURALIST. (VoL. XXXVI.

50 FYTI |

ra us um

» "E

` Tr

35 [ii

Tit

- Li

"5 atl

" Li

b d M

LLII. [T H

ELEL "T

mu a

E EI.

" 35 » i5 5 a

Fic. 16. — A ster p thatde

modes = 32, 34; 39, 41, 43, 493 9 = 5.716510 + .1

uhl. Bract curve of 658 heads. Mean — 44.044072 + .150314 ;

06288.

No. 422.] QUANTITATIVE STUDY OF VARIATION 133

60)

S t

^ |

E T1

~ |

gs HL H

[| | T

i! | T

30 t

8 t

* i

5 t1

x 1]

20| | tT

a he See =

i5 HH

g TILITLELLLTLA4

10 HTHIIPHTIIITEERÉ IEEE

- jJ 1l id dd tt 1-1

Hia bil -LLEEHH-HHHHC

HETTHETRITE rH

7 HH HTT

itl

i5 20 E] p 35

Fic. 17. — Aster prenanthoides Muhl. Ray curve of 658 heads.

mod

Mean = 28.037993 + -107021 ;

es = 27, 31, 33; T = 4-070971 = .075675

134 THE AMERICAN NATURALIST.

[VoL. XXXVI.

u 3

Fic. 18. — Aster prenanthoides Muhl. Disk curve of 658 heads.

€

lasses doubled. Mean =

50.297872 + .165889 ; modes 48-49, 52-53 ; € = 6.310315 .117301.

No. 422.] QUANTITATIVE STUDY OF VARIATION 135

a fs = mt MNSENE LILLI T

WI ros

N| — co [4€ [40 —

a GINERE e s Sr te [Es ES RS [ES SS e ps [s

— Lip na

| | po E

— E Mex

~ Co| il |

POP eo -— M ds Ir

—|— Geo & z 1g

P |I E - [

m ci — Nie — w Sil]

Sur POOF CFS eS Sl de

E =|= © gam ca, A

Oj é wa

-laella =|!

— NNN I I1

m E sum Dmm

E -— =E N m= S

— js =

bl nw co EE

i — =

eu = =

— —_ Sea E

sed 3 uri To - Ur

ied ennt — — — u

— — wu

| =

| Sls

pee v

= co

= we 2

gad e

Fic. 19. — Aster p thoides Muhl. Correlation surface for 658 heads. Rays subject and bracts

relative. p= .776834 + .007820.

THE AMERICAN NATURALIST. [VoL. XXXVI.

emee

3711

PII

ETAR

2|3

o[sij52]sa |54 55|56

5/1/31

-P--

47148

Lidl

| 13]2

e| — || IN IN| |

9401918243]

teat eeathieea. Aas eee

ipa

HR-

ETTET PP TTETTI

Hitt E ERR E BN NH

Fic. 20. — Aster prenanthoides Muhl

- Correlation oo Rays subject

+ 008042.

and disk florets relative. p= .770316

No. 422.] QUANTITATIVE STUDY OF VARIATION 137

25 - [T

20| La

BITS

$52 Hi

HRS i

m ITI Mi

of T T id

B MILL |

i T d

t HET

a Hiii

5 TIT T

HT TM

! IT TE

HT TH

3 TEL TET

! p 50 60

p 3) lj 5i él

Fic. 22

40 50 60 70

Fic. 21. 30

7 ELEELE] 25

L1 a

20 San ee 20

EETTIT u PPEIT

10 TANER 10 ELIT

EELELIL. a2 8 Sage

— = —_

5 INI E] 3T TT *

30 zy) 60

3 4 Ši 6i 30 ug 59

FIG. 23. Fic. 24.

Classes

Fic. 21. — Aster prenanthoides Muhl. Bract curve of 117 heads collected September 27.

doubled. Mean = 0256 +. o; mode = 49-50; 4 = 5.524237 + -243626.

G. 22 ster p : thoi z Š Muhl act cu curve of 143 heads collect on September 30. Classes

č 723. I

G. 23. — Aster prenanthoides Muhl. Braet curve of 1

doubled. ean = 43.834532 + tah .840; modes, 44-45;

Fic. 24. — Aster enanthoides Mu h Bract curve. rs 176 he —€— grate

doubled. Mean = 41.92045 modes -34s P o»: 49-50 4.8896

I 38 THE AMERICAN NATURALIST. [Vor. XXXVI.

i. A

TI SN

10 |l

iil

5 |i

Lil

21 31 4

Fic. 25

der dw d EJ d

& si TT arem

—— oe ae ay

; ELA aa aa Nae

—À — 4—— a ———À——

ARA San

5 ase aS 1:

Ree if

| HEU

10 iol | T] |

cee i

os ia EE RA

I metet n

: 30 : 20 30

21 3 uu 2i 3l iH

« 39.

Fic. 25. — Aster ebrei Muhl. eras curve of 117 heads collected on September 27. Classes

doubled 547; mode — 26-27, 32-35; Lid -9858 ha 173749.

Fie. 26. — A e Prenante tit Mobi. E curve of 143 wee 30. Classes

oubled. Mean = 28.706293 = .201316; m 30-333 € = 3.569141 + n ic

. FiG. 27. — Aster prenanthoides Muhl. ted curve gt 139 kad collected on October 4. Classes

doubled. Mean = 28.251798 + .200320; modes = 24-25, 28-29, 32-33 ; € = 3.501476 + . 141641.

No.422] QUANTITATIVE STUDY OF VARIATION 139

4 OO

HHH " EH :

ee aR Li

LLLI.

viii

FFH- TETTE

Se ee ' Liu

Pug ET 0 in rl

"-—- HEI

ETT Bim

mg eu Lu Tj

HH H

BE oe 5 LL

S Th T1

ELIT EI

FT LEE itl

lh 1l

30 40 50 60 70

hi u 5) 6l 71 8l

W Fic. 29

pi E

~ r.i

55 ni 25

20 =n Gam

TTE. 20 ee n ee Gee ee —4—

Bey g L—1—3 cuna = Se 8 ee ee

aaa ta Ge 5 LIE. E IEEE LÀ EJ

SLET :

5 4 — $f n n a

FIT = L—— = = L 18 1 1 1 — 331—193

LIII) LERLI. a! Bis FR ea ee

t arii

of Tit wo —

Uhl Oe ee ee. eee

rp TERI LIS 1 1] Enana

b Ll Mu NE ok DO GS E Gs Ke oe es

Lhi

LIM | aur

Lil "

PX. P D 5

a à " uo 50 a

Fic. 28. Fic. 30.

Fic. 28. — Aster prenanthoides Muhl. sind curve of 176 heads —— on October 8. Classes

oubled. ean = 26.335237 + .153066; mode = 26-27; 010607 + -108234

Fic. 29. — As gí T Muhl. Disk curve of 117 ai Per on a Sepii 27. Classes

oubled. Mea 7356 + .248547 ; modes s 1; o= 3.985839 + -175749-

llected on September 3o. Classes

n= 8547; m = 56-57

Fic. 30. — Aster b Muhl. Disk curve of r43 heads co

do ubled. Mean = 51.713286 + .281728; modes = 45-46, 517525 7 = 4-994779 = .199211-

140 THE AMERICAN NATURALIST. (Vor. XXXVI.

ege

E S in S i

FiG. 31. — Se Ari des M

fis oides Muhl. Disk curve of 139 heads collected on October 4. Classes

49.158273 + -279452 ; mode = 49-50; o = 4.884653 + . 197602.

z SZ

2 a S & ^ = v

Fic. 32. — A ster prenanthotdes Muhl. D

e isk curve of 176 poss woes on October 8. Classes

doubled. Mean = 45.778409 + .242280; mode = 45- = 4.777197 + .171318.

No. 422.] QUANTITATIVE STUDY OF VARIATION. 141

ES FT E

[ X;-8-8-2 TH 1-65-34-3-2-1 0 0 [

hd 33|34/35|36 37 3853 |40 |41 [4243

4 -EEEEELEHEHHAHUHHHHHH-HH

z i EIITILLDELELETLLLLLE

z l SUELE ES IIEUTEFELTELELLE- 4 ITI

i T222 ga ITI as Sa ddl

far [ 2|1[2|1! l Hae SUMP Re

» BE 2|! fl He Seek

X L2 ili l MGR H R A E E E A

- tl ja} {au m oe ee

1 j

—

LELI

i| | |

"rli

S2R2 8

ELELILAI Edd]

SRS

ELLEL IM

ELI

‘Bae se

—S] So St |

Fic. 33. — Aster dio cet Jes Muhl. Correlation surface for 117 heads sings on September #7.

Rays subject and bracts relative. p= .855944 + -0122

CLEC

3013313413 37:38 [39 [90111 [2113 |] u5]86 [47 4514915015115 | -

M EI

|] li ta

Ena E

A CHEGA at a

8 [20 Bs CLERI,

M |] LIE

"I mp nu gu dtl Ed.

T LI LH 112 b ribs. Uc

| Th Pa | | iat |21|.— 2|1|1 DE ed

BE 8 we Enr | Lb Lobo

- a i| 12/6 rli gm

Be LIST ee 211i LI

ae oe ot ume 2l! 2 L Laud

ae 2| lati 1212] | wes

ee re l 1| 1312 apn

SEPE N: me PE M UE 3lii211131211[2] tei SS

OT Ur peer

E LIT MT E. n ELI.

LI EU TTIPIULL. 5 TT

[I ap ; sas

LI LL [1 Hiii.

Fic. 34. — Aster prenanthoides Muhl. Correlation surface for 143 heads collected on September 30. »

Rays subject and bracts relative. P= 833702 + .012701.

142 THE AMERICAN NATURALIST. [Vou. XXXVI.

E: |

-BRM -3-8-7-6 3-8 —3-23— ] l ii

30/3 1]32)33134|35|36/37|38]39]40 [41 [42143044145 1464714849150151 1525358

- [2 | LI

tal Di M dl

22 ai

(1-435 13 Zi itt iJ is j

1-8 29] | | [| ay aan 6113/1] 1

| | 33 (25 2| [hi] li

- (26 TART

hte ie 1/2 512) 1 |

LL ] 21113|3|212 l

|| jp ps Litt [2|2]2/]1]2/2

] 30 i211] 12] in

| RBI iE HI2[21]102] L1 du

)m 2| |2/2]2/^| |2| 1| | |

LEN LIE. iii21115li

Ll] B i| Dn

| j

7 Bol |i

Fic. 35. — Aster prenanthoides Muhl. Correlati

Rays subject and bracts relative.

yf £.

P = .798642 + .01513

139 heads collected on October 4.

15133.

2S EIETITI

| XH- -&-3-À-i- I 9 |

LI. 25[ojai 35/36 87]38 39 0 t fu2 3 |a] t5 fc 108|49 |50 51

ded

G28 [i

| [oe jel 2E

=F i| [ila =

E sE 2|2|1|2] shit ji

Tia 2| |2| [ili

[7 let 1|2[1 2] iZi

ies 2]1| |I

LL 2|211 2|311

LIN i l 16|5|15] |i

ME EE | |4[2|4]2 [111110 i

[I I| [1| [2i5|3|2]1

BL ] i] 11112111 |

LAEN | Ii 312 i

La l l

-.]9 |]. | I i

| | [d 1

Fic. 36. — Aster prenanthoides Muhl. Correlation

Rays subject and bracts relative.

aT a £,

p = .803092 + .o1

176 heads collected on October 8.

3443

No. 422.] QUANTITATIVE STUDY OF VARIATION. 143

| | | |

ee: ERRE 3333433112313 IRENE RE

x p——— sues sels saissteni esee eode 771727374 | |

BEAS i | Lil

HE l 1

| [38 124 l 2| |!

|_| = 1121 |! l i

- |26 l 2i 131111 eee

=3 [27] [ MAREE itt

=2 128) Hilt Lp iui

3 pA am LIO JEN

1301 1 | |

|_| p Bl 12] |! = ili l

[1 B2 tie] Init

L| 2 133 3I leat 2|!

T BIX I 1| fal [0221] 1212131 |

|| p 135) Series l i

LI B l 1 2

Edd l LE

EL L l l L3

| | B BS x GRRE RERBRES ao) ee

————1—1 t—31 F—1—31 L--— p p

L1 TITLE Bu

Fic. 37. — A ster given s Mu face for 117 heads collected on September 27.

hl.

subject and | disk florets relative. p= .573693 + .033347-

ESSERE x " | s

H 3chs7 else fao fuia 55 [67 GG H

| ic NZ kad PEREL] — E

| | = Ed pf ft 19 rm

OPen E pp pp} A

| ie {2 ike a MA ppt L-L—IAÀ7M3M3—

Rar) nee Ao

|_| 7 12 Ed LEE - E deli

E 79 23 LI d LLLI

MEE: LE E LI T Ed

L3 ps LLL TL I-II

| [2 pel l H2 Z | ]!] e 5———

GEA i i 1 LEFELIL LLLI

es Z |

w y m" á [RR LA dL LE =

BINE [DM | l EET ELLLL

LR i | i3itiiia| Di LI LLLLL

L| BI» TT T1 nii pit nisi ] BL LLL TL.

[TN a3 UTITITLLLH 2| U2 e

) E bod Io dE ee ae a

[-| T7 B6 a i ITILLET:

i A HHHH

wee See we

Vic. 38. — Aster prenanthoides s Muhl. Correlation surface for 143 heads collected on September 3°

Rays subject and disk flore ets relative. p=.358909 + 5.

144 THE AMERICAN NATURALIST. [VoL. XXXVI.

E 12-801 6 0-8 3-72 3178 3 3 21-73 3-2] 6 4 1 2 3 ] 1 3i

as 25/26/27128.25130131/32/33134135136137138139170l 11 [u2 |u3| qu ws uel segfsojs salsa sss s7 salse Gd dou

~ Hd

= 1

d I |

Be l Vii [pat Dif D2

3 Har 12 hi 12! Hi 1 Hane Oe ay

BE 23) [i] gata [i

EE EE 1/3

Lit i 3|1] 13 1 1i

LL: MEID 1

1 DAN eA

1 1i3[1 | I

ull 113]1i1j3]| |!

LILI [11313111212 l

2| |i i i

] HEI i i

| "030 E RS Ree

l FEILLI

| CETINI

ji i Rene

Fic. 39. — Aster prenanthoides Muhl. Correlation surface bus 139 "— collected on October 4.

Rays subject and disk florets relative. P = .353102 + .043050.

E | HHHH

Be: 31/32/33] 39/35/3637]38 39 uo [i1 [42 9344 45]u6|47 8/49/5015 | [52153154155

LIS l IIT ||

aS LH 3-4 |i]

EE RR l [ eS

mg iT [21 ]- 12 3[3| | |2 i

p p l 3|1|1| (HILH

l- 111[1|2)412/13]2 2

BE 3|1| |1[3|2|2 2

M d l Le Rn 2

ES 2| |3|3|1]9]8[1[2|212|3|2 | 1 l

Eu iili 2| iji LII.

ee LI HEn iSieret filet | | 1

TB Bo iti | felze | T 12/2

iM; Eo prei nnl

Id LAIT I

E E l |

| HH

L1 HA m

Fic. 40. — Aster PESEE Muhl. Correlation surface for 176 "ead case on October 8

ys subject and disk florets relative. p= 545208 + ;

No. 422.] QUANTITATIVE STUDY OF VARIATION.

145

TABLE F. — Constants OF Rays or Four PICKINGS OF

ASTER PRENANTHOIDES

Munr.

| 117 CAPITULA,

143 CAPITULA, |

139 CAPITULA, |

| COLLECTED COLLECTED | COLLECTED

| SEPTEMBER 27. | SEPTEMBER 30. | OCTOBER 4.

pec T

Mean |^ 30.769230 | 28.706293 28.251798

| | | 24-2

26- |

Mode { mise | 30-33 . | 1 28-2

| 32-33

A. D. 3.353057. — 2.9917 36 3.012680

es ccu 3-985839 | 3-569141 3.501476

FEAD + 2.688448 + 2.407386 | + 2.361745

P. E. M. + .248547 | + .201316 | + .200320

RE. + .175749 | + .142351 | + .141641

CV; 12.953945 | 12.4333306 | 12.393818

176 CAPITULA,

| COLLECTED

OCTOBER 8.

26.335227

| 26-27

| + .108234

| 11.431865

The variations of the disk in the four collections may be

compared in the following table.

TABLE G. — CONSTANTS OF Disk FLORETS OF Four PICKINGS OF

ASTER PRENANTHOIDES MUHL.

117 CAPITULA, 143 CAPITULA, 139 CAPITULA, 176 CAPITULA,

COLLECTED COLLECTED COLLECTED COLLECTED

SEPTEMBER 27. | SEPTEMBER 30.

Mean 56.427350 | 51713286 | 49.158273 | 45.778409

Mode 1 56-57 f 45-46 49-50 45-46

60-61 51-52

A. D. 3-353057 3.925179 3.619481 3.750581

ce pM 3.983839 4-994779 4.884653 4-777197

P. E. A. D. + 2.688448 + 3.368978 + 3.294698 + 3.222219

P. E. M. 4+ .248947 | + .281728 | + .279452 | + -242280

FEY. + .175749 + .19921I + .197602 + .171318

C. X. 12.953947 9661212 | 9.936585 10.435481

4. Correlations. The correlation surfaces for rays and bracts,

in the four collections, are shown in F igs. 33, 34 35» and

36, and for rays an

d disks in Figs. 37, 38, 39, and 40. The

coefficients of correlation are arranged for comparison acum

following table.

146 THE AMERICAN NATURALIST. [Vow. XXXVI.

TABLE H. — CORRELATION OF Rays AND BRACTS AND OF Rays AND

Disk FLORETS IN Four PICKINGS OF ASTER

PRENANTHOIDES MUHL.

117 CAPITULA, 143 CAPITULA, 139 CAPITULA, | 176 CAPITULA,

OLLE COLLECTED COLLECTED | COLLECTED

SEPTEMBER 27. SEPTEMBER 3o. | OCTOBER 4. | OCTOBER 8.

SRM Bes EE EN ae ea EE : See See

Rays and | p | 855044 | 833702 798642 | 803092

Bracts (P.E. p| + .012237 + .012701 $ 015133 | + .013443

ee ins coge Setar eer cerdo AORA EA

Rays and p -573693 358909 353102 -545208

Disk E. nid + .033347 + .042145 + .043050 + .012157

Inspection of this table shows that the highest degree of

correlation between the parts was found at the beginning of the

flowering season, and that there was a constant fall for both

bracts and disk florets until near the end of the season, when

the coefficient of correlation again slightly increased. The

increase in the coefficient of correlation between rays and

bracts in the last collection was less than the « probable error ”

of the determination, but that between the rays and disk florets

had increased until its value was only .0284 less than in the

first collection.

V. Discussion or RESULTS.

The species which have been chosen for this study are quite

distinct, and although Aster puniceus L. and Aster prenanthotdes

Muhl. belong to the same division of the genus, Aster nove-

anglié L. is more distant from these, and Aster shortit Hook.

belongs to a division which is separated in a marked degree

from the others. Considering this wide separation, it is rather

remarkable that there should be so close an agreement in the

number of bracts in the four species. The lower limit of

range varies from 28 to 30, the upper limit from 49 to 64, and

the means from 36.8 to 44.5. With the bracts so nearly con-

stant, there is a range in the mean number of rays from 14 in

A. shortit Hook. to 42.8 in A. nove-anghe L., and yet the

degree of correlation between rays and bracts is high in all

No. 422.] QUANTITATIVE STUDY OF VARIATION. 147

the species studied, the coefficient of correlation between them

ranging from .549 + to .802 +.

The fact that the correlation between bracts and rays was

found to be greater in every case than that between rays and

disk florets gave rise to the question, What is the relation

existing between bracts and rays? After a careful study of

the position of the rays with reference to the bracts, it seems

that the rays are axillary to the bracts, and that typically a

nearly constant proportion of the bracts in the capitula of

. a species produce rays in their axils, the rest remaining empty.

The material studied showed the mean number of empty

bracts in each head to be as follows: A. sortii Hook. 22.8,

A. prenanthoides Muhl. 16, A. puniceus L. 7.9, and A. nove-

angle L. 1.2.

The number of species here-studied is too small to permit

the derivation of laws covering so large a genus as Aster, but

in these four species the degree of imbrication of the involucral

bracts was apparently in direct proportion to the number of

empty bracts. In A. shortii Hook., which has so many empty

bracts, the scars left by their removal occupies the convex

surface of an inverted cone, the base of which served as the

receptacle for the comparatively small number of florets; while

in A. nove-anglig l., where almost every bract has its ray,

the scars formed a narrow ring about the broad receptacle.

In the former there was considerable difference in form and

size between the minute outer bracts and the inner ray-bearing

bracts, while in the latter all the bracts were very much alike in

In a number of capitula of A. nove-anglie L.

size and form.

Some of the more

there were found more rays than bracts.

marked cases of this kind were examined and the rays distinctly

seen to form a scattering second row within the full outer row

which is typical of Aster; in other words, some of the disk

florets developed ligulate corollas.

The cue several writers that statistical s

will prove valuable in taxonomic work is not sustained by the

results set forth in this paper. It is obviously impossible to

describe a species by means of the variability constante = y

such manner as to allow the classification of individuals man P

148 THE AMERICAN NATURALIST. [Vov. XXXVI.

possess but single variates of the kind used in determining

those constants.

The above-stated results in J. puniceus L. make obvious the

same truth in regard to species such as those of Composite, in

which the individual may have a sufficient number of variates

to give a good frequency polygon and constants with small

“ probable errors.” The wide differences between the three indi-

viduals of A. puniceus L. show that the variability * constants "

for individuals are only in a measure less variable than the

characters upon which they depend.

The study of the successive collections shows another phase

of the subject of variability which would materially affect the

value of statistical methods in taxonomic questions relative to

the Composite. In these successive collections there was a

continuous decline in the numbers of bracts, rays, and disk

florets, and a continuous change in the position of means and

modes. These results could have been in no way dependent

upon unnatural conditions induced by the clipping of the heads,

since even the last to bloom were well-developed buds at the

time the first collection was made; and had there been a change

brought about in this way, it must have been in the opposite

direction, since it is a well-known fact that the removal of the

earlier flowers gives increased vigor to later ones.

This continuous change of means and modes during the

blooming season is suggested as a possible explanation of

Lucas's (98) results upon Chrysanthemum leucanthemum Asi,

other than that of difference of locality. He found that there

was a marked difference between the results obtained from

material collected at Yarmouth and Grand Pic, Nova Scotia,

and that collected later at Milton and Cambridge, Mass. The

earlier material from Nova Scotia showed the mean on 24.389

and the principal mode on 22, while the later collection from

Massachusetts had the mean on 21.61 and the mode on 21.

Ludwig (00), in commenting on these results, attributes

their deviation from his own Observations to the scantiness of

Lucas's material, while Lucas implies, though he does not say

it definitely, that the difference is a local one. In the light of

my observations on 4. prenanthoides Muhl. it seems a fair

No. 422.] QUANTITATIVE STUDY OF VARIATION. 149

question whether the difference may not have been in part

due to the difference in the time at which the collections

were made.!

Although the material of A. prenanthoides Muhl., upon which

this study of successive collections is based, is even less than

that used by Lucas, the constancy of the fall in numbers of all

the parts can leave little doubt as to the essential correctness

of my results. As a few days makes a marked difference in

the condition of these variable characters, different seasons

may likewise be found to give different results upon material

collected from the same individuals.

As the capitula which terminate the axes in the Asters are

the first to bloom, my observations on the decline in the

number of parts in the heads of A. prenanthoides Muhl. bear a

close relation to those of Burkill (95) on Caltha palustris L.,

Ranunculus arvensis L., R. bulbosus L., R. Ficaria, and Tha-

lictrum flavum, where he arrives at the conclusion that the

“ position of the flower on the axis affects the sexual organs, if

they vary." ?

Ludwig (00) remarks concerning this, that “weil die Zahl

der Carpelle, ebenso wie die der anderen Blütentheile und der

Blütenzahlen in der Inflorescenz von der Stellung an der Axe

abhángig ist, wird man Mischcurven erhalten, wenn man die

Zahlungen auf sámmtliche Organe der einzelnen Pflanze, nicht

auf einzelne gleichwerthige Organe der verschiedenen Pflanzen

1 Since this was written, W. L. Tower has investigated Chrysanthemum leu-

canthemum L. with reference to this point, and found that there is a continuous

decrease in the values of mean and modes from the beginning to pa end of the

flowering season. His results will appear in pment vol. I, n

? In his studies on the plants named, and others in which Pn uL of sta-

mens and carpels vary, he finds that *the earlier formed flowers on the plant

carry more stamens or more -— or perhaps more of both organs, than those

formed later in the season,” and that “flowers holding any position of advantage

on an inflorescence, 7.e., terminal on a cyme, or at the base of a raceme, even if

not maturing earliest, carry more stamens or carpels or more of both, than in the

other flowers of the same inflorescence.”

Finally, he finds that in forms such as Ste//aria media, and fe

which have a long flowering season, there is a kg decrease pares

of stamens and carpels during the flowering season, re of a

on the inflorescence, — a result which is closely "pudiicied by my work on

prenanthoides Muhl.

150 THE AMERICAN NATURALIST. [VoL. XXXVI.

ausdehnt, und zur Bestimmung der constanten Mischcurve der

Art ist eine sehr grosse Anzahl von Zahlungen nóthig."

I agree with Ludwig that a large number of counts is

necessary for the determination of the constants in material of

this kind; but how great must it be? It is to be noted that,

in his tables representing the variation in number of achenes

in the heads of Ranunculus acris L., he gives the results of two

sets of observations, each consisting of counts of 1000 heads.

The maxima of the two multimodal curves formed, alternate

with each other, and their summation gives but few maxima

where they ought (?) to be, — the material is insufficient ; yet he

makes virtue of the fact that the summation of Lucas’s data,

only 831 counts, presents maxima upon the series of Fibonacci.

This series of Fibonacci, which is of recognized importance

in the phyllotaxy of flowering plants, should be accepted with

caution as the key to all variation among plants. The members

of the series, along with Ludwig's “ Unterzahlen," which are

made up from the numbers of the Fibonacci series by multipli-

cation or addition, —¢.g., 10 = (2 X 5), 29 = (8 + 21), etc., —

include so large a proportion of all the smaller numbers that

many modes must fall on or near one of them, even if there be

no fundamental relation existing between this complex series

and the number of floral parts or other organs under considera-

tion. To account for modes which do not fall on any of these,

Ludwig creates the ‘‘Scheingipfel,” which is formed by the

overlapping of curves having their modes on adjacent numbers

of the Fibonacci-Ludwig complex. Thus, if the maximum

falls upon 9, it is a “ Scheingipfel" formed by the union of

curves having maxima upon 8 and 10; if it fall upon 11, it is

made up of curves having maxima upon 10 and 13, etc. It is

evident that such a scheme will furnish an Pep Apamon f of

almost any condition which might arise.

7 In conclusion, it needs to be said that there remains much

s to be done in determining the many causes of variation. The

quantitative study of variation shows only the existing condi-

tion of the material studied. It deals only with results, and if

there has been no intentional selection of material, it indicates

nothing as to the causes which have brought about those results.

No. 422.] QUANTITATIVE STUDY OF VARIATION. I5I

While deprecating the selection of material on false bases, I

take it that intelligent selection of material existing under

known conditions is more certain to give valuable results than

at-random selection, which, while sure to select, ignores the

conditions under which the selection is made.

Until we know the causes of variation and how great degree

of variation may be produced by how slight causes, all general-

izations based upon limited observations should be accepted

with caution. Granting that observers have no preconceived

notions as to what they ought to find, the conditions found by

each investigator are true of the material upon which he worked,

but how true his general conclusions are, further investigation

alone can tell.

If every valley and hillside is to have its own place-mode and

every day and every season is to bring forth a changed condition

in the variable characters of its living forms, only a universal

collection of material covering a long period of years can give

us true constants which may be verified by a repetition of the

process.

VI. SUMMARY.

Quantitative studies were made upon the bracts, rays, and

disk florets of four species of Aster growing at Yellow Springs,

Ohio. i

A close correlation was found between bracts and rays, and

attributed to the fact that the rays are axillary to the bracts.

The degree of imbrication of the bracts was observed to bear

a relation to the number of empty bracts.

Curves and “constants” were determined for the material

of the four species studied. “Constants " determined for sev-

eral individuals of A. puniceus L. growing in identical surround-

ings indicated how great variations may exist in the variability

“constants” of individuals.

Studies upon successive collections from a single group of

specimens of A. prenanthoides Muhl. showed that the number

all decrease continuously from

of bracts, rays, and disk florets

the beginning to the end of the flowering season, and that the

152 THE AMERICAN NATURALIST,

character of the curves and the position of their means and

modes likewise change continuously.

I wish to express my thanks to Prof. W. L. Tairas for his

kindly interest and helpful suggestion during the prosecution

of these studies, and also for the revision of the manuscript.

YELLOW SPRINGS, OHIO, "

June 25, 19or.

BIBLIOGRAPHY.

BURKILL, I. H.

1895. On Some Variations in the Number of Stamens and Carpels.

Linnean Society Journal of Botany. Vol. xxxi, pp. 216—245.

DAVENPORT, C. B.

1899. Statistical Methods with Special Reference to Biological Varia-

tion. New York, John Wiley & Sons. 59 pp., 10 tables, 28 figs.

LUCAS, RF. €.

1898. Variation in the Number of Ray-Flowers in the White Daisy.

Amer. Nat. Vol. xxxii, No. 379, pp. 509-511. 2 figs.

Lupwic, F.

895. Ueber Variationskurven und Variationsflachen der Pflanzen.

Botanisches Centralblatt. Bd. lxiv, pp. 1-8, 33-41, 65-72,

97-105. 2 Tafeln

Lupwic, F.

1896. "Weiteres über Fibonaccicurven. Botanisches Centralblatt.

Bd. Ixviii, pp. 1-8. t Taf.

Lupwic, F.

1896. Eine fünfgipfelige Variationscurve. Berichte deutsch. Bot. Ges.

Bd. xiv, pp. 204-207. 1 fi

Lupwic, F.

1898. Die pflanzlichen Variationscurven und die Gauss'sche Wahr-

scheinlichkeitscurve. Botanisches Centralblatt. Bd. Ixxiii,

pp. 241—250, 289-296, 343-349, 374-379. 1 Doppeltaf.

Lupwie, F.

Ueber Variationscurven. Botanisches Centralblatt. Bd. Ixxv,

PP- 97-107, 178—183. 1 Doppeltaf

Lupwic, F.

1900. Ueber Variationspolygone und Wahrscheinlichkeitscurven. Bei-

hefte zum botanischen Centralblatt. Bd. ix, Beiheft 2, pp. 89-

Ht 1 8

STRONG, R. M.

1901. A Fen enin Study of Variation in the Smaller North-American

hrikes. Amer. Nat. Vol. xxxv, No. 412 (April, 1901),

»- 271-298. 8 figs.

EDITORIAL COMMENT.

In order that our summaries may correspond, so far as possible,

to the calendar year, * The Quarterly Records of Gifts, Appointments,

Retirements, and Deaths” will hereafter appear in the numbers for

February, May, August, and November. Under the heading ** Educa-

tional Gifts " we have included all donations from private individuals

for distinctively educational purposes, so far as they have come to

our notice. We have included, therefore, not only gifts to schools

and colleges, but those to libraries and museums, since in many cases

itis otherwise impossible to draw a sharp line. We have not included

in our summaries appropriations by the national, state, or local gov-

ernments for educational purposes, nor have we included the formal

transfer by Mrs. Stanford to the Leland Stanford Junior Univer-

sity of securities and other property, with an estimated value of

$30,000,000, the largest single gift for educational purposes in the

history of the world. Yet, omitting these, the gifts for the year

190r, as catalogued in our pages, foot up the enormous sum of

$43,233,035. These recent educational gifts, increasing in amount

as the years go by, are of immense importance in any estimate of

the future of education in America, and in the opinions of some are

not an unmixed blessing. Certainly, so far as the financial basis is

concerned, the United States will soon stand ahead of any country in

the world, but the question cannot help suggesting itself, Will it

advance as rapidly and to such prominence in intellectual matters as

it does in the material side of education?

While speaking of our Record it may not be out of place to say

that since the American Naturalist came into the hands of the

present management its notices of appointments, resignations, and

deaths has been the most complete published. In these years 1717

personal notes have appeared under these three headings.

153

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Mechanism and Vitalism. — Under the title of Mechanismus und

Vitalismus, Prof. O. Bütschli has published in brochure form his

address before the International Zoólogical Congress which met in

the summer of 19or in Berlin. Bütschli points out that the modern

thinkers and investigators who stand for the doctrine of vitalism, and

who are often referred to as “ Neovitalists," do not in reality uphold

anything fundamentally new, since there is no important distinction

between the old and the new vitalism. Both the old and the new

doctrine rest on the assumption that life and life-processes cannot be

understood, or at least not entirely understood, except as the outcome

of a special principle, or force, or of a “peculiar something," which is

not present in inorganic or rather dead substances. The new doc-

trine of vitalism goes further, perhaps, in maintaining that the purely

causal mechanical point of view of living phenomena is also as cor-

rect as the teleological, but even this is not a real departure from the

older view, since the latter also expressed itself causally in the sense

that the postulated vital force, that was supposed to account for the

phenomena of life, acted according to the causal formula.

Bütschli begins by defining as briefly as possible his own general

standpoint in regard to the theory of knowledge. A few pages are

given up to the discussion of the Ego and the Object. It is not clear

why the author should introduce his subject by such a thread-worn

metaphysical discussion, which is likely, in our opinion, to discourage

and disappoint the reader at the start, but the mantle of metaphysics

falls on the seventh page, and the author returns to his real theme.

Bütschli states that by “mechanism,” as applied to the organism,

he does not mean simply the kind of mechanics that deals with

motion and with equilibrium, but rather the conception of the organ-

ism on the bases of regular sequence of cause and effect in the

same way in which we account for inorganic changes. “A purely

mechanical ption is impracticable even in inorganic phenomena."

There follows an explanation of that most evasive of German words,

“Auslösung,” and its relation to causal phenomena. Bütschli then

NOTES AND LITERATURE. 155

proceeds to give a neat and convincing account of what is meant by

a “descriptive” science and points out how a number of modern

critics have misinterpreted the term. In their desire to show that

all science is only descriptive they have failed to discriminate

between orderly sequences, such as night following day, and neces-

sary sequences, such as the explanation of the alternation of night

and day as the outcome of the revolution of the earth on its

axis, etc.

After discussing whether the simplest organisms — bacteria for

instance — might be accounted for as the outcome of a physico-chem-

ical accident, Biitschli asks: If this is possible, can the same assump-

tion account for the highly complicated organism? This leads toa

discussion of what is meant by “chance” or “accident.” Bütschli

points out that one of the chief peculiarities of living things is their

power of reproducing other living things like themselves, so that if a

given form once arose by chance, its continuation does not any longer

depend on chance, since by its own nature it reproduces that special

group of **accidents " that brought it into existence. The argument

leads naturally enough to Darwin's hypothesis of the origin and sur-

vival of chance variations. Bütschli affirms his belief that up to

the present no better hypothesis has been advanced to explain the

adaptation of organisms to their environment. There follows an

admirably clear analysis of what we mean by adaptation. It would

lead too far to enter into this discussion, but we cannot refrain from

expressing great admiration for the clearness and ability with which

the subject is handled. :

Pflüger's “teleological causal law" is skillfully divested of its meta-

physical covering. Bütschli points out that the same law is equally

applicable to a steam engine with a regulator. Cossmann's recent

argument, in which he attempts to demonstrate a special “ biological "

sequence of causes and effects in organisms as contrasted with the

sequence in the inorganic world, is severely criticised and its fallacy

exposed. T

Bütschli points out that Driesch's demonstration of vitalism rests

on a very doubtful assumption.. If it could be shown, as Driesch

claims, that the reorganization of a piece of an egg or of an adult

into a new whole with proportionate parts isa phenomenon peculiar

to living things, then Bütschli admits that Driesch might make good

his position, but that this is true is by no means proven to be the

case. Asa parallel inorganic phenomenon it is pointed out that »

drop divided in two forms two new drops. Again, if a drop of some

156 THE AMERICAN NATURALIST. [VoL. XXXVI.

substances is drawn out into a cylindrical thread, the latter will,

under certain conditions, break up into a series of drops of definite

size and of definite distances from each other. If two such cylin-

drical threads of unequal size are treated in the same way, they will

form drops proportionate to the original sizes of the threads, etc.

Here Biitschli claims we find an analogy to the tripartite division of

the archenteron of the gastrula of the sea urchin. I may add that a

still more striking parallel is to be found in the behavior of the “fluid

crystals” described by Lehmann. If a portion of one of these is

pinched off, it shows from its optical behavior that it has assumed

the crystal condition characteristic of the original whole.

Bütschli concludes : The old and the new vitalism alike empha-

size the presence of the unsolved riddles of biology and express a

doubt as to their solution on mechanical principles. They teach us

nothing about the organism, since the very premises of the vitalistic

argument rest on the assumption of an ultimate orderly action that

is in itself beyond our comprehension. Therefore, we may well say

that we can only grasp those parts of the phenomena of life that we

can interpret by means of physico-chemical principles. ^ yy M.

Biometrika, *a journal for the statistical study of biological

problems," makes its first appearance with the number for October,

190I. Its aim is to serve “as a means of collecting under one title

biological data of a kind not systematically collected or published in.

any other periodical,” and of spreading such a knowledge of statis-

tical theory as may be requisite for scientific treatment of the data

collected. The editors are “in consultation with Francis Galton,"

Professors W. F. R. Weldon, Karl Pearson, and C. B. Davenport.

Biometrika is published in Cambridge, England, at the University

Press, a sufficient guarantee that the excellent form given to the

initial number will be maintained. An excellent portrait of Darwin,

from the Pinker statue at Oxford, forms the frontispiece. An edi-

torial by Francis Galton is followed by papers by Professor F. Lud-

wig, Miss M. Beeton. and Professors Karl Pearson, W. F. R. Weldon,

and other well-known students of variation.

No. 422.] NOTES AND LITERATURE.

“I

ZOOLOGY.

Biogeographical Regions. — A valuable contribution to biogeog-

raphy has recently been published by Jacobi. The author has

accepted the modern views on geographical distribution, and espe-

cially the fundamental idea that the present distribution does not cor-

respond, in many cases, to the present conditions of life, but has

often its origin in the past, and indicates conditions prevailing in

former geological periods. He points out that the best zodgeo-

graphical divisions proposed by previous authors have not covered

all cases, and cannot do so, because the past conditions were often

directly the opposite to the present ones. Nevertheless, he tries to

give a scheme that is intended to unite past and present conditions,”

and selects Lydekker’s division in three realms (Arktogza, Neogza,

and Notogza) as the most appropriate, since he believes that it

accounts best for the distribution of mammals and birds from the begin-

ning of Tertiary times.

Aside from the question whether it is necessary at all to have any

biogeographical realms or regions, we cannot agree with this idea

that biogeography ought to unite past and present conditions into

one scheme ; indeed, in many cases it is directly impossible to do so,

since we do not see any way to reconcile connection and discon-

nection. And in most cases it would amount just to this, to bring

under one head certain parts of the earth’s surface which are now

connected, while they were formerly disconnected, — or vice versa.

Believing this to be an impossible task, we have always advocated

another method of investigation, namely, the attempt to establish

the present conditions of life (not the actual distribution of animals

or plants) and to divide the earth into regions accordingly. These

regions refer only to the present time, and by comparing this scheme

with the actual distribution of animals, those cases which do not

agree with it are at once revealed. This method calls our attention

to those facts in distribution which need special investigation and

explanation, and in most cases we shall be able to account for them

1 Jacobi, A. Lage und Form biogeographischer Gebiete, Zeitschrift der

er für Erdkunde zu Berlin, Bd. xxxv, Heft 3 (1900), pp- 147-238.

2 pls.

2 The same idea has been advocated by Prof. H. F. Osborn (Science, April 15,

1900, p. 563), who says: “This, then, is our E to connect i: -

tribution with distribution in past time and to gie which will be in

harmony with both sets of facts."

158 THE AMERICAN NATURALIST. [Vor. XXXVI.

by supposed changes in the conditions of life that have taken place

during the earth’s history.

The investigation of instances of the latter kind forms a large

part in Jacobi’s paper, and he has collected valuable material which

tends to show that certain parts of the earth’s surface, in their fauna

and flora, possess a uniformity which is inexplicable by the present

conditions. He calls those parts “areas of dispersal” (Ausérei-

tungsgebiete) and indicates them on his map (Pl. VII). There are

fifteen of them :

Arabian. 11. Philippinian.

1. Greenlandian. 6.

2. Lusitanian. 7. Indo-African. 12. Southern Japanese.

3. Mediterranean. 8. Antarctic. 13. Siberian.

4. Sarmatian. 9. Papuan. 14. Beringian.

5. Iranian. to. Farther Indian. 15. Central-American.

Of these, the 4th (southern Russia and Turkestan), the 5th

(Persia), the roth (Farther India), and the rsth (Central America)

are situated on continents and do not present any remarkable

features, since they are not opposed to the present conditions.

The 2d, connecting England with western France and Spain, the

3d, connecting the Mediterranean countries, the gth, connecting

New Guinea with Australia, the 11th, connecting the Philippine

Islands with each other and with Formosa, the 12th, connecting

South Japan with Korea and China, the 13th, connecting North

Japan with Siberia, and the 14th, connecting Siberia and Alaska,

are well known and have been generally accepted as well established.

The chief interest centers in the remaining areas of dispersal,

namely, the rst (connection of East Greenland with Spitzbergen,

Norway, and Scotland), the 6th and 7th (connection of East Africa

and India, partly by way of Abyssinia and Arabia, partly by way

of Madagascar and the islands of the Indian Ocean), and the 8th

(connection of South Africa, Australia, New Zealand, and South

America with Antarctica). Indeed, none of these connections is

new to science, and some of them have been repeatedly discussed

lately, but it is interesting that Jacobi's studies have led him also

to the assumption of the former existence of these very important

biogeographical relations, which can only be explained by the theory

of a former connection of the respective parts by land. In the dem-

onstration that such conditions must have existed in former times,

and in the collection of known facts as well as introduction of new

ones, which tend to support this assumption, lies the chief value of

No. 422.] NOTES AND LITERATURE. 159

this paper, and thus it will be of great use to any one who proposes

to study these highly interesting zoógeographical questions.

A. E. O.

The Apogonoid Fishes of Japan. — Jordan and Snyder continue

their monographic reviews of the various groups of Japanese fishes

with an account of the cardinal fishes, or Apogonide. Seventeen

species are described, most of them being figured. Six of these are

new, one new genus, Telescopias, being recognized. The authors

have overlooked the fact that Dr. Günther has substituted the generic

name of Synagrops for Melanostoma, which is preoccupied. D. 3. f.

Jenkins on Hawaiian Fishes. — In the Bulletin of the United States

Fish Commission, Dr. O. P. Jenkins continues his studies of the very

rich collection of Hawaiian fishes made by him in the summer of 1889.

Fifteen species are described and figured as new: Sphyrena helleri,

Sphyrena snodgrassi, Anthias fuscipinnis, Aphareus flavivultus, Eupo-

macentrus marginatus, Chromis velox, Chetodon mantelliger, Chetodon

sphenospilus, Ostracion camurum, Ovoides latifrons, Tropidichthys jacta-

tor, Eumycterias biteniatus, Scorpenopsis cacopsis, Parapercis notostigma,

Brotula marginalis. Later investigations of the Hawaiian Commission,

of which Dr. Jenkins is a member, show that Chetodon mantelliger is

the original Chetodon miliaris of Quoy and Gaimard ; Parapercis

notostigma has been recently and earlier described as Percis schauin-

slandi by Steindachner. DSL

Seale on Hawaiian Fishes. — In an * Occasional Paper of the

Bernice Pauahi Bishop Museum of Polynesian Ethnology and Natural

History,” Mr. Alvin Seale, curator of ichthyology, describes six new

species of fishes from Honolulu, illustrating them with photographs

of the type specimens.

These species are: Epinephelus quernus, Novaculichthys tattoo,

Serranus brighami, Balistes fuscolineatus, Monocanthus albopunctatus,

Thalassoma berndti (misprinted “ berendto ").

Of these the Serranus brighami seems to be related rather to

Etelis than to Serranus. D. 5. J.

Starks on the Synonymy of the Fish Skeleton. — In the Pro-

ceedings of the Washington Academy of Sciences, Mr. Edwin Chapin

Starks gives a comparative study of the names applied to the

bones of fishes. This will prove a great convenience to students

160 THE AMERICAN NATURALIST. [VOL XXXVI.

of osteology, as giving a clue to the labyrinth of names due to the

premature assumption of homologies between the fish skeleton and

that of man.

There are now few fields in zodlogy so little worked and at the

same time so repaying as that of the comparative osteology of the

bony fishes. Most anatomists treat the group as though all bony

fishes were alike in their osteology.

The paper is illustrated by plates of the skeleton of the « Striped

Bass," the best drawing of the fish skeleton yet published. These

are by Mrs. Chloe Lesley Starks. BEST

Eigenmann on a New Psenes from Newport, Rhode Island. — In

the Bulletin of the United States Fish Commission, Dr. C. H. Eigen-

mann describes a new oceanic fish from the Gulf Stream off New-

port, under the name of Psenes edwardsi. It was found, as is often

the case with other nomeid fishes, under a Portuguese man of war.

The diagnosis of the family Nomeidz is amended by Dr. Eigen-

mann. ‘The group possesses, like the Stromateidz, denticles in the

throat, and, according to Eigenmann, it differs only in the larger

number of the vertebrae. But in Stromateidz, as in Nomeidz, the

vertebra are in increased numbers, 30 to 36. I know at present no

real difference between the two families. D. 8. .

Eigenmann on the History of the Young Squeteague. — In the

Bulletin of the United States Fish Commission, Dr. Eigenmann gives

a useful study of the development of the young weakfish, or sque-

teague, Cynoscion regalis. Ds

Nishikawa on the Development of the Japanese Anchovy. — In

the Journal of the Japanese Fisheries Bureau, Mr. T. Nishikawa gives

a similar study of the development stages of the common anchovy of

Japan, Engraulis japonicus Schlegel. By some error, Mr. Nishikawa

accredits the name japonicus to Houttuyn. Houttuyn’s Atherina

Japonica, however, was not an anchovy but a sardine. Dsi

. Boulenger on the Classification of the Trachinoid Fishes. — In

the Annals and Magazine of Natural History, Dr. G. A. Boulenger

has a very valuable study of the osteology and relationships of the

group of fishes called Trachinoidea.

The family of Trachinidz was established by Dr. Günther in 1861,

for spiny-rayed fishes, with perfect ventrals, short first dorsal, and

lacking the special traits of other related groups.

No. 422.] NOTES AND LITERATURE. 161

The family was evidently a provisional one, and several writers,

notably Dr. Gill and Dr. Bleeker, have noted the incongruity of its

members and suggested the removal of certain of its subdivisions to

other groups. Nevertheless, in default of any study of the skeletons,

these families have been kept in some sort of association by subse-

quent writers. The study of the skeletons shows plainly the neces-

_sity of a complete revision of the assemblage. This has been done

by Dr. Boulenger. :

Dr. Boulenger shows first that in Trachinus the scapular fenestra,

as in the codfishes, lies between the scapula (hypercoracoid) and the

coracoid (hypocoracoid) instead of piercing the former, as in ordi-

nary fishes. This character is shared by Notothenia, Chznichthys,

Parapercis, Eleginops, Harpagifer, Trichonotus, Callionymus, and

their relatives. All these lack the supraocular lamina, which is

present in Trachinus. Trichonotus and Callionymus show other

osteological characters, which separate them as families, although

not invalidating their general trachinoid relationship. The genus

Percophis agrees closely in osteology with Trachinus, but the scapular

fenestra is entirely within the scapula as in the percoid fishes.

Bembrops, Chimarrhichthys, and Leptoscopus agree with N otothenia

in the absence of a subocular lamina, but differ in having the fenestra

within the scapula. These forms show affinities with the Batrachoi-

dide. Boulenger refers them to Leptoscopide.

Dactyloscopus, which has reduced ventrals and the pectoral arch

of Clinus, is regarded as a true blenny, notwithstanding its apparent

likeness to Uranoscopus. Gillellus, Dactylagnus, and Myxodagnus

will doubtless go with it.

Uranoscopus and its allies (Anema, Ichthyscopus, Kathetostoma,

Ariscopus) have also the scapular fenestra in the scapula, but show

a number of other osteological peculiarities. They are, however,

unquestionably trachinoid in general relationship. E

The group Trachinoidea in Boulenger's view, therefore, includes

the following families : Trachinidz, Nototheniide, Percophide, Lep-

toscopide, Uranoscopide, Trichonotidz, and Callionymidz. All

these agree with the Blenniidz, Batrachoidide, Ophidiidz,, and Gadida

in which the fin rays are frequently not

in having jugular ventrals,

Te uborder of

of the normal number which is I, 5. For the division or suborder

fishes thus characterized, containing these families e thei allies,

Dr. Boulenger proposes to revive the old name “ Jugulares. due

The remaining genera referred to Trachinidz or to Trachinoidea

show no real affinity with Trachinus, Callionymus, and Uranoscopus.

162 THE AMERICAN NATURALIST. [VoL. XXXVI.

They have not jugular ventrals and should be removed to other parts

of the system.

The Chiasmodontide (including Chiasmodon, Pseudoscopelus,

and apparently Champsodon) have the ventrals rather abdominal

than thoracic, not being connected with the pectoral arch, or in

Champsodon joined by ligament only. These may be Percesoces,

but that is very unlikely and their real affinities are doubtful.

The Trichodontidz (Trichodon and Arctoscopus) are percoids,

most nearly related to the Latrididz.

The Sillaginide are, as supposed by Cavier, most nearly allied to

the Sciznide.

Most of the remaining genera agree closely in skeletal characters,

notwithstanding their variations in external appearance, and their

actual relationships are altogether percoid, approaching closely to the

genus Plesiops, which Boulenger ranges among the Serranidz of the

Anthias group. These constitute the family Pseudochromididz, com-

posed of Pseudochromis, Cichlops, Opistognathus, Latilus, Caulola-

tilus, Malacanthus, Bathymaster, and their allies. This family seems

to the writer still. very heterogeneous. Bathymaster has a greatly

increased number of vertebre, Pseudochromis has two lateral lines,

and Opisthognathus differs superficially in many ways from Latilus.

Cepola, not mentioned by Boulenger, must lie near this group as a

distinct family, Cepolide. Pinguipes differs from all this in lacking

the supraocular lamina. It forms a distinct family, Pinguipedide.

Dni

Jordan’ on Distribution of Fishes. — In Science, Dr. Jordan has

an extended account of the origin of the fish fauna of Japan, with

deductions of general application from the relation of the Japanese

fauna to those of other regions.

He finds no evidence from the fishes of a direct connection between

Japan and the Mediterranean, and no evidence of the submergence

of the Isthmus of Suez. In the large identity of genera, and the

divergence of species on the two sides of the Isthmus of Panama,

he finds evidence of former submergence (perhaps Miocene) but

none during the lifetime of the present species. He does not find

in the wide distribution of the Antarctic fresh-water troutlike genus,

Galaxias, certain evidence of the former union of South America

and Australia in Antarctic Continent, but would accept the theory

on geological evidence. i

No. 422. NOTES AND LITERATURE. 163

In a note in a subsequent number of Science, Dr. A. E. Ortmann

claims the existence of adequate geological evidence of the former

extension of the continent “ Antarctica.” In this case the distribu-

tion of Galaxias would be easily explained, but it could be conceiv-

ably explained without it. Dr. Ortmann notes also evidence of the

faunal union of Japan with Europe when the climate of Siberia was

much warmer than now. This evidence is drawn from the distribu-

tion of Crustacea. The distribution of the fishes does not, however,

yield evidence of this kind. D. S. J.

Jordan and Snyder on the Puffing Fishes of Japan. — In the

Proceedings of the United States National Museum, Jordan and Snyder

continue their monographic reviews of the fishes of Japan, treating

of the gymnodont fishes, or puffers. Twenty-seven species are

described, belonging to eight genera. Four new species are figured,

besides several previously known. The authors unite the genus

Lagocephalus with Spheroides, finding a continuous series from one

extreme to the other. In like manner, Ovoides is merged into

Tetraodon. ` ' DEI

Kerr on the Paired Limbs of Vertebrates. — In the Proceedings

of the Cambridge Philosophical Society, Mr. Kerr discusses the ques-

tion of the origin of the paired limbs in vertebrates. He finds the

view of Balfour and others, that these limbs had their origin in a

lateral fold, without adequate support in fact or in theoretical

considerations. :

The view of Gegenbaur, that they arose from modification of the

gill septa separating gill slits, he also criticises unfavorably.

As a provisional hypothesis he brings forward the theory, already

foreshadowed by others, that the vertebrate limbs are modified exter-

nal gills. The close association of the fore limbs and gills shown in

Mr. Kerr's plates of the young Lepidosiren, in another paper, seems

to lend color to this theory. RSI

Notes on Fishes. — Professor Alfredo Dugès. of Guanajuato has

recently sent a bottle of little fishes taken in the very hot spring at

Ixtlan, in the northwestern part of the Mexican state of Michoacan.

These belong to the species described by Woolman as Gambusia

infans. Itis a valid species, distinguished by its small size and plain

color among other things, but the original description 1s at fault in

164 THE AMERICAN NATURALIST. [VoL. XXXVI.

placing the beginning of the dorsal opposite that of the anal. As

usual in this group, the dorsal is considerably beyond the front of

the anal fin. i$]

In the American Naturalist for March, 1901, I published a note in

regard to the planting of fish in Crater Lake, Oregon, in the summer

of 1900, by the Rev. Edwin Sidney Williams of Saratoga, Cal., who

was of the opinion that fish had not previously existed in the lake.

Mr. J. S. Diller of the United States Geological Survey has

recently informed me that he visited the lake in July last and saw in

it a number of fish, ranging in size from six to thirty inches, the

largest ones in many cases being white upon the back or other parts

of the body as if diseased, and on this account being readily seen.

The smaller ones were in good condition. The fish were spotted like

the large trout of the Klamath Lake region. None of them were

captured. Mr. Diller had no difficulty in getting them to take grass-

hoppers or white pumice when thrown into the water.

From the large size of some of these fishes it would appear that

they existed in the lake prior to Mr. Williams's visit. They were

doubtless results of some earlier plant from the Klamath.

i D. S. J.

About March ro, 1897, the State Fish Commission of California,

through Mr. Norman B. Schofield, assistant, planted 855,000 young

fry of the quinnat salmon in Paper Mill Creek, the chief tributary of

Tomales Bay.

As this stream has never contained any salmon, and is open to

observation for its length of twenty-five miles, this operation gave

especially good opportunities for the observation of the young fish.

They soon dropped down from the stream in which they were

planted, tail first, salmon fashion, and in forty-five days were found

in considerable numbers in brackish water. Some of the young

salmon were taken in April at Marshall, fifteen miles down the bay.

In June they disappeared entirely.

The next year, 1898, two million additional salmon fry were placed

in Paper Mill Creek.

Recently, about Nov. 1, 1901, four and a half years after the first

planting, an adult male quinnat salmon weighing about seventeen

pounds was sent to me from Mr. A. D. Hochfort of Point Reyes,

as one of a large run of strange fish seen for the first time in Paper

Mill Creek. IT DSI

No. 422.] NOTES AND LITERATURE. 165

BOTANY.

Chamberlain's ** Methods in Plant Histology.’’ !— The thin vol-

ume at hand is based on a series of elementary articles in the /ournal

of Applied Microscopy dealing with methods of studying the finer

structure of plants. It is, therefore, a discussion of methods rather

than a treatise on plant histology. Chapters dealing with reagents,

methods of killing and fixing, staining, sectioning, and mounting

deal with the processes named. The directions are given in a clear,

straightforward style, and numerical data appear where most desirable.

The preparation of reagents is described fully enough, and the

absence of citations of indefinite quantities and time limits, which

often renders useful books less useful than they might be, is

commendable.

'The second part deals with the most useful methods of preparing

material, from the lowest to the highest groups of plants, and special

methods and devices for difficult types of plant life are described.

Some of the main features to be looked for are indicated, mainly as

a means of judging of the success of the preparation.

Although in no sense a complete work, one rendering Lee's Vade

Mecum or Zimmermann's Microtechnique less necessary, most teachers

of botany not themselves primarily histologists will find this volume

a very useful addition to their shelf of ready reference books.

KH E

Notes. — Very substantial evidence of activity in the botanical

laboratory of the Imperial University of Tokyo, Japan, under the

direction of Dr. M. Miyoshi, professor of botany, is at hand in the

shape of several papers published in the Journal of the College of

Science during the current year.

K. Saito? presents an anatomical study of the most important

plant fibers of Japan, giving attention especially to those derived

from bast cells. The shape, dimensions, markings, contents, and

wall structure are indicated, and the reactions of the walls to the

most important reagents stated. Several points of interest bearing

on the developmental history of certain of these fibers have been

1 Chamberlain, Charles J. Methods im Plant Histology. Chicago, University

g ; eed gens ee E "Studien über wichtige Faserpflanzen Japans mit

besonderer Berücksichtigung der Bastzellen, Journ. Coll. of Sci, Imp. Univ. Tokyo,

vol. xv, pt. iii (1901 x pp- 395-462. 2 pls.

166 THE AMERICAN NATURALIST. |. [Vor. XXXVI.

worked out. The work closes with a key for the microscopical

identification of the fibers of Japan.

S. Kusano? reports investigations made to determine the amount

of transpiration taking place in evergreen trees of Japan in winter.

He finds that the giving off of water from the foliage at no time

actually ceases, even though the minimum temperature (in Hondo)

falls to a point several degrees below zero. The time of least trans-

piration is, however, found to coincide with this minimum, falling

in the month of January. Since photosynthetic activity has been

shown by Miyake not to come to a complete standstill in winter in

the latitude concerned, Kusano concludes that the abundance of

evergreen trees in Japan is chiefly due to its favorable climate.

H. Hattori’ has studied the action of copper sulphate on certain

plants during considerable periods and agrees with those who have

previously investigated the effects produced by copper salts in finding

that it is extremely toxic in its action. Amputated conifer twigs,

seedlings, pot plants, and molds were used in his experiments.

Little that is entirely new has been developed, but a number of

interesting things are reported. Among others is the fact that

copper sulphate in a solution containing 0,000,001 per cent of the

salt is not harmful to corn seedlings cultivated in it for considerable

periods. The capacity of the soil for fixing considerable quantities

of salt supplied to it in solution accounts for the greater amount of

copper endured by plants grown in pots of earth. Molds grown in

copper containing culture media were found to be stimulated by

minimal amounts of the metal, 0,004 per cent to 0,008 per cent

being indicated for Aspergillus and Penicillium, respectively.

T. Inui? has studied the lower plant organisms connected with the

preparation of awamori, an alcoholic, whisky-like beverage brewed

in the Luchu Islands (situated between Formosa and the Kiushu

Islands).

Boiled rice or millet is inoculated from material of former cultures

and, after sufficient growth has been made first on the grain in a

! Kusano, S. Transpiration of Evergreen Trees in Winter, Journ. Coll. of Sci.,

imp. iq ere vol. xv, pt. iii (1901), pp. 313-366. 1 pl.

? Hat H. Studien über die Einwirkung des Kupfersulfats auf einige

PUn telis Coll. of Sci, Imp. Univ. Tokyo, vol. xv, pt. iii (1901), pp. 371-

3094 TN.

.$ Inui, T. ANIM. über die niederen Oryininnes welche sich bei der

;Zebostung des iibubclidus Getiankes “ Awamori” betheiligen, Journ. Col.

4f N me ioa vol. mae iii (1891), pp. 465-478. 1 pl.

No. 422.] NOTES AND LITERATURE. 167

moist atmosphere and afterwards in a more watery preparation, the

alcoholic product is obtained by distillation. The author has isolated

several new forms from the material used, — Aspergillus luchuensis,

related to Aspergillus wentii Wehmer; Aspergillus perniciosus, near

the latter; and Saccharomyces awamort. He reports the presence of

a species of Monilia and of Saccharomyces anomalus, which latter

form produces the characteristic aroma.

Professor Miyoshi * makes a second contribution to his study of a

mulberry disease called “‘ Ishikubyo,” and comes to most interesting

conclusions. The injury is due to a lack of proper balance in the

tissue development of the plant. The leaves manufacture more

carbohydrates than the conducting system is able to withdraw dur-

ing the usual time. The woody tisSue of affected plants in all parts

is also underdeveloped, and a diminished water transpiration results.

This account is the more interesting since physiological derangements

are not often traced back to ill-balanced tissue formations.

In this connection it may be noted that Suzuki, investigating this

disease in the light of Woods's work on the “ mosaic disease " of the

tobacco plant, has come to the conclusion that in the mulberry, as in

the tobacco, the trouble is due to an overproduction of oxydizing

enzymes accompanied by a more or less advanced state of starvation.

Another article by Professor Miyoshi? reveals an extremely perfect

means for securing spore dissemination in a new tree-inhabiting

Japanese lichen named by the author Sagedia macrospora. The

rounded sporocarps lie imbedded in the thallus until ripe, when,

through the weakening of the tissues surrounding them, they separate

from the thallus as free rounded masses. On absorbing water, the

sporocarp expands and opens out by means of the hygroscopic prop-

erties of the walls, exposing the hymenium. The spore-containing

sacs rupture, thus freeing the spores. The fixation of the free

sporocarp to the bark of the tree is secured by the sticky outer sur-

face. The sporocarps by repeated moistening and drying can ‘be

made to open and close many times, even after killing the living

parts by heat or other means.

n über die Schrumpfkrankheit (“ Ishikubyo”)

1 Miyoshi, M. Untersuchunge t

: Journ. Coll. of Sci, Imp. Univ. Tokyo, vol. xv,

des Maulbeerbaumes. II. Bericht.

pt. iii (1901), pp. 459-464- | E

? Miyosh : : "M. : Ueber die Sporocarpenevacuation und darauf erfolgendes

usstreuen bei einer Flechte, Journ. Coll. of Sci, Imp. Univ. Tokyo, vol. xv,

pt. iii (1901), pp. 367-370. 1 pl.

168 THE AMERICAN NATURALIST. (VoL. XXXVI.

Dr. Robinson’s admirable presidential address before the Botanical

Society of America, on * Problems and Possibilities of Systematic

Botany," is printed in Science of September 27.

In the Annals of Botany for September, A. H. Church suggests

that the substitution of a logarithmic spiral in place of the Archime-

dian spiral for the genetic and secondary spirals in phyllotactic studies

refers this complex subject to a simple relation to the distribution of

energy in the growing points.

Two additional works by Rafinesque — a Florula Lexingtoniensis

and Zhe American Florist — have been unearthed by W. J. Fox in

the library of the Academy of Natural Sciences of Philadelphia,

as appears from a note by him in Science of September 27.

An interesting addition to the literature of myrmecophilous plants

is a paper presented at the meeting of the British Association last

summer by Yapp. The ant galleries of two Malayan species of Poly-

podium are described. An abstract of the paper appears in Mature

for October 17.

Papers on the fruit and seed structure (by Lonay) and leaf anatomy

(by Goffart) of Ranunculacez are contained in the Mémoires de la

Société Royale des Sciences de Liége, Vol. III, issued in July.

From a study of the anatomy and embryogeny of Nelumbo, H. L.

Lyon concludes, in the current volume of Minnesota Botanical Studies,

that the Nymphzacez should be transferred to the monocotyledonous

group Helobiz.

Robinia neomexicana is figured in the Tidskrift of the Svenska

Tradgaardsforeningen for July-August.

Numerous popular varieties of Begonia semperflorens are figured

from photographs in Die Gartenwelt of September 28.

Several Canadian gentians of the section Crossopetale are

reviewed by Holm in the Ottawa Naturalist for November.

In Part II of Vol. VII of the Contributions from the United States

National Herbarium, Mr. O. F. Cook adduces reasons for believing

in the American origin of Cocos nucifera.

Scirpus supinus and its North American allies are reviewed by

Fernald in Rhodora for October.

Mr. Eaton’s ninth paper on Equisetum, in the Fern Buletin for

October, deals with E. litorale.

No. 422.] NOTES AND LITERATURE. 169

George H. Curtis has several lists of Kansas diatoms in the Zran-

sactions of the Kansas Academy of Science for 1899—1900.

The flora of the Azores receives an important addition through the

publication in Vyt Magazin for Naturvidenskaberne of a paper on the

fresh-water diatoms of the archipelago, by Holmboe. The species

are European rather than American, and are believed to have been

introduced by adhering to migratory birds.

Professor Hitchcock publishes a list of additions to his * List of

Plants in my Florida Herbarium" in Vol. XVII of the Zransactions

of the Kansas Academy of Science.

Native Kansas plants adapted to cultivation are discussed by

Grace R. Meeker in Vol. XVII of the Zransactions of the Kansas

Academy of Science.

The Journal of Applied Microscopy for October contains an illus-

trated article on the botanical laboratory and garden of the Imperial

University at Tokyo, by Miyake.

An excellent portrait of the late Professor Cornu accompanies the

September number of the Bulletin de la Société Botanique de France.

The Fern Bulletin for October contains a portrait of Thomas

Meehan.

Trabut begins an illustrated article on caprification as practiced in

Algeria, in the Revue Horticole of November 9.

A Flora of the Presidency of Bombay, by Theodore Cooke, is in

course of publication by Taylor & Francis of London. The familiar

sequence of orders of Bentham and Hooker is followed, and the

single part thus far issued reaches into Rutacez.

An extensive report on a botanical survey of the Dismal Swamp

region, by Thomas H. Kearney, forms the concluding number of

Vol. V of the Contributions from the United States National Herba-

rium, issued under date of November 6.

The plants of western Lake Erie are considered as to their eco-

logical anatomy and distribution by A. J. Pieters in a paper sepa-

rately printed from the Bulletin of the United States Fish Commission

for r9or.

Vol. III of Professor Greene's Plante Bakeriame begins with a

fascicle of thirty-six pages devoted almost exclusively to the eme

tion of new species from Mr. Baker’s collections of 1901, on the

Gunnison watershed of Colorado.

170 THE AMERICAN NATURALIST. [VoL. XXXVI.

Four miscellaneous phanerogamous species from Colorado are

described by Osterhout in the Bulletin of the Torrey Botanical Club

for November.

Notes on the vegetation of the Cape Nome region, Alaska, by

Collier, are included in the account of the Brooks reconnaissance of

Seward Peninsula, recently published by the United States Geological

Survey.

A monograph of the genus Sorbus, by Hedlund, constitutes No. 1

of Vol. XXXV of the Kongliga Svenska Vetenskaps-Akademiens Hand-

lingar.

The volume of Botaniska Notiser for 1901 contains (pp. 33-72,

83-106, 155—158) a discussion by Hedlund of the forms of Aes

rubrum in the broad sense.

Under the name of Pilea heptaphyllus, Dr. Rarmíez describes and

figures a representative of the Papayacez, in No. 1 of the current

volume of Anales del Instituto Médico Nacional, of the city of Mexico.

An article on the preparatory fermentation of cacao, by Preyer,

published in the Z»openpflanzer for April, is reprinted in transla-

tion in No. ro of the Boletin del Instituto Fisico-Geografico de Costa

Rica.

Various diseases of cacao and sugar cane and bud variation of the

latter are discussed in No. 3 of the current volume of the West Indian

Bulletin, of Barbados.

The chayote, Sechium edule, and its varieties are the subject of

Bulletin No. 28 of the Division of Botany of the United States

Department of Agriculture, by Mr. Cook, special agent for tropical

agriculture.

“Plant Breeding,” an illustrated essay by Professor Hays of

Minnesota, is published as Buletin No. 29 of the Division of

Vegetable Physiology and Pathology of the United States Depart-

‘ment of Agriculture.

Economic studies of Rhizophora mangle, Avicennia nitida, and

Eriodendron occidentalis are published in No. 2 of the current volume

of Anales del Instituto Médico Nacional, of Mexico, which also con-

tains an article, by Martinez del I Campa e on plants used in that

country as dum mor

: A bot hlet on the he Howe and the manufacture

dis of rubber from them in Brazil, pte M Me decise br

No. 422.] NOTES AND LITERATURE. 171

Sr. J. Barbosa Rodrigues, director of the botanic garden of Rio de

Janeiro, under the title 4s ZZeveas ou Seringueiras.

In No. 1197, “ Advance Sheets," of Consular Reports is reprinted

from the Venezuelan Herald an article by Dr. L. Morisse on rubber

culture and manufacture in Venezuela from species of Hevea.

Dr. Peckolt is publishing a series of articles on the medicinal

plants of Brazil in the Berichte of the German Pharmaceutical

Society, and an alphabetic list of Portuguese and Tupi names for the

economic plants of the same country in the Pharmaceutical Archives.

Stem and bark anatomical studies of Hamamelis (by Jensen) and

Myrica (by Krembs) are published in the Pharmaceutical Archives

for July.

From No. 3 of the Bulletin du Jardin Impérial Botanique de

St.-Pétersbourg it appears that 33,697 species and varieties of plants

were cultivated in that establishment last year. Some of the princi-

pal groups grown under glass are represented as follows : ferns, 798 ;

orchids, 1433; cacti, 748; palms, 402; cycads, 60; conifers, 567;

heaths, 186; aroids, 585; bromeliads, 420; succulents, chiefly agaves

and aloes, 991. The plants grown in the open air comprise: trees

and shrubs, 1240 species and varieties; perennials, 4385; and annu-

als, 1410. Over 37,000 persons visited the plant houses during the

year.

It may not be generally known that an account of the government

gardening in the District of Columbia is each year included in the

Report of the Chief of Engineers. The volume for rgoo contains a

list of the woody plants cultivated about the White House.

Sr. Rodrigues, director of the botanic garden of Rio de Janeiro,

has begun the publication of a series of French Contributions du

Jardin Botanique de Rio de Janeiro in quarto pamphlet form.

The tendency of some German investigators to confine their

examination of the literature of subjects they investigate to publica-

tions by their fellow-countrymen is not inopportunely, if somewhat

caustically, commented upon by Dr. MacDougal in Torreya for

November. :

An excellent portrait of the late Thomas Meehan is published sae

Meehans’ Monthly for December, and a biogr aphic sketch in the

January number. |

Vol. XXVI of the botanical Bihang till Kongliga Svenska Vetenskaps-

Akademiens Handlingar, forming a thick volume, contains a wide

172 THE AMERICAN NATURALIST.

range of subjects, handled in the characteristically excellent Scandi-

navian fashion.

The Ohio Naturalist for November contains the following botanical

articles: Tyler, * Geophilous Plants of Ohio," II ; Kellerman, * Ohio

Fungi Exsiccatz," with reprint of original descriptions ; Schaffner,

“The Maximum Height of Plants," III.

Notes on the geotropism of fungus stipes, extrusion of the gametes

of Fucus, and adaptation of Spartina polystachya to environment are

published by E. B. Copeland in Torreya for November, in an account

of the last season’s work at the Cold Spring Harbor laboratory.

In Part II of the current volume of Proceedings of the Academy of

Natural Sciences of Philadelphia, Dr. Harshberger discusses the limits

of variation in plants, and Mr. Meehan contributes some observa-

tions on the upbending of mature wood in trees.

The anatomy of the conducting tissue of style and stigma is

being considered by Guéguen in current numbers of the Journal

de Botanique.

Eleanor E. Davie has compiled from the writings of the late

W. H. Gibson an attractive little book on Blossom Hosts and Insect

Guests, which, fully illustrated by the author’s admirable drawings, is

published by Newson & Co. of New York.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS, AND DEATHS.

EDUCATIONAL GIFTS.

Akron (Ohio) Public Library, $70,000, conditional from Andrew Carnegie.

Alfred University, $1000, by the will of E. P. Barker.

Amherst College, $1000, by the will of E. P. Barker.

Barnard College, Columbia University, a conditional gift of $200,000, from

John D. Rockefeller.

Berea (Kentucky) College, $25,000, by the will of Mrs. George L. Stearns.

Butler (Indiana) College, $20,000 from Mr. and Mrs. E. C. Thompson of

Indianapolis.

Bryn Mawr College, a conditional gift of $230,000, from J. D. Rockefeller.

Carnegie Institution, of Washington, D.C., $10,000,000 from Andrew

negie.

Carnegie Polytechnic Institute, of Pittsburg, $1,000,000, from Andrew

Chicago University, $1,250,000, from John D. Rockefeller.

Columbia University, $12,000, from Dean Lung, for the Chinese depart-

ment.

Drury (Missouri) College, $8000, from E. A. Goodnow of Worcester,

. Mass.

Harvard University, nearly $100,000, from an anonymous donor, for a new

building; $5000 for the Peabody Museum of Archzology, by the will

of Mrs. S. D. Warren ; $50,000, from Mr. T. Jefferson Coolidge, for

physical research.

Kenyon College, $150,000, from various donors.

Manchester (New Hampshire) Institute of Arts and Sciences, $80,000, by

the will of Mrs. Charles E. Balch of that city.

Metropolitan Museum of Art in New York, over $5,000,000,

Jacob S. Rogers.

Montana Biological Station, $250, from Senator W. A. Clark.

Nashville, Tennessee, $100,000, from Andrew Carnegie, for a public library.

New York Botanical Garden, $3000, from Misses Oliva and Caroline

Phelps Stokes. d

New York Historical Society, $130.000. by the will of Miss C. B.

De P ;

New York University, $20,200, from various persons, for a fellowship ;

$5000, from Miss Helen M. Gould, for a museum of pedagogy.

173

by the will of

174 THE AMERICAN NATURALIST. [VoL. XXXVI.

Northwestern University, $15,000, from an anonymous donor.

Oberlin College, $300,000, from various sources, thus securing the condi-

tional gift of $200,000 from J. D. Rockefeller.

Palmer (Iowa) College, $30,000, from Mr. F. A. Palmer; $20,000, from

_ other sources.

Princeton University, $2500 a year, for five years, from George A. Armour,

for the classical seminary ; $15,000, from the estate of John Sayre.

San Juan, Puerto Rico, $100,000, from Andrew Carnegie, for a public

library.

Sheffield Scientific School of Yale University, $6000, from Edward B. Page,

to found scholarships.

Springfield (Missouri) Public Library, $50,000, from Andrew Carnegie.

Syracuse University, $4000, from Mr. George C. French.

Tufts College, $50,000, and land valued at $40,000, by the will of Mrs.

George L. Stearns.

Tuskegee Institute, $25,000, by the will of Mrs. George L. Stearns.

Union Christian (Indiana) College, $30,000, from Mr. A. B. Palmer of

ew York.

University of Pennsylvania, $25,000, from Messrs. Keasbey and Matteson ;

$5000, from John F. Wentz ; $10,000, from the Frazer family ; $20,000,

from Dr. George Woodward, for a fellowship in physiological chemistry.

Vassar College, $30,000, from Miss Helen M. Gould, for scholarships ;

$110,000, from J. D. Rockefeller, for a new dormitory ; $2000, from

other sources.

Washington University, $100,000, from Mrs. James F inney How.

Wesleyan University, $125,000, from various donors for buildings.

Williams College, $5000, by the will of Mrs. S. D. Warren.

Worcester Polytechnic Institute, $30,000, from four trustees: S. Salisbury,

C. H. Whitcomb, C. H. Morgan, and C. G. Washburn.

Yale University, $2500, from Mr. B. F. Barge.

APPOINTMENTS.

Wilhelm Arnell, lector in natural history in the University at Upsala. —

Archibald A. Atkinson, assistant instructor in biology in the University of

Oregon. — Dr. Georg Bitter, docent for botany in the Academy at Münster.

— Prof. M. Büsgen, professor of botany in the school of forestry at Han-

nover-Münden. — Dr. Carl Busz, professor of mineralogy in the Academy

at Münster.— Dr. M. Caullery, professor of zoólogy at Aix-Marseilles.

— H. S. Davis, instructor in vertebrate zoólogy in the Washington Agricul-

tural Experiment Station. — Frank S. Earle, assistant curator of the col-

lection of fungi of the New York Botanical Garden. — Dr. Géza Entz,

professor of zoólogy in the University at Budapest. — Dr. J. M. Flint, pro-

fessor of anatomy in the University of California. — Willy Foy, director of

‘aed 1

Cologne. — Dr. Elisha H. Gregory, Jr.,

No. 422.] GIFTS, APPOINTMENTS, RETIREMENTS. 175

demonstrator of anatomy in the University of Pennsylvania. — W. Hammer,

assistant in the Austrian Geological Survey. — Dr. Charles M. Hazen, pro-

fessor of biology in Richmond (Virginia) College. — Mr. J. M. Hillier,

keeper of the Museum of Economic Botany at Kew. — Mr. H. T. A. Hus,

assistant in botany in the University of Amsterdam. — Henry M. Huxley,

Hemenway fellow and assistant in anthropology in Harvard University.—

J. T. Jenkins, lecturer on biology and geology in Hartley College, South-

ampton, England. — Dr. H. Joseph, docent for zoólogy and comparative anat-

omy in the University at Vienna. — S. Killermann, professor extraordinary of

anthropology, zoólogy, and botany in the episcopal school at Ratisbon. —

Dr. E. H. Kraus, instructor in mineralogy in Syracuse University. — Gustav

Krause, district geologist of the Prussian Geological Survey. — Dr. A. A.

Lawson, assistant in botany in Leland Stanford Junior University. — Dr.

A. B. Macallum, professor of physiology in the University of Toronto. —

Dr. A. Pelikan, professor of mineralogy and petrography in the German

University at Prag. — Dr. R. G. Perkins, lecturer on bacteriology in the

medical school of Western Reserve University.— J. M. Prather, instructor in

biology in the University of Cincinnati. — Dr. K. Theodor Preuss, director's

assistant in the royal museum at Berlin. — William Riley, instructor in

entomology in Cornell University. — Dr. V. Schiffner, professor extraordi-

nary of botany in the University at Munich. — Dr. Conrad von Seelhorst,

professor of botany in the University at Góttingen. — Dr. Hans Solederer,

professor of botany in the University at Erlangen. — Dr. Julius Stoklasa, pro-

fessor of plant production in the Bohemian technical school at Prag. —

Dr. E. Stolley, professor of geology in the Braunschweig Technical

School.— Dr. Richard P. Strong, director of the government biological

laboratory in Manila. — Dr. Roland Thaxter, professor of cryptogamic

botany in Harvard University. — Dr. Victor Uhlig, professor of geology in

the University at Vienna. — Dr. F. C. Waite, assistant professor of histology

and embryology in the medical school of Western Reserve University. —

Dr. Thomas L. Walker, professor of mineralogy and petrography in the

N. Whitford, assistant in botany in the University of Chicago. —

H. Winckler, first assistant in the Botanical Garden at Breslau. — Dr. Hans

Winkler, docent for botany in the University at Tübingen. — Dr. A. Smith

Woodward, keeper of geology in the British Museum. — Oscar Zeise, land

geologist of the Prussian Geological Survey.

RETIREMENTS.

J. R. Jackson, from the keepership of the Museum of Economic Botany

at Kew, after forty-three years of service. — George Nicholson, from the

I 76 THE AMERICAN NATURALIST. [Vor. XXXVI.

the Museum at Demerara, British Guiana. — Dr. Henry Woodward, from

his position as keeper of geology in the British Museum. — Dr. John Young,

from the chair of natural history in Glasgow, after thirty-five years of

service.

DEATHS.

Lugui Maria D’Albertis, the ornithologist and explorer, in Sassari, early

in September. — Dr. F. Arnold, lichenologist, in Munich, August 8, aged

73. — F. J. Birtwell, ornithologist, by an accident, in New Mexico, June 28.

— David Carnegie, botanist and explorer, on the Middle Niger, Nov. 27,

1900, aged 30.— Albert Nelson Cheney, ichthyologist, at Glens Falls,

N.Y., August 18. — Dr. J. H. Chievetz, director of the Anatomical Mu-

seum at Copenhagen. — Professor Miguel Colmeiro, director of the botanical

gardens at Madrid, aged 86. — Dr. Carl Cramer, professor of botany in the

Polytechnicum at Zürich. — William Doherty, entomological and ornitho-

logical collector in Nairoba, East Africa, May 25. — Dr. James Foulis,

anatomist, in Edinburgh, October 17.— Prof. Dr. Robert Hartig of

Munich forestry station, October to, aged 62.— Prof. Alfonso Herrera.

in Mexico, Jan. 27, 1901, aged 67. — Dr. Federico Horstman y Cantos,

professor of anatomy in the University of Havana. — Dr. Georg Jablo-

nowski, assistant in the Anatomical Institute in Berlin, September 28. —

Josef Bernhard Juch, cryptogamic botanist, in Constance, August 14,

aged 83. — Clarence King, director of the United States Geological Sur-

vey, 1878-1881, December 24, at Phoenix, Arizona. — James Walker Kirkby,

geologist and student of fossil ostracodes, in Levin, Scotland, July 30,

aged 66. — Dr. Arthur Kónig, professor extraordinary of physiology in the

University at Berlin, October 26, aged 45.— Prof. A. A. Kowalevski,

formerly professor of zoólogy in the University at St. Petersburg, Novem-

22.— Dr. Albrecht von Krafft of the Indian Geological Survey in

Allahabad, in September. — Jacob Heinrich Krelage, a Dutch botanist,

December 1, aged 76. — Henri Lacaze-Duthiers, the eminent French zoólo-

gist, July 21. — L. Liener, botanist in Constance, in May. — Rev. Hugh

Alexander Macpherson, a Scotch zoólogist, aged 43. — Prof. M. Marcher,

director of the Agricultural Experiment Station at Halle, Oct. 19, 1901. —

Mr. Thomas Meehan, the well-known botanist, at Germantown, Pa.,

November 29, aged 75. — Charles F. Mohr, botanist in Asheville, N.C

July 17. — Dr. H. Müggenburg, dipterologist in the Zoólogical Museum in

Berlin, July 3. — Count Emil Neubauss, student of Lepidoptera, April 21,

aged 57. — Henri Philibert, professor of botany in Aix, May 14, aged 79.

— Dr. Max Reess, formerly professor of botany in the University at

Erlangen, in Klingenmünster, September 14, aged 56.— Louis Schneider,

botanist and entomologist, in Philadelphia, aged 65. — Dr. L. Serrurier,

formerly director of the ethnological museum in Leiden, in Batavia,

July 7. — Gene B. Simpson, student of fossil Polyzoa and a well-known

paleontological artist, at Albany, N.Y., October 15. — Dr. Henry Spencer

No. 422.] GIFTS; APPOINTMENTS, RETIREMENTS. 177

Smith, of London, October 29, aged 88. — Hermann Strecker, entomologist,

at Reading, Pa., November 30, aged 65. — Dr. Peter Cormack Sutherland,

formerly government geologist of Natal, in Durban, Nov. 30, 1900,

aged 79. — Ralph Tate, professor of natural history in the University at

Adelaide, Australia. — Dr. Wilhelm Tomaschek, professor of geography in

the University at Vienna, September 9, aged 60. — Dr. A. A. Tokasy, head

of the physiological laboratory at Moscow. — William West, student of

fresh-water algae, in India, of cholera, aged 26. — Lionel L. Wigglesworth,

ornithologist, in Viti Levu, June 7, aged 37.

PUBLICATIONS RECEIVED.

(Regular exchanges are not included.)

BARTHOLOMEW, E. Ellis and Everhart’s “ Fungi Columbiani": Alphabetical

Index, Centuries I-X V. — BEECHER, C. E. Studies in Évolution: Y

Charles Scribner's Sons, 19or. xxii +638 pp., 34 pls. — CHURCH, A. H.

the Relation of Plyllotasjs to Mechanical Laws. Pt. i, Construction by Orthog-

onal Trajectories. Oxford, Williams & Norgate. 8vo, 78 pp., 10 pls., 34 text-

figs. — CoLE, G. W. Bermuda and the Challenger Expedition. Printed for

private distribution. Boston, 190r. 16 pp. — COMSTOCK, J. H., and KELLOGG,

The Elements of Insect Anatomy: an Outline for the Dee of Students in

Entomological Laboratories. Third edition, revised. Comstock Publishing Co.,

Ithaca, N.Y., 1901. 145 pp. II figs.— GErTsINGER, E. C. A New Theory of

Biology. Reprinted from the Medical Times, November, 1901. 17 pp.— Gov-

ernment Museum and Connemara Public Library: Report on the Administration

Jer the Year r900—rgor. — MoRGAN, T. H. Regeneration. Columbia University

Biological Series, No. 7. New York, The Macmillan Company, 1901. xii+316

66 figs. $3.00. — United States Geological Survey. Twenty-first Annual onic

ae Pt. i, xi+608 pp.; pt. vi, 656 pp.; pt. vi (continued), xi4-634 pp.

ALLEN, J. A. The Proper Generic Names of the Viscacha, TE and

their Allies. Proc. Biol. Soc., Washington. Vol. xiv, pp. 181— 182. — ALLEN, J. A

Note on the Names of a Few South American Mammals. Proc. Biol. Soc.,

Washington. Vol. xiv, pp. 183-185. — BAKER, F. C. Some Interesting Molluscan

oree Trans. Acad. Sci., St. Louis. _ Vol. xi, No. 8, pp. py Pl. XL

KS,

Wash. Acad. Sci. Vol. iii, PP- 541-546. — BLASDALE, W. C. nne to

the Mineralogy of California. Buz. Dept. Geol., Univ. of Cal. Vol. ü, pp. 327-

348. — BUCKLEY, E. R. The Clays and Clay Industries of Wisconsin. Bull.

Wis. Geol. and Nat. Hist. Surv., No. 7, pt. i. Economic Series No. 4. — CHAP-

Vol. i, No. 11, October, 1901. 24 pp., numerous figs. — Cor, W. R. The Nemer-

576, text-figs. 62-66. — DALL, W. H., and Simpson, C. T. The Mollusca of

Porto Rico. Bull. U.S. Fish Com. for rgoo. Vol. i, Pp. 351-524, Pls. LIII-

LVIII. — ÉAKLE, A. S. Mineralogical Notes. Bull. De ept. Geol., Univ. of Cal.

Vol. ii, pp. 31 5-326, Pl. IX. — EIGENMANN, C. H. Description of a New Oceanic

Fish found off Southern New England. Contributions from the Biological

Laboratory, U.S. Fish Com, Woods Hole, Mass. — EIGENMANN, Cn Inves-

— into the History of the Young Squeteague. Bull. U.S. Fish Com. for

Ear Bana. een M aem, ME rubei

PUBLICATIONS RECEIVED. 179

Lake, Montana, s Ragged Mee 1899. Trans. Amer. Micr. Soc. Vol. s

pp. 63-8o, Pls. X-XVII. 1901.—— FELT, E. P. Scale Insects of Im

tance and List of the nif in eia Vork State. Bull. New York State Vend

No. 46. Pp. 291-377. 15pls. — GRANT, U. S. eeen Report on the Copper-

Bearing Rocks of Douglas County, Wisconsin. E Wis. Geol. and Nat. Hist.

Surv., No. 6 (second edition). Economic Series No. 3. — Harcrrt, C. W.,

edo C. G. The Alcyonaria of Porto Rico. pet U..S. Fish Com. Pa 1900.

Vol. ii, pp. 265-287, Pls. I-IV. — HUNTER, S. J. On the Production of Artificial

Parthenogenesis in Arbacia by the Use of Sea Water concentrated by Evapora-

tion. Amer. Journ. of Phys. Nol. vi, pp. 177-180. November, 1901. — IKEDA, I.

Observations on the Development, Structure, and Metamorphosis of Actino-

trocha. Journ. Coll. Sci., Imp. Univ. Tokyo. Vol. xiii, pp. 507-592, Pls. XXV-

XXX. 1901.— JENKINS, O. P. Descriptions of Fifteen New Species of Fishes

from the Hawaiian Islands. Zu. U.S. Fish Com. for 1899. Pp. 387-404. 1901.

— JORDAN, D. S., and Snyper, D. O. A Review of the Gymnodont Fishes of

Japan. Proc. U.S. Nat. Mus. Vol. xxiv, pp. 229-264.— Linton, E. Parasites

of Fishes of the Woods Hole Region. Bull. U.S. Fish Com. for 1899. Pp. 405

492, Pls. I-XXXIV. 19or.— MERRIAM, C. H. Description of Twenty-three

New Harvest Mice (Genus PEE Se Proc. Wash. Acad. Sci. Vol. iii,

Pp. 547-558. 1901.— MERRIAM, C. H. n New Mammals from Mexico,

including a New Genus of Rodents. Proc. Wak Acad. Sci. Vol. iii, pp. 559-

563. 1901. — MERRIAM, C. H. Preliminary Revision of the Pumas (Felis concolor

group). Proc. Wash. Acad. Sci. Vol. iii, pp. 577-600. — MoHm, C. Notes on

the Red Cedar. Bull. U.S. Dept. Agr. Dept. Poe No. 31. 37 pp» 13

text-figs., 3 pls. — NEEDHAM, J. G. and BETTEN, C. Aquatic Insects in the

Adirondacks. Bull. New York State Museum, No. 47. 1901. Pp. 383-612. 56 pls.

— OBERHOLSER, H.C. Seven New Birds from Paraguay. Proc. Biol. per Wash-

ington. Vol. xiv, pp. 187-188. — Oscoop, W. A New White-Footed Mouse

from California. Proc. Biol. Soc., Washington. Vol. xiv, pp. 193-194. — PIETERS,

. J. The Plants of Western Lake Erie, with Observations on their Distribution.

Bull. U. S. Fish Com. for 1901. Pp. 57-79, Pls. XI-XX.— Price, O. W. Prac-

tical Forestry in the ii ern Appalachians. Reprint from the Yearbook U.S.

Dept. for rgoo. Pp. 357-368. 6 pls.— RICHARDSON, H. Papers from

the Hopkins Stanford Meta os Expedition, 1898-1899. VI. The Isopods

Proc. Was, ol. iii, pp. 565-568. 190r.— RiDpGWaYv, R. Th

xxxi--715 pp. 20 pls. Washin Papers

from the Harriman Alaska fates XXV. The Alge. Proc. Wash. Acad.

Sci. Vol. iii, pp. 391-486, Pis, XLIII-LXITI November, 1901. — SMITH, H. I.

Bull. U. S. Fi Fish Com. for peas:

Dredging and Other Records of the U. d Fish Coaitalecton Steamer ross, with

Bibliography relative to the Work of the the Vessel. U.S. Fish Com. Report for

1900. Pp. 387-562, Pls. I-VII. — TRELEASE, W. The Progress made in Botany :

during the Nineteenth Century. Trons Acad. Sci., St. Louis. Vol. xi, No. 7s

180 THE AMERICAN NATURALIST.

pp. 125-142. November, 1901. — VAUGHAN, T. W. The Stony Corals of the

Porto Rican Waters. Ju//. U.S. Fish Com. for rgoo. Vol. ii, pp. 289-320,

Pls. I-XXXVIII. — WarLiCH, C. A Method of ings Egg PEREAS for

Use of Fish-Culturists. U.S. Fish Com. Report for 190 Pp. 185-194. 1 pl.—

WHEELER, H. J. Commercial Fertilizers. £u. an Island Agr. Exp. Sta.,

No. 81. Pp. 111-112. October, 1901. — WHITE, D. Two New Species of

Alga of the Genus Buthotrephis in the Upper soci of Indiana. Proc. U.S.

Nat. Mus. Vol. xxiv, pp. 265-270, Pls. XVI-XVII

The American Antiquarian and Oriental Journ Vol. xxiii, No. 6. — The

American Journal of Science. Ser. 4. Vol. x o. 72. December, 1901. — The

American Museum Journal. Nol. i No. tr. ease: 1901. — Anales del Museo

Nacional de Montevideo. Tome ii, pp. 417-492. — Annales de la Société Entomo-

logique de Belgique. Tome xlv, pt. xi.— The Botanical Gazette. Vol. xxxii,

No. November, 1901. — Botantsches Centralblatt. Bd. Ixxxviii, Nos. 5-10. —

Bulletin of the Johns Hopkins Hospital. Vol. xii, No. 12 October, 1901.

Bulletin T v ipd deem Peces Vol. xii, No. 128. — podes of the Torrey

Botanical I November, 19o1. — The Canadian Ento-

mologist. erg xxxiii, ee. 12.— final of the Cincinnati Society of Natural His-

tory. Vol. xx, No.1. 1901. — Linnean Society of New South Wales. Abstract

of € Aug. 2 28, Sept. 25, Oct. 4: 1901. — Nature novitates, 1901. Nos.

—'The Ohio Naturalist. Vol. ii, No. 1.— Proceedings of the Nebraska

Dae t Union at its Second ous Metu I9OI. IOI pp., 10 pls. — Fro-

ceedings of the Philadelphia Academy of Natural Sciences, 1901. Pp. 513-544. —

The Procession, a Magazine of Science and General Interest. Los Angeles.

Vol. i, No. 1. — Revista wd de Historia Natural. Año v, Nos. 8, 9. — Rhode

Island Agricultural Experiment Station. Fourteenth A annad Report. Pt. ii. —

Science Gossip. Vol. viii, nal gt. December, 1901. — Zoologischer Anzeiger.

Bd. xxiv, Nos. 655, 657-658.

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THE

AMERICAN NATURALIST

Vor. XXXVI. March, 1902. No. 423.

A REMARKABLE OCCURRENCE OF THE FLY,

BIBIO FRATERNUS LOEW.

JAMES G. NEEDHAM.

Nor far from my home in Lake Forest there is a forty-acre

patch of woodland pasture that has been enclosed but not

grazed for a number of years. It has become well overgrown

with a thick blue-grass sod. Save for a few young hawthorns

in scattered clumps, there is no underbrush. The open wood

consists mainly of white oaks. Black oaks were originally

abundant but have been cut down, and now the sod is thickly

dotted with their rotting stumps. Around the bases of these

stumps, where the sod closely enwraps them, is the larval home

of Bibio fraternus.

I had seen but few specimens of this fly until the spring of

1901, and was wholly unacquainted with its immature stages

and with its larval habitat. It chanced that on walking across

this pasture in April I kicked over a rotten stump, and, where

a lateral root pulled out from under the sod, there in the fine

black soil that had resulted from the complete decomposition

of the bark were a large number of dipterous larvae, which

later, when reared, yielded imagoes of this species. a |

181 : d

182 THE AMERICAN NATURALIST. [VoL. XXXVI.

Nothing appears to have been written of the species since

Loew penned the original description. In fact, the immature

stages seem not to have been described or figured for any

American species of the family Bibionida. Mr. D. W. Coquil-

lett, who determined my imagoes for me, having assured me

that under the existing circumstances even a slight contribution

to the knowledge of thesé things would be welcomed by dipter-

ologists, T offer heréwith some observations on the habits of

this species and a description of its immature stages.

Habits. — My acquaintance with the habits of the species

began as indicated above. On May 5 I collected a large num-

ber of grown larvz and a few pupz about different stumps and

placed them, with a quantity of their native soil, in a breeding

cage. Imagoes began to appear on the 13th and continued

transforming until the 20th. On the 18th, when my cage was

full of fresh imagoes, I went out to the pasture but could find

none there. In the earth were plenty of pupæ, but no more

larvae. Perhaps the development of those in my cage was

hastened by the higher temperature of the laboratory in which

they were kept. About May 22 the first few imagoes were

observed at large. On the 24th I discovered, on walking across

the pasture, that the grass was full of them. A score could be

swept up at each stroke of the net; but none were flying, for

the air was chill. On the 26th, 28th, and 3oth, visiting the

pasture again I saw the great swarms of them which I now

come to describe.

I was not prepared for any exhibition of aérial activity by

what I had seen in my breeding cage. There the imagoes

seemed hardly able to walk, to say nothing of flying. They

were very sluggish ; they climbed the sides of the cage weakly,

and tumbled about even in walking, and were handled without

any thought of their getting away. Even in the field on the

24th I had picked them up with my fingers and examined them’

unconfined under a lens. But on these three later days warm

sunshine had stirred them to unexpected activity. They were

: flying i in countless numbers in the more sheltered places in

the woods and were actively climbing about in the branches

of Ene ecu MM an aces in pairs on the

No. 423.] . REMARKABLE OCCURRENCE (‘OF THE FLY. 183

leaves... The flight:of the males is long sustáined;: -Both males

and females, and especially the latter, have- some -difficulty-in

getting launched. | They climb to a leaf top, balance carefully,

stretch their wings and try them:before letting go, —

swing free into the air.

I have rarely come upon a scene of greater animation than a

sheltered hollow in this wood presented. Theré was the tndu-

lating field, clad in waving grass and set about with the pale-

hued springtime foliage of the white oaks; there. were the

flowering. hawthorns ; and there were the myriads. of. Bibios

floating in the sunshine, streaming here and there like chaff

before sudden gusts and swirls of air, All the spiders’ webs

E B

A, larva of Bibio fraternus Loew., lateral view. B, pery view of head of female

pupa. C, ventral view of male p

in the bushes were filled with captives ; little groups of ants

were dragging single flies away to their nests, and once I saw

overhead a chestnut-sided warbler, perched on a bare bough

directly in a stream of passing flies, rapidly pecking to right

and to left, persistently stuffing his already rotund maw,

I stood in one spot and swept the air with my net for half a

minute and obtained 128 male Bibios and 5 females : ratio, 25: 1.

I swept the grass a few times with my net and thus obtained

127 males and 372 females : ratio, 1: 2.2. The discrepancy of

ratios is due to the fact that a comparatively small proportion of

the flies were inthe air. I counted.a number of flies I could see

resting on the grass in several small areas wide apart and found

the counts averaged 15 Bibios per square foot; and there were

here in one place forty acres of such Bibio territory! There can

` 1 Professor Herbert Osborn notes the of Bibia albipennis in phe-

nomenal numbers in Iowa, in Jnsect Life, vii iii (1891), p. 479.

184 THE AMERICAN NATURALIST. [VoL. XXXVI.

be no doubt that this obscure and little-known fly is a factor of

considerable importance in the ecology of such a situation.

Immature Stages. — I now describe the larva and the pupa,

and accompany the descriptions with drawings by Miss Maude

H. Anthony.

Larva.— Length 13-14 mm. ; diameter 1.5 mm. Body cylindric as far

as the 9th abdominal segment, or with slightly greater depth in the thorax ;

head well developed, exserted, about equal in length to the body segments,

except the first, which is a third longer than the others ; there is a distinct

Y-shaped suture on the top of the head; a pair of sete are inserted just

before and another pair just behind the clypeal suture ; eyes wanting ; four

setze on the epicranium each side, — two at the edge and one above and

one below ; there are lesser setze on the maxilla externally. The lacinia

of the maxillae and the mandibles are elongate pyramidal, with low teeth on

their truncated apices. There are two pairs of thoracic and eight pairs of

abdominal spiracles: the foremost is situated at the rear of the prothorax

and is larger than those following, except the last ; the second pair is at the

front margin of the metathorax; the next seven pairs are just before the

middle of abdominal segments 1-7 ; those of the 9th abdominal segment

are very large and each is divided by a median groove. Each of the middle

body segments is completely encircled by a row of about a dozen tubercles,

which is slightly oblique upon the sides and in which the dorsal tubercles

are usually longer than the ventral, and there are a few tubercles out of

line at the sides in the vicinity of the spiracle ; on the 1st segment behind

the head there is another row of eight tubercles, on the dorsum between

the spiracles, and on this segment the tubercles are all longest at the sides ;

on segment 8 of the abdomen the dorsal tubercles are more elongate and

the ventral ones more reduced ; segment 9 is depressed conic, tapering

posteriorly, with the large spiracles inserted in anterior emarginations of

a brownish, chitinous, saddle-shaped plate, which covers most of the dor-

sum of the segment; behind this plate arise two pairs of filaments (elon-

gate tubercles), the median pair as long as the segment is thick, the external

pair a third shorter.

Skin granulate, amd invested with adherent dirt particles.

Pupa.— Length 7.5 mm. ; breadt h of thorax 2.1 mm., of abdomen

1.5 mm. Body iiss, whitish, with low appressed head, short, thick

thorax, and straight abdomen whose sides are parallel except at its abruptly

tapering apex.

. Head fiat, with short antenna cases which extend hardly more than

halfway across the eyes, with well-marked paired jaw cases, and with

an ocellar tubercle at the rear of the head above, low in the male, high

ah poniienpln el eee E:

eyes).

No. 423.] REMARKABLE OCCURRENCE OF THE FLY. 185

Prothoracic spiracles low, not elevated above the thoracic dorsum; no

respiratory horns or trumpets.

Ninth abdominal segment terminating on the dorsal side in a pair of

stout divergent triangular processes, that are about as long as the segment

which bears them.

The rather generalized larva is characterized by an unusually

well-developed head, simple mouth parts, rings of tubercles on

all the segments, those of the two hindmost segments elongate,

becoming filaments, showing out of what material the borders

of the tipulid caudal respiratory disk have been made. The

pupa is characterized by the unusual brevity of the antenna

cases, which reach but halfway across the eyes, distinctly

paired jaws, low prothoracic spiracles without respiratory

trumpets, horns, or tubes, and a naked, spineless skin.

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AN UNUSUAL OCCURRENCE OF DINOFLAGEL-

LATA ON THE CALIFORNIA COAST.

HARRY BEAL TORREY.

DiscoLorations of the sea due to the presence of minute

organisms, and often many miles in extent, have been seen in

various parts of the world, along the shore and in the open sea.

Deep-water sailors are familiar with such appearances, which

they call * whale food," or “brim.” The cause of the color is

not always the same organism, though perhaps most frequently

some species of Dinoflagellata, in enormous numbers.

This phenomenon was observed at San Pedro, Cal. during

the present summer, and, according to the testimony of old

residents, for the first time in that region. Something of the

sort is said to have been noticed in Tomales Bay, some five

hundred miles farther northward, in the early seventies. These

are, I believe, the only known instances of its occurrence on

the California coast; so.that its rarity alone seems to demand

a recording of the few notes taken. during this last appearance.

It was first noticed on July 7 as a red streak off the mouth

of San Pedro Harbor. During the next few days it approached

the shore, changing its shape and dividing into several patches,

each many acres in extent. On the 16th it reached the shore,

and was the cause of a most unusual display of phosphorescence,

which reappeared many evenings in succession... The phosphor-

escent water was found to contain vast numbers of a peridinian

which I have referred tentatively to the genus Gonyaulax (see

description below), and which gave it its red color during the

day. Under the microscope Gonyaulax responded with a flash

of light to any marked disturbance. of the water. containing it.

At this time Noctiluca, so abundant later on, was not noticed.

Wherever the ocean waters were roughened the wavelets were

crested with green-blue light. In the quiet waters of the harbor

the fishes, pelagic worms, and crustaceans traced trails of fire.

187

188 THE AMERICAN NATURALIST. [Vor. XXXVI,

On the 20th, four days after the red (characteristically a

muddy vermilion) streak had reached the shore, a most sicken-

ing odor arose from the water along the beach. On the 2rst it

was almost unbearable. During the night, on a beach about

four hundred feet long, a large number of animals were left

by the tide. Among them were several hundred holothurians

(Trachostoma arenata), several specimens of two species of

sting ray (Mylobatis californicus and Urolophus halleri), two

species of guitar fishes (Azzobatus productus, 3 individuals ;

Platyrhinoidis triseriatus, 3 individuals), two cestracionts (Gyro-

pleurodus francisci), two dogfishes (Galeus californicus), a red

perch, a large number of smelts, and several octopi. The

fishes and octopi were dead, but many of the Trachostoma

lived for several days. Many Pinnixa faba had crawled out of

the cloaca of the holothurians, on which eco are normally

parasitic.

On the 22d more Trachostoma were left on the beach. Some

days later the odor had almost disappeared, but the water in the

harbor and along the beaches was a brilliant vermilion. It

was at this time that on two succeeding mornings a number

of specimens of the blindfish (ZygA/egobius californicus) were

thrown up, both alive and dead. Then for two days came a

number of Zevila crassatelloidis, many Petrolisthes cinctipes and

Cancer antennarius. Finally, great numbers of Hippa analoga

were thrown up, most of them alive but apparently debilitated

and unable to dig as is their habit. They were mostly of large

size.

This wholesale destruction was entirely unprecedented for

the summer. The height of the tide and breakers were but

subordinate causes at best; for, although the holothurians and

selachians appeared: after a higher tide and stronger sea than

usual, as many of the former came up on a beach protected by

a breakwater as on the beach shown in the photograph (Fig. 1).

And as the tide fell from day to day, the animals did not cease

to appear. The Typhlogobius appeared when the tide was

moderate, with practically no sea running. This is interesting,

not only because Typhlogobius usually lives, undisturbed by

breakers, in holes and crannies among therocks, which must

No. 423.] OCCURRENCE OF DINOFLAGELLATA. 189

have been in this instance some distance from the beach, but

also because it has never before been reported from San Pedro.

The Gonyaulax served here as a means of discovery.

The above facts have been enumerated because they exhibit

in a striking way a natural cause in process of eliminating the

unfit of several and diverse species, which are resisting with

various degrees of persistence and success. Whether this elimi-

nation is promiscuous, or correlated with variation from the type

Fic. 1. - Beach covered with dead animals, mainly holothurians with several selachians in

of the species, is a question for whose solution material has been

collected by Dr. F. W. Bancroft of this laboratory.

The “red water" occurred for two hundred miles at least

along the coast, from the region of Santa Barbara to San Diego,

and extended several miles to'sea. It was still present around

San Pedro on September r. Wherever it was seen the fisher-

men reported a scarcity of food fishes. The small harbor fishes

seemed to be unaffected by: it, and the tiny invertebrates of the

plankton were abundant in it. - Noctiluca appeared in great

numbers toward the end of July, and devoured Gonyaulax with

190 THE AMERICAN NATURALIST. [Vor. XXXVI.

avidity. There were more copepods in the red streaks than

in the intervening bluer water. I was not able to see any

Gonyaulax in them, however.

Gonyaulax undoubtedly produced its harmful effects by dying

in enormous numbers, the putrefactive changes thus occasioned

polluting the water and giving rise to the stench already men-

tioned. It died most rapidly along the water's edge, thus affecting

the littoral animals especially.

When placed under the microscope for examination in a drop

of water, many Gonyaulax soon became quiescent, and the pro-

toplasm drew away from the cuirass, forming a spherical cyst,

which was often discharged by a rupture of the cuirass along

any sutural line. - Others died outright and disintegrated, leav-

ing only the transparent, colorless cuirass. This tendency to

die readily was noted long ago by Darwin for a species of

dinoflagellate found in swarms off me western coast of South

America.

With this pines abundant species other species of Dino-

flagellata were found: Ceratium furca, C. fusus, C. michaelis,

Peridinium divergens, Prorocentrum micans, Polykrikos sf.,

Dinophysis Sp-s Noctiluca sg., and certain swarmspores resem-

bling those of Peridinium ovatum. The most abundant of these

was Noctiluca, toward the end of July, but it did not begin to

approach Gonyaulax in numbers. Where there was one Noc-

tiluca there were hundreds, if not thousands, of the other form.

The latter was present in the tow as early as June 17, in

company with, and about as abundant as, the above-named spe-

cies. All of these species increased during July, but Gonyaulax

outstripped the others remarkably. The cause of this increase

of one species over these closely related ahd associated species

is not clear. I did mot see a single case.of fission, but the

individuals varied so greatly in size as to suggest rapid repro-

duction, probably by encystment and spore formation, though

no spores excepting the large Peridinium (?) swarmspores were

seen. Noctiltica was sporulating at this time. It. is possible

that the immense patches of Gonyaulax . were brought in by

currents from: ner water. Nothing, however, -was seen of

them throughout the season around Santa Catalina Island,

No. 423.] OCCURRENCE OF DINOFLAGELLATA. IQI

some twenty miles to sea. And if this were so, the cause of

the rapid reproduction would still remain obscure.

The boundaries of the red streaks were quite sharply marked,

although the water between streaks often contained Gonyaulax

in abundance. Temperatures and determinations of salinity

were taken at the surface and at the bottom of both red

streaks and water entirely free of Gonyaulax. The. readings

averaged the same in all cases for the same depths. Soundings

with a rather crude water bucket indicated that Gonyaulax was

present approximately in as great abundance at the bottom at

the depth of six fathoms as at the surface.

The key to the problem lies, I believe, in the chemical consti-

tution of the water, since, aside from a sensitiveness to chemical

Fic. 3.

Fic, 2. Gonyau lax, showing both

Fic. 3. — Gonyaulax from upper cg sou arrangement of anterior plates.

stimuli which.'it possesses in common with all living things,

Gonyaulax is holophytic, if its color is a safe criterion. Zumstein

has recently shown that the chlorophyll-bearing Euglena may lose

its color under appropriate conditions and change its method of

feeding in consequence to the saprophytic type. The C. michae-

lis and Polykrikos sf., as well as the Peridium (?) swarmspore,

found with Gonyaulax, were colorless. Not only, then, within the

limits of one family of protophytes do two methods of nutri-

tion exist, but in a single species, and possibly in a single

life cycle. But the question of the source of the food of marine

organisms is too vast a subject to:be attacked with the scanty

materials at hand. Whether- Gonyaulax is nourished by inor-

ganic or organic solutions, and whether a variation in food supply

is the explanation for its unusual abundance, are problems to be

solved by future experiment.

192 IHE AMERICAN NATURALIST.

I have called the dinoflagellate with which these notes are

chiefly concerned Gonyaulax, because in the first place it

approaches that genus very closely, and in the second place

such a small amount of the literature of the group is accessible

to me that I do not feel warranted in erecting a new genus

for it. It differs from Gonyaulax as described in Engler and

Prantl's Die natürliche Pflanzenfamilien only in that in the

anterior moiety of the cuirass there are six intermediate plates

instead of five, and four end plates instead of three.! It may be

described as follows (Figs. 2, 3):

Body divided by the equatorial fun into two subequal

halves, each of which is a low cone, the posterior one truncated.

Anterior cone formed by six large intermediate plates meeting

four much smaller unequal end plates in an angle somewhat

less than one hundred and eighty degrees. Posterior cone of

five large intermediate and one large end plate, with a narrow

accessory plate next the longitudinal furrow. Equatorial furrow

slightly oblique, deep, with toothed edges. Longitudinal fur-

row broader posteriorly than anteriorly, extending from the edge

of the posterior end plate to the apex of anterior cone. All

plates regularly bossed. Sutural lines raised into high, narrow

ridges.

The equatorial flagellum alone is active in ordinary swimming,

producing a rapid rotary and forward movement. The other,

which trails behind during such progress, is responsible for the

spasmodic jerkings occasionally seen:

The size of the cuirass varies from 25 » in diameter by 29 &

in length, to 53.4 by 56 p. Cysts average 24 p in diameter.

They are almost spherical, with granular contents of a brown-

green color. : Cuirass itself colorless.

ZOOLOGICAL LABORATORY, UNIVERSITY OF CALIFORNIA,

September, 1901.

. 1Since the manuscript was sent to press I have found that Bütschli (Bronn's

Klassen und deus has noted a similar discrepancy between the original

description of Gonyaulax and hie on cpeerations, which e eads me to suspect

er the eres oe Lm ird dace pon

REGENERATION IN HYDRACTINIA AND

PODOCORYNE.

ANNAH PUTNAM HAZEN.

In a paper entitled “ Experiments in Regeneration and in

grafting of Hydrozoa,”! Peebles has given the results of some

interesting experiments with Hydractinia and Podocoryne.

These two hydroids are found at Woods Hole on the shells

inhabited by hermit crabs. Each colony consists of three

different kinds of individuals, namely, the nutritive, the repro-

ductive, and the protective (a low cone-shape growth extend-

ing over the hydrorhiza is considered by some authors to

constitute a fourth class of individuals). The experiments

described by Peebles were confined to the first three classes.

The well-marked characteristics of the different kinds of indi-

viduals make it possible to distinguish between them even

when the polyps are immature or incompletely regenerated.

The knoblike tentacles of the reproductive polyps appear dur-

ing regeneration earlier than the reproductive organs, and the

long, slender tentacles of the nutritive polyps can be easily

distinguished from those of the protective individuals, which

contain nematocysts. Peebles found that polyps freed from

the hydrorhiza would live for days in small dishes containing

sea water and quickly regenerate lost parts. _ Pieces of the stalk

of these three classes of polyps, from which the heads had

been cut, regenerated new heads of the same kind as those

that had been removed. Small pieces cut at different levels

from the stock of nutritive and protective polyps regenerated

at one end a head like the one removed. Not infrequently a

head appeared at both ends of a short piece, Without exception,

the regenerated heads had the same characteristics as those that

had. been removed, and no constant variation in time or capacity

for regeneration at the different levels was observed.

1 Archiv für Entwickelungsmechanik der Organism, Bd. x, Hefte 2, 3.

194 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Nutritive polyps cut from the hydrorhiza were grafted by

Peebles in pairs, either by their oral ends (after removing both

heads) or at the aboral ends (with both heads attached), or the

aboral end of one individual was grafted-to the oral end of

another, from which the head had been removed. In all cases

the line of union between the two components was ‘soon

effaced, the digestive cavities became continuous, and a com-

plete union seemed to be effected. In all the experiments in

grafting ‘nutritive polyps, nutritive heads were regenerated at

one or at both ends, and often also in the region of the graft.

_ The union of different kinds of individuals was also successfully

accomplished as follows: The aboral end of a nutritive polyp

was grafted to the aboral end of a protective polyp,— the

direction of the two components being thus reversed. After

a few hours the nutritive head was removed, leaving a portion

of its stalk attached to the aboral end of the protective polyp.

The head which regenerated at the free cut surface showed

markedly the characteristics of the nutritive individual and

did not seem to be in the least affected by union with the

protective polyp. Similar grafts were made with nutritive

and reproductive polyps, followed by a removal of the repro-

ductive head. Here, again, the regeneration of a reproductive

head at the end of the stalk showed no modification due to

the union with a nutritive polyp.

The results of these experiments by Peebles suggested the

following questions: first, Would the fixation by means of

stolons of one of the components of the graft alter the result?

second, Would the kind of regeneration that took place be

affected by the relative orientation (with respect to the polar-

ity) of the pieces? With these questions in mind, I carried

out the following series of experiments at the Marine Biolog-

ical Laboratory during a few weeks in July and August, 1901.

Both Hydractinia and Podocoryne were used, and the method

of grafting was similar to that employed by Peebles. It was

found to be of importance to use fresh material since that

kept for several days in the aquarium: regenerated more

sd and less bir) than material’ | just brought in. A

few

repeated

n Lu

No. 423:] HYDRACTINIA AND. PODOCQR YNE. 195

and similar. results obtained, although most: of the grafts were

made. by joining-the-aboral end ofa reproductive polyp to the

oral end. of a nutritive stalk, the head of the latter having been

previously removed. .In this way. the direction of the two

components was the same, instead.of reversed, as in Peebles’s

experiments. |The pieces were allowed to remain. undisturbed

for a short time after grafting, or until, the tissues hàd become

united, so that the reproductive head could be cut off without

danger of separating the grafted pieces, Better results seemed

to be obtained. if the head were removed

within a few: hours after union, so that

the process of regeneration started in the

first night after the graft had been made,

although satisfactory results were sometimes

obtained when the cut was made on the

second day. Fig. 1 represents a typical

experiment of this kind. -An immature Fic.:— x45. Theaboral

reproductive polyp (7) was selected and ee ee eee

grafted at its abóral end.to- tbe oral end.. wagafted:to:the oral

end of a nutritive stalk

of a nutritive stalk (z.) from which the head - (4) August 3. A few

had been removed, The tissues joined, and

a few hours later the reproductive head was was cut off, as indicated

removed, leaving the two components Of ` next day a reproductive

nearly equal size. Within twenty-four hours

a complete reproductive head had regener- reproductive stalk.

ated (rh.), which showed no modification due (Dawn Aususts) De

to the Wise of the stalk of the nutritive oral end of each irdi-

polyp.

When the union of the two soliponeets was incomplete and

a part of the cut surface of the nutritive stalk was exposed

at the side, tentacles or a whole nutritive head developed at

this place, as shown in Figs. 2, 3, and 4, zz. This occurred

frequently, as the nutritive polyps are often much larger than

the reproductive polyps, and the latter, unable to cover the

whole cut surface, unite with the nutritive only at one side.

This did not prevent.the body cavities of the two. eens

from becoming continuous. The portion. of the ‘re cti

stalk (7.) used in the experiment shon, È in Pe 2 was s taken

196 THE AMERICAN NATURALIST. [Vor. XXXVI.

from the oral region of a young individual, while the nutri-

tive component (z.) came from the aboral region and shows

a piece of a spine (s.) attached to its base. Although, in this

case, the nutritive component was rela-

tively much larger than the reproductive

stalk, a head regenerated on the latter

which showed no apparent influence of

the larger component but possessed only

the characteristics of the reproductive

head.

Another ex-

periment is

shown in Fig.

3 in which the

nutritive stalk

(n.) was many

times larger

than the small

component

taken from a

reproductive

polyp (7). The

latter was cut

(s.) was attached ^ d the a from the abo-

(Drawn August 5

ral region of

an adult individual from which most

of the generative buds had been

removed.! Here, again, the union was

complete, and again the regenerated

head showed no effect produced by

the larger component, but the new

growth corresponded to what would

probably have appeared on a repro-

ductive stalk alone from which the a) amt shi te (ut)

head had been removed. ven Moon ey d

Many similar experiments were made, from which these

have been selected as typical. Considering that these different

1 Two generative buds were left, as shown in Fig. 5 Z.

No. 423.] HYDRACTINIA AND PODOCORYNE. 197

individuals are organically connected, it seemed at the out-

set possible that, under certain conditions where a complete

union was effected, the subsequent regeneration might be

modified. This seemed especially likely to occur if the

nutritive.polyp, which is the more vigor-

ous and the less specialized of the two,

formed the larger component. Repeated

attempts were made to leave as small a

portion of the reproductive stalk as pos-

sible attached to the nutritive compo-

nent, but, with one exception, none were

obtained in which the regenerated head

showed the influence of the nutritive

polyp. The result is. the. same when | ` e |

the direction: of the two components is |

reversed, as in Peebles's experiments,

and when it is the same, as in most of

my experiments (see Figs. 1, 2, and 3).

All regions. of the stalk

from which the components

are taken seem equally un-

able to influence the sub-

sequent regeneration or to

be influenced by the union.

It was also found that the

regeneration following a

graft of this sort is no

more easily influenced Frc. 4.—x 45. The aboral end of a reproductive polyp

when the reproductive a ta ee ove bed Ga ei ci Ja ly 21,

component is taken from Soror o oe doe ca ue WA vise

an immature polyp than WR stalk and cages tentacles A age = the

when it is taken from an (at) had also developed on the aboral end of the nutri-

tive stalk attaching it to the disk. (Drawn July 23.)

adult.

"An interesting and solitary exception to these results is

illustrated in Fig. 4. A nutritive polyp (#.) and a reproduc-

tive polyp (r.) were grafted as in the preceding cases, and on

the following day the reproductive head was removed, leaving

only a small part of the reproductive stalk attached. A day

198 THE AMERICAN NATURALIST. [VoL. XXXVI.

later a nutritive head (7/.) had regenerated on the end of the

reproductive component and, at the same time, stolons (s7.)

had pushed out from the nutritive polyp, fastening it to the

bottom of the dish.. As this was the only instance in which

a reproductive polyp produced a nutritive head, it was prob-

able that a new condition had appeared and. affected the

regeneration. What this condition may have been was not

satisfactorily determined, but a possible explanation was sug-

gested by comparison with the results of another series of

experiments carried on at the same time. These will be

described. below.

Peebles showed that pieces of the stalk of nutritive polyps

cut free from the colony and allowed to remain undisturbed at

the bottom of a shallow dish of water sometimes sent out long

branching stolons, the original stalk becoming absorbed and

indistinguishable from the newer growth. * After several days

new individuals began to appear, growing up at right angles

to the stolons, just as the different individuals of a colony

grow from the hydrorhiza. The formation of new polyps con-

tinued for several weeks when the experiments were brought

to an end. As many as ten to fifteen new polyps were pro-

duced by one piece of the stalk. These polyps were inva-

Fic. 5. — Represents a small portion of a colony which had. developed from a nutritive polyp,

showing a new nutritive polyp (s), a reproductive polyp (+), and stolons (sż.). The original

pelyp had been abserlied., colony was started July 24. The drawing was made August 3.

The disintegration of the colony began August 8.

riably nutritive individuals, characterized by their long slender

tentacles.” I repeated this experiment and also used pieces

of nutritive polyps as well as pieces of the stalk. The

growth of a large number of colonies followed. New nutri-

tive polyps were always the first to appear on the stolons,

No. 423.] HYDRACTINIA AND PODOCORYNE. I99

but after the colony was well formed a few small reproductive

polyps also grew out from the stolons at occasional intervals

(Fig. 5). -Although the colony lived for several days or a week

after the production of the reproductive polyps, no signs of

Fic. 6. — Represents a luctive polyp which t from the hydrorhiza July 20. Several

generative buds were removed; two were left (g.). The reproductive tentacles were

replaced by four nutritive tentacles (nt). Two small knoblike wen () are seen

below the patritive tentacles

t) g l end, fastening the individual to the disk. pets ee

developing generative buds appeared. Indeed, the life of the

colony seemed spent, the stolons became more slender and

broke apart, and the polyps died, one after another. During

the period of growth the stolons often anastomose and two

adjacent colonies become united. Such colonies are most

easily started if whole polyps are cut off close to the hydro-

rhiza. A small piece of a polyp will lie on its side on the

bottom of the dish, often producing new hydranths and chang-

ing its form, until it disintegrates, but rarely forms stolons.

In the same way reproductive polyps were removed and

kept in dishes, with the hope that they would also form

colonies. They sent out stolons much more slowly than the

nutritive polyps did, and although a large number of individu-

als formed stolons, many of them died before new polyps had

developed. It is probable that others, which seemed more

vigorous, might have developed new polyps had more time

been allowed them. In one dish in which half a dozen repro-

ductive polyps had formed stolons a curious change was

observed. The stolons had grown rapidly, and the hydranths

200 THE AMERICAN NATURALIST,

which had borne the small knob characteristic of the repro-

ductive polyp showed, in three days, a remarkable. develop-

ment of tentacles like those on a nutritive hydranth, as

shown by z£, Fig. 6. Two small knobs (£.) are. present

below the ring of tentacles, and these may— or may not —

represent those of the original reproductive hydranth. A

few large generative buds had been removed. from the polyp,

but two remained as seen in Fig. 6, g. The polyps seemed

vigorous, but possibly the presence of the generative buds

caused an early disintegration. At all events, they all died

before any new polyps were formed. by the stolons. What the

condition was which brought about this change was not deter-

mined, but it seemed possible that the formation of stolons

might be connected with the. development of nutritive tentacles.

The growth of stolons i in this case (Fig. 6) was more rapid than

in that of other reproductive individuals in which the nutritive

tentacles did not develop. In other instances the reproductive

heads may have lost the power of developing the long tentacles

when the stolons were late in appearing. A clearing up of

this point might throw light on the experiment described above

(Fig. 4), in which a piece of a reproductive stalk grafted to a

nutritive polyp regenerated a nutritive head. The reproduc-

- tive head had not been removed until the day following the

graft, and the stolons developed at the same time that the head

was regenerated. The regeneration of the nutritive head might

have been influenced by the developing stolons.

It gives me pleasure to acknowledge my indebtedness to

Prof. T. H. Morgan for — suggestions during the course of

this work.

SMITH COLLEGE,

NORTHAMPTON, MASSACHUSETTS.

ON THE STANDING OF PTEROPUS HALDEMANI

HALOWELL.

JAMES A. G. REHN.

In 1846 Halowell described a species of Pteropus from West

Africa as P. haldemani (Proc. Acad. Nat. Sct. Phila., Vol. III,

p. 52); this species has generally been overlooked. Several

years ago Mr. Paul Matschie of the Berlin Museum fiir Natur-

kunde resurrected the species (Sttzungs-Bericht der Gesellschaft

naturforschender Freunde zu Berlin, 1899, Nr. 2, p. 30) and

made the rather startling assertion that it was based on a young

example of Hypsignathus monstrosus H. Allen (Proc. Acad. Nat.

Sci. Phila., 1861, p. 156). In his recent monographic work

(Die Megachiroptera des Berliner Museum fiir Naturkunde, p. 42)

Mr. Matschie repeats his previous views. As Mr. Matschie did

not examine the types of either P. haldemani or H. monstrosus,

his conclusions were formed entirely from a description which

gave no idea of the cranial or palatal characters.

The types of both species are contained in the collection of

the Academy of Natural Sciences of Philadelphia, and on exami-

nation the writer finds that Mr. Matschie's conclusions are not

sustained by the specimens.

The type of Pzeropus haldemani consists of an alcoholic skin

from which the skull had never been detached, and on removal

the latter is found to be quite typical of the genus Epomophorus,

and in no way closely related to Hypsignathus. The molars are

low and slightly grooved as in Epomophorus and lack the lateral

cusps characteristic of Hypsignathus. The skull and the exter-

nal aspect of the head exhibit no trace of the extraordinary club-

like development observed in the last-mentioned genus, which

would surely be appreciable even in a young individual. — .

The structure of the palate agrees exactly with Epomophorus

gambianus Ogilby (Proc. Zool. Soc. London, 1835, p. 100) as

201

202 THE AMERICAN NATURALIST.

figured by Dobson (Catalogue Chiropt. Brit. Mus., Plate II,

Fig. 3, a), and Pteropus haldemani can, without hesitancy, be

considered synonymous with Ogilby’s species.

ACADEMY OF NATURAL SCIENCES,

PHILADELPHIA, Pa.

THE COLORS OF NORTHERN POLYPETALOUS

FLOWERS.

JOHN H. LOVELL.

AcconniNG to the later systems of classification the Apetalze

and Polypetalae form a single subclass, the Choripetalze. In the

apetalous families of eastern North America, which have been

considered in an earlier paper, there are 175 green, 89 white, 51

yellow, 45 red, and 24 purple flowers. The 1217 polypetalous

plants have 140 green, 410 white, 333 yellow, 84 red, 193

purple, and 57 blue flowers. The northern Choripetalz, then,

contain 315 green, 499 white, 384 yellow, 129 red, 217 purple,

and 57 blue flowers. Of the 92 families belonging to the Chori-

petale, 47 contain green, 52 white, 45 yellow, 28 red, 39 purple,

and 5 blue flowers. The much greater abundance of species

and families with green, white, and yellow coloration, as well as

the less specialized structure of the flowers, points to these

colors as more primitive or more easily developed than red,

bright purple, or blue. In certain genera, however, small dull

red and purplish flowers are evidently derived directly from the

primitive green.

The order Ranales includes the Nymphzacez, Magnoliacez,

Ranunculacez, and six other families of less importance. The

simpler species of this order have the organs of the flowers

spirally arranged, separate and distinct, and the stamens and

pistils indefinite in number, as in Ranunculus. From the domi-

nating character of the Ranales, says Engler, it is clear that the

other orders of this series diverge from the Ranales in various

ways, some following one direction of development, some another.

The ancestral form of the angiospermous flower would appear

to have been a branch, or part of a branch, with carpophylls at

the end, followed by androphylls, and then by the primitive

1 The Colors of Morc Monocotyledonous Flowers, And Nat., vol. xxxiii,

p. 493; The Colors of Northem Apetalous Flowers, Amer. Nat., vol. xxxv, p. 197.

203

204 THE AMERICAN NATURALIST. [VoL. XXXVI.

leaves destined to form petals, sepals, and bracts. No such

inflorescence now exists, but an approach to it may be found in

the Magnoliaceze. In Magnolia the floral bud terminates the

secondary branches, of which it is only a prolongation. The

oblong flower of the Magnolia has the receptacle prolonged and

the parts of the perianth, the stamens, and the pistils, spirally

arranged and indefinite in number. In the arrangement of the

plant families in a lineal series, those flowers which have the

organs separate and which resemble most closely the proangio-

spermous floral branch are regarded as the simplest; while the

highest rank is assigned to those which have the organs modified

and consolidated, as in the Orchidaceze and Compositze.

In the Nymphzacez, or water lilies, we meet with dicotyledo-

nous plants resembling monocotyledons in the structure of

the leaf and rootstock. The analogous arrangement of the fibro-

vascular bundles in a Nymphaea and a Tradescantia has been

excellently figured by De Bary. The successive steps in which

simple and distinct carpels may become compounded are also

well shown by the genera of this family. In the primitive genus

Nelumbium, also in the oldest forms found in a fossil condition,

the simple carpels are contained in little pits in the large fleshy

receptacle ; in Cabomba and Brasenia the oblong carpels are

borne on the receptacle, but are separate ; in the fossil genus

Anoectomeria of the Middle Tertiary the carpels are partially

united ; in Nymphza the union is complete ; and finally, the

culmination is reached in Victoria regia, where the compound

ovary is inferior. Of the eleven northern species four are white,

five yellow, one red, and one purple. Yellow coloring was

probably very early developed, as it is the color of the primitive

` Nelumbo lutea and of the three native species of Nymphza

(Nuphar). Throughout the summer the numerous broad shin-

ing leaves and large yellow flowers of Nymphaa advena, the large

yellow pond lily, form a well-known characteristic in the vegeta-

tion of ponds and slowly moving streams. The six sepals are

unequal : the three inner are large and bright yellow; the three

outer and smaller are wholly or partially green exteriorly, though

the upper half is often yellow, while the interior lower portion

No.423.] NORTHERN POLYPETALOUS FLOWERS. 205

truncate, wedge-shaped bodies, which serve as honey glands.

The honey is freely secreted on the outer side near the top, the

nectariferous portion being orange yellow. The fruit is scarlet.

I have collected upon the flowers in Maine four Diptera, two

Coleoptera, and one small bee, 7ZaZictus nelumbonis, which con-

fines its visits to this flower. Great numbers of the small fly

Hilara atra revel in the pollen. The single species of Cabomba

is white with a yellow base, and the northern species of Castalia

(pond lily) are pure white tinged with pink, or deep pink in a

variety growing in eastern Massachusetts. In the famous

Victoria regia of the Amazon the outer petals are white and

the inner crimson, and in Nymphea cerulea of the Nile the

flowers are blue. The leaves of several genera are strikingly

bicolored, being light green above and violet or reddish purple

below. According to Kerner the anthocyanin, or purple color-

ing matter on the lower side of the leaf, can arrest the rays of

light (which would otherwise escape into the water) and change

them into heat and make them useful to the plant.

The largest tree flowers known belong to Magnolia. ‘Their

effect in early spring is grand beyond description, illuminating

the whole landscape and filling the air with their rich perfume."

The red flower of M. campbellit of Himalaya is ten to eleven

inches broad, while 77. macrophylla of the Southern States has

flowers ten inches in diameter. Of the five species of the

Magnolia four are white and one greenish yellow. M. macro-

phylla is white with a purple center, while Zzriedendron tulipi-

fera, tulip tree, is greenish yellow without and yellow within.

Magnolia grandiflora is fertilized by rose beetles or Cetoniz.

At night the white flowers are 5° to 10° C. above the surround-

ing air, and thus afford warmth as well as ‘food.

Both the Nymphzeacez and Magnoliacez produce very large

_and conspicuous flowers, which are pollinated by the smaller

bees, flies, and beetles. While these splendid showy flowers

are, no doubt, of benefit for attracting the attention of insects,

yet it is evident that they could be produced only by large trees

and vigorous herbaceous plants. The water lilies grow in situa-

tions where the soil is charged with nitrogenous matter, and the

framework of the plant requires a minimum of mechanical

206 THE AMERICAN NATURALIST. [Vor. XXXVI.

support.. The Magnolias are stately trees, growing in rich

woodlands. Both families have very large leaves. The primitive

color of the flowers»was probably green, as it still is in Magnolia

acuminata, ‘Transition stages from green to yellow occur both

in Liriodendron and in Wymphea advena. In other instances,

as in certain species of Magnolia and Castalia, they became

white at a very early stage, and in certain species subsequently

changed to red. In the case of M. macrophylla, where the

white flower has a large purple center, the entire flower may

have been purple, as in ///ictum floridanum, or anise tree, or it

may be a deposit of pigment resembling that found on the inner

side of the sepals of Vymphea advena.

In a part of the species of the Ranunculacez the petals are

wanting ; in a part they are small and transformed into hollow

nectaries; while in others they are regular and conspicuous.

The ancestral stock from which the various genera have diverged

doubtless possessed petals, or phyllomes, corresponding to this

whorl; but they had already been lost by some generic lines

when insects began to visit the flowers. Of the ninety-seven

species, six aré green, twenty-six are white, thirty-eight are

yellow, three are red, thirteen are purple, and eleven are blue.

Conspicuousness is insured by the sepals in Clematis and

Caltha; by the petals in Ranunculus; by both the petals and

sepals i in Aquilegia ; and by the numerous stamens in Thalictrum,

Actzea, and Cimicifuga. In Thalictrum the white or lilac fila-

ments are broad and petaloid. The flowers are visited infre-

quently by flies and the short-tongued bees.

Of the apetalous flowers, Caltha palustris and C. flabellifolia

have a yellow, and the aquatic C. natans a white, calyx. A

change of locality may induce a change from yellow to white,

as Anemone alpina on the Central Alps bears chiefly sulphur-

yellow flowers, but in the eastern limestone Alps its flowers

are always white. In our native species of Anemone the sepals

are green, white, or purplish, but florists offer scarlet and blue

varieties. The anthocyanin displayed by the underside of the

sepals of A. nemorosa, especially in bud, probably serves to con-

vert light rays into heat ; this plant blooms in early springtime.

The sepals of Clematis ochroleuca are green, of C. virginiana

No.423.] NORTHERN POLYPETALOUS. FLOWERS. 207

white or greenish white; but in most species of Clematis the

color is purple. By hybridization a great variety of hues have

been produced, including blue, red, cream, and yellow. The

calyx of Hepatica is white, rose, or blue. Kerner remarks that

the flowers, which appear as soon as the snow leaves the ground

in open woods, by their blue color present a strong contrast

against the yellow-brown leafage, whereas in green meadows

they would scarcely be seen. The occurrence of blue sepals is

rare and is perhaps nowhere so well shown as in this family.

From this brief survey of the coloring of the calyx it is evident

that its capability for developing a wide range of colors equals

that of the corolla. Though commonly green and only protec-

tive, it may give rise to any color.

In other genera of the Ranunculacez the petals are present

but are transformed into nectaries. In Eranthis and Helleborus

they take the form of a trumpet ; in Isopyrum they are spoon-

shaped, in Nigella lamp-shaped ; ; in Coptis they resemble a hood,

in Aquilegia a horn of plenty; and in Delphinium the two

upper sepals and petals are both spurred. Tubular petals have

also been observed in Ranunculus repens by Masters.! This

tendency is inherent in the flowers and not induced by the

agency of insects, and has greatly aided in the production of

the different genera. The sepals of all these flowers are con-

spicuous.. In Helleborus viridis the calyx is yellowish green,

and in H. niger, ^or Christmas rose, which blooms in winter, the

flowers are at first white, turning pinkish, and then green.

Trollius laxus, which grows in the shade of dense swamps,

has greenish-yellow flowers, while 7. europeus has bright yellow

sepals. Coptis trifolia, a woodland plant, has small white flowers

in springtime.

The species formerly included in the genus Ranunculus, but

now segregated into several genera, have the petals large and

regular. The nectar is secreted in a little pit near the base.

The thirty-one species of Ranunculus native in the northern

states are yellow, or whitish in R. xivalis, or occasionally in

R. acris in autumn. A wide range of colors is displayed by

KR. asiaticus, including yellow, red, purple, and nearly black.

1 Vegetable Teratology, p- 23.

208 . THE AMERICAN NATURALIST. [Vor. XXXVI.

R. viridiflora is green, with scarlet edges. In R. auricomus the

petals are sometimes wanting, and ‘their place is taken by

the sepals with their bright yellow limb. . In Ficaria ficaria

(R. ficaria) the ‘petals are yellow, or red fading whitish. The

white flowers of Batrachium, or water crowfoot, show that they

are descended from ancestral yellow forms by retaining vestiges

of this color as honey guides at the base of the petals. The

cultivated Adonis annua, or pheasant’s eye, has showy orange

or red flowers, to which the black anthers offer a marked con-

trast. The most highly specialized regular flowers of this family

belong to Aquilegia, or columbine: They may be regarded as

the culmination of the buttercup type. The sepals are regular

and petaloid, while the petals are prolonged backward into a

hollow spur. The intermediate stages between the nectariferous

pit of the buttercup and the hollow petal of Aquilegia are

excellently shown in the extremely variable petals of A. auri-

comus: - Aquilegia canadensis produces scarlet flowers, which

are yellow inside and rarely all over, or occasionally they are

white. There are two other species in the northern flora which

exhibit a similar coloring, Lonicera sempervirens and Spigelia

marylandica, and the former is sometimes. yellow throughout.

A. canadensis is visited by bumblebees and humming birds. The

European A. vulgaris is blue, purple, or white.

The two zygomorphic genera, Delphinium and Aconitum, are

the most recent in their origin, and represent the highest stage

of development. attained by the Ranunculacezm. The sepals are

petal- like... The flowers are blue or revert to white, or, in Acozz-

tum reclinatum, are regularly white. High specialization, polli-

nation by bees, and. probably blue color, are here correlated.

Masters gives an instance in Delphinium peregrinum of perfectly

regular flowers having five sepals and five oblong stalked petals,

and also in oe the tubular petals may be repoa by flat

.ones.

At oe period ie insects began to visit the y S of the

Ranunculacez, the different genera were in a transition state

and only partially. differentiated. It is evident that one effect

of their visits was to flowe . This result

ONCE Ts SP F

has been reached in different v ; in tl i geni era, according

cw NAE o

No. 423+] NORTHERN. POLYPETALOVS FLOWERS. 209

as the pigments have been deposited .in the sepals, petals, or

stamens: But while insects have been instrumental in develop-

ing bright coloration, the particular colors of the different species

have been very largely determined by other conditions. The

yellow of Caltha palustris and Ranunculus has probably been

preceded by no other color than. green. These flowers are

visited by numerous flies, beetles, and the less specialized bees.

Syrphida are abundant, a family of flies with many species,

characterized by yellow markings, which would indicate that

yellow was especially attractive to them. Müller states that he

has seen Eristalis intricarius hover over the yellow flowers of

Caltha, as the males hover over the females, then suddenly settle

to suck honey or feed on the pollen. He also suggests that

their love of yellow may have arisen from their visiting flowers

of this color, and that sexual selection may have been guided by

this taste. The white flowers of Batrachium have evidently

passed through a yellow stage, but in other genera white or

purple has been developed directly from the primitive green, as

in Clematis. The blue flowers of Hepatica, Aquilegia, Delphi-

nium, and Aconitum, which are very attractive to bees, appear

to have passed through a yellow or white stage. The organs of

the flowers of the Ranunculacez, according to Masters, more

frequently revert to leaves than do those of any other family

except the Rosacec. = .

The Berberidaceze, Menispermaceze, Calycanthacez, and Lau-

racez contain but few species. The "shrubs of Berberis, or

barberry, have the wood as well as the calyx and corolla yellow.

The flowers are attractive to bees and flies. The foliage in

autumn is yellow‘and crimson. ` The three white-flowered species

of the Berberidacez occur in woodlands and bloom in May. The

Menispermaceze are woodland vines with small white flowers in

panicles, or green in Calycocarpum, where the petals are wanting.

The small flowers of the Lauracez are apetalous, but the calyx

is yellow. The species are aromatic shrubs and trees.

The original color of the Papaveracez, or poppy family, was

probably yellow. The sap also of many genera is yellow or red.

The beautiful flowers of Papaver are yellow or scarlet, with

a darker center. If the flowers contained nectar, this central

210 THE AMERICAN NATURALIST. [VoL. XXXVI.

marking would be regarded as a honey guide, but as they are

nectarless, and are visited by Andrenidz and Diptera for pollen,

it must have a different significance. It may be due to the

more abundant nutrition received by the central portion. In

the cultivated variety called Shirley the edges of the flower are

white and the center is crimson. If a plant receive a check

during growth by transplanting, the flower may revert to pure

white. Such color contrasts, however, render the flowers more

conspicuous. According to Kerner, honeybees do not visit

scarlet flowers, either because they do not distinguish this color,

or because it is unpleasant tothem. Ina garden in front of the

house where he lived the scarlet geranium, Pelargonium zonale,

and the narrow-leaved willow-herb, Epilobium angustifolium,

were in bloom at the same time and not far apart. He observed

that butterflies visited both indiscriminately ; but the honeybee

never paused in its flight over the scarlet flowers, though it

frequently sought the red-purple flowers of the willow-herb.

The flowers of Papaver are one to three inches broad, and hybrids

are sometimes twelve inches in width. The flowers of Argemone,

or prickly poppy, are also several inches in width. A. mexicana

is yellow, rarely white, and A. alba is regularly white. San-

guinea canadensis, bloodroot, has a white flower and red sap; $

but the flower is sometimes pinkish, as is indicated by the name

“red Indian paint.” The genera Stylophorum, Glaucium, and

Chelidonium have yellow sap and yellow flowers. x

A gfoup of irregular flowers formerly placed in a separate

family, the Fumariaceæ, are now included in the poppy family.

The species are fertilized by the long-tongued bees and flies.

The heart-shaped pendulous flowers of Bicuculla (Dicentra),

Dutchman’s breeches, are white or pink. The smaller flowers

of Capnoides (Corydalis) are pale yellow, while the larger are

bright yellow. | C. sempervirens is pink with yellow tips, and in

bud is greenish white, while the European C. solida has the

entire flower red. The influence exerted. sis insects upon the

pecie colors is very uncertain.

-"Fhe Cruciferae, ike the Umbelliferee, fomi: a very natural”

family ; th sely resemble each other and differ chiefly,

as Müller states, in the number and position of fhehopes inde

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 21I

and in the situation of the anthers in relation to these and to

the stigma. Many genera show evidences of retrogression in

the small white flowers and in the regularity with which self-

fertilization occurs. Pringlea antiscorbutica, which grows on

the stormy shores of Kerguelen Land, where no winged insects

can exist, since they would be swept into the sea, has reverted

to wind-fertilization. In sheltered places the. petals, which

usually are wanting, are frequently present, and “on the same

raceme some flowers may possess only a single petal, others two,

three, or four ; and the petals, though usually of a pale greenish

color, are occasionally adorned with purple." This family is

much less attractive to insects than the Umbelliferz, and is

visited chiefly by flies and the less specialized bees, such as

Andrena and Halictus, and by a few beetles and Lepidoptera.

Some flowers have no recorded visitors.

Of the 113 flowers, 2 are green, 54 white, 46 yellow, 1 red,

and ro purple; 20 of the 37 genera contain white and 16 yellow

flowers, but only 4 genera contain both yellow and white.

The white flowers are usually small, or even minute, and self-

fertilized as in Subularia (awlwort), Lepidium (cress), and Bursa

(shepherd's purse). The yellow flowers are larger, as in Sinapis

and Brassica, and frequently show a tendency to change to

white. In Rhaphanus (radish) the wild species, A. raphanis-

trum, is yellow, changing to white; while the garden radish,

R. sativus, is white or pink. In Roripa (nasturtium) the two

aquatic species are white, as well as the horse-radish, which

grows wild along streams; the other species are yellow. In

Draba four of the species are yellow and seven white, and

D. nemorosa is yellow fading to whitish. In other species the

calyx changes from green to yellow. It would seem probable

that in these genera the white-flowered species have been derived

from yellow progenitors ; yellow appears to have been very early

developed, . and was not improbably the original color of this

family.

The white flowers very frequently show a tendency to turn

pink or purplish, and three genera contain both purple and white

flowers. Malcolmia maritima:has pink-red flowers, changing to

violet purple. -This transition is beautifully illustrated by the

212 THE AMERICAN NATURALIST. [Vor. XXXVI.

rhizome of Dentaria bulbifera, which when taken from the earth

is as white as ivory, but, if placed in a glass of water and exposed

to the light of the sun, in a few days turns to a deep violet.

The cultivated varieties of the genus Mathiola (stock, or gilly-

flower), from Europe, present a wide range of colors, — as white,

yellow, red, violet,.and blue. The colors of Iberis (candytutt)

are white, red, and purple; and the marginal flowers of this

genus and .of Alyssum and Dentaria have the outer petals

enlarged, as occurs in the Umbelliferze. -= Hesperis tristis has

dark-coloréd flowers, visited. at night by insects, which are

attracted by the scent... |

The inflorescencé is in umbel-like racemes, which in fruit

become greatly elongated. The central green buds and the

surrounding flowers form a nearly flat surface and present a

variety of color contrasts. Kerner has described several differ-

ent classes. In Draba verna and. Thlaspi. rotundatum the

green buds in the center are surrounded by two colored rings, —

an inner one of small white flowers and an outer one of brown

or purplish fruits, to which the petals, now enlarged to twice

their original size, closely adhere. A variegated effect is thus

produced. In Thlaspi alliaceum and T. arvense the flowers of

the inner ring are white, but in the outer ring the green sepals

have changed to yellow, while the fruit remains green. In a

third group the inner ring is composed of flowers with colored

petals, which in the outer ring have faded to white. In Draba

aizoides there is a change from yellow to white, and in Arabis

cerulea from blue to white. A fourth group is represented by

A: thionema grandiflorum, in which the upper and under sides of

the petals are differently colored. The flowers of the inner ring

present the white upper side of the petals, but in the outer ring

the petals have changed their position so that the red underside

is now visible.

While the flowers of the Cruciferae show that conspicuousness

is correlated with the visits of insects, they furnish little evidence

that particular colors have been evolved by the selective tastes

of special insect groups. Both yellow and white flowers are

ee xi Syrphidæ, and they are also very frequent visitors to

the inconspicuous but scented flowers of Lepidium sativum.

No. 423] NORTHERN POLYPETALOUS FLOWERS. 213

The blue flowers of Arabis cærulea are likewise sought by flies,

while the nocturnal flowers of Hesperis tristis are dark-colored

and dependent upon their fragrance. The parti-colored appear-

ance of the flower clusters and the changes of color of individual

flowers would indicate that the guests were a miscellaneous group

of insects, with the color sense but feebly developed. How

comparatively unimportant the color of a flower may become is

well illustrated by Lepidium sativum. The small white flowers

can be seen only a short distance, and in rainy weather do not

expand. Yet they possess a strong scent, and Miiller found

them visited more frequently by insects than any other crucifer.

Throughout the species of this family the petals have not been

extensively modified either in form or color.

The flowers of the Capparidaceze resemble those of the

Cruciferze both in form and color. Polanisia graveolens (clammy

weed) has purplish sepals and stamens, and whitish petals.

There are three species of mignonette (Resedacez) in the

northern states. The flowers are especially attractive to small

bees of the genus Prosopis. These bees, which have an aro-

matic odor, are coal black, marked with yellow. They are

doubtless attracted by the sweet fragrance, which can be per-

ceived at a long distance. Reseda luteola (yellow weed) has

greenish-yellow flowers and is cultivated for its yellow dye.

R. alba has white flowers. In R. odorata the receptacle is at

first yellow, changing to orange red ; the petals are white or red,

and the anthers red. The perfume is intense. On flowers

growing in his window Müller collected five species of Prosopis,

besides other insects. The strong scent of these flowers probably

compensates for their small size and greenish colors.

The Sarraceniales contain two remarkable carnivorous families,

the Sarraceniacez, or pitcher-plants, and the Droseracez, or

sundews. The leaves, which are. adapted to trap insects, are

more conspicuous than the flowers. Sarracenia purpurea has

purple-veined leaves and large purple flowers, while S. fava has

green leaves and yellow flowers. The walls. of the orifice, and

the hoods, or lids, of the pitchers are the most highly colored

parts. Leaves without ascidia or pitchers are green, The

upper part of the pitcher of .S, laciniata. is pure white, marked

214 ` THE AMERICAN NATURALIST. | [Vor. XXXVI.

with a network of dark red veins. Of the five northern species

of Drosera, four have white and one purple flowers. The red

glandular leaves are far more prominent than the flowers. In

the common D. rotundifolia the small white flowers open at

midday, one or two at a time, and are visited sparingly by flies.

The order Rosales includes some thirteen families, differing

but little from each other. The Saxifragacez, according to

Engler, form the center of development, while the Rosaceze and

Papilionaceze are regarded as sister families. The primitive

character of many of the genera appears in the indefinite num-

ber of the stamens and pistils and their separate insertion upon

the receptacle. The order, however, exhibits an advance upon

the Ranales in that the carpels are more often united and the

ovary inferior.

Of the Crassulacez, or orpine family, two species are green,

two white, five yellow, two red, and two purple. The two green-

flowered species have small flowers and grow in wet places, and

in the case of Penthorum sedoides (ditch stonecrop) the petals

usually are wanting. The genus Sedum (stonecrop) contains

white, yellow, red, and purple flowers, with the honey readily

accessible to short-lipped insects. In S. acre the leaves are

yellowish green and the flowers yellow. While the blossoms

are small they are produced in such profusion that they completely

cover the plants, which grow in dense tufts and are sometimes

called “golden moss.” In S. telephium the petals are purple

and often the entire plant. The honey is more deeply concealed

than in the preceding species, and the flowers are in broad cymes,

which are conspicuous and facilitate insect visits. In the species

of Sempervivum the honey is still more deeply hidden and can

be reached only by long-tongued insects. The cymose flowers

are showy pink or purple. The petals of Sempervivum wulfentt

are sulphur yellow with a purple base, and are pollinated by

bumblebees. This purple coloring Miiller believed to be a rem-

nant inherited from a purple-flowered ancestor, from which the

sulphur-yellow form, which is unlike the primitive yellow form

of Sedum, had been developed by the selective influence of

bumblebees. This color change is not improbable, for the petals

of Arnebia cornuta, when they expand, — with dark

No.423.] MORTHERN POLYPETALOUS FLOWERS. 215

purple spots, which on the third day fade entirely away, leaving

the flowers bright yellow. The colors of the flowers of the

Crassulaceze are often correlated with the colors of the stems

and leaves.

The flowers of the Saxifragaceze are small and arranged in

racemes, cymes, and panicles, or solitary in Parnassia. The honey

in most species is easily accessible, and the visitors are chiefly

flies, which appear to have influenced the development of the

coloring, To these insects white and yellow, marked or dotted

with yellow, red, or purple, are thought to be especially attractive.

The relation of Diptera to the flowers of Saxifraga is shown in

the accompanying table compiled from Miiller and Knuth.

i SI ed Dır- | Hymenop-| Co reop- LEPIDOP- Wena:

: x TERA. TERA. TERA. TERA. e

S. rotundifolia .| White .....| 14 I — de 15

S elles. ey White La is 12 2 I I 16

S. aspera | White ....:. 2 — m Po 2

S oye ...|IwHe..... 9 2 2 on 13

Sater i ) White ss. 65 II 5 10 9t

d ME Hee ass Whit V. ay 15 3 3 3 24

A eerie. ...| Wie s 4 I — — 5

a ’

SS. muscoides . . | Greenish yellow 6 I I I 9

w”

S. aizoides. . . . | Golden yellow .| 85 20 8 13 126

-—

S. oppositifolia .| Purple ..... 4 = I 3 8

ee 216 41 mr -a.T

ak a TÉ

The genus Saxifraga is highly interesting because of the

colored dots upon the corolla of many of the species. The white

corolla of S. rotundifolia is sprinkled with round dots, the outer -

216 THE AMERICAN NATURALIST. > [Vor. XXXVI.

of which are intense: purple red; the inner are yellow, and the

anthers white. The snow-white petals of .S. sze//arzs are beset

with ‘purple. dots and adorned with two orange-yellow spots.

S. aspera and S. bxyoides are white, with numerous shining

yellow dots. S. aizoides has large golden-yellow flowers, marked

with numerous orange-red: dots; the nectaries and anthers are

also yellow. This is the most conspicuous species of the genus

and attracts 126 insects. S, oppositifolia has the honey more

deeply concealed and is carmine or ‘purple, and is diligently

visited by butterflies. S. Azreu/us, of Labrador, is bright yellow

with scarlet spots. In S. michauxii the three largest white

petals have a pair of yellow spots at the base, but the two smaller

are unspotted. The petals of S. geum are white, with a yellow

spot at the base and several smaller purplish spots in the middle.

As evidence that these markings are pleasing to Diptera, Müller

states that he saw many specimens of two drone flies, Sphegina

clunipes and Pelecocera scevoides, before sucking honey or eating

pollen, poising before the dotted flowers of S. rotundifolia as if

delighted by their appearance.

In his AZgeub/umen Müller has tabulated his observations

upon the relations of Diptera to the different colors of flowers.

Most anthophilous species and families of flies made a much

larger number of visits to white and yellow than to red and blue

flowers. The Bombylidez, which are suctorial only, showed a

preference for red and blue to white and yellow in the proportion

of 75 to 25; and the genera Volucella and Rhingia, of the

Syrphidz, showed a similar inclination in the proportion of 7744,

to 22,5. The less specialized Diptera were by far the most

= common on white and yellow; but as the proboscis increased

in length and the species confined themselves more strictly to

flowers, the percentage of visits to red and blue flowers increased

from 14,1; in the Muscidz to 294, in the Syrphide, and to 75

in the Bombylidz. The AEA as flower visitors surpass all

other Diptera, both in numbers and importance. The percentage

of visits to yellow and white was 69,5, and to red and blue

flowers was 2945. The species marked yellow, whether short

or long tongued, were two to three tioesashbamant t ae white

Meg cnn h > flov None of

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 217

which do not restrict themselves to flowers, were observed on

red and blue blossoms. Due consideration should. be given to

the limitation of the visits of flies by the form of the corolla,

by scent, by the acuteness of the species in finding the honey,

as well as by its adaptations and habits, and to the fact that in

the case of several families the above conclusions are based on

comparatively few observations.: .

Chrysoplenium alternifolium has no petals, but the sepals are

bright yellow and the disk is also yellowish. | Like the species

of Saxifraga, it is visited chiefly by flies. The petals of Par-

nassia are white, with greenish veins. |. palustris is described

as a “deceptive flower.” It contains numerous yellow glandular

bodies which appear like minute drops of honey, by which flies,

especially Syrphidze, are deceived. The flower contains a small

amount of honey. In Hydrangea the cyme is made conspicuous

by the enlargement of the calyx of the marginal flowers. In

H. hortensia, cultivated from eastern Asia, the flowers are nearly

all neutral and enlarged, and at first are green, changing directly

to pink or purple. “The variable character of Hydrangea

flowers," says a writer in the Journal of Horticulture, “when the

plant is grown under certain conditions, has for many years

been a sort of horticultural puzzle, which is still far from being

satisfactorily solved. Plants with bright pink flowers, and those

with flowers of a tolerably good blue, are not infrequently met

with in positions near each other and apparently in soils exactly

alike." A change of color in the flowers from pink to blue is

sometimes caused by a change of soil, as the removal of a plant

from a peaty soil to one of opposite character. The addition of

iron to the soil frequently appears to produce the desired color

change, though time is always required, the blué coloring not

developing until the second year. Saxifraga pennsylvanica and

Heuchera americana have green petals. Several species of

Saxifraga are proterogynous, and the flowers in passing from

the female to the male stage increase greatly in size.

The Grossulariaceze (gooseberry family) contain only a pe

genus, Ribes. Of the northern species four are green, six white,

one yellow, and two'greenish purple. — o of the species are

greenish in part. In“ Ribes. lpi th e flowers are green

218 THE AMERICAN NATURALIST. [VoL. XXXVI.

and the male greenish yellow and much more conspicuous,

though of no larger size. By this device insects are induced to

visit the male, or pollen-bearing flowers, first. The petals of

R. sanguineum change from white to pink, and of R. aureum from

yellow to carmine. This color change, says Müller, also occurs

in several species of Fuchsia and Lantana and enables the more

intelligent bees to economize time by determining instantly those

flowers which no longer contain nectar.

There are four green, thirty-five white, thirty-nine yellow,

thirteen red, and four purple flowers in the Rosacee. The green

flowers are small and apetalous, as in Alchemilla; the white

flowers are very generally tinged or tipped with red, and

vary from small in Spirzea to large in Rubus. Of the twenty.

genera in the northern states, twelve contain white flowers. Of

the four species in Spirza, three are white and one, .S. tomentosa

(hardhack), is rose, or rarely white. The flowers contain nectar

and attract numerous flies, beetles, and Hymenoptera. Beetles

are very frequent visitors to the small white flowers of Aruncus

aruncus. The genus Rubus contains seventeen species, all of

which are white except the purple R. odoratus and the pink

R. arcticus. R. strigosus, or the wild red raspberry, has small,

erect white petals, and the visitors are much fewer than to the

blackberry, R. villosus. Though the flowers of the blackberry

are also white, their increased conspicuousness secures them a

much larger company of visitors, which is not far from one

hundred. The petals are broad and flat, and the panicles large

and numerous. Dalibarda repens, a woodland plant, produces

both cleistogamic and open white flowers which are seldom

fertile. The common field strawberry blooms in May and June,

when the white blossoms contrast with the green meadow, and

the growing grasses are as yet too short to conceal them.

Of the twenty-four species of Potentilla, two, P. arguta and

P. tridentata, are white and are probably derived from yellow-

flowered ancestors. They both grow in dry, rocky places, The

other twenty-two species are yellow and are visited by small

bees and flies. The prevalence of yellow coloring in this primi-

tive genus would indicate its early development in the rose

family. — M —— À —ÀÁ

No. 423} NORTHERN POLYPETALOUS FLOWERS. 219

strawberry), which is certainly closely allied to the line of the

field strawberry. | Wa/dsteinia fragarioides, or the barren straw-

berry, which in leaf habit has the aspect of Fragaria, is likewise

yellow. The simpler species of avens are yellow or white, while

those with the honey concealed are purple. In Geum rivale

(purple avens) the petals are purplish orange and the calyx

brown purple. The honey can be reached only by long-tongued

bees and flies, such as Bombus and Rhyngia. In Comarum

palustre (marsh cinquefoil) the petals are purple and the sepals

also inside. The plant grows in swamps and is visited by flies

and less frequently by bees. The primitive color of both these

species seems to have been yellow. The purple-flowering rasp-

berry, Rubus odoratus, is however derived from white-flowered

progenitors, for in New York at Pine Hill, Ulster County, a

white-flowered form occurs, and the white species of this genus

are frequently reddish or purple.

The ten species of Rosa in the northern states are rose or

pink, varying to white. Though the flowers contain no honey,

their color and fragrance attract many insects to the ample store

of pollen. Several species have odors so peculiar that they may

be recognized by them alone. The theory of Delpino that the

distribution of this genus was determined by the range of certain

families of Coleoptera has been disproved by the observations

of Müller and others who have found bees very frequently upon

the flowers. Beetles may often be collected, some species of

which devour the petals bodily. The flowers of Rosa canina

. gain increased conspicuousness by always turning towards the

sun. In Germany they are visited by eight beetles, six bees,

and two flies.

The family of the Rosacez is very prolific in hybrids, and

more than two hundred have been observed in nature, Innu-

merable hybrids of Rosa have been produced under cultivation

which display countless combinations of yellow, red, and white,

—as white tinged with yellow or pink, yellow fading to white

or shaded with rose, or pink changing to white, rose shaded with

yellow, or intense coloring such as crimson, carmine, and scarlet.

In one variety on the same root there are produced every inter-

mediate shade between white and red, and in another the colors

220 THE AMERICAN NATURALIST. | [Vor. XXXVI.

vary from yellow to crimson. There is no blue rose, but a

white variety of Rosa rugosa from Japan has a bluish tinge.

Blue does not, indeed, occur in this family. While red is

common, none of the Rosacez are adapted to Lepidoptera.

The visitors are a miscellaneous company of flies, beetles,

and Hymenoptera.

This family exhibits a marked tendency in stem, leaf, bud,

flower, and fruit to develop reddish coloration, — a tendency

which is probably due to the chemical constitution of the sap.

The smaller and less specialized Rosaceæ are yellow and white,

and are visited by a variety of short-lipped insects. The increase

of the white flowers in size and conspicuousness is usually

attended by red coloration. Owing to the chemical constitution

of the nutritive fluid, probably to its acidity (for when the petals

of a rose are treated with ammonia they become blue), there has

been no opportunity for the development of blue coloration by

insects. With the enlargement of the perianth and the increased

flow of sap, red tints have tended to appear by process of oxida-

tion. The correlation of red coloring with an increased flow of

the sap is well illustrated by the galls of the wild rose tree,

which are often “as rosy as the rosiest apple." An abnormal

flow of sap is caused to the part stung by the insect, and red

coloration is due to the action of light, for it is of no service to

the plant. Again, when the flowers of Crat@gus coccinea are

stung by the gallfly the different organs all become red, and the

change in coloring is accompanied by an increase in size. In

some instances, according to Darwin, red colors indicate greater

vigor on the part of the plant, and I have also observed that the

dwarfing of red flowers under cultivation may cause them to

revert to white. ©

There is nothing more beautiful in the vegetation of the tem-

perate zone than an orchard laden with expanding blossoms.

The great quantities of flowers form billowing banks of white-

ness, tinged with rose and flecked with the vivid green of the

unfolding leaf buds, from which exhales the well-known’ sweet

fragrance of the: apple blossom. Of the Pomacez, or apple

mol twenty-seven species. are white and five red or partially

are reg and usually clustered. —

No. 423.] NORTHERN POLVPETALOUS. FLOWERS. 221

of Sorbus (mountain ash) and Crataegus (thorn) are very attract-

ive to beetles, though often visited by flies and bees. The pear

and apple are sought by fewer beetles and depend chiefly upon

Hymenoptera and Diptera. The native species of apple are

rose or pink, but readily change to white. The so-called bloom-

less apple has small green petals resembling sepals.

Twenty species of the Drupaceze, or plum family, are white,

and one, the peach, is pink. The blossoms of many plums

appear in early spring in advance of the leaves and often com-

pletely wreathe the limbs. The flowers are visited by a great

number of Andrenidz, and in less than two hours I collected

one hundred and twenty specimens on a Japan plum, and these

were but a small part of those present.

The Mimosacez, Czesalpiniaceee, Krameriacez, and Papiliona-

cea are often united in one family called the Leguminose,

because the simple pistil becomes in fruit a legume. The

Mimosa family connects the rose family with the Papilionacee.

It is confined chiefly to the tropics, where the species are very

numerous. The flowers are small, perfect, and regular, with the

stamens distinct as in the rose family, or monodelphous as in the

Papilionaceze. In the northern species the colors are white,

yellow, or pink. The filaments are long and ciidimncian and

are more conspicuous than the petals.

The pea, bean, clovers, vetch, and a whole host of leguminous

allies are grouped together in the Papilionaceze, —a name derived

from the fancied resemblance of the flowers to a butterfly.

Occasionally perfectly regular flowers occur by reversion, as has.

been observed in Laburnum. For the most part, nine of the

ten stamens unite to form a tube, at the bottom of which lies

the honey, if present ; four of the petals interlock around this

tube, while the fifth, called the “standard,” is broad and erect

and bright colored to attract the attention of insects. The

flowers are fertilized by bees, which rest upon the wing petals,

bracing the head against the standard and bringing the ventral

side of the body in contact with the stigma and pollen. The

irregularity of the flowers is due to their horizontal position and

the unequal strains to which the petals are subjected. In the

case of certain species bees alone are able to depress the keel

222 THE AMERICAN NATURALIST. [VoL. XXXVI.

and obtain the nectar, and in their absence the flowers fail to set

seed. Slight imperfections frequently permit flies and. butterflies

to steal the honey without rendering any service in return ; and

it has been suggested that the numerous species may be due to

the efforts of the plants, metaphorically speaking, to remedy

these defects.

The inflorescence is mainly in heads and racemes, but the

effect of the individual flower cluster is often magnified mani-

fold by the massing of the plants. In worn-out fields the vetch,

Vicia cracca, often takes entire possession of the soil, forming

large patches of purple blue. Similar effects are attained by the

clover and wild lupine. Of the 197 species in the northern

states, 39 are white, 33 are yellow, 13 red, 88 purple, and 24

blue. The predominance of blue and blue purple are believed

to be due to the preference of bees for this color. The well-

known experiments of Lubbock have shown that the honeybee

can distinguish between colors. Scarlet, fire red, and all lurid

colors are avoided by the honeybee. Blue, violet, and red are

most attractive, followed by various shades of purple, yellowish

white, and white.

An examination of the genera in which more than two species

occur shows that they are rarely monochromatic; one or more

species are usually differently colored from the rest. In Bap-

tisia three species are white, two yellow, and one blue; in Trifo-

lium (clover), four species are white, three yellow, four red, and

three purple; in Psoralea two species are white, six purple, and

three blue; in Astragalus seven species are white, three yellow,

twelve purple, and one blue; and in Meibonia (tick trefoil), two

species are white, one red, and nineteen purple. It is probably

more advantageous in these genera for a part of the species to

be of one color and a part of another than for all to be blue.

When species are closely allied bees tend to visit them indis-

criminately, as may be observed in the buttercups and golden-

rods. But even in these cases they exhibit a preference to keep

to a single bom and would be greatly aided by differences in

the coloring.

. . Many of the flowers of the Papilionaceze i in fading undergo a

change of position and color, in the white clover the white

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 223

central flowers contrast with an older outer ring of rose-colored

flowers. In the yellow clover the newer flowers contrast with a

ring of chestnut brown. In Vicia cracca the older flowers bend

downward and turn from violet blue to dark purple; while the

purple flowers of Desmodium become green in withering.

Striking color contrasts are also presented by the individual

flower. The wings of the white corolla of Vicia faba (bean)

are marked with two black eye-spots, and in the sweet pea and

cultivated lupines the combinations of color are innumerable.

Astragalus vesicarius has yellow blossoms in the Tyrol but

violet on the limestone mountains of Hungary.

Nine families belong to the order Geraniales. In the genus

Geranium of the Geraniacez the larger and more conspicuous

purple flowers are visited abundantly by insects and have nearly

or quite lost the power of self-fertilization. The smaller flowers

are paler or white, attract few insects, and self-fertilization regu-

larly takes place. Geranium pratense has been seen to produce

on the same plant, when cultivated in a garden, both white and

blue flowers. G. robertianum, or red robin, has ribbed red-

purple petals and, "notwithstanding its disagreeable odor, is

sought by bees as well as flies and beetles. In the pink flowers

of Erodium cicutarium the upper petals are marked with dark

lines, which serve as pathfinders. According to a series of

forms figured by Knuth these markings vary greatly, from a

few lines to spots and markings on all five petals? In the genus

Pelargonium from the Cape of Good Hope, this office of the

upper petals becomes very highly developed; P. tricolor has the

three lower petals white and the two upper crimson, each with

a dark spot at base. The flowers of the Geranium always turn

towards the sun. 4

Most of the northern species of Oxalis of the Oxalidacez are

yellow and possess an acid juice, but O. acetosella, which grows

in open woodlands, has large, pretty white flowers, veined with

pink. It is sometimes called * wood sour,” as druggists obtain

from it salt of lemons. The plant is of social habit, and the

flowers are quite conspicuous, yet are rarely visited nd flies,

lDarwin. Animals and Plants under Domestication, vol. i, p- 404-

2Knuth. Handbuch der Blutenbiologie, Bd. i, p. 1

224 THE: AMERICAN NATURALIST. [VoL. XXXVI.

beetles, or bees. Cultivated species of Oxalis are red or rose.

O. flava is yellow, edged with: red; and ©. versicolor is white

above and red beneath, so that the blossoms are white in the

sunshine, and red when rolled up in the shade.

Like the Papilionacez, the flowers of the Polygalacez stand

horizontal, and the petals are more or less united into a tube

with a carina and ale. The largest-flowered species is the

fringed Polygala, P. paucifolia; which in spring produces beau-

tiful rose-purple crested blossoms, "with an occasional white

variety. It is attractive to butterflies as well as bees. On the

Alps, Müller found one species of Polygala fertilized entirely by

butterflies. Most of our species have very small flowers, which are

either greenish purple, or yellow changing to green in drying.

In the capability of the leaves to develop bright colors, and

in the minute and reduced flowers attractive to Diptera, the

Euphorbiacee, or spurge family, resemble the Aracez, though

it has not the remarkable adaptations for fertilization of. Arum.

The spurge family is of immense size and of very wide geograph-

ical distribution. The flowers are minute and have undergone

much reduction. They are usually apetalous, and the entire

perianth may be wanting, as in Euphorbia, where a single stamen

represents a flower, and the flower cluster with its colored invo-

lucre was mistaken by the older botanists for a single flower.

The genus Euphorbia is attractive to flies, though also visited

by beetles and Hymenoptera; and in response to their visits the

inflorescence in certain species possesses bright colors, honey,

and a honey-like scent. The colors of many northern species are

green, but a part are white, yellow, rose, or red. On £. . cypa-

rissas, the common cypress spurge, naturalized from Europe,

and which has escaped from cultivation, the bracts are yellow.

In the Alps, Müller collected twenty-one flies, one beetle, four

Hymenoptera, and three Lepidoptera on the flowers, and in

middle Germany he found many more Coleoptera and Hymenop-

tera. The genera Ricinus and Croton are extensively cultivated

for the tropical aspect of the magnificent mode Rino com-

munis is anémophilous.

In the order Geraniales, to which the families, just considered

belong, and in the succeeding. order, the Sanders, the flowers

No. 423-] NORTHERN. POLYPETALOUS FLOWERS. 225

mark an advance upon the Rosales in that the cyclic arrangement

prevails and there is an imperfect union of the carpels. The

families of the Sapindales differ widely and have been divided

into numerous subseries. |The Empetracez, Buxacez, and

Limnanthacee are represented in the northern states by only

four species. The flowers are small, and probably partly

primitive and partly reduced.

The Anacardiaceze and a number of allied families are of much

interest, as the inflorescence has been but little modified by

insects. The species are mainly trees and shrubs with green,

white, and greenish-yellow flowers, which are visited by- Diptera

and the smaller and less-specialized Hymenoptera. The Ana-

cardiaceze, or sumac family, have small, regular flowers in dense

panicles. Both the sepals and petals are present, but the flowers

are greenish or greenish yellow, though sometimes tinged with

red as in the smoke tree.: They secrete honey and are attractive

to flies and Andrenide. On the smoke tree, Cotinus cotinus

(Rhus cotinus), there have been collected six flies, one beetle,

and ten Hymenoptera. . The species tend to become dicecious,

and Cofzmus cotinus (Rhus cotinus) shows all transitional stages -

between staminate, hermaphrodite, and pistillate flowers. While

the visits of insects have not developed bright colors, the leaves

of Rhus radicans and the fruit of Re aromatica are red in

autumn, and the wood of several species is orange yellow.

R. glabra sometimes has the whole or a part of the flower

cluster changed into-small leaves.

The various species of Illicaceze, or holly, have small white

flowers, with freely exposed honey. . The berries of Merx verti-

ciliata are bright red, or rarely yellow, or even white. |The visit-

ors are similar to those of the preceding family. The Celastraceze

is composed of trees and "abes with small, — green, yellow,

and purple flowers. |

Five species are. green, | theod edis ind one nti in the

Aceracez, or maple family. The green flowers of Acer saccha-

rinum are without petals:. In A. rubrum (red maple) the diœ-

cious flowers are crimson, and-the visitors are Andrenida and

Diptera. There is no reason to:suppose that the crimson color-

ing has been developed by insects, for.the entire flower, the leaf

226 THE AMERICAN NATURALIST. [Vor. XXXVI.

buds, the young leaves, and the twigs are of this color, while the

bark yields a purple dye. The flowers appear in early spring

before the leaves and were formerly anemophilous. A. spicatum

blooms later, and the compound racemes of greenish-yellow

flowers are large and erect. Miiller states that dull-yellow

flowers are avoided by beetles, but I have observed many

beetles, as well as bees, upon the inflorescence of this species.

Closely allied to the maple, but of more recent origin, is the

genus /Esculus. The flowers seem to have possessed special

capabilities that led to their adaptation to bumblebees. Among

ornamental trees few present a more stately and splendid appear-

ance when in blossom than the common horse-chestnut, sculus

hippocastanum. It has large white flowers in crowded panicles,

with the petals marked with yellow, which in a few days changes

to orange and then to crimson. I have observed the honeybee,

four species of Bombus, and one Andrena as visitors. Of the

four other species of this genus three have yellow petals, and

one, ZEsculus pavia, bright-red flowers an inch in length.

Impatiens biflora, of the Balsaminacee, is orange yellow,

spotted with reddish brown. One of the petals forms a spurred

sac. August IO I examined a large number of flowers ; none of

the spurs were perforated, and they were visited legitimately

by Bombus vagans, which made from seven to twelve visits per

minute. August 23 and.27 I found hundreds of the flowers

perforated, and both honeybees and bumblebees stealing the

nectar. If the Impatiens, fitly called *touch-me-not," could

speak, what a protest it would utter! Various Diptera are

attracted to the outside of the sac by the bright colors.

The colors of the Vitacez, or vine family, are green, closely

resembling the foliage, and depend entirely upon their strong

fragrance to attract insects. The inflorescence is in dense pani-

cles. The calyx is minute, with the limb nearly obsolete. The

green valvate petals form a hood over the stamens and never

expand, but fallaway by separating at the base and coiling spi-

rally upward. The odor, which resembles that of mignonette,

can be petes at a long distance. “In a journey up the

anube,” says Kerner, “through the part of the valley called

the Wastan, wih i neta slopes, fund the ai of the

No. 423.] WORTHERN POLYPETALOUS FLOWERS. 227

whole valley, even that above the water, so filled with the scent

of vine flowers that it seemed almost impossible they should be

so far off. And yet the nearest vines on the banks were one

hundred yards above the water and at least three hundred yards

from the boat. Afterwards I found, when wandering through

the vineyards, that the smell of the flowers close at hand was

much weaker than at a distance, and was forced to the paradox-

ical opinion that with increasing distance and diffusion over a

wider area the scent does not diminish but waxes stronger.”

Under cultivation the species are usually perfect, but when

growing wild are mostly unisexual. Cross-fertilization, accord-

ing to Knuth, is occasionally effected by the wind. As visitors

to the flowers of Vitis vinifera there have been observed the

honeybee, and various species of Halictus ; while of Coleoptera,

notwithstanding the green flowers, there have been enumerated

twenty-one species. Müller's statement that “beetles are only

or mainly attracted to flowers by bright colors” does not agree

with the many species observed on the flowers of the vine.

Union of the carpels prevails in the order Malvales, which

includes the two families Tiliacez and Malvacee. A part of

the genera with the carpels distinct, or but slightly united, still

preserves a more primitive stage in the evolution of the flower.

As in the vine family, the flowers of the linden, or lime tree,

depend on their strong scent, rather than upon their coloration,

to attract pollinators. The greenish-white flowers of Tilia

americana are in small drooping, cymose clusters, sheltered

beneath a floral bract. The thickened concave sepals secrete

and contain the honey, which a pubescent fringe prevents from

escaping. A high value is placed upon honey made from the

linden ; in some localities the leaves are also covered with a copi-

ous secretion of a sweet liquid, and sugar has been made from

the sap. The strong scent, which is more noticeable at a short

distance than close to the tree, is very attractive to the honey-

bee, many of which I have seen at work on the flowers in Maine.

On T. ulmifolia in Germany Müller and Knuth collected eleven

flies and seven bees. The absence of beetles I attribute not

to the dull color of the petals, but to the inaccessibility of the

drooping inflorescence, covered both by the leaves and the floral

228 THE AMERICAN NATURALIST. [Vor. XXXVI.

bract. The leaves of a part of the species are bicolored, green

above, and white, because densely woolly, beneath.

One-half, or thirteen, of the northern species of Malvacez

are pink or red, four are red purple, five yellow, and four white.

No other polypetalous family has. so large a percentage of the

flowers red. Both in temperate and tropical regions the blos-

soms are remarkably large and showy, and are usually yellow or

red. The chief agents in intercrossing are bees. Malva rotundi-

folia has white flowers ‘striped with pink, and slender pink stig-

mas, while 77. moschata has either pink or white blossoms. ' The

marsh mallow, A/thea officinalis, well known for the use of the

mucilaginous root in confectionery, has pink flowers an inch

broad; and A. rosea, the hollyhock of the garden, displays

white, yellow, rose, crimson, purple, and black hues. A single

species of Abutilon with yellow flowers has become naturalized

from Asia. In South America the natural fertilizers of Abutilon

are humming birds. The genus Hibiscus consists of splendid

flowers of immense size. In color they ‘are yellow, red, or

white ; the yellow forms have a purple or blackish eye, and the

red a dark center... The common cotton, Gossipium herbaceum,

one of the most valuable of nature’s plant productions, has yellow

petals, and the floral and extra-floral nectaries attract many

insects: as well as the ruby-throated humming bird.

Within the order Parietales the placentz are parietal; “and

the floral evolution,” says Engler, “has already reached very

complicated floral types." The Theacez (tea family) is mainly

of tropical distribution, consisting of shrubs or trees with large

solitary white flowers. Three species occur in the southern

states. The handsome white flowers of Stwartia malachodendron

have purple filaments ; in-S. pentagyna the petals are cream color

and the sepals redditi outside. To this ay end the tea

plant and the Camellia. -

The Hypericaceae have died ulottivat flowers, as

twenty-two species are yellow and two red. The larger flowers

of Hypericum are bright or orange yellow, as Z7. ascyrum, while

the smaller are often pale yellow. |The leaves, sepals, and petals

of. ce salo pice gta vengo dots.

H. perfi . Europe, contains no honey, ‘but

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 229

the large, black-dotted, bright-yellow flowers attract many flies,

which feed on the pollen. The smaller-flowered species are

rarely sought by insects, and self-fertilization is possible through-

out the genus. The two small, reddish-flowered species belong

to the genus Triadenum. The petals of 77zadenum virginicum

are rose colored, with deeper veins, and do not exceed the calyx ;

the honey glands are orange and the anthers bright yellow

with white filaments. The flowers are very sparingly visited by

insects, and after repeatedly watching them I have observed

only two bees and three flies. The stem, the capsule, and often

the leaves are a deep crimson, as is often the case in Hypericum.

The change in the petals from yellow to red is probably corre-

lated with the red coloration of the plant rather than due to

insect selection. In Hypericum perforatum, which frequently

has the foliage crimson colored, the yellow petals are tinged

with rose in the bud, which fades away as the flower opens. In

Epilobium palustre reddish flowers are more common when the

stems afé red than when they are green.

In the Cistacez, or rock-rose family, the genus Helianthemum

(frostweed) has large yellow flowers, which throughout the day

follow the sun. They are ephemeral, but the plants remain in

blossom for a long time, as there is a succession of flowers, as

in the crucifers. . There is no honey, but the pollen attracts a

miscellaneous company of insects. As is frequently the case

in pollen flowers, pollen falling from the anthers is not lost but

lodges in the concave petals. The small greenish flowers of

Lechea (pinweed) are self-fertilized.

The violet family in the northern states contains 7 white,

6 yellow, 4 purple, and 17 blue flowers. About 300 species of

this family and 150 of the genus Viola have been described,

Müller regards yellow as the original color of the violets, The

small, short-spurred V. dzfora, which in the Alps is fertilized by

flies, has yellow flowers; the large-flowered V. tricolor, variety

alpestris, presents all stages in the passage from yellow to blue.

« Many plants have flowers which are yellow throughout; in

others the flowers are yellow when they open but change gradu-

ally to blue ; and in others the change to blue occurs immediately

after opening, or even before." V. calcarata, which has a spur

230 THE AMERICAN NATURALIST. [VoL. XXXVI.

from 13 to 25 mm. in length and is fertilized by Lepidoptera,

chiefly by Macroglossa stellatarum, displays, according to Kerner,

in the meadows of the western Alps a blue corolla, and a yel-

low corolla in the eastern Alps. Darwin transplanted a large,

uniformly colored, dark-purple variety of the pansy while in full

flower, and it subsequently. produced smaller flowers, with the

lower petals yellow. The white forms appear to be derived from

blue progenitors, for the former are purple veined and may be

tinged with violet. Blue species of Vjola which often have

white flowers are V. palmata, V. obliqua, and V. odorata, and

in various stages V. canadensis. Pathfinders among white

northern polypetalous flowers are of infrequent occurrence.

The most prominent instances are Saxifraga michauxii, with

three of the unequal white petals marked with yellow ; Oxa/zs

acetosella, white with reddish veins; Parnassia, with greenish

veins ; Magnolia macrophylla, ZEsculus hippocastanum, Lespedeza

hirta, and L. capitata display purple spots. In the white violets

the purple honey guides are highly developed, which may be

best explained by supposing them descended from blue-flowered

ancestors with darker veins similar to existing species.

The beautiful and richly variegated varieties of V. tricolor, the

garden pansy, are due partly to selection and partly to hybridi-

zation. The corolla may be pure white, yellow, red, blue,

purple, or black; or there may be combinations of yellow and

blue; yellow and red; yellow, blue, and white; and blue and

white. According to Strasburger, in the pansy the cells of

the epidermis of the petals contain both violet sap and yellow

granules. “The striking diversities in color presented by dif-

ferent parts of a given petal depend wholly upon combinations

of these two elements of color, — namely, violet sap and yellow

granules."! Places which are devoid of both these elements are

white, as the light is refracted and reflected by the intercellular

spaces containing air. When sections of white roots of a violet

plant are exposed to the air they change to violet ; and the leaves

of V. cucullata when grown at alpine altitudes become yellow.

While the violets vie with the roses in popular favor, they do

not prove attractive im an equal degree to insects. In MaineI ©

1 Goodale. ee NE i

No. 423.] WORTHERN POLYPETALQUS FLOWERS. 231

have rarely observed visitors to the white and blue varieties, but

on the yellow V. rotundifolia, which blooms early in May, I

have seen many bees belonging to the genera Bombus, Nomada,

and Andrena. Besides the conspicuous flowers, which are often

unfruitful, many species produce cleistogamic flowers, in which

the petals are reduced to. mere scales, but the green sepals

remain essentially unchanged. Occasionally the violets bloom

a second time in late fall. "n

The failure of the yellow plastids to develop and the predomi-

nance of the colored cell sap produce blue flowers, and the non-

development of both color elements results in white blossoms.

In these color changes other ecological factors are more impor-

tant than insects. For instance, of the yellow-flowered species

one is visited by flies, another by bees, and the either yellow or

blue V. calcarata is adapted to Lepidoptera.

The passion flowers are mostly natives of South America and

are fertilized by humming birds and bumblebees. The calyx

of the common Passiflora caerulea remains green until it has

attained nearly its full size, when it changes to blue and white;

the petals are white; the outer corona consists of several rows

of blue filaments banded with white, but the inner corona is

smaller and unmarked. Fritz Müller considered the corona to

be of service in detaining small insects and keeping them caged

for humming birds.

The Cacti, of which over a thousand species are natives of

America, are especially abundant on the sandy plains of Mexico.

The nearest living representative of the ancestral stock of this

family is the genus Pereskia, which still possesses leaves of the

usual form. “The earliest derived line was Opuntia. From

the primitive Opuntia forms the columnar Cereus line was

derived, with its numerous generic branches and diverse habits.

Low down upon the columnar Cereus line the Echinocactus line

branched out, which gave rise later to Mamillaria and still later

to Anhalonia” (Botanical Gazette, Vol. XXVII, No. 3, p. 228).

The flowers of the Cacti are solitary and sessile, with the sepals

and petals and stamens numerous and spirally arranged. In size

they are usually large and showy, as in Cereus grandiflora, queen

of the night, where they are 20 cm. in diameter. In Pereskia

232 THE AMERICAN NATURALIST. [VoL. XXXVI.

the flowers are white or yellowish; in Opuntia yellow, or in

O. opuntia and O. humifusa yellow with a reddish center; in

Cereus in the nocturnal. forms white, in the diurnal species red

or crimson; in Echinocactus yellow, white, and crimson; in

Mamillaria yellow, pink, white, and purple; and in Anhalonia

white, rose, and purple. The spines of the Cacti are often very

beautiful objects and exhibit a great variety of color, as white,

black, yellow, red, and purple. The stems, while usually green,

are frequently nearly blue; the fruits are green, red, and purple.

The Thymeleacez and Eleagnacez in the northern states

have no petals, but the calyx is v; enlarged; prolonged into a tube,

and colored yellow, red purple, or white. In Europe Daphne

mezereum has the red-purple flowers visited by long-tongued

bees, flies, and butterflies ; in the Alps the white or red flowers

of D. striata are strong scented in the evening and are visited

by moths. —

The petals are wanting in the single green species of the

Lythracez, or loosestrife family, Didiplis diandra, which has

minute solitary flowers. The loss of the corolla is doubtless

due to retrogression. In Ammannia the flowers are small and

the purple petals fall away as soon as they expand, and in the

southern A. latifolia they are wanting. The purple flowers of

Lythrum vary from small to large. The eighteen possible ways

of fertilization in the red-purple trimorphous flowers of Lythrum

salicaria were made the subject of a long series of laborious

experiments by Darwin, which resulted in breaking down the

last barrier between species and varieties. It is a singular fact

that the pollen of the longest stamens in this species is green,

and of the middle-sized and shortest, yellow. The legitimate

fertilizers are long-tongued insects. In Parsonia, or Cuphea as

it is better known, the flowers are pink or crimson, and i in a cul-

tivated species from Mexico, C. platycentra, there are no petals,

but the tubular calyx is bright vermilion with a violet border.

The Melastomacez are represented in North America only

by the genus ; Rhexia, with handsome purple or yellow flowers.

The fma is most abundant in South America. In Heeria

a part of the stamens are devoted to rendering the flowers

conspicuous and a pen to producing pollen. - Cyanophyllum

No. 423.] NORTHERN POLYPETALOUS. FLOWERS. 233

metallicum, from Central America, has magnificent leaves, purple

beneath and metallic blue above.

In the Onagracez, or evening primrose family, there are three

green, fourteen white, twenty-four yellow, ten red, and six purple

flowers. The three small green flowers belong to Ludwigia,

and the petals are either minute or absent. Many of the white

flowers show a tendency to change to pink or red. The large

white flowers of the three northern species of Anogra all turn

pink with age. Several species of Gaura also change to red.

G. coccinea is red, turning scarlet in fading. The two species

of Circæa (enchanter’s nightshade) are delicate, colorless little

plants, , which grow in damp, shady woods and have small white

flowers, fertilized by flies. The yellow flowers are in part diurnal

and in part nocturnal as in Onagra biennis. Yellow is a much

rarer color than white in flowers expanding at night.

With a single exception the flowers of Epilobium are red or

red purple. Æ. angustifolium, called **fireweed " (as the plants

spring up in abundance in newly cleared or burnt lands), has

very conspicuous flowers in long, terminal racemes. The sepals

and petals are purple red, except the lower sepal, which is white,

and rarely the whole flower reverts to white. The pollen is

greenish purple, and the stems and seed vessels are purple.

Honey is secreted by a green, fleshy ring at the base of the

style. Bumblebees are the most important agents in intercross-

ing and will visit as many as thirty-seven flowers in a minute.

I have collected on this species twenty-one Hymenoptera, five

Diptera, three Lepidoptera, and two beetles. In the smaller

species of Epilobium, such as E. palustre and E. lineare, the

small flowers are reddish or white, both kinds occurring on the

same plant. When the stems are purple the flowers are more

often red than when the stems are green. There are few vis-

itors, and self-fertilization regularly occurs. The only northern

white species of Epilobium is Æ. a/Pigum, which has small

white flowers which are regularly self-fertilized. In Æ. %irsu-

tum the four white stigma lobes form a cross on the red field

of the petals.

The exotic genera deserve special mention. In Zofezza

racemosa, from Mexico, there are on two of the petals dry,

234 THE AMERICAN NATURALIST. | [Vor. XXXVI,

shining, yellow bodies, which appear like drops of honey and are

deceptive to flies, like the mock nectaries of Parnassia. The

species of Fuchsia are confined chiefly to the shady forests of

Central and South America. Both whorls of the perianth are

highly colored, the calyx and stamens crimson or scarlet, and

the petals purple or red. The flowers are pendulous and visited

by humming birds. Crimson or scarlet flowers are not common

where there are no humming birds. Kerner calls attention to

the rarity of scarlet in south Europe compared with its frequent

occurrence in tropical America, where in the primeval forests

there are a great number of scarlet or fire-red species of Bego-

nias, Fuchsias, Lobelias, Erythrinas, Salvias, and other crimson

blossoms, which are surrounded by humming birds. Many

hybrids of Fuchsia have arisen under cultivation. The earliest

white-sepaled form was produced in England in 1822, while the

first white corolla was secured in 1853.

In the Haloragidaceze, or water-milfoil family, the petals are

small or wanting and the flowers are wind-fertilized. The

species are mainly aquatic.

The last of the polypetalous orders is the Umbellales, which

includes the Araliacee, Umbellifere, and Cornaceze. The

flowers are small and densely aggregated in umbels, cymes,

and panicles. In the ginseng family, or Araliaceze, the flowers

are white or greenish. In open, sunny thickets many plants of

Aralia hispida grow together and produce numerous umbels

of inconspicuous flowers with small white petals. The honey

is abundant and freely exposed, and I have collected upon the

flowers eighty-two visitors. Bees, with the exception of the

honeybee, are not common; but the less specialized Hymenop-

tera, such as ichneumon flies and wasps, are numerous. Not-

- withstanding their want of bright colors, the flowers are very

attractive to the butterfly Argynnis aphrodite, several of which

may often be seen at work on the same plant. Though 4. race-

mosa may be found growing but a short distance from A. hispida,

its habitat is within the precincts of shady woods, where the

greenish flowers attract a much smaller circle of guests. The

Lene — of A. eter: pen orap are also visited

, v insects 7 A slight

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 235

importance in attracting insects, compared with its sunny,

sheltered location and easily accessible supply of honey.

Few plant families have attained a form of inflorescence so

well adapted to insure cross-fertilization as the Umbelliferz.

To this family belong the caraway and carrot, the wild parsnip,

the water hemlock, and parsley, — plants growing luxuriantly by

the roadside, along the river, and in the meadow. There are

about 1600 species, mostly confined to the temperate zone.

The flowers differ very little in structure, and the species can

be identified only by the mature fruit. The small flowers gain

conspicuousness by aggregation, and by standing in the same

horizontal plane afford a convenient landing place for insects,

and admit of rapid fertilization. The honey is secreted in a thin

layer by the fleshy disk surrounding the style. The number

and variety of the visitors surpass those of all other families.

In Germany there have been collected on the caraway 55, on

the wild carrot 61, and on the wild parsnip 118 insects. Prob-

ably the number of visitors to many species exceeds 200 ; while

there is thus ample provision for intercrossing, self-fertilization

is in most species prevented by the anthers and stigmas matur-

ing at different times. Admirable simplicity and perfection are

here combined.

There are 58 white, 16 yellow, 1 purple, and 3 blue flowers.

As in the Cruciferz, which have also very uniform flowers, white

and yellow predominate, red and purple are more rare. The

16 yellow flowers belong to 11 genera, only three of which con-

tain more than one species. In his Fertilization of Flowers

Müller enumerates 7 Diptera and 7 Hymenoptera collected

on the dull-yellow flowers of Pastinaca sativa; 4 Diptera

and 4 Hymenoptera on Bupleurum falcatum ; 15 Diptera and

31 Hymenoptera on Anethum graveolens ; and remarks, “So the

dull-yellow flowers of this plant, P. sativa, like those of Bupleu-

rum and Anethum, are visited only by Diptera and Hymenop-

tera, not by beetles.” On page 574 he adds: “ All dull-yellow,

dirty-yellow, brownish-yellow, yellowish-white flowers, Bupleu-

rum anethum, Pastinaca, Rhus cotinus, Galium mollugo, Ruta,

Neottia, Euonymus, Euphorbia, Adoxa, Alchemilla, are entirely

or almost mary. avoided by beetles. The only apparent

236 THE AMERICAN NATURALIST. (VoL. XXXVI.

explanation of these facts is that beetles are only or mainly

attracted to flowers by bright colors. If this explanation is

correct, dull yellow must be an advantageous color for plants

with freely exposed honey, protecting them from injurious guests."

Subsequent observations of Müller himself showed that Bupleu-

rum falcatum was very frequently visited by the beetle Mordella

pumila. In Schlesien, Loew observed on Anethum graveolens

5 species of beetles; numerous beetles were observed in the

Tyrol by Schultz on Rhus cotinus, and on Euonymus both

Schultz and Knuth observed beetles. The writer has found

beetles very frequent visitors to the dull-yellow flowers of Acer

spicatum. These illustrations need not be carried further, as

it is evident that more extended observations have disproved

Müller's generalization that dull yellow excludes the visits of

beetles. i

In the Umbellales both yellow and white have probably been

derived directly from the primitive green. The involucre of

Cornus canadensis changes from green to white, and the petals

of many species of Cornus are green in the bud but become

white as the flowers expand, while in the Umbelliferæ greenish-

yellow flowers occur. In Apium one of the species is greenish

yellow and two others are white. In Sanicula, S. gregaria has

yellow petals exceeding the calyx and bright-yellow anthers, and

S. marylandica has very small greenish-white petals and anthers.

In Peucedanum three species are yellow and one white. White

flowers may in some instances have been derived from yellow,

as it has been shown that the yellow petals change to white in

individual flowers in the Cruciferee. As regards attractiveness

to insects the yellow flowers of the Umbelliferae do not appear

to possess any advantage over their white competitors.

In Thespium two of the species have yellow flowers, and in

the third they are purple, with a common yellow variety. Many

of the white species tend to become pink or reddish. In Daucus

carota the central flower of the umbel, and frequently of the

umbellets, is purple, and I have often seen the entire umbel

pinkish. The coloration of this single flower can, of course, be

of no advantage to the plant in attracting insects, and Darwin

— a relic of a former ancient condition. I am

No.423.] NORTHERN POLYPETALOUS FLOWERS. 237

inclined to believe that it is due rather to the chemical constitu-

tion of the soil and nutritive fluids and the action of light. The

terminal flower receiving the greatest amount of sap would be

the first affected, then those of the umbellets, and finally the

entire umbel. The color changes of many flowers appear to be

due to such causes rather than to the influence of insects. In

Low Germany Pimpinella magna is white, and in the more

intense light of the Alps pinkish. Both rose-colored and yellow

flowers have been found on plants of Eriogonum ovalifolium

growing in silver-ore localities. Chemical analysis showed that

the rose flowers contained arsenic, which was not present in the

yellow. As a stimulus in nutrition may intensify the color, so

a check in growth may cause it to revert to white.

Increased conspicuousness of the umbel is gained in some

genera by the enlargement of the outer petals of the marginal

flowers, as in Heracleum. White bracts subtend the white

flowers of Astrantia, and yellowish bracts the dull-yellow flowers

of Bupleurum, and in Eryngium the flower stalk is colored.

The Cornaceze include many ornamental shrubs valued for

their bright-red bark, their variegated leaves, the masses of

handsome flowers, and the coral and blue berries. The flowers

are chiefly white, but in the European Cornus mascula are

yellow. The forty or more small white flowers of the herba.

ceous bunchberry Cornus canadensis, gain conspicuousness by

an involucre of four white bracts, sometimes tinged with red. I

have also seen the leaves partially white. This species yields

very little nectar, which can be detected only by close examina-

tion. The blooming season lasts for over a month, and the

flowers are produced in the greatest profusion. I have watched

these plants long and carefully, and have enumerated thirty-six

visitors ; but not once have bumblebees been seen to visit the

blossoms, — a fact I attribute to the scarcity of honey. The

shrubby species contain a more abundant supply of nectar.

The cymes are large and very numerous, and not infrequently

are sought by bumblebees. On C. alternifolia I have collected

twenty-eight insects. There. can be no doubt that the quantity

and flavor of nectar is an element in the limitation of insect

visits, but one that has received much less attention than

238 .THE AMERICAN NATURALIST. | [Vor. XXXVI.

coloration. C. florida is provided with an involucre that is

usually white, but varies to pink or red. The fruit is scarlet

and the leaves are bright red in autumn.

SUMMARY.

I. Throughout the Choripetale, with few exceptions, con-

spicuousness is correlated with fertilization by insects. In the

grape family the flowers depend upon their strong scent rather

than upon bright coloring. In many genera it is possible to

arrange the species in a progressive series, in which there is an

advance from inconspicuousness, few. visitors, and self-fertiliza-

tion, to many visitors, great conspicuousness, and the loss of

thé power of self-fertilization. Pigments may be developed in

all the organs of the inflorescence, as bracts, stems, sepals,

petals, stamens, and pistils.

2. The green flowers of the Polypetale are small, and the

petals are frequently wanting. The white and yellow flowers

vary from small to large, are the most common, and contrast

more strongly with the foliage than purple or blue. Of the

seventy-one polypetalous families, forty-three contain white,

forty-one yellow, and twenty-nine both kinds of flowers. White

flowers are most common in families or genera containing shrubs

and trees, small flowers aggregated in a dense inflorescence, and

nocturnal flowers. Dark nocturnal flowers are strongly scented,

Yellow flowers are more commonly herbaceous and are most

abundant in the same families as white flowers, unless the

species are shrubs or trees.

3. There is no evidence of the preference of beetles for

flowers of any particular color. They do not avoid dull yellow.

They are most common on small, white-clustered flowers with

easily accessible honey and pollen. Diptera visit most frequently

white and yellow flowers, but as they become more specialized

and restrict themselves to flowers the percentage of visits to red

and blue flowers increases. . They appear to find a parti-colored,

mottled, or dotted inflorescence, as in the Cruciferae and Saxi-

fragaceze, attractive. Carrion flies eru malodorous lurid-

Pepe or r flesh-colored flowers.

No. 423.] NORTHERN POLYPETALOUS FLOWERS. 239

4. The changes of color and their sequence in individual

flowers are noteworthy. Green changes to white (Cornus), to

yellow. (Thlaspi, Cardamine), to red (Hydrangea), to purple

(Clematis), to violet (Cobzea) ; white changes to green (sepals of

Helleborus niger), to yellow (Lantana), to red (Dianthus, Hibis-

cus mutabilis), to blue (many large blue flowers remain white

until nearly ready to expand) ; yellow changes to white (Draba),

to red (/Esculus) to blue (Myosotis); red changes to blue

(Venetus and many Boraginacez) ; violet and blue may turn

purplé, green, or white in fading. The tendency of green,

white, and yellow to change to red or blue is much stronger

than the reverse.

5. The floral colors are often correlated with the colors of

the stems and leaves, as in Sedum. The foliage of the plants

with white flowers is, as a rule, paler than when the flowers con-

tain pigments. The development of bright colors in autumn

leaves presents a series of color changes, which are in part par-

allel to those which occur in flowers. With the disappearance

of the chlorophyll the leaves become whitish, yellow, or red,

according as the cells contain no pigment, or solid yellow gran-

ules, or red pigment dissolved in the cell sap. The leaves of

many plants are yellowish green, due to the presence of a yellow

pigment. Green, yellowish-green, and greenish-yellow flowers

contain chlorophyll, and though usually small are occasionally of

large size. Many white and yellow flowers are derived directly

from the primitive green. White is usually a structural or opti-

cal color due to the unequal reflection and refraction of light by

the intercellular air spaces and the cells devoid of pigment.

White flowers are a less tax upon the energies of the plant.

Flowers of all colors may revert to white, which is commonest

in nature and most true to name under cultivation. If with the

disappearance of the chlorophyll there is an insoluble yellow

pigment in the cells, the petals are a pale yellow, and with its

increase change to bright yellow or orange. The development

of anthocyanin, or red pigment, dissolved in the cell sap, changes

white flowers to red and yellow flowers to scarlet. With a

decrease in the acidity of the cell sap the red flowers become

blue. Müller's observations led him to the conclusion that the

240 THE AMERICAN NATURALIST. [VoL. XXXVI.

honeybee prefers blue, violet, various shades of purple and red,

to white and yellow, and avoids scarlet and lurid colors.

6. The formation of pigments is effected by the chemical

composition of the soil, by altitude or the intensity of light, by.

latitude, and by the absence or presence of moisture, as well as

other ecological forces. The particular coloration of flowers is

largely a chemical problem.

No. 423.] WORTHERN POLYPETALOUS FLOWERS. 241

THE COLORS or NORTHERN POLYPETALOUS FLOWERS.

: ; z ti. :

ORDERS, FAMILIES. E E E a E i: E

OTRITILMAIEIM LG

(| Nymphaacee .... 4 I II

Ceratophyllacem ..| 1 I

Magnoliacee . t£] 3 6

. |] Anonsceio 65. yes 1 1

Ranales , . . . . 4| Ranunculacez 61 951331] ST et eel OF

= Berberidace 3 3 I 7

Menispermace I 2 3

Calycanthacex 2 2

Merete NUR 2 4 6

Papaveraceze EE uu a 23

e a aT wl6l. r1 to 113

Puteo re Capparidaceze s " 2 I I 7

: (| Resedacee...... FT g 3

Acar are a a { Sarraceniaceæ I I 2

psi Droseracez ..... 4 I 5

(| Podostemacez I 1

Crassulacee..... 2 2 5 2 2 13

Saxifragaceze (5 2 4l 35 6 3 43

Grossulariacez .. 4 6 I - 13

Hgmamelidacez Ei .xli 4 3

Platanacee ..... I I

Rosales ..... j| Rosacem ....... 4i 341 30 1 4 95

Pomacee ...... 27 5 32

Drüpacem ...... 20 d.d 21

Mimosacee ... 3 I 2 6

Czesalpiniaceze 2 I 7 I II

ime i . I I

L| Papilionacee . . 39 | 33 | 13 | 881 24 | 197

(| Geraniacem ..... I 41 y [ rt

ali Bi... rT ee I 8

inacem ....... I 6 2 9

Zygophyllacez 2 2

Geraniales ... 4 ee *l.a 3

Simarubacez 3| ok 1

lygalacem .. ... 11.31 321 81 16

Euphorbiacez 3|.» sl: | 44

itrichaceze uf. 4

242

THE COLORS oF NORTHERN. POLYPETALOUS FLOWERS.

THE AMERICAN NATURALIST.

- (Continued.)

co ee: ET da

ORDERS. FAMILIES. E z E á = E s

[4 w 5| 2 A o

O [Ff Boe me fe poet

(| Empetracee . .... 2 =

Bixhcee^ 21. ri I I

Lininanthacez I I

Anacardiacee .. 7 I 8

Cyrillacee ...... I I

: PCat 0.2.1.1 IO IO

Sapindales: «-- 3] celastracese |.: . : 3 I 2 6

Staphyleacee ...: I I

Aceratee ...... 5 3 I 9

lippocastanacez . I 3 I 5

Sapindacee ..... 2 2

\| B: saminacez 2 2

Phannal ... { Rhamnaceze Pa AT 5 3 8

- Vitacee ~... 13 13

Siidviles TOT Tiliacee ......- 3 3

pr Malvacee ...... 4 f] 34 4 26

[| Théaceg —.. es 3 3

Hyperi . 22 2 24

Elatinacez ...... 4 4

Parietales . . . - ?| Cistacee....... 9 5 14

Violacem ....... , 7 6 41357 34

$ rac F I I 2

- cM HU AEE s T 3 5

Cactaces. . 2.55 i? Feal i 12

hymeleacez . . I I 2

ERAN 3 3

Lythracee...... 2 10 12

| Melastomacee . . . LL. 4

Onagra La Pad 431 70) mT |. 6 57

Trapaceze S DIT I I

Haloragi ud ti 6] 13

Avaliace® ..... so gr 6

Umbelliferæ . . 5| 16] si 4| 3

Comaezg ......| 3| 9| t|. 1 | 14

Total... . .... [140 | 410 1333 1-84. } 1903 | 57 | 1217

NOTES AND LITERATURE.

ZOOLOGY.

Shipley and MacBride's Zoólogy.! — It may be said in beginning

that this is among the best of text-books for elementary college

classes in zoólogy. It is fresh, clear, well arranged, and fairly accu-

rate. In its first twelve pages it outlines the basal facts of zoólogy,

or better of morphology, including short statements concerning life,

protoplasm, and the theory of evolution, and then begins the sys-

tematic portion of the volume. The Protozoa come first; then, in

order, the Coelenterata, Porifera, and Ccelomata, — this latter divi-

sion ending with man. As a result, the flatworms, rotifers, nema-

todes, etc., appear in the most unexpected place, — the end of the

volume. There does not appear any hint that there is a question

as to the validity of the ccelom or its use as a basis of classification.

In the treatment of each group the authors follow the well-known

English model of giving, first a detailed description of some form

selected as a type, then a general statement, followed by too brief

and inadequate an outline of classification.

Good as the book is as a whole, we find here and there parts

which cannot have our full acceptance. In the earlier portions the

authors have succeeded well in their attempt, as avowed in the

preface, to produce “an elementary treatise on zoology which

could readily be understood by a student who had no previous

knowledge of the subject,” but in the later portions, and especially

in the treatment of the vertebrates, they fall far short of their

ideal. No student, for instance, could gain from these pages,

without outside assistance, any adequate idea of the skull or the

nephridial system. As the whole work seems so well done, it is

hardly just to publish a long list of questionable or erroneous state-

ments which would seem to contradict the good opinion already

expressed; and yet, directing attention to some of these shortcom-

ings may lead to their correction in the subsequent editions, which

will certainly follow. —

1 Shipley, A. E, and MacBride, E. W. Zodlogy, an Elementary Text-Book.

New York, The Macmillan Company, 1901. xxi + 632 pp.

243

244 THE AMERICAN NATURALIST. [Vor. XXXVI.

In dealing with the arthropods we object to the violent divorcing

of the closely allied Crustacea and arachnids by shoving the insects

between them. The conception of a crustacean head composed of

five segments is unwarranted, and the retention of the group Myria-

poda, in view of the work of Pocock, is indefensible, while the

statement (p. 193) that Limulus occurs on our Pacific coast is

erroneous. We do not believe that ZZzx aspersa is acclimated in

this country to the extent implied on page 198, and on page 221 the

terms “squid” and “cuttlefish” are transposed. On page 241 there

is an erroneous explanation of the way in which the starfish opens

clams and mussels, and we are not inclined to accept the statement

(p. 221) that there are over five hundred species of Unionidz in

North America. Certainly many of the species enumerated by

Conrad, Lea, and others are not entitled to any such rank.

In their account of the vertebrates the authors have attempted to

carry too far homologies between the true vertebrates and the other

chordate forms. For instance, the anterior head cavities of the

shark cannot be compared to the proboscis cavity of Balanoglossus ;

and, by the way, what do the authors mean (p. 293) when they rele-

gate the term “ Balanoglossus ” to the category of popular rather than

generic nimes? On page 332 there is an erroneous conception of

the pharynx, no portion of which is stomodeal; the idea of a gono-

tome (p. 343) has been vigorously combated, and certainly, in view

of the comparatively recent researches of Rabl, the pronephric duct

should not be described (pp. 342, 352), without a question, as ecto-

dermal in origin. It certainly harmonizes with some theories, but —.

Not all Dipnoi (p. 371) have two lung sacs, while some Czecilians

(eg., Ichthyophis) do have a ductus Botalii (p. 437). Again, it is

certainly questionable whether Gadow's conception of the structure

of a vertebra (p. 378) is correct; and the statements (p. 330) con-

cerning the optic nerve need revision in the light of the results of

Keibel, Assheton, and others, for these nerves are clearly not the

stalks uniting the primary optic vesicles with the brain, but true

nerves which replace them. There is also a complete crossing of

the fibers in the lower vertebrates. The division of birds into Ratitze

and Carinate (p. 497) should not be kept up in this year rgor. In

view of the statement that the vertebra: of mammals *have no cup-

and-ball articulations with one another" (p. 510), we should advise

an examination of the cervicals of a horse, and (p- 404) would call

attention to the fact that several amphibians have amphiccelous

vertebrae. E LR ee

No. 423.] NOTES AND LITERATURE. 3 245

Here and there the English used is not beyond question, being

sometimes inelegant, sometimes confused. The term “ urinogenital ”

has been cast into limbo by Huxley (Zife and Letters, Vol. II, p. 63),

but language is a dangerous thing to discuss. So much depends

upon the point of view, and. our authors have occupied. a peculiar

position for seeing things, as we learn from the legend of Fig; 200,

which represents the right half of a lamprey *seen from the inside

of a female specimen." The mechanical part of the work is well

done, and the illustrations, mostly zinc etchings, are clear, and: as a

rule bring out well the points made in the text. K.

Notes. — Wallengren (/enaische Zeitschrift für Naturwissenschaften,

Bd. XXXVI, pp. 165-180, Taf. VII-VIII) gives a brief but interesting

account of the innervation of the proboscis of some of the polychate

worms. The sense cells always occur in the papillz of the proboscis,

never on the surface between papilla. In Nephthys and Phyllodoce

they are radially arranged within a papilla, but in Glycera and Goniada

they are grouped into multicellular organs, and single sense cells do

not occur. In all cases the distal ends of the sense cells pierce the

superficial cuticula, and in Glycera at least they end externally in a

brushlike expansion. Proximally the fiber from each cell body either

passes undivided to the central organs, as in Nephthys, or gives off

branches at the base of the papilla, forminga subcutaneous nerve

plexus as in Glycera. No observations were made on the functions

of these organs.

The ganglion cells of the electric lobes in the Torpedo are said

by Hatai (Journal Cincinnati Society of Natural History, Vol. XX,

pp. 1-12, Pl. I) to show an apparent fibrillar structure suchas that

described long ago by Max Schultze. Bundles of fibrils seem to enter

the cell from the dendrites as well as from the neurite. This appear-

ance, however, is due to the form of the protoplasmic reticulum,

the meshes of which are much drawn out in the regions of the cell

processes, and does not indicate the presence of true nerve fibrilla.

While the bundles of fibrille as described by Schultze are thus.

explained away, those discovered by Bethe and Apàthy belong to

another category and probably are dun pin Gradations from

the ordinary mesh work to the mesh > have

been described by Hatai in the’ spinal ganglion cells of the ‘white fat.

An ingenious method of recording égg development for the use’

of fish culturists has been devised -by °€: Wallich (United States:

246 THE AMERICAN NATURALIST. [Vor. XXXVI.

Fish Commission Report for 1900, pp. 185—194). It is well known

that fish eggs develop much more rapidly in warm water than in

cold, and when, as is often the case, the temperature of the water

at a hatching station varies from day to day, it is often difficult to

estimate the stage of development reached by a given lot of eggs.

This difficulty can be. overcome by the use of the temperature unit

system. By a “temperature unit" is meant one degree Fahrenheit

above 32° F. for one day. Thus 36° F. for a day is equivalent to

four temperature units. In any lot of eggs the stage of development

is recorded by adding the temperature units to which they have been

subjected since fertilization. The success of such a system depends

upon the uniformity of its results. As judged from the experiments

on salmon at the government station at Baird, Cal., the method gives

a close approximation to uniformity. In something over fifty lots of

salmon eggs, in which the incubation period varied from forty-eight

to ninety days, the greater number of eggs hatched. at about nine

hundred temperature units, the extremes being 874 and 940. The

utility of this method, not only to fish culturists but also to embry-

ologists, is evident.

C. E. Beecher has contributed to the series of Yale Bicentennial

Publications a volume entitled Studies in Evolution (New York,

Charles Scribner's Sons, 1901). The volume is made up mainly of

reprints selected from the publications of the Laboratory of Inverte-

brate Paleontology at Yale, and contains studies on the origin and

significance of spines, on the structure and development of trilobites,

and on the development of brachiopods. Most of the papers have

already appeared in various journals, but their collection into a

single volume will be welcomed by advanced students, not only as

evidence of the work done at Yale, but as an indication of the new

fields into which modern paleontological research is making its way.

The stony corals of the Porto Rican waters have been reported

upon by T. W. Vaughan (United States Fish Commission Bulletin Jor

I900, Vol. II, pp. 289-320, Pls. I-XXXVIII) About twenty-five

species are recorded and beautifully illustrated by process plates from.

photographs. | |

C. W. Hargitt and C. G. Rogers give an account of the Alcyo-

naria of Porto Rico (United States Fish Commission Bulletin Jor 1900,

Vol II, pp. 265-287, Pls. I-IV). Some twenty-five species are

reported, of which five are new. dun "oio

No. 423.] NOTES AND LITERATURE. 247

A descriptive catalogue of the mollusks of Porto Rico, with

a check list of species, has been prepared by W. H. Dall and

C. T. Simpson (United States Fish Commission Bulletin Jor 1900,

Vol. I, pp. 351—524, Pls. LIII-LVIII). In all, about 650 species

are recorded, of which forty-two are new. The report is illustrated

by excellent figures, and has a good index.

BOTANY.

The Yearbook of the Department of Agriculture.! — The grow-

ing size, complexity, and activity of the national Department of

Agriculture has led to attempts to simplify its publications by the

establishment of series accredited to the several divisions and more

or less closely pertaining to the nominal work of the latter; and a few

years ago the secretary had the happy idea of confining the annual

volume — before that called a Refort—to an administrative and

scientific summation of the year’s work, under the title Yearbook.

Difficulty appears to have been experienced in carrying out this idea,

however, and the Yzaroo£ has once more approximated its former

size and character, notwithstanding the voluminous annual aggregate

of bulletins and circulars published from the several divisions. In

addition to the report of the secretary and an appendix of over two

hundred pages, collectively constituting the report proper, the Year-

book for ri9oo includes a biographic sketch of the late William

Saunders, for many years connected with the care of the grounds

and plant houses of the Department, and no less than thirty special

articles, on such subjects as Smyrna fig culture, date culture, prac-

tical foresty in the southern Appalachians, etc. ,-——all of interest

and value. T.

Britton's Manual.*— The most potent means of i impressing ideas

or facts on the public mind lies in their concrete presentation in such

form as to insure ready accessibility and invite frequent reference.

When, in 1892, the American botanists assembled at

adopted a dification which involved extensive changes

1 Yearbook of the United States Department of Agriculture, 1900. Washington,

Government Printing Office, r90r. 888 pp., 87 pls., and numero us figs.

?Britton, N. L. Manual of the Flora of the Northern States and Canada.

New York, Henry Holt & Co., 1901. x + 1080 pp.

248 THE AMERICAN .NATURALIST. [Vor. XXXVI.

in the Latinized hames of our higher plants, they insured the most

rapid possible introduction’ ‘of the proposed changes by taking steps

for the compilation and publication of ʻa check list in which these

changes should appear; and the frequent adoption of the names

approved by the authors of that list, in general and local botanical

papers, and their subsequent application through the large /Wustrated

Flora of Britton and Brown, have made them generally familiar, — to

the pleasure of some and the great regret of other botanists, both at

home and abroad. With the changes in names have also come a

change in the way of viewing species and increased attention in the

field to the divisibility of species which scarcely admitted of a satis-

factory knowledge from herbarium material.

To meet these needs and changed conditions the ///ustrated Flora

was prepared; and yet it was too bulky and, notwithstanding its

remarkable cheapness, too costly for the fullest desirable uses.

Because of these reasons, Dr. Britton, the active editor of the Fra,

set to work to prepare a more condensed manual with the same pur-

pose and limits as the Fra, and it now appears in a convenient

and —for this class of book — attractive form, and is undoubtedly

destined to have a wide sale and to serve a good purpose. That it

will lead to a recognition of many valid species of the region covered,

not recognizable by use of the familiar Manual of Dr. Gray, is cer-

tain, and that it will ultimately replace the latter in general use is not

improbable, though the present writer would recommend the use of

the two conjointly, rather than of either by itself. T.

Vegetable Pathology.' — Almost from the installation of the first

agricultural experiment station, the diseases of plants have been a

. fruitful and legitimate subject for investigation, and the selection of a

station botanist has more than once hinged on his fitness to become

the plant pathologist at the same time. Yet the efforts of investi:

uen have been T turned to the — of the life histories of

fow::3 tlia: es with: the

details E. E aled any ud the green pneus us marks a distinct

advance in. knowledge. and. points the way for the establishment of

~ Ward; "H. Marshall - P PAN agree NM. vae

: e go PP-

No. 423:] NOTES AND LITERATURE. 249

an art of plant therapy, if this is ever to be possible or worth the

establishment.

The Flora of Tennessee.! — No living botanist is as familiar with

the interesting plants of Tennessee as the venerable Dr. Gattinger,

who, like the late Dr. Mohr of Alabama, has spent many years in

field study and now brings together the results of his work. Ten-

nessee is a state of diversified topography, and the systematic list

is preceded by a discussion of the factors which influence plant

distribution. Like Dr. Mohr, Dr. Gattinger adapts himself to the

Neoamerican practices in nomenclature and to the phylogenetic

classification of the Germans, though not without a word of protest.

Whether or not he be followed in his philosophic ideas, they contain

the kernel of much that is good.

The Grasses of Iowa.*— In the prefatory note to this book

Professor Calvin, the state geologist, tells us that the law creating the

Iowa Geological Survey provides for the publication of bulletins on

subjects of economic interest relating to the natural history of the

state, and this consideration of one of the most important economic

groups of plants forms the first of such bulletins. ‘The topics treated

are: the general structure and physiological characters of Graminez ;

purity and vitality of grass seed; cereals; fungus and bacterial

diseases of grasses; the pastures and meadows of Iowa; weeds

of meadows and pastures; chemistry of foods, and feeding; and

lawns and lawn-making in the state. Professors Pammel and

Weems are well equipped by training and opportunity for the con-

sideration of these topics, and it is understood that the system-

atic discussion of the group, for which Professor Lamson-Scribner

will be largely responsible, is to constitute a second volume com-

plementary to the one now used.

Chlorophyll.?— No. 10 of the biologic series of Scientia presents

in a manner not too technical for the layman the main facts concern-

ing the chlorophyll function in plants as carried on under various

external conditions. The structure, as wellas the function, under-

goes adaptive modifications in response to these conditions, and the

lGattinger, Augustine. Zhe Flora of O and a Philosophy » Botany.

Nashville, 1901:

? Pammel, L. H. ems J B., and Lassen Sodbuer: F. The Grasses of Towa.

Des Moines, fool.’ 525 pp.; 2

-3 Griffon, Ed. desiit, cbletvphsiliette et la structure des. plantes,

Scientia, Biologie, No. 10. Georges Carré et C. Naud, Éditeurs. . Paris.

250 THE AMERICAN NATURALIST. [Vor. XXXVI.

author here traces these effects of the environment. Means of meas-

uring photosynthetic activity described in the first chapter are applied

to the study of this form of activity in saprophytes and parasites and

to the effect. of light, temperature, moisture, and mineral salts on

this function in green plants having a normal nutrition. p H.T.

Our Forests. — Popular interest in the preservation of forests is

too often a matter of extremes, so diverse reasons as pure emotion or

simple business considerations leading to the wish for their protec-

tion, while private greed is ever encroaching on them in the most

ruthless fashion. A most attractive presentation of the subject from

the zsthetic standpoint is made in a little book by G. F. Schwarz.’

A pleasingly written commentary on a few selected trees representa-

tive of our deciduous and coniferous woods is followed by chapters

on forest adornment, the distribution of American. forests, the char-

acters of the broad-leaf and coniferous forests, and the artificial

forests of Europe. "Throughout, the book is ‘well. illustrated by

process cuts. It is neither botany nor forestry, but a delightful

presentation of the beauties of nature as exemplified in forests,

equally correct in treatment when viewed from: either of these

special points of view, and calculated to draw increased attention

to the desirability of holding as tenaciously às possible to what is

still left of this most charming phase of nature. T.

The Annals of the Calcutta Garden.?— Like its predecessors,

this volume is not only well brought out, but of permanent value, and

the authors, Messrs. King, Duthie, and Prain, are to be congratulated

on it. - The plates are lithographed, and an excellent photogravure

representing A/bizzia Richardiana serves as frontispiece. T.

Notes. — In the Bulletin of the Torrey Botanical Club for Denada

Miss Eastwood describes several new Delphiniums from California.

A paper on seed coats of certain species of Brassica, including

good anatomical figures, by A. J. Pieters and V. K. Charles, is pub-

lished as Bulletin 29 of the Division of Botany of the United States

pio toes of Agriculture.

1 Schwarz, G. F. Forest Trees and Forest Scenery. Ner York, The Grafton

Press, 1901. xiv -- 183 pp.

? Annals of the Royal Botanic Garden, Calcutta. Vol. ix; pti A Second Cen

tury of New and Rare Indian Plants. CPI Ee Ranga) Sdcetaein) P

Igor. 82 pp., 94 pls. Zrigs —

No. 423.] NOTES AND. LITERATURE. 251

Hefte 11-12 of Abtheilung I of the 1901 volume of the Botanische

Zeitung consists of a paper by Hannig on the septal wall of the fruit

of Cruciferze.

Professor Hildebrand contributes biological notes on Jeffersonia

diphylla and Apios tuberosa, as well as other species not of our flora,

to No. 8 of the current volume of Berichte der deutschen botanischen

Gesellschaft.

Schlotterbeck and Eckler publish a paper on fruit structure and

development of — floridanum in the TERHROPEHCNI Arthives

for November.

The Celastracez of China are undergoing revision by Loesener in

Engler's Botanische Jahrbücher, which, as — of -— is for the

most part devoted to African plants. :

The development of the seed and the differentiation of the pro-

tective testa in several Sapindacez is discussed by Guérin in the

October number of the Journal de Botanique.

A monograph of the leguminous genus Dorycnium, by Rikli, is dem

lished in Heft 3 of Engler's Botanische JaArbücher, Vol. XXX

A case of parthenogenesis in Alchemilla is described by Murbeck

in Vol. XXXVI of Acta Universitatis Lundensis, which also contains

an article by the same author on chalazogamy; in connection with a

further study of the same genus.

In Zorreya for December Mr. Small publishes a new Texan m

under the name Prunus eximia.

Spirea millefolium is figured in Curtis’s Botanical Magazine for

December.

Those accustomed to seeing Cereus peruvianus as it is orsnuioa)

cultivated in plant houses will be interested in an open-air portrait of

it as grown at Monte Carlo, = by Becker in Die Gartenwelt

of December 21.

Opuntia monocantha, O. glauca, arid O. engelmanni, which prove

hardy in the botanical garden of the University of Cambridge, are

figured in the Gardeners’ Chronicle of December 7.

An illustrated account of the cacti found to be hardy in Giessen

is given by Rehnelt in Die Gartenwe/t of December 14.

- The affinities of the Valerianacez and Dipsacez are nones by

Hock in Vol. XXXI, Heft 3, of the Botanische Jahrbücher. ;

252 THE. AMERICAN. NATURALIST. [Vou. XXXVI.

In the. number of the. Proceedings of the Washington Academy of

Sciences issued December 6, Mr. Coville separates from the genus

Cassiope the species which have been known as C. oxycoccoides,

C. stelleriana, and. C. hypnoides, proposing for the first the generic

name Arcterica, and for the last two the generic name Harriman-

ella, in honor of Mr. E. H. Harriman, of whose Alaskan party

Mr. Coville was a member.

In Rhodora for November Walter Deane indicates a form Zuco-

carpum for Vaccinium pennsylvanicum; a similarly named form for

V. corymbosum atrococcum and a form chiococcum for V. canadense are

indicated for New England.

The inelegant compound inflorescence often produced by the

Edelweiss under cultivation, and which is then the despair of gar-

deners, is the subject of an illustrated article by Brunotte in the

evue Générale de Botanique of October 15.

The four United States forms of Leontodon are reviewed by

Fernald in Rhodora for December.

In Rhodora for December Mr. Ames describes an artificially pro-

duced hybrid of Lobelia inflata 9 x L. cardinalis 3.

Euphrasia, as it occurs in North America, is reviewed by Dr. Robin-

son in Rhodora for November. ` Æ. williamsii, E. randii, E. randii

Jarlowii, and E. americana canadensis are new names.

The west-coast Solanums of the group of S. umbel/iferum are

revised by Parish in No. 5 of the current botanical volume of Zro-

ceedings of the California Academy of Sciences.

For various species commonly placed in Bartonia or Mentzelia,

Cockerell proposes the generic name Hesperaster, in Zorreya for

December.

Hefte 5 and 6 of Engler’s Das Pflanzenreich, devoted respectively

to Rafflesiacez and Hydnoracez by Sohns-Laubach, and Symplo-

cacez by Brand, were issued November E.

Anatomical studies of certain Australian Podalyriex, by Hühner,

constitute Heft 3 of the current volume of the Beihefte zum botan-

A paper by G. N. Collins, on seeds of commercial saltbushes, con-

stitutes: Bulletin 27 0f the Division of Botany of the. United States

No: 423.] NOTES AND LITERATURE. = =" 253

-À synopsis of thé palms of Puerto Rico, with much extra-limital

matter, by O. F. Cook, is published, with illustrations, in the October

Bulletin of the Torrey Botanical Club.

Habit and detail figures of /wbea spectabilis are published in the

Revue Horticole of January.

In the Bulletin de la Société Botanique de France, Nos. 5-6 of the

current volume, Gagnepain publishes a revision of the zingiberaceous

genera Mantisia and Globba, as represented in the herbarium of the

museum at Paris. Several new spécies are described and figured.

In the Revue Horticole of December 16 a new Pitcairnia, P. miche-

liana, from Mexico, is described and figured by André.

As No. 4 of the current botanical volume of Proceedings of the

California Academy of Sciences, is published a revision of the genus

Calochortus, by Carl Purdy.

In the November Bulletin of the Torrey Botanical Club, Dr. Rydberg

monographs the species of his orchidaceous genera Limnorchis and

Piperia, occurring north of Mexico.

The southwestern species of Sisyrinchium are passed in review

by Mr. Bicknell in the October Bulletin of the Torrey Botanical Club.

Prof. H. Marshall Ward’s handbook of grasses, recently issued

from the University Press of Cambridge, is a convenient little work,

with keys based on vegetative characters, leaf anatomy and fruit, as

well as the more usual flower and inflorescence characters.

A paper by Holm on new anatomical characters for certain

Graminez is published in Vol. XI, Heft 2, of the Beihefte zum

botanischen Centralblatt.

One of the most important recent cytological papers is by Dr.

Webber on spermatogenesis and fecundation of Zamia, presented

as a thesis for the doctor’s degree at Washington University last

season, and now printed as Bulletin No. 2 of the Bureau of Plant

Industry of the United States Department of Agriculture,

A paper by Worsdell on comparative anatomy of the Cycadacez

forms Part II of the current volume of Transactions of. the Linnean

Society of London.

A. paper on the red cedar, ptéputed dy Di Mohr aisha woke

his death, has recently been published as JBuHetin No. 3r of the

Division of Forestry of the United States Départment of Agriculture;

254 THE AMERICAN NATURALIST. [Vor. XXXVI,

A specimen of Araucaria. cookii, coning in the temperate house

at the Kew Gardens, is figured in the Gardeners’ Chronicle of

November 30.

The New England form of Lycopodium complanatum is iius var.

flabelliforme by Fernald in Rhodora for November.

B. D. Gilbert publishes a note on Lycopodium tristachyum in

Torreya. for October.

J. H. Faull. contributes an account of the anatomy of Osmundacez

to the Botanical Gazette for December.

The account. of the mosses of the * Belgica" expedition to the

extreme South, by Cardot, occupies a quarto pamphlet of 48 pages,

illustrated by 14 plates, published by the Belgian government.

Pseudoleskea artarie and Leskea obscura are comparatively described

by Thériot in a paper reprinted from the Recueil de la Société Havraise

d' Études Diverses for 19or.

An extensive and well-illustrated paper on Alaskan alge, by De

Alton Saunders, constitutes No. 25 of the “ Papers from the Harri-

man Alaska Expedition” in course of publication in the Proceedings

of the Washington Academy of Sciences:' The catalogue includes 380

species, of which 240 are new to Alaska. Nine new species and one

new genus, (Pleurophycus, Setchell and Saunders) are described.

Lemaire, who has been studying the micro-chemistry of the sheath

of the blue-green alge, states, in the recent numbers of the Journal

de Botanique, that in Anabzena, Nostoc, and some other genera the

mucilaginous sheath consists of a pectic compound, which, in the

case of some species of Scytonema and Phormidium, is associated

with another substance, which he calls “ schizophycose,” the latter, in

some species of Scytonema, Foot, etc. > Peng further associated

with cellulose.

The Diatomacez of the Hull district are listed and Roiited in

No. 4 of the Transactions of the Hull Scienti tfic and Field Naturalists

Club, issued in December.

The cultivation of mushrooms is anally effected by the use : of

spawn either grown in mushroom beds or derived from compost

piles or similar sources. In 1896 Costatin and Matruchot published

interesting details of methods of securing pure and unquestionable

spawn or mycelium of a few forms, and in the Revue Générale de

Botanigüe of November r5 last they further describe their method of

obtaining pure commercial cultures of Zricholoma nudum

No. 423:] NOTES AND LITERATURE. 255

The modes of preserving pileate fungi without loss of their natural

colors are reviewed by Lutz in the December Bulletin de la Société

Mycologique de France.

Heft 9 of Schimper’s Botanische Mittheilungen aus den Tropen con-

sists of Alfred Moller’s studies on Brazilian phycomycetous and

ascomycetous fungi, and is fully and well illustrated.

No. 44 of the Korrespondenzblatt des Naturforscher- Vereins zu Riga

contains a list of the twelve truffles occurring in the Baltic district.

In No. 8 of his serial Mycological Notes, Mr. C. G. Lloyd figures

several species of Scleroderma, Geaster, and Mycenastrum.

Dr. Thaxter's epoch-marking'work on Laboulbeniacez furnishes.

the text for a popular article by Mr. Massee in the November number

of the Journal of the Quekett Microscopical Club.

The genus Fusarium, which includes several species growing on

dead animal matter of various sorts, receives an interesting addition

under the name F. eguinum, described in Science of December 6 by

Novgaard as occurring in the hair follicles and sébaceous glands of

horses in Oregon.

A monograph of the genus Aspergillus, by Weimer: is sepiräiely

published from the Mémoires de Ta — de io adams et d' Histoire

Naturelle de Genève, Vol. XXXIII. `

In the Bulletin of the Torrey Botanical Club for December, Pro-

fessor Arthur publishes descriptions of a considerable number of

new Uredinez.

The effects of Rhizoctonia on the potato plant are discussed in

Science for December 6, by F. M. Rolfs.

The fungi causing spot diseases of the cherry, etc., are discussed

in an illustrated article by Aderhold in the December number of

Landwirtschaftliche Jahrbiicher.

Leaf diseases of the currant are considered by Stewart and Eustace

in Bulletin No. 199 of the New York Experiment Station.

PETROGRAPHY.

** Perknite,” a New Rock Term. —Turner,! in a recent article,

suggests the use of the term * perknite ” as the designation of coarse-.

grained lime-magnesia rocks composed essentially of monoclinic

1 Journ. Geol., vol. ix (1901), p. 507-

236 THE AMERICAN NATURALIST. [VoL. XXXVI.

pyroxenes and amphiboles. The group of the perknites would include

pyroxenites, hornblendites, and hornblende-pyroxene aggregates. it

would include also websterites and similar aggregates in which mono-

clinic amphiboloids predominate. The augitites and. some of the

hornblende-picrites are the corresponding effusive types. Analyses

of some of the author’s characteristic perknites follow :

SiO, AlO; FeO; FeO MgO CaO Na,O K,0 H,O Various

2 , ` .

I. 46.28 19:54 9.91 2.21 1.89 :

II. 48.04 7.82 2.01 9.32 13.33 13.01 .69 -48 3.07 2.74== 99.8t

HI.. 48.63 8.33 2.91 3-90 21.79 13.04 db 23 2.81. ¥,46== 700. 13

IV. 50.80 3-40 1.39 8.11 257 12.31 tr tr .52 +73 = 100.03

M... 83.25 2.80 69 5-93 19.9! 16,22 +19 tr 29° * o>. 09.98

Viv) 53:25 1.04 1.44 7.92 20,78 13.12 JI :07 1,01 .87 = 100.47

I. Hornblendite. Dyke near Silver Peak Village, Esmeralda County, Cal.

II. Amphibole-pyroxene rock. Abundant in Mariposa County, Cal.

III. Perknite (peridotite). Belchertown, Mass.

IV. Pyroxenite. Johnny Cake Road, Baltimore, Md.

V. Websterite. Mt. Diablo, Cal.

VI. Websterite. Oakwood, Cecil County, Md.

Shonkin Sag; A Study in Differentiation. — Shonkin Sag is a

flat laccolite rising from the plains southeast of the Highwood Moun-

tains, Montana. It has afforded Weed and Pirsson ! excellent oppor-

tunities for the study of the differentiation of a magma in place, since

it is dissected by a stream valley formerly occupied by a branch of

the Missouri River. Examination of the sections exposed to view

indicate that the igneous portion of the laccolite consists of concen-

tric shells of leucite-basalt, skonkinite, a transition rock composed

of groups of large augite crystals and long slender foils of biotite

with white feldspathic material between, and syenite, with the last-

named rock in the center. The various rock types grade into one

another without discernible breaks between them, ‘The facts suggest

to the authors “that the body of the magma forming the laccolite

must have been injected as a whole, in a homogeneous condition,

and that the rearrangement and formation of the various parts fol-

lowed within the mass itself.... The first stages of cooling’ and

crystallization against the outer envelope of sedimentary rocks was

relatively more rapid than that affecting the inner portion, and

resulted in producing the outer porphyritic shell.” There was,

further, a gradual withdrawal and concentration of feldspathic mate-

rial toward the inner portion of the mass and an enrichment of the

outer zone with lime, iron, ‘and ‘magnesia, thus producing the concen-

c dro.

No. 423.] NOTES: AND LITERATURE. 257.

Basic Rocks in Maryland. — A series of acid and basic rocks

intrude the gneisses of northeastern Maryland, and these in turn are

cut by extensive dykes of granite and pegmatite. The principal

types are biotite granite, diorite, tonalite, norite, hypersthene-gabbro,

pyroxenite, peridotite, and serpentine. The order of eruption seems

to have been norite and gabbro, diorite and granite. The last two

rocks are connected by gradation phases, and this is thought to be

evidence that their difference in age is not great. The peridotites

and pyroxenites are mainly younger than the norites and gabbros,

but some of the pyroxenites are apparently peripheral phases of the

norite,

The succession taken as a dol t is supposed to * furnish an exam-

ple of the occurrence of several rock types which represent the facies

of a single magma and unite to form a geological unit."

The serpentines were derived partly from the peridotites and

partly from the pyroxenites. In the latter change the hypersthene

and diallage first alter into fibrous hornblende, and this later passes

into the serpentine. The area is very similar in its essential features

to the Delaware area of gabbros described by Chester and the Balti-

more area described by Williams. The distinguishing features of

the present area are (1) the abundance of diorite, (2) the eompara-

.tive rarity of gabbro-diorite, and (3) the great abundance of t

tite and granite dykes.!

Rock Structure. — The study of a series of andesitic and rhyolitic

rocks that occur as extrusives in the Great Basin region suggests to

Spurr? that the differences in structure of igneous masses may be

brought about by slight changes in conditions under which crystalli-

zation took place. Among the differences which affect structure may

be mentioned slight changes in the rate of cooling. Textural varia-

tions are less common in acid than in basic extrusives because in the

latter the viscosity varies more rapidly with rapid cooling than in

the former. The more important structures are not characteristic of

particular rocks, but are the functions of the relation between viscos-

ity and cooling rate. Structure therefore cannot, according to the

author, be made the principal element in rock classification.

1 Leonard, A. G. Amer. Geol. (1901), p- 135.

2 Journ. Geol., vol. ix (1901), p. $86. `

258 THE AMERICAN NATURALIST.

CORRESPONDENCE,

To the Editor of the American Naturalist :

SIR : — In that very interesting and not unimportant book Memoirs

of Odd Adventures, Strange Deliverances, etc., in the Captivity of John

Giles, Esq. . . . written by himself, describing his captivity among the

Indians of New Brunswick in 1689-98, occur many quaint but truth-

ful remarks upon the habits of common animals. But among the

others occurs this: |

Of the Tortoise. lt is needless to describe the fresh-water tortoise,

whose form is so well known in all parts: but their manner of propagating

their species is not so universally known. I have observed that sort of

tortoise whose shell is about fourteen or sixteen inches wide. In their

coition they may be heard half a mile, making a noise like a woman wash-

ing her linen with a batting staff. They lay their eggs in the sand, etc.

I have asked several students of these animals for information

about the noise here mentioned, but none of them seem to know it.

Giles (or more properly Gyles) is usually so accurate in his observa-

tions that I can hardly believe he was wholly mistaken in this case.

Can any of the readers of the American Naturalist throw any light

upon the subject?

The immediate reason for my interest in the matter is an inquiry

from my friend Mr. Victor H. Paltsits, of the Lenox Library, New

York City, who is editing and exhaustively annotating a new edition

of Gyles's Memoirs, soon to be published by Dodd, Mead and Co.

of New York. W. F. GANONG.

SMITH COLLEGE, NORTHAMPTON, Mass.

(No. 422 was mailed February 13.)

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VOL. XXXVI, NO. 424 — APRIL, 1968 —

AMERICAN

NATURALISL

A MONTHLY JOURNAL mee

DEVOTED TO THE NATURAL SCIENCES :

IN THEIR WIDEST SENSE

CONTENTS

The American Naturalist.

ASSOCIATE EDITORS:

J. A. ALLEN, PH.D., American Museum of Natural History, New York.

E A. AN DREWS, PH.D., Johns Hopkins University, Baltimore.

L :

STOCK, a

JILLIAM M. DAVIS, M.E., Harvard University, Cambridge.

ALES HRDLICKA, M.D., Vew York City.

D. S. JORDAN, LL.D., Stan, Univer.

-CHA

tanford ersity.

S A. ALAS PH.D., University E Cali "eie Berkeley.

EEDHA

IHE

AMERICAN NATURALIST

VoL. XXXVI. April, 1902. No. 424.

HOMOPLASY AS A LAW OF LATENT OR

POTENTIAL HOMOLOGY.

HENRY FAIRFIELD OSBORN.

My study of teeth in a great many phyla of Mammalia in past times has con-

vinced me that there are fundamental predispositions to vary in certain directions ;

that the evolution of the teeth is marked out beforehand by hereditary influences

which extend back hundreds of thousands of years. These predispositions are

aroused under certain exciting causes and the progress of tooth development takes

a certain form SORTE into actuality what has hitherto been potentiality.

Science, N.S., Vol. VI, No. 146 (Oct. 15, 1897), pp. 583-587.

IN previous communications, as shown in the above quota-

tion, I have spoken of the * potential of similar variation," as

covering cases of the independent evolution of identical struc-

tures in the teeth of different families of mammals, especially in

relation to the homologous “ antecrochet " and “ crochet ” folds

in the teeth of horses, rhinoceroses, and we may now add, of

titanotheres (Osborn, '94, p. 208). In the present commu-

nication I propose to treat somewhat more fully of the same

phenomenon, as a special form of homology which has been

clearly defined by Lankester in 1874 as homoplasy, but into

which poleoptelogy h has. brought the idea of “ potential. n

259

260 THE AMERICAN NATURALIST. (VoL. XXXVI.

THE Broap SIGNIFICANCE OF ANALOGY.

We are familiar with the classic distinction of analogous

organs as having a similarity of function: analogy (Owen,

'43, p. 374), “a part or organ in one animal which has the same

function as another part or organ in a different animal";

Lankester (70): * Any two organs having the same function

are analogous, whether closely resembling each other in their

structure and relation to other

parts or not; and it is well to

retain the word in that wide

sense." Analogous organs

may or may not be homolo-

gous. “Analogy” is there-

fore an extremely broad and

comprehensive term, and it

appears that we must include

under it all cases of the similar

evolution of organs either of

common or of different origin

due to similarity of function.

For example, the * analogous

variation" of Darwin, the

pe ag insti ge wat roit n homoplasy" of Lankester

B, Hyracotherium.. Not believed tobe genet- 1N part at least, the “ conver- -

eme leneng dh ipo Pro on ea genz" of German writers,

l the “homomorphy” of Für-

bringer, the “heterology,” “parallels,” and « parallelism " of

Hyatt, of Cope ('68, also Origin of the Fittest, p. 96), of Scott,

and of most. American writers, are all illustrations of analogy

and may be very misleading as to homology.

As Scott observed in 1896, * Parallelism! and convergence

of development are much more general and important modes

| 1 The term “parallelism” was employed by Cope in his essay of 1868 on the

CNN of Genera” (reprinted in the Origin of the Fittest) in two quite different

No. 424] LATENT OR POTENTIAL HOMOLOGY. 261

of evolution than is commonly supposed. By parallelism is

meant the independent acquisition of similar structure in forms

[2.e., animals] which are themselves nearly related, and by con-

vergence such acquisition in forms [7.z., animals] which are not

closely related, and thus in one or more respects come to be

more nearly alike than were their ancestors."

The term **homoplasy ” (Lankester) has been long used by

the writer and others in a somewhat similar sense, but it is

not equivalent either to “ parallelism” or “convergence.” As

will be seen below, the fundamental idea is different, because

homoplasy always involves homology, while parallelism and

convergence may or may not involve homology.

ANALOGY IN EVOLUTION.

Analogous Variation (Darwin). Similar congenital variation in

more or less distantly related animals and plants.

Parallelism. Independent similar development of related animals,

plants, and organs. _

Convergence. Independent similar development of unrelated animals,

ringing them apparently closer together.

ANALOGY.

Homoplasy (Lankester) (? Homomorphy, F ürbringer). Independent

imilar development of homologous organs or regions giving

rise to similar new parts.

In brief, analogy embraces similar changes due to similar

adaptation in function both in homologous and in non-homol-

ogous organs, both in related and in unrelated animals.

Tue LIMITED SIGNIFICANCE oF Howorocy.

Owen ('43, p. 379), Lankester (70), and Fürbringer have

especially defined and elaborated the very ancient concep-

tion of homology, as employed by Oken, Geoffroy St. Hilaire,

and Vicq d'Azyr: homology (Owen, '43), “the same organ in

“On the Mode of Evolution in the Mammalia " ('91, pp. 363-367), “ parallelism ”

is used in a very broad sense as affecting the skeleton and teeth, on the principle

“that identical modifications of structure, constituting evolution of types, have

supervened on distinct lines of descent,” as embracing not only single characters

but whole series of them.

262 THE AMERICAN NATURALIST. (VoL. XXXVI.

different animals under every variety of form and function ”;

homogeny (Lankester, '70) : * Structures which are genetically

related, in so far as they have a single representative in a com-

mon. ancestor, may be called homogenous.” E. B. Wilson

(95, pp. 101-124) has shown that the comparative anatomical

test of homology is more reliable than the embryological.

Gegenbaur ('98, pp. 23—25) has given a full presentation of the

distinctions as the basis of comparative anatomy ; in his recent

great work ('98, p. 23) he presents the matter in terms which

may be briefly analyzed with the usages of other authors, as

follows :

I. HOMOLOGY, GENERAL: as of vertebrze and limbs.

I. HOMOTYPY: as of opposite limbs, eyes, kidneys, etc.’

?. HOMODYNAMY: (in part the “general,” in part the “serial,”

homology of Owen; the ‘meristic” homology of Bateson).

Corresponding limbs, parts, segments (e.g. the humerus and

femur) on the same side of the body.

3. HOMONOMY: parts which are in the same transverse axis of the

; y, or on only one section of the longitudinal axis; ez.,

the rays of the fins of fishes, the single fingers and toes of

the higher vertebrates are homonomous organs.

—

. HowoLocy, SPECIAL: (the “« homogeny " of Lankester).

1. COMPLETE HOMOLOGY of elements which have retained their rela-

tions unchanged, as of single bones from the Amphibia to the

Mammalia. :

2. INCOMPLETE HOMOLOGY, as of organs which have either gained

new parts or lost certain of their parts.

4. defective, as in comparison of fins of teleosts and of selachians.

b. augmentative, as in the heart of cyclostomes and of the higher

^ vertebrates. VEM ;

c. imitative, as where different vertebræ connect with the ilium

and become sacral. `

Hi ‘Homomorpny (Fürbringer): from these homologies certain structures

are to istinguished as homomorphic which are more or less

similar to each other but stand in no phylogenetic connection.!

_Homomorphy comes nearest, as we understand it, to the “ homo-

v»: Pplasy” of Lankester, but the latter term has the priority of

£t

* ^3 Literally translated from Gegeabagn 0.

No.424] LATENT OR POTENTIAL HOMOLOGY. 263

DISTINCTION BETWEEN HomoGENous AND HÓMOPLASTIC

ORGANS.

In the strictest sense, special or genetic homology, the

*homogeny" of Lankester, is the only absolute homology.

For example, in all four-limbed vertebrates, or Tetrapoda

(Credner), the first and second phalanges of the tibial digit or

hallux are homogenous ;

the earliest tetrapods

had such phalanges, so

far as we can judge from

both paleontology an

embryology, and all

others are derivatives.

But suppose we should

discover that these two

phalanges had originated Ps. 2.— Ideal embryonic ground plan of rhinoceros molar,

independently in several s d r

different classes of vertebrates, and were not derivatives ;

should ad then be considered analogous or homologous?

* Again," says Lankester (70), “it may perhaps be admitted

that the common ancestors of the Osseous Fishes and Mam-

malia had a skull of decidedly undifferentiated character, with

a much less amount of

differentiation than is

observed in the skulls

of either of these

groups. It is only in

so far as they have

parts represented in

the common ancestor

that we can trace Zo»o-

re iae het ei Mem nme rey i these routs;

‘and yet the homology

of a vast number of bones in the skull of the two is discussed

and pointed out." Suppose, accordingly, that in the formation

of dermal roofing bones in different orders of fishes a pair of

bones corresponding in position to the parietals should arise

264 THE AMERICAN NATURALIST. | [Vor. XXXVI.

independently, or that in the evolution of the teeth cusps should

arise independently having the same form and position, — what

criterion should be applied? All such structures are habitually

regarded as homologous, yet it is apparent that they are not

derivatives of each other and therefore not homogenous or

homologous in the strictest sense.

Such cases of independent evolution of apparently homolo-

gous organs I recently proposed! to signify as potential, or latent

homology, borrowing the term “latent”’ from Galton as indica-

tive of a germinal rather than of a patent or adult character,

and the physical term “potential” as expressing the innate

power or capacity to develop a certain organ. But my col-

league, Prof. Edmund B. Wilson, pointed out to me that such

cases were almost exactly covered by the original defint-

tion of the word *homoplasy" by Lankester ('70, p. 42), as

shown in the subjoined quotations from his essay:

When identical or nearly similar forces, or environments, act on two or

more parts of an organism which are exactly or nearly alike, the resulting

modifications? of the various parts will be exactly or nearly alike. Further,

if, instead of similar parts in the same organism, we suppose the same

forces to act on parts in two organisms, which parts are exactly or nearly

alike and sometimes homogenetic, the resulting correspondences called forth

in the several parts in the two organisms will be nearly or soci d alike.

I propose to call this kind of agreement homoplastic or Aomoplasy? . .

What is put forward here is this : that under the term « homology," belong:

ing to another philosophy, evolutionists have described and do describe two

kinds of agreement, — the one, now proposed to be called “homogeny,”

depending simply on the inheritance of a common part ; the other, proposed

to be called * homoplasy," depending on a common action of evoking causes

or moulding environment on such homogenous parts, or on parts which

for other reasons offer a likeness of material to begin with.

Homology thus includes i Hope

Homogeny.

It follows that subsequent writers, including myself, have mis-

used the term “homoplasy,” confusing it with “ parallelism "

lina communication before the National Academy of Science, Nov. 13, 1901.

? Ttalics are min:

3 At this time pcan accepted Herbert Spencer’s Lamarckian views. Sub-

erasiaka coe ee

No.424.] LATENT OR POTENTIAL HOMOLOG Y. 265

and “convergence,” which; as we have seen, may affect

absolutely non-homologous structures. Homoplasy should be

confined to structures in which there is an element of homology.

Independently of Lankester (that is, not familiar with his

paper) I had therefore reached a similar conclusion through

years of observation in paleontology. I would now like to

expand an idea which he also lightly suggested in 1870 in

the words, “or on parts which for other reasons show a likeness

of material to begin with.”

Tue Law or HOMOPLASY AS IN PART IDENTICAL WITH

DEFINITE OR DETERMINATE VARIATION.

As observed in the evolution of the teeth especially, homop-

lasy appears to be of very great importance, not on the tech-

nical grounds of uniformity in nomenclature, but because it

seems to coincide with the principle of definite or determinate

evolution, a principle which may be of wider application."

Homo, showing independent or

+h R, 1

p F

ti af the hi ne. ky fi

y : ^y,

From the time of the “ Origin of Species ” it has been admitted

that evolution, so far as it depends upon variation, is not in

every possible direction, but is limited to certain changes,

the expression of certain hereditary or constitutional causes

which we do not in the least understand. The evolution of

the teeth of mammals enabled me in 1889 to give many con-

crete illustrations of this principle and to show that variation

is hardly the proper term to apply to rudiments which do not

arise in a variable but in a fixed manner.

1 See especially the correspondence of Darwin and Asa Gray; also Osborn,

The Palzontological Evidence for the Transmission of Acquired Characters,

Nature, Jan. 9, 1890; the Orthogenesis and Orthoplasy of Eimer, Lloyd Morgan

aldwin ; Baldwin’s Dictionary of Philosophy and Psychology, vol. i, p. 243-

266 THE AMERICAN NATURALIST. . [Vor. XXXVI.

It appears that. von Waagen suggested the term “ mutation ”

for immeasurable variations somewhat similar to these. Scott

in 1891 (91, p. 388) pursued the idea further in the following

striking passage: “ These facts.at least suggest the possibility

that individual variations are not incipient species, but that the

Fic. 5. — Superior molars of primates. 4, Adapis; B, Hyopsodus; C, Notharctus. Showing

homoplastic cusps, Ay, m/, ps, ms, mts.

causes of transformation lie deeper, and act with more or less

uniformity upon large numbers of individuals. It may, per-

haps, be the outcome of future investigations, that while varia-

tions are generally due to the union of changing hereditary

tendencies, mutations are the effect of dynamical agencies

operating long in a uniform way, and the results controlled by

natural selection. While this may be true, a great many facts

must be gathered in its support, before it can be regarded as

more than a suggestion." Scott subsequently, in his article

“Variations and Mutations,” expanded this idea: * Bateson's

results, as compared with those of paleontology, confirm this

distinction in many significant ways and emphasize strongly

the difference between variation and that steady advance along

definite lines which Waagen called mutation." This paper in

turn is said to have influenced de Vries's recent work, Die

Mutationstheorie.

It is a singular coincidence that the human teeth were

selected by both Empedocles and Aristotle to test the “ survival

of the fittest" versus the purposive or teleological theory of

evolution. I pointed out in the papers above referred to

(Osborn, '89, pp. 561—566; '90) the significant fact that new

cusps of the molar teeth do not appear at random, but at

certain definite. points; that they are at first so minute that

Nov424.], LATENT OR POTENTIAL HOMOLOGY. 267

they can barely be perceived, so that it is difficult to theoreti-

cally assign them a survival value in the struggle for existence;

that the mechanical or Lamarckian explanation is the only one

which can be offered!; I laid the chief stress, however, not

upon the mechanical explanation, but upon definite or determi-

nate origin, and this has been confirmed by the subsequent

study of thousands of teeth in different families of mammals,

The still more significant fact that this definite and determi-

nate evolution was proceeding independently in à great many

different families of mammals did not at the time impress itself

so strongly upon my mind.

If molar teeth are found independently evolving in exactly

similar ways in such remote parts of the world as Switzerland,

Wyoming, and Patagonia, it is obvious that the process is not

governed by chance but represents the operation of some

similar or uniform law deduced from the four following

considerations:

Firstly, the teeth differ from all the other tissues and organs

of the body in being preformed, beneath the gum.? Unlike all

other organs they are not modified, improved, or rendered more

adaptive by use; on the contrary, after the first stage of wear,

Fic. 6. — Superior molar of Merychippus, showing styles fs, ms, mits, and conules pl, mi,

h lastic with th f th wholly 1 lp i t Fig. 5; E.

homopias

the longer they are used the more useless and less adaptive

they become. Thus, new structures in the teeth do not first

appear as modifications (as distinguished from congenital

1 Ryder and Cope confidently advanced the mechanical explanation ; it is not

without grave difficulties, owing to the lack of an heredity theory.

of this fact was first pointed out to me by Prof. E. B.

Poulton of Oxford. sitaitiug: vigiiniebal s1 MIU

268 THE AMERICAN NATURALIST. |. [Vor. XXXVI.

variations) in course of life, as is so often if not invariably the

case with new structures in the skeleton. New cusps, folds,

crests, and styles are invariably congenital. Thus, of all organs

of the body the teeth most exclusively and purely represent

the current of stirp, germinal, or constitutional evolution.

Secondly, the teeth are, nevertheless, among the most pro-

gressive organs in the body. Whereas the adaptation of the

skeleton, among the mammals at least, is by a constant loss

or numerical reduction of parts, the adaptation of the teeth |

is by a constant addition and modeling of parts (Osborn, '88,

pp. 1067—1079).

Thirdly, according to the present paleontological evidence

many of the different families and orders of mammals diverged

from each other at a time when they possessed three cusps on

the upper molar teeth and from three to five cusps on the lower

molar teeth. This being the case, only the cusps comparable

in different orders of mammals with these original three upper

and five lower cusps are derivatives or homogenous.

Fourthly, it follows that the new cusps of the teeth fur-

nish an example of Noquopler independent of the individual

modification.

Thus, we may say that in the teeth at least omoplasy

involves a law of latent or potential homology, without profess-

ing to understand what is its significance.

We should, a priori, expect that if additional cusps were

added independently in different families and orders of mam-

mals in different parts of the world, under highly different

conditions, the teeth of the higher Mammalia would present

very great diversity. As a matter of fact, the new cusps in

different families are absolutely uniform up to a certain limit.!

In the twenty-three orders of placentals and in the seven mar-

supial families, many of which are adaptively equivalent to

orders, the independently developed fourth to eleventh cusps

of the upper molars, if so many are developed, are uniform

and may be termed homologous; the eight cusps and folds

succeeding the original homogenous three arising, if at all,

1The excess of this limit is. in multituberculism, or es where

cuspules are indefinitely multiplied.

No. 424.] LATENT OR POTENTIAL HOMOLOGY. 269

at similar points and presenting a latent homology or homoplasy.

The record in the upper molar teeth stands thus :

HOMOLOGY.

AEE v S TT PAG -ETTE

HOMOGENY ‘ HOMOPLASY

Cusps or folds which are or may be

independently developed in dif-

Primitive three cusps common to

ali ferent orders.

ammals.

Protocone ypocone

aracone Metaconule

Metacone Protoconule

—- arastyle

— Hypostyle

This expresses the comparison of mammals as a whole.

Within many of the orders, such as the Perissodactyla, which

arise from six cusped ancestors, the homology is different.

HonoLocv.

eS a (rite NI. een

HOMOGENY HOMOPLASY

Protocone Parastyle

Paracone Mesostyle

Metacone Crista

Hypocone Crochet

Protoconule Antecrochet, etc.

Metaconule diss

The elements to which these terms are applied are best

exemplified in the molar teeth of some of the primitive horses

(Fig. 6).

The teeth are by no means the only structures which evolve

under this principle, the skull, vertebral column, and limbs also

evolving under it more or less completely; but the teeth afford ,

a singularly beautiful illustration of it because they exclude

individual modification.

270 THE AMERICAN NATURALIST. [Vor. XXXVI.

The chief object of this communication is to enforce the rec-

‘ognition of homoplasy as something which must be accounted

for. These homoplastic cusps do not arise from selection out

of fortuitous variations, because they develop directly and are

not picked from a number of alternates. Neither: does it

appear that the mechanical-inheritance theory, if granted,

would produce such a remarkable uniformity of result. We

are forced to the conclusion that in the original tritubercular

constitution of the teeth there is some principle which unifies

the subsequent variation and evolution up to a certain point.

Herein lies the appropriateness of Lankester's phrase, “a like-

ness of material to begin with.”

Philosophically, predeterminate variation and evolution brings

us upon dangerous ground. If all that is evolved in the Ter-

tiary molar tooth is included in a latent or potential form, in

the Cretaceous molar tooth we are nearing the embottement

hypothesis of Bonnet or the archetype of Oken and Owen.

Embryologists have recently gotten ‘into the same dilemma,

and my colleague, Wilson, has proposed to drop the idea

“homology” altogether and substitute the idea “equivalent.”

In the present case, however, I think we have to deal with

homology or, more strictly, with a principle intermediate between

homology and analogy. |

In a paper recently read before the American Morphological

Society (December, 1901), this author has urged the necessity

of adhering as closely as possible to the historical standard in

the embryological study of homology, and of avoiding the use

of the term *homology" when this standard is not available. He

therefore suggests for descriptive purposes the use of the non-

committal terms “ equivalent ” and * homoblastic," the former

being applied to embryonic structures of like fate (i.e, giving

rise to homologous parts), the latter to those of like embryonic

origin. The only decisive test of the homology is historic

community of derivation (é.e., homogeny).

.424] LATENT OR POTENTIAL HOMOLOGY. 271

BIBLIOGRAPHY.

Corr, E. D. On the Origin of Genera. Proc. Acad. Nat. Sci.

Phila. October, 1868.

Corr, E. D. The Origin of the Fittest. 8vo. New York.

GEGENBAUR, CARL. Vergleichende Anatomie der Wirbelthiere. 8vo.

Leipzig, 1898.

LANKESTER, E. R. On the Use of the Term * Homology " in Modern

Zoólogy and the Distinction between Homogenetic and Homoplastic

Agreements. Ann. Mag. Nat. Hist. Ser. 4, vol. iv, pp. 34-43.

NEuMAYR, M. Die Stämme des Thierreiches. 4to. Pp. 60-61.

Vienna, 1889.

Osporn, HENRY F. The Evolution of Mammalian Molars to and

from the Tritubercular Type. Amer. Nat. Vol. xxii (December,

1888), pp. 1067-1079.

OSBORN, HENRY F. The Paleontological Evidence for the Trans-

mission of Acquired Characters. Amer. Nat. Vol. xxiii (1889),

pp. 561-566

Osporn, HENRY F. Evolution and Heredity. Biological Lectures,

1890, Marine Biological Laboratory, Woods :

OSBORN, HENRY F. The Rise of the Mammalia in North America.

Proc. Amer. Assoc. Adv. Sci. Vol. xlii (1893), pp. 187-227.

Owen, RICHARD. Lectures on the Comparative Anatomy and Physi-

ology of the Invertebrate Animals, delivered at the Royal College of

Surgeons in 1843. 8vo. London, 1843.

Scorr, W. B. On Some of the Factors in the Evolution of the

Mammalia. On the Osteology of Mesohippus and Leptomeryx,

with Observations on the Modes and Factors of Evolution in the

Mammalia. Journ. of Morph. Vol. v, sis hi ae pp. 379-402.

Scott, W. B. On Variations and Mutations. Amer. Journ. Sci.

Vol. xlviii (November, 1894), pp. 35 qp

Scott, W. B. Palæontology as a Morphological Discipline. Biologi-

cal Lectures, 1895, Marine Biological Laboratory, Woods Hole

Pp. 43-61. Boston, 1896.

DE VRIES, HuGo. Die Boo donéibeorie: Versuche und Beobach-

tungen über die Entstehung von Arten im Pflanzenreich. 8vo.

Leipzig, 1902.

WiLsoN, E. B. The Embryological Criterion of Homology. Bio-

logical Lectures, 1895, Marine Biological Laboratory, Woods Hole.

Pp. 101-124. Boston, 1895.

NEW INSTANCES OF COMPLETE DIVISION OF

THE MALAR BONE, WITH NOTES ON

INCOMPLETE DIVISION.

ALES HRDLICKA.

Tur malar bone in man, and also among lower mammals, is

found occasionally to present a partial or a complete division.

The complete malar divisions are in general rare; the incom-

plete ones are rare in most of the species of the lower mam-

mals, but are quite common in man. Both varieties of the

divisions occur with unequal frequency in the different races

of man, and, seemingly, even in different localities in the same

people.

The first case of a complete malar division in man was

described in 1779 by Ed. Sandifort.1 Since then the anomaly

has been observed among most of the principal branches of the

whites and also among the Japanese, Ainos, Chinese, Buriats,

Bashkirs, Burmans, natives of the Malay Archipelago and of

the Philippine Islands. The most important contributions to

the literature of the subject are those by Breschet,? Wenzel

Gruber? and Virchow*; other authors who occupied them-

selves with malar divisions are Amadei, Flesch, Riccardi,’

1 Observat. Anat. Pathol. Lugd., Batav. 1779. Vol. iii, 8, p. 113. Cit. Gruber.

2 Recherches sur différentes piéces osseuses du squelette de l'homme et des

tional publications: on the subject in Arch. f. Anat. u. Phys., 1873, p. 208; 1875,

p. 194; a p. 230; and in Arch. f. pathol. Anat. u. i ING 1877, Bd. Ixix, p. 382 ;

and 1879, B d. Ixxvii, p. 113.

4 Über die ethnologische Bedeutung des os malare bipartitum, Monatsherichte

d. k. preuss. Akad. d. Wiss. Berlin, 1882, p. ag

5 Arch. p. P antropol. e P etnolog., 1877, p. 1

6 Über das zweigetheilte Jochbein, Vero. d. phys.-med. Ges. in Wirelérg,

1877; pp- 51, 52.

7 Suture anomale dell’ osso malare, Arch. 2. 7 antropol. e P afa tomo viii -

(1878); pp. I et seq.

274 THE AMERICAN NATURALIST. (Vor. XXXVI.

Hilgendorf,’ Tarenetzkij? Calorij? Matiegka," etc. In other

mammals cases of complete malar division have thus far been

found in two Cercopithecit sabei (Laurillard,? Breschet ?) ;

four Mycetes seniculi (Laurillard,. Breschet) ; in one. Mycetes

niger (Breschet); in an orang (Flesch®); in a hippopotamus

(Breschet); and in a Phascolomys wombatus (Gruber ?).

The complete malar divisions present some variety. In a

very large majority of the cases, both in man and the lower

mammals, the malar was found divided into two portions by a

suture running antero-posteriorly from the malo-maxillary to

the zygomatic articulation. In the orang described by Flesch

the posterior extremity of the anomalous division terminated

on the temporal border of the malar bone, a short distance

anterior to the upper end of the zygomatic suture. In two of

Calori's cases the posterior termination of the anomalous suture

was situated respectively at the base and above the middle of the

ascending portion of the temporal border. The second of these

cases shows much similarity with one I have at hand, in which

the malar division is due to fracture (v. the end of this paper).

‘Testut reports a case ? in which the division ran in a curve

from the ascending portion of the orbital border to the middle

of the malo-maxillary suture. Spix,!* Breschet,? Riccardi,’ and

Ruggeri,” finally, observed a division of the malar into three

portions, the location of which differed in each instance.

Of incomplete malar divisions there have been described

thus far only horizontal ones in which the suture connects with

either the zygomatic or the malo-maxillary suture.

8 Arch. f. patholog. Anat. und Phys., Bd. lxx (1879), pp. 113 et seg.

? Beiträge zur Craniologie der Ainos und Sachalin, Mém. de / Acad. nbi Sci.

de St Pétersbourg, Sér. 7, tome xxxvii (1890), p.

u le anomalie dell' osso zigomatico, etc., Mer. d. R. Accad. d. Sei. del"

Zstit. d Bologna, Ser. . 5, tomo iii (1893), p. 415.

u Prvni Lebka z janl s “Os Malare Bipartitum,” Véstnik Král. České Spoled.

Néuk, vol. xxviii, pp. 1 Prague, 1899. Also in Germ

12 In Cuvier’s Leg. FW npe 2d ed., tome ii (Padi, 1837), pp. 385, 386.

Cit. Gruber.

13 Traité d’ Anatomie Humaine, toii i, p. 174. . Paris, 1889.

M Cephalogenesis s. capitis ossei MORAN eic et eit Ss kra

' 1815), p. te Cit. Graber, enr

35 Un zigomatico tripartito, jc Serin. di Pivetiqie tomo

(1897), p- 460; refer. Central. f. Anthropol., Bd. iii (1898), p. 20. Cit. de

No. 424.] DIVISION OF THE MALAR BONE. 275

The presence of a malar division is generally held to signify

a non-fusion of the original centers from which the bone

developed. It is not yet definitely settled whether there are

usually two or three of these centers. According to Breschet

the malar bone develops in the majority of cases from one, in

a smaller number of instances from two, and occasionally from

three centers of ossification. Gruber was of a similar opinion.

Rambauld and Renault, on the other hand, advanced the theory

that the malar develops always from three points of ossification.

In the words of these authors (Origine et dévelopment des os

(Paris, 1864), pp. 161, 162, Atlas, Pl. XIII, Figs. 4, 5), **nous

l'avons (l'os malaire) toujours vu s'ossifier par un seul centre,

mais composé de trois points." ‘Ces points... apparaissent

de trés bonne heure et se réunissent trés-vite." “Vers le

milieu, et plus souvent à la fin du second mois, l'os malaire

est composé de trois points." The subject is much in need of

further embryological investigation.

Nothing is known as to the immediate causes of the Anomaly

and but little as to its signification.

The meaning and importance of the incomplete divisions

will be considered later.

New TESTAMENT OF COMPLETE MALAR DIVISION.

The new cases of complete malar division which came under

my observation comprise one instance of the anomaly in a

white adult woman, one in an adult male Peruvian (prehistoric),

a bilateral division in an orang and in a -

Lagothrix, and a unilateral one in a Cerco-

pithecus, a Mycetes, and a bear. In detail,

these cases are as follows:

Case I (Fig. 1). — The first specimen con-

sists of a part of the face, including the

right malar, of a white adult female, found

by me among the unidentified bones in Professor Hunting-

ton's osteological collection in the College of Physicians and

Surgeons (Medical Department, D MNA New

York City.

Fic. 1

276 THE AMERICAN NATURALIST. [Vor. XXXVI.

The malar bone shows remnants of an anomalous division,

which separated the frontal process from the body of the

bone.

The division runs in the line of the posterior or horizontal

position of the temporal border of the malar and consists of a

row of small, irregular perforations, with a larger, round fora-

men in the middle. It is plain, on both the anterior and

posterior surfaces of the bone, that the line of openings is

the remnant of a former complete separation of the frontal

process.

There are no signs of injury. The part of the face with the

left malar has not been recovered. The right malar shows no

further abnormality.

The malar division in this specimen is analogous with the

superior of the two divisions in the case described by Riccardi,

and is considered by Matiegka, whom I informed of the case,

to represent a non-union

of the superior of three

centers from which the

i bone developed.

j Case 2 (Fig. 2). Malar

= Division in a Peruvian. —

This is the first instance

of a complete malar divi-

sion found among aboriginal Americans. It is the only skull

with a complete division that I found among 492 old-Peruvian

and over 2000 other Indian crania from various localities. The

specimen is one of the Bandelier collection, in the American

Museum of Natural History, New York City. -

The skull is that of a male of between forty and fifty years

of age. It is of a moderate size, and shows nothing extraor-

dinary except a moderate artificial frontal compression and a

persistence of the metopic suture.

The left malar presents a complete and. patent, horizontal,

serrated suture, which divides it into a narrow lower, and large

upper, portion. The right malar shows in a similar location a

0.5 cm. long, straight, anterior, anda 0.7 cm. long, slightly

serrated, posterior, incisure. | |

Fic. 2.

No. 424] DIVISION OF THE MALAR BONE. 277

The two malars differ considerably in size, the left one being

higher, but narrower, than the right one.

Lert. RIGHT.

Highest point of fronto-malar suture vertically downward

to the anomalous suture, 4.2 cm. —--—

Highest point of fronto-malar suture vertically downward to

lower edge of the malar, 5.2 cm. 4-4 cm.

A line between the most cpi point of the malo-maxil-

lary and the zygomatic su 37 cm. 3.9 cm,

A line between the most iliis ien of the malo-maxillary

and the zygomatic suture, 2.7 cm. 3.2 cm.

This excess in height and deficiency in breadth of the fully

divided malar when compared to its mate is in full accordance

with previous observations on this point. A relative narrow-

ness of the inferior of the two portions of the divided malar is

also more the rule than the exception.

The incisures in the right malar have apparently influenced

the growth of that bone but little. The evidence of the

measurements is against their equivalence with the complete

division on the right. If these incisures are remnants of a

complete division, the measurements indicate an early union

of the middle portion of the malar.

The presence of the frontal division has ipie» no relation

to that in the malar bone. A persistent metopic suture is

present in 97 of the 492 Peruvians, without there being another

case of a complete

malar division ; there

is also in these skulls

no excess of malar

incisures.

Case 3 (Fig. 3). A

Bilateral Complete Ln i

Malar Division in an R

Orang. — The skull is

that of a young orang and is a part of a mounted skeleton in

Professor Huntington’s museum in the College of Physicians

and Surgeons, New York City. Besides the malar divisions

the specimen shows nothing unusual.

The malar sutures are in every way much alike on the two

sides. They run each with a moderate double curve from near

Fic, 3.

278 THE AMERICAN NATURALIST. [Vor. XXXVI.

the middle of the malo-maxillary to very near the superior ter-

mination of the zygomatic suture. Both divisions are slightly

serrated. -

The right malar bone is slightly higher than the left one,

but the frontal process and to a less extent the body of this

latter is broader.

The situation of the malar sutures in this case is similar to

that in the orang described by Flesch; the posterior termina-

tion of the division in Flesch’s case lay, however, still some-

what higher, terminating on the temporal border of the malar,

slightly anterior to the zygomatic suture.

Case 4 (Fig. 4). A Unilateral Complete Malar Division in

a Cercopithecus callitrichus.— Among the forty skulls of Cerco-

pitheci in the zoological collection of the

American Museum of Natural History, New

York City, there is one with a complete and

one with a partial malar division.

The skull with the complete division belongs

to an adult female, C. callitrichus (No. 13,923),

and shows besides the malar also a bilateral parietal division.

The malar division is found on the left only ; there is no

trace of a similar anomaly on the right side.

The malar suture begins anteriorly at a point between the

lower and middle thirds of the malo-maxillary articulation, runs

backward parallel with the inferior border of the malar, and

terminates in the curve of the temporal border of the bone,

0.8 cm. anteriorly to the upper end of the zygomatic suture.

The divided malar is in whole higher than the undivided one

(l. 2.9 cm., r. 2.4 cm.), and its frontal process is broader; but

the bodies of the two bones are of very nearly the same

dimensions. |

The lower separate portion is relatively narrow.

_ Neither in this nor in any of the preceding cases is there an

extension of any process of bone from the zygoma to the

maxillary.

Fic. 4.

The case of a partial malar division in a Cercopithecus (Fig. 5)

may be conveniently described in this connection. The skull

showing this anomaly is that of an adult animal of unknown

No. 424.] DIVISION OF THE MALAR BONE. 279

sex and variety. It is free from other abnormalities. The

division occurs in the right malar bone. It begins distally in

the deepest portion of the curve of the temporal border of the

bone, is somewhat curved, well serrated,

and extends forward to the middle of the

body of the bone, It is equally pronounced

on the anterior and posterior surface of the

malar. The divided bone is externally very

slightly higher than the other malar, and

its body is slightly narrower.

Case 5 (Fig. 6). — The skull of an adolescent male Lagothrix

humboldti (specimen No. 6333, American Museum of Natural

History, New York) shows a bilateral, antero-posterior, com-

plete malar division. Anteriorly the division begins on each

side slightly below the middle third of the maxillary border of

the bone and from this point runs horizontally backward, to

terminate on the temporal border of the malar. The posterior

termination of the division reaches to within one millimeter of

the superior portion of the zygomatic suture on the left and to

within two and a half millimeters of the same on the right side.

Both the anomalous sutures are serrated. They show both

much better dorsally than ventrally. In this specimen there

is on each side but a slightly incomplete temporo-maxillary

arc. The large malar foramina (regular in these monkeys) are

situated well above the divisions.

The two malars are very nearly

of equal size, except anteriorly,

where the right bone is slightly

higher.

The skull shows no other

anomalies, nor anything pathological. In three other skulls of

monkeys of the same variety the malars are normal.

— Case 6 (Fig. 7). — Among twelve skulls of Mycetes, in the

American Museum of Natural History, there is one (No.

14,169) which shows a complete unilateral malar division.

The skull with a complete division is one of an adoles-

cent male M. alouata, and is otherwise in every respect

normal. ihi i den

Fic. 5.

R PAR.

Fic. 6.

280 THE AMERICAN NATURALIST. [Vor. XXXVI.

The division affects the left malar. It begins anteriorly

slightly above the lower third of the maxillary border of the

bone and runs, with some serration, backward, very nearly par-

allel with the inferior border of the malar, to the

zygomatic suture near its superior termination.

Ventrally the division is covered by a broad

temporo-maxillary arc.

The right malar is normal. It is both lower

"eT and narrower than the left bone, and even the

zygomatic arch is lower on the right.

MEASURES.

LEFT. RIGHT.

Height along the maxillary border, 2.0 cm. 1.85 cm.

Width at middle (width minimum), 0.6 cm. 0.55 cm.

Height of temporal process vertically,

from the superior end of the zygo-

matic suture, 0.8 cm. 0.65 cm.

In a skull of Mycetes, in the same collection, there is in

each temporal border of the malar, slightly anterior to the

zygomatic suture, a short incisure, directed forward.

Besides those already mentioned I have examined, for malar

divisions, the skulls of 6 Lemurs, 5 Nictipitheci, 31 Cebi,

IO Ateles, 1 Szlenus vetur, 16 Cercocebi, 1 Senmopithecus,

54 Macaques, 4 Mormons, 1 Innuus, 2 Cynopitheci, and 16 Cyno-

cephali, —all in the collection in the American Museum of

Natural History. Among these specimens there is none with |

complete and only a few with incomplete malar divisions. Of

these latter only the following one deserves to

be mentioned in this place.

Cynocephalus olivaceus (No. 10,747 ; Fig. 8).

— The left malar shows a faint and only

dorsally visible, incomplete, serrated suture,

running from a. point a very little above the

lower third of the maxillary border of the oe

bone to the malar foramen. The length of the suture is 1.1 cm.

The right malar shows traces of a similar division. The two

malars are almost exactly equal.

No. 424.] DIVISION OF THE MALAR BONE. 281

This case differs from any thus far on record, The exact

nature of the division is uncertain.

Case 7, Unilateral Malar Division in a Bear. (No. 4282,

American Museum of Natural History ; Fig. 9). — The division

affects the left malar of a young American black bear (incisors

and canines not yet replaced by teeth of the second dentition).

Dorsally only the superior third of the division is open, the

other portion of it being secluded but still traceable ; ventrally

the anomalous suture is patent in its entirety.

The division begins anteriorly considerably below the middle

of the malo-maxillary articulation and runs in almost a straight

Fic. 9.

direction upward and slightly backward, to the base of the

frontal process. The suture shows a slight serration.

The frontal process of both the malars is partly detached.

The right malar shows a small anterior incisure in a location

corresponding to the antero-inferior terminus of the division

on the left. Finally, there are on the left one small bone and

two larger, oblong ones, and on the right one large, oblong

osside, intercalated between the antero-inferior two-thirds of

the malar and the superior maxilla.

The two malars show but little difference in size.

The above specimen is one of twenty young adolescent and

young adult skulls of bears of the same variety preserved in

the American Museum of Natural History. No other of these

skulls presents a complete malar division similar to that in

No. 4282; several of the specimens, however, show an anterior

malar incisure, visible on the internal surface of the bone.

Besides this almost all the specimens present more or less of a

separation of the frontal process of the malar. In four of the

1$ A similar bone has been found in man by wg (Arch. f- Anat., Physiol,

etc. (1873), p- 195, Pl. V, Fig. 1). See also Bresc

282 THE AMERICAN NATURALIST. [VoL. XXXVI.

skulls the separation of the process is complete, in two bilater-

ally and in one on one side. The following figure shows one

of these instances ; in this case there is not only a complete

separation of the frontal process, but also a. superior incisure,

running in a similar direction to the anomalous suture in the

young bear (No. 4282; Fig. 10).

Separations of the frontal process, such as the one just

pictured, have been observed in the seal and walrus (Meckel,

Pander, d’Alton, Hallman, Kostlin, cit. Gruber; the condition

is present in a number of young seals in the collection in the

American Museum of Natural History) ;

they remind one of the superior division

of the malar in Riccardi’s case and the

one similarly situated in the first case

reported in this paper.

The division of the body of the malar bone is, as shown in

the bear (No, 4282), independent of the separation of the frontal

process. This latter appears in the bears as an P CPD si of a

comparatively late formation.

It is interesting to note that the nice termination of the

malar suture, situated somewhat higher in the orangs, is placed

still higher, relatively, in most of the specimens in lower

monkeys and other mammals. This condition is very marked

in the Cercopithecus and especially in the young bear. The

cause of this lies, it seems to me, not so much in a different

situation of the centers from which the malar develops, as in

the different tilting of the bone, and in the differences in the

temporal process, in man and the mentioned animals. The

human malar bone is less tilted than that in bears, and its tem-

poral process is considerably shorter and relatively broader than

that of apes, monkeys, and bears, as well as other mammals.

FIG. 10.

PARTIAL MALAR DIVISION.

The ; anterior and particularly the posterior portion of the

malar shows occasionally a partial division. These. divisions

occur mostly in the form of lineal incisures, but occasionally

also as serrated sutures. They differ much in b si in some.

No. 424.] DIVISION OF THE MALAR BONE. 283

cases not exceeding oné or two millimeters, in others extending

to near the middle of the malar, In man they run as a rule in

an antero-posterior direction, beginning distally from either the

zygomatic or the malo-maxillary suture.

The posterior divisions are in the average more pronounced

and much more frequent than the anterior ones. An ante-

rior and a posterior partial division may coexist in the same

bone, and there is a tendency, though there are exceptions, to

similar divisions on the two malars.

There is a considerable difference in the frequency of these

partial malar divisions in different peoples. They are found in

the adults of both sexes, but grow more rare as age advances ;

they are, according to my data, somewhat more frequent in

early than in adult life.

These malar incisures are well known, but their meaning is

still somewhat doubtful. Gruber, who was the first to give

these divisions close attention, considered them as morpho-

logically distinct from the complete separations. In the words

of this author (Virek. Arch. Bd. LVII, 1879, p. 117), “os

sygomaticum bipartitum is a term applied only to that malar

bone which is fully divided into two portions and persists thus

divided after the second month of the foetal life, as well as

after birth ; or to that malar, which was originally fully divided,

but in which the anomalous suture became later (in adult life),

in consequence of synostosis, more or less occluded.” ‘When

[ebid. | in a foetal malar bone, through a hindrance in develop-

ment (Bildungshemmung), the streaks between the orbital and

maxillo-zygomatic portion of the bone do not on one or both sides

ossify, and this defect of ossification persists, in the form of an

incisure, into advanced age, leaving thus a sort of a sutura men-

dosa, in that case one evidently cannot speak of a divided malar."

This apparently clear and rational theory was three years

later opposed by Virchow.! This author expresses himself

(p.251) as “inclined to recognize the majority of all malar

incisures, particularly the posterior Ones, às real remnants of

an old transverse malar suture"; but (p. 252) “the interpre-

tation of a posterior incisure will be the more doubtful, the

higher the situation of the incisure." . - unicus SEL

284 THE AMERICAN NATURALIST. . [Vor. XXXVI.

Both Gruber’s and Virchow’s opinions gained adherents, and

in the absence of decisive embryological investigation the matter

is still unsettled.

The significance of the partial malar divisions can be actually

demonstrated only by studies on the embryo. Nevertheless,

the examination of older subjects brings out a few points from

which, it seems to me, it is possible to draw a few deductions

having a bearing on the nature of these divisions.

Such points are the size of the incisures, their somewhat

greater frequency in childhood than in adult life, and the lack

of correspondence in frequency between the incomplete and

complete divisions in childhood in general or in the adults of

different ethnic groups.

All the points mentioned speak against an exact equivalence

of the partial and complete malar divisions and for Gruber’s

theory. Were all or a majority of the incisures remnants of

formerly complete separations of the malar, it is reasonable to

suppose that we should, even if rarely, meet with a remnant of

the separation that would extend over more than a half of the

bone ; and it would be logical to expect a greater frequency of

complete divisions in early than in later life and in those tribes

or races in whom the incisures are more common than in others.

As these conditions are not realized we must search for another

explanation for the incisures. This explanation can be found,

as already pointed out by Gruber, in the analogies of the devel-

opment of the malar with other cranial bones, particularly the

occipital, or the parietal.

The parietal bone will, I think, serve the demonstration even

better than the occipital. The bone develops, as was shown by

Toldt and Ranke, from two superimposed centers of ossifica-

tion, — a very. similar process to that which undoubtedly takes

place in the body of the malar, though this bone may possibly

develop from more than two original points. The two centers

of the parietal have but a very brief separate existence. They

grow in all directions and very soon coalesce, Their union

takes place at their reciprocally most advanced portions, in the

middle, the parts anterior and posterior to this point of fusion

e D 3) |

pa by a more or less V-shaped space or fissure.

No. 424.] DIVISION OF THE MALAR BONE. 285

This fissure, as the bone grows, gradually closes at its proximal

extremity, but becomes extended, by the new bone, at its distal

end. The result of this is that even as late as at the birth of

the human foetus the parietal bone shows a more or less marked

posterior, and rarely also an anterior, partial division, exactly

analogous with, it seems to me, though almost generally less

persistent than, the malar incisures. The parietal division may

persist even up to adult life and then the analogy between the

same and a malar incisure is still closer. The difference

between this form and a complete division (or a remnant of

such) in either the parietal or the malar is that in the simple

incisure and fissure we have before us only a retardation of the

union of the distal portions of the original segments of the

bone, while in the complete division (or its remnant) we are

confronted with a case of a complete non-union and separate

development of these segments. The two effects may be pro-

duced, and probably are produced in such a case as that in the

Peruvian skull with a complete malar division on one and two

incomplete ones on the other side, by different degrees of the

same cause ; nevertheless they cannot be considered identical.

The cause which induces a separate development of all or

some of the centers of ossification of a bone must be potent

and be present extremely early ; a persistence of a fissure might

be due to a weaker cause of the same nature, but possibly also

to later causes of differing kind. The complete divisions deserve

fully the term “anomalies,” or “atypical formations," while the

partial divisions come more under the category of developmental

defects or retardations.

The partial malar divisions are not attended by similar con-

ditions in other bones of the skull. They also do not seem to

stand in any relation with the metopic division ; specimens

with the frontal suture in my Peruvian series have, upon an

additional inspection, shown no greater. percentage of malar

incisures than other skulls in the same collection.

There is nothing in the appearance of the partial divisions,

except, perhaps, their size, which would allow us to differentiate

them from remnants of complete malar sutures. Should a

division extending beyond the middle of the bone be found, it

286 THE AMERICAN NATURALIST. [Vou. XXXVI.

would then be safe, I think; to consider it a remnant of a partly

occluded complete suture. Tarenetzkij would distinguish the

incisures from remnants of complete divisions by the greater

height of the malar in which a complete division had existed

than that of its mate, but such a criterium, though theoretically

seemingly correct, meets in practice with numerous difficulties.

Not many malars, even though entirely free from divisions, are

exactly equal. The presence of partial divisions, especially

when large, is probably not, without influence on the size of

the bone. Where a complete suture had existed but became

early subject to partial obliteration, the size of the malar was

undoubtedly less affected than in the cases where the suture

remained patent until more advanced life. Finally, where a

bilateral complete division existed, if obliteration on the two

sides proceeded simultaneously, the malars would be liable to

be of a very nearly equal size, and the indications of a complete

suture would be absent; or, had the occlusion proceeded with

much difference in time, the two malars would be unequal, and

we should be predisposed to consider only the incisures in the

larger malar as the remnants of a complete suture.

Actual measurements bear out well the uncertainty of basing

any conclusions upon the difference in the size of the malars.

Taking at random one hundred skulls of Peruvians of both

sexes, in whom both malars were well preserved and the malar

articulations were free from any signs of occlusion, I measured

the dimension of the malar mostly affected by a transverse

division, namely, the height of the body of the bone. The

measurements were taken with a sliding compass at the middle

of the bone, parallel with the malo-maxillary suture. Among

the one hundred skulls there were seventy-two in which the

malars showed no division, sixteen in which both malars showed

a posterior incisure,? four in which the right, and seven in which

the left, malar presented a posterior incisure, and one in which

both malars showed both posterior and anterior incisures, The

results of the measurements, which hardly need spre comment,

will be found i in the table on the opposite. page.

1 Tarenétzkij.- Mim. de l'Acad. Imp. de Sei. de St. 5e EUM tome xxxvii,

No. 13 (1890), p. 39. 18 Only lineal incisures above 1 mm. in length considered.

287

DIVISION OF THE MALAR BONE.

No. 424.] :

LARGER BY LARGER BY LARGER BY LARGER BY LARGER BY | TNCS BY

0.5 MM.“ 1 MM. 1.5 MM. 2 MM. “2.5 MM; S ‘= 3 MM,

EQUAL. ni e oe e

1) Malars with no division . . . . .| 29 6 5 10 na 3 2 033 6 | ale 2 $ o.

2) Both malars with a posterior incisure. | 3 3 a o 3 3 o 2 o 1 "bo Ee 3|.

3) Right malar with a posterior incisure . I o o I o o o o o I I o o

4) Left malar with a posterior incisure . | 3 o 2 1 o o o o I o o bo o

5) Both falari with an anterior and a i

posterior incisure . . o I o o o o o o o o o p o

RÉSUMÉ : ;

EQUAL. Lert MALAR HIGHER. |RiGH T Marar Hiner.

ry. ee 29 24 19

A Te 3 9 4

Sh so) RF see I 2 I

4). ^. eee 5 Ua I 4:

goce! > o 1 o ;

288 THE AMERICAN NATURALIST. (VoL. XXXVI.

The incomplete malar divisions occur in whites, according to

various authors,” with the following frequency :

Gruber found the posterior incisure among 4000-5000 mainly Russian skulls in

28, or 8.5 to 10.7% and among 50 skulls of children and embryos in 7, or 14%.

Meyer found the incisures among 517 German skulls in 20, or 3.9%; among

2 Russian skulls in 3, or

Tarenetzkij found the incisures among 45 French skulls in 2, or 4.4%; among

416 Russian skulls in 28, or 6.7%.

Anutschin (source and kind of incisures not stated) in 1.277.

Matiegha found among 300 Czech skulls the posterior incisures in 22, or 7.3%; in

anterior incisures in 6, or 2.0% of the cases.

All the authors who examined various ethnic crania for the

partial malar divisions (Meyer, Sarasin, Tarenetzkij, Anutschin,

Koganei, etc.) found them to be more frequent in almost all

other peoples (Australians excepted, Anutschin) than in the

whites,

In the human skulls that I have examined for this condition

the partial malar divisions were distributed as shown in the

tables on pages 290 and 29r.?!

The data given on these pages show plainly :

I. A considerably greater frequency of partial malar divi-

sions in the American aborigines than in the whites. This

excess in the partial is not associated with any corresponding

excess in the complete divisions.

2. The divisions among the American aborigines occur

relatively more frequently in the females than in the males and

most frequently in children; however, the differences in the

percentages in these three groups are not excessive.

3. The posterior divisions greatly preponderate in frequency

over the anterior ones.

4. Both the anterior and posterior incisures are much more

frequently bilateral than unilateral.

in Pat after Matieghka.

2 Has unusually low percentages with all the groups examined — apparently

— — — recordin ng

No. 424.] DIVISION OF THE MALAR BONE. 289

No malar incisures were found in the Australians and Poly-

nesians. Among the various American tribes, the largest pro-

portions of the incisures were found among the Peruvians and

the Southern Utah Cliff Dwellers, but in general the data on

the Americans, or at least those on the better represented

tribes, show a considerable agreement.

In mammals below man malar incisures appear to be very

rare. However, I found some instances of the condition in

monkeys and a bear. The cases in monkeys, besides those

mentioned, are as follows :

Cynocephalus porcarius (No. 10,277, A. M. N. H.). — The left

malar shows dorsally a marked anterior incisure, beginning

in the maxillary border of the bone a short distance above

the antero-inferior angle and directed obliquely upward and

backward.

The skull of a young Cynocephalus (No. 135, C. P. and S.)

shows on each side an anterior malar incisure somewhat similar

in location and direction, but smaller, particularly on the right,

to that in the preceding case.

In a Cercocebus mona (No. 5025, A. M. N. H.) a 0.5 cm.

long, horizontal incisure divides slightly above its middle the

temporal process of the left malar.

The subject of partial malar divisions could be finished here

were it not for two other forms of incisure which are met with

on the bone.

The first of these incisures, already mentioned, is the one

which occurs at the base of the frontal process of the malar in

some animals. It is common in younger American bears ; it is

also occasionally found in seals and other pinnipeds, but is

rarely seen in other classes of mammals. This division is a

remnant of a complete separation of the frontal process of the

malar; no analogous incisure has been found thus far in man,

apes, or monkeys.

The second special form of partial malar division seems to

occur only in man. It consists of a more or less marked, ver-

tical, or more often somewhat oblique, frequently bilateral

incisure, which runs downward from some part of the malo-

frontal suture. So far as I can find, this incisure has not yet

THE AMERICAN NATURALIST. [Vor. XXXVI.

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DIVISION OF THE MALAR BONE.

No. 424.]

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292 THE AMERICAN NATURALIST. . [Vor. XXXVI.

been described. I have met with it in the skulls of whites

as well as in those of Indians. It is a rare condition,

much more rare in the Indians than the divisions in the pos-

terior portion of the malar. Thus far I have found no speci-

men which would offer an explanation of

the significance of this superior incisure.

Occasionally, as in the specimen illustrated

in the accompanying figure, it is in a curv-

‘ing line with several malar foramina.

In my examination for malar divisions I

came across three specimens which may prove

of interest in connection with this paper.

In looking over the few bear skulls in the collection in the

College of Physicians and Surgeons (New York), I found one

with a peculiar anomaly present on both sides. Both malar

bones in this specimen show a partial separation of the frontal

process, but no other division. There is, however, on each

side a complete separation, by a previous, well-serrated suture,

of that portion of the zygomatic process of the temporal bone

which covers the temporal process of the malar (Fig. 12).

The two other specimens are human malars (Nos. 1068 and

1069, College of Physicians and Surgeons, New York), one of

which shows the marks of a complete, the other those of an

incomplete, vertical fracture. Both of these malars

present a pronounced depression

over their tem- _

poral portion.

Fic. 11.

Fig. 12:

Fic. 13. Fic. 14.

Some force was undoubtedly applied to this point, and the malar

bone was fractured over and along the line of the internal malo-

maxillary articulation. The border of the maxillary articular

surface acted as a fulcrum over which the bone broke. There

is no callus formation, and both cases. bear some resemblance to

a malar suture (Figs. 13 and 14).

No. 424.] DIVISION OF THE MALAR BONE. 293

In connection with the search for malar incisures in Indians

two other points were noted, namely, the lineal, antero-posterior

groove on the ventral surface of the malar (Gruber) and the

temporo-maxillary arc. The groove was found to be almost

constant, while the arc is generally more or less incomplete.

NOTE.

While my paper on malar divisions has been in the hands of

the printer, the American Museum of Natural History has

received two skulls of adult African lions, one of which shows a

partial malar suture.

The suture is in the left malar; the right malar has been

fractured (post-mortem), and its posterior half is wanting. Dor-

sally, the separation runs sagittally from a point slightly pos-

terior to the angle of the

temporal border of the

malar and a somewhat

greater distance anterior

to the zygomatic suture,

over more than half of

the malar, after which it

bends downward and ter-

minates; the division is

somewhat serrated and

contains five small Wor-

mian ossicles. Ventrally, the malar suture runs in a serrated

manner a short distance forward and unites with the boundary

of a large Wormian bone, the anterior extremity of which

reaches to a point slightly posterior to the dorsal termination

of the suture.

The malar shows no signs of any recent or old injury, and the

division is undoubtedly congenital. It is probably a remnant of

a complete malar suture, the anterior extremity of which became

subject to an early synostosis. The curving anterior end can

be explained by a simultaneous excess in growth of the anterior

portion of the superior, and the posterior portion of the inferior,

part of the malar.

Fic. 15.

294 THE AMERICAN NATURALIST.

The frontal process of the bone shows in its summit a vertical

incisure.

There are eighteen other lion skulls of various ages in the

Museum, but in none of these is there any indication of a

sagittal malar division.

COLLATERAL BUDDING IN ANNELIDS OF THE

GENUS TRYPANOSYLLIS.

HERBERT PARLIN JOHNSON.

AsipE from the marvelous development of an extensively

branching stock in Sy//is ramosa, the only mode of asexual

proliferation hitherto known among annelids is the linear stolo-

nization so well exemplified by species of Autolytus, Myrianida,

Procerastea, Nais, and a few other genera of small chaetopods.

Previous to the discovery of the extraordinary forms about to

be described, the production of numerous co//ateral buds from

a definite and circumscribed proliferating region has never, it

is safe to say, been even imagined as among the possibilities of

annelid development.

As briefly announced more than a year ago,! two species of

Syllidze living on our Pacific coast have been found to produce

sexual zooids by collateral budding. These I have studied as

fully as the scanty material permits, and the results, though

far from complete, are, I trust, of sufficient interest to warrant

their publication, particularly as the obtaining of more material

is a matter of great uncertainty.

Both species undoubtedly belong to the genus Trypanosyllis

as defined by Malaquin,? and both are unusually large for

syllidians. The larger, for which the name Z77ypanmosyllis

ingens is proposed, is a veritable giant of its kind, measuring

13 cm. in length, exclusive of the tail buds, and 6 mm. in width.

The only specimen of this rare species I have seen was given

to me by Prof. Harold Heath of Stanford University, to whom

my grateful acknowledgments are due. It was found by him

under a stone between tide marks, at Pacific Grove, California.

1Johnson, H. P. A New Type of Budding in Annelids. Paper read at the

eleventh annual meeting of the American Morphological Society at Baltimore,

Dec. 27, 1900. An abstract was published in the Biological Bulletin, vol. ii, No. 6

(1901). .? Malaquin, A. Recherches sur les Syllidiens, p. 72. Lille, 1893.

295

296 THE AMERICAN NATURALIST. | [Vor. XXXVI.

The worm is much flattened in the horizontal plane, measuring

only 2 mm. in its greatest dorso-ventral diameter. The para-

podia (Fig. 1) are very diminutive for the size of the trunk, and

the somites are so short that the stout dorsal cirri (inasmuch

as their thickness exceeds the length of the somites) form an

alternating series. The somites number nearly 500.

As Trypanosyllis ingens is undoubtedly new to science, a

diagnostic description, with figures sufficient for its identifica-

tion, is given. It is unfortunately too rare to serve as a

BiG); — Anterior: Separadas rypanosyllis in, The palpi, antennæ, median cirrus

peristomi and all the d cirri, och cus one enti, are broken off at the basal

sui. x A ., peristomial cirri ; d.c., dorsal ci

subject for study of collateral budding. It occurs in none of

the numerous gatherings of Pacific coast Polychzta that came

into my hands during six years' residence in California; and

Dr. Heath, who has done a great deal of collecting at Pacific

Grove and vicinity at all seasons of the year, informs me that

he has found only this specimen.

Trypanosyllis loge Sp. nov,

Form elongated, broad, much depressed ; dorso-ventral differ-

entiation very slight, dorsum slightly convex, ventral surface

plane, longitudinally bisected by a narrow median welt, sharply

bounded by two fine parallel grooves ; extremities evenly and

abruptly rounded off; slightly more tapered posteriorly than

anteriorly ; somites very short, crowded, about 476 in number.

No. 424.] THE GENUS TRYPANOSYLLIS. 297

General coloration tawny ; parapodia, dorsal cirri, and ante-

rior extremity, purple; caudal buds, from presence of ova, rich

yellow.

Prostomium (Fig. 1) trapezoidal, its length two-thirds of its

width, distinctly bilobed, the lobes scroll-shaped, with a deep

median notch at posterior border; eyes 4, on anterior scrolls,

connate. Palpi, antenna, and median cirrus, all wanting in

the type specimen.

Peristomium somewhat broader than the prostomium, enclos-

ing the latter posteriorly and on the sides; with dorsal and

ventral peristomial cirri (Fig. 1, .¢.).

Parapodia uniramous, short, crowded, with

bases of the dorsal cirri interlocking (Fig. 1,

d.c.); dorsal cirri (Fig. 2, d.c.) thick, coiled,

articulated, rather rapidly tapering towards

the tip; ventral cirrus tongue-shaped or fusi-

form ; setze (Fig. 3) 11-13 in each foot, with

Fic. 2.

Fic. 2.— Foot of 7rypanosyllis NC = about the middle of - length. ac., aciculz

(three); d.c., do ; Pa., Ventra

tral cirrus.

Fic. 3. — Seta of the same, in profile. Si Fic. 74 — Seta, sexual so of the ame. X 467.

toothless hooked appendage ; aciculz (Fig. 2, ac.) 3-5, their

distal ends extending into two finger-like lobes, one usually

much thicker than the other, at the dorsal tip of the foot.

Mouth large, its transverse diameter equal to width of pro-

stomium; bounded by 7 crenulations. Somites 1-8 curved

backward around math, as in Anrhe tan and Pal-

myridæ.

298 THE AMERICAN NATURALIST. (VoL. XXXVI.

Proboscis short and small (2 mm. long), thin-walled, bounded

posteriorly by a double ring of fleshy papille, 9-10 in each

ring, those of the outer circle more than twice as long as the

inner ones.

Esophagus short (3.5 mm.), and of small diameter (1 mm.)

for the size of the animal, with usual chitinized lining, which

terminates in a “trephine ” anteriorly. CEsophagus surrounded

Fic. s.— Seven somites from the middle region of a female sexual zooid of Trypanosyllis

ingens, viewed from the dorsal aspect. With two exceptions (d.c.) the dorsal cirri are

broken off. The rudimentary alimentary canal (a/.c.) is indicated ; also the ova on the

left side and in one parapod on the right. The longitudinal muscle bands (/.77.) mark the

bases of the parapodia. In clove oil. x 19.

by 9-10 glands, of wavy or crenulate outline, opening near the

papille, their blind ends extending back a short distance on

the proventriculus.

Proventriculus also small for size of the animal, 6 mm. long,

1.5 mm. in greatest transverse diameter, slightly club shaped,

thickest anteriorly, tapering gradually to posterior end; two

well-defined raphes. No coeca were detected.

Intestine with extensive paired diverticula in every somite

back of the proventriculus.

Sexes distinct ; sexual products, so far as known, developed

only in a cluster of collateral buds, growing from a definite

proliferating region near the posterior end; probably also in

the parental somites posterior to the budding zone.

Length, 130 mm.; greatest transverse diameter, including

parapodia, 6 mm.; dorso-ventral diameter, 2 mm.

No. 424.] THE GENUS TRYPANOSYLLIS. 299

. Habitat, Pacific Grove, California; between tide marks.

The form seemingly most nearly allied to 7. ingens is Tryp-

anosyllis gigantea (M’Intosh) Ehlers of the Southern Ocean.

The latter is but little inferior in size, measuring, according to

M'Intosh,! 90 mm. in length and 7 mm. in breadth. The form

—— PN

; a7 a Bi

i n. C * ^ dia.

ep- - 4

ts a

tm. JT / La

$ E =

Fic. 6.— Transverse section, female zooid of T. ingens, right half. A portion of the dorsal

body wall (indicated by dotted lines) is broken away. The plane of th ti ters

ight. aZ.c., rudimentary alimen!

the septum on the rig men tary ; d.v., dorsal blood vessel ;

v.v., ventral ditto; mes., dorsal mesentery ; £7., problematic glandular structure (external

: Sei erse muscle ;

extremity) ; #.c., nerve cords ; /.m., longitudinal musculature; £.»»., transv m

ef., epidermis. x roo.

Lo?

Lda a

"uu. ee n ES

Fic. 6 A.— Problematic gland-like tube, from right side of another section of the same series,

with portion of body wall adjacent. The external extremity (aperture?) is toward the

right. The tube is laid open by the section, showing ends of columnar cells. ring

as in Fig. 6. x roo. yo Bie

is very similar, the parapodia of both have multiple aciculæ

(6 in T. gigantea), and the sete of the.two species are prac-

tically identical. There is nothing, however, in M’Intosh’s

description or in his figure of a cross-section to indicate that

-1M'Intosh, W.C. Challenger Reports. The Polychaeta, vol. xii (1887), p. 193-

300 THE AMERICAN. NATURALIST: [Vor. XXXVI.

T. gigantea has the great intestinal diverticula so conspicuous

in 7. ingens and in 7. gemmipara (Fig. 16). . The prostomium

of 7. gigantea is also of simpler form.

Unfortunately the. delicate tail buds of this formalin- -pre-

served specimen were somewhat macerated and broken in

transportation ; most of them had fallen off, and probably some

of them were missing, when the specimen came into my

hands. They number about 30, and are all at about the

same advanced stage of development. Each bud exhibits the

external characters of the stock. Its somites have nearly

the same proportions ; its parapodia, with their diminutive cirri

and setz (Fig. 5), are of the same pattern, with the slight

difference that the appendages of the setze have a small tooth

and minute serrations on the concave side, while these are

absent in the setae of the stock (cf Figs. 3, 4). The average

length of the buds is 5 mm., their breadth 2 mm., their

thickness 0.5 mm. The number of their somites is approxi-

mately 25.

Each zooid is divided into a ERA series of coelomic spaces

by the intersegmental dissepiments and a minute median

tubular structure, which I interpret as a rudiment of the ali-

mentary canal (Figs. 5, 6, a/.c.). The coelomic spaces, includ-

ing even the parapodial cavities (Fig. 5), are packed full of

eggs, which possess very large nuclei and nucleoli, the latter

usually vacuolated (Fig. 6) ^ No ova have been found in the

parent stock. The longitudinal muscular system of the bud is

not highly developed (Figs. 5, 6, Zm.), and no circular muscu-

lature was detected. | Circular muscles are very slightly devel-

oped in the stock. There is a slender transverse muscle

extending from the lateral body wall to the nerve cord (Fig. 6,

£m.) This, however, I suspect is not so much a separate muscle

as a portion of the very extensive system of septal muscles.

A. dorsal and a ventral blood vessel (Fig. 6, dv, v.v.) are

seen close to the digestive tract. The ventral vessel with its

coagulum. is apparently of uniform diameter throughout its

course, while the- dorsal vessel is segmentally constricted

(Fig. 6 "pe a constricted. € No lateral. branches

have been detected

No. 424.] THE. GENUS -TRYPANOSYLLIS. 301

A pair of problematic. organs, possibly much-modified

nephridia, are. present in each segment. These organs are

tubular, with rather narrow lumen, their thick walls composed

of columnar, much-vacuolated cells (Fig. 6.4), and no opening

has been seen at either extremity. They occupy a ventral

position, the internal extremity being directed towards the

median line, and the outer extremity obliquely towards the

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"Fic. 7.— Caudal buds of Trypanosyllis gemmipara, dorsal aspect. The Reon qiie

pornn nf the intaati pe 3 ? 6X h EE * ti

of the stock. — x 46.

ventral body wall, which it penetrates as far as the epidermis.

It is known that in several species of syllidians the segmental

organs serve as genital ducts, and for this purpose undergo

enlargement at-sexual maturity.! Such a function would seem '

to be impossible in this instance when we compare the diameter

of the ova with the diameter of the tubes. _

1 Ehlers, Emst. Die Borstenwiirmer (1864), p- 205.

302 THE AMERICAN NATURALIST. [Vor. XXXVI.

A pair of ventral nerve cords (Fig. 6, n.c.) are conspicuous

in transverse sections, in which the cut ends of the nerve fibers

appear, surrounded at intervals by a few darkly stained bodies,

the nuclei of ganglionic cells. The ganglia, one to each seg-

ment, can be plainly seen in a zooid cleared with glycerine.

The nerve cords, although in direct contact with the epidermis,

are plainly not fused with it.

No head or eyes were detected; but as these structures are

obviously present in the zooids of the closely allied Trypano-

syllis gemmipara, failure to find them in the present species is

almost certainly due to the broken condition of the material,

the tips of the buds being especially liable to injury.

The place of origin of the buds is near the posterior extrem-

ity of the stock. Probably, as in 7. gemmipara, it is about the

length of a zooid, or twenty somites, in front of the pygidium.

The proliferating region, although now destitute of buds, has

been sectioned, and shows merely a mass of undifferentiated

tissue pervaded by a radiating system of muscle fibers, which,

in all probability, passed into the zooids, as they certainly do in

T. gemmipara.

The rudimentary and apparently functionless condition of

the alimentary canalis noteworthy. Although impossible, from

the defective material, to tell whether a mouth and anus are

really absent, it is highly probable that they are.

Trypanosyllis gemmipara.

The other species with collateral buds was found in a collec-

tion of Polychaeta gathered in Puget Sound and vicinity by

Nathan R. Harrington, since deceased, member of the expedi-

tion sent out by Columbia University in the summer of 1896.

It has been described and figured in the writer's report! on the

Polychzta of that expedition under the name Trypanosyllis

gemmipara. |

Although much smaller than the preceding, 7. gemmipara is

nevertheless very large for a syllid. The Puget Sound speci-

men measures 68 mm. in length and 3 mm. in width. Like

1 Johnson, H. P. The Polychata of the Puget Sound Region, Proc. Boston

Soc. Nat. Hist., vol. xxix (1901), No. 18, p. 405, Figs. 72-76.

No. 424.] THE GENUS TRYPANOSYLLIS. 303

T. ingens, and in fact most, if not all, species of this genus,

T. gemmipara is much flattened horizontally, being only I mm.

in dorso-ventral diameter. The somites number about 300, and,

as in T. ingens, their width many times exceeds their length.

The tail buds, 50 in number, present all stages of develop-

ment. They form a tuft arising from the ventral aspect near

the posterior extremity, and superficially resemble a bunch of

gs mI

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Fic. 8. — Caudal buds, ventral aspect, of T. gemmifara. The young buds are seen on the left

of the figure. x 26.

ferns (Figs. 7, 8). Neither as regards position nor order of

development do they obey the laws of bilateral symmetry.

The whole cluster is bent to the right, making an angle of

about 25° with the longitudinal axis of the worm. Only 13 of

the buds have reached an advanced stage of development;

these are elliptical in shape, rather thin in proportion to their

other dimensions, and possess the chief external characters of

304 THE AMERICAN NATURALIST. [VoL] XXXVI.

the parent. Their somites are 20 to 28: in number; their

small but well-developed parapodia aré furnished" with monili-

form dorsal cirri and minute setze of the same form as those of

the parent. The average length of a mature bud is 2.5 mm.;

7197.

a at

err S xx

d98pk de

Fic. 9. — Frontal h b Tc Soemnitiri s showing cdi nohis iei

—— -e se of development The z zooids

of the yrs e,m

rd are marked à

, nearl: sagittal plane) : The youngest bul

5 indicates a bud in which Segmentation is s just Wpuniug i &, 63, buds in .

à al ready well

x developed ; 4-56, buds

———

buds progressively - older ; br, brain ; veh first dorsal cirrus Zn, `

muscles ; ny ave CPt; Ss sperm ce

the bresdih, retuned? Bertipodis is 1mm. The: bud tapers

more towards the caudal than ‘towards the cephalic extremity.

The anal cirri are neg and oe bre Bow de of Pa aiden :

cirri (Figs. 7, 8). ^

No. 424]

THE GENUS TRYPANOSYELLIS.

F1G..9.4. — Anterior extremities of zooids marked 45 in Fig. 9, showing, at two different stages,

th *

the ids than that

e Anlagen of eye and b

zooi

rain as they appear in a section further from the median plane of

the ventral longitudinal

d Shirer Lettering

asin Fig.9. x 200.

represented in Fig. 9. T e extension of

: 1 1:E L gre Pi A ages

TAN

sia AR M

dimer

ASRA

(X - e

T chua

" : Y CAN f

T^ (np

: NL 3 j e) wg

n & x i 4E) X .

Wid- SET EES E $ ard: C. d. x LN ps , bu

9 B. — Transverse s

lower level than Fig.

left are. cut obliquely ; in one

9. The MH - in transverse rows

of these (2D, segmentation of the m

Afi x è To ee Ta

306 THE AMERICAN NATURALIST. [VoL. XXXVI.

Only advanced buds are visible from the dorsal aspect

(Fig. 7). These completely cover the young buds on the ven-

tral side and overlap so that the heads of all the advanced

buds are concealed... Occupying the most. dorsal position

amongst the advanced buds is the attenuated and rapidly

diminishing caudal extremity of the stock. It is much like the

buds in general aspect, and further resembles them in contain-

ing sperm cells, which are absent in all portions of the stock in

front of the proliferating region. It differs from the buds in

possessing (1) a heavily ciliated continuation of the alimentary

canal (Fig. 7, aie);

(2) the anus, and

(3) differs in lacking

cephalization. In

every respect except

that it contains

sperm cells, it re-

sembles a regener-

ated posterior

extremity. It pos-

sesses twenty -four

somites, including

the pygidium. The

proliferating region

is therefore this

number of somites

in front of the pos- -

terior extremity. As

ANN) Fig. 7 shows, the

ges o. — Frontal section of a portion of a full-grown budof attenuated posterior

A asclcule (ec), muscles Yay en coer a Mas almost

glands (27). The intersegmental dissepiments and median ag much the appear-

strand (s.) are indicated. x 75. *

ance of being an

appendage to the budding zone as any of the mature buds.!

It is worthy of note that into this region the deep intestinal

s This attenuated posterior extremity is. probably a permanent feature. I have

found it in every entire speci Xf T. gemmipara I have collected on the Cali-

fornia coast, but the number collected is small. i

No. 424. ] THE GENUS TRYPANOSYLLIS. 307

diverticula do not continue. The diverticula of this portion of

the alimentary canal are very shallow, and were not detected in

the entire specimen (Fig. 7).

The attachment of the buds to the proliferating area (invisi-

ble in the entire specimen) is by a short unsegmented pedicle

at the cephalic extremity of each bud (Figs. 9, 9 A, 17, 7).

There is no distinct prostomium. The

head is merely the eye- and brain- Lm

bearing first segment, which shows its =

primitive character by also carrying

parapodia equipped with dorsal cirrus,

setz, and acicule (Fig. 9, d.c.'). This

fact is significant, because it is exactly

what we find in the stolons of not a

few syllidians, among others those of ‘3))

Trypanosyllis krohnis ES

The eyes, apparently only two ‘©

(although each may represent two, |

completely fused), are large and highly \ vw

developed, as is often the case in Fic. 11.— Frontal section at a lower

stolons. There is a well-defined lens tie ep rcd

(Fig. 15, 4), a “vitreous body” (v) dinal muscle (Ln, and parapodial

$ glands (g/.). x 75

composed of the rods of the retinal

cells, and a very thick, cup-shaped retina, the inner layer of

which contains a small quantity of brown pigment (7.), but not

dense enough to conceal its structure. .

The buds are destitute of an alimentary canal, unless the

merest rudiment exists in the form of a median strand (Figs.

10, 12, s.). Certainly there is no tubular structure such as

occurs in the bud of 7. ingens. There is no mouth, anus, or

blood vessels. The zooid is sharply divided by the dissepi-

ments and an incomplete median partition into a double series

of coelomic spaces. That the right-and-left coelomic chambers

of the same somite are sometimes continuous is clearly shown

by the sperm masses in the more posterior segments repre-

sented in Fig. 10. This seems to be the case only in the ven-

tral portion of the chamber. The sperm masses do not

- 1 See Malaquin, of. cit., Pl. X, Figs. 18, 19.

L--

a AN df Ifa

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p i

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308 THE AMERICAN NATURALIST (Vou. XXXVI.

completely: fill the spaces, and do not ‘seem in any’ case to

occupy the cavities’ of the parapodia. “Spermatozoa were hot

observed in any of the buds, indicating that even the oldest

are not fully ripe.

The. parapodia ‘are fans: equipped viti dorsal: and ventral

cirri, setze, a pair of aciculze, — the latter projecting deeply into

the sperm masses (Fig. ro, ac.); — and the usual musculature.

Occupying a large portion of the cavity of each parapodium

are pouch-shaped glandular: structures of unknown function.

Each is composed of a few very large vesicular cells, and lias a

good-sized lumen which appears to open near the base of the

parapodium, on the ventral side (Figs. JO, rij 35; pi). “NG

actual opening, however, has

SP dm been found, and. the mouth

inte seems entirely blocked by col-

umnar epidermal cells (Fig.

13). These glands occur in

the segments of the attenuated

caudal region, where they have

the same size and aspect as in

the buds. In the parapodia

—Á— E of the stock — at least those

fis wall marie oed, UNE NNNM of immediately in front of the

fibers and ganglionic cells. x 100 budding zone —they are also

found. Without much doubt these structures are homologous

with the parapodial glands (glandes pédieuses) known to be

present in Zrypanosyllis celiaca and in species of other genera

of Syllidae! Their function is the secretion of mucus. From

their histological aspect, this is genie likewise the functión

of the above-described glands.?

‘There are two large and conspicuous ventral nerve cords,

with paired ganglia in every somite (Figs. 11, 12, Lc. eg).

Proportionately to the size of the bud, the nervous system is

much larger in this species than in 7: ingens. As accurately

as — be made out from the single series of mo" iN

; See Malaquin, p isal M sur ja Syllidiens, P. sr PL vir, Fig. "a ERR

: Da is interesting to find that these ese glands are not "v relatively, but Tat

lately, 1 larger in the egm nts and in the bud: tock.

No» 424.] -- THE GENUS ITRYPANOSYLLIS. 309

nerve cords run forward to the cephalized extremity and there

innervate the eyes and a darkly staining mass of ectodermic

cells which probably functions as brain (Fig. 14; cb)... Although

the nerve cords are plainly separate from the epidermis, as

shown by a cross-section (Fig. 12, #.¢.), the. brain is no more

than a thickened portion of the epidermis, and fully confluent

with it. As clearly shown in one section (Fig. 14), the optic

nerve comes from the ventral cord, and not from the brain.

“8S.

Fic. 13. — Two pedal (mucous?) glands of bud of T. gemmifara, in longitudinal section. The

lumens of the glands are bounded by stippled bands; the gland cells (g/.c.) are highly

vacuolated; the epidermal cells at mouth of glands are columnar. #.#2., transversal mus-

cles; £.s., intersegmental dissepiments ; e/., epidermis. X 525.

When viewed from the ventral aspect the cluster of buds

presents an interesting condition of things (Fig. 8). On the

right, at the base of the cluster, is a group containing about

25 very young buds (Fig. 8), as yet showing no segmen-

tation, but each with two distal processes which are the

Anlagen of the anal cirri. As may clearly be seen in a trans-

verse section of the group (Fig. 9 B), the buds are arranged

in rows, partly transverse and partly longitudinal as regards

310 THE AMERICAN NATURALIST. [Vor. XXXVI.

the long axis of the stock. In a graduated series proceeding

from the left side of the cluster of youngest buds are more

advanced buds, the largest approaching the mature condition

(Fig. 9, 6—2°). At an early stage the anal cirri attain a nota-

ble length and become articulated.

As shown in a frontal section (Fig. 9) Dauh the pro-

liferating region, the buds sprout from a mass of undifferen-

tiated tissue traversed by muscle fibers which are continuous

9$ 92,9 *

ayers ® s s

Fic. E Fides section of a portion of "m same bud as shown in Fig. ro. z., nerve to

cb.); vd 7., Optic nerve; 7., na, eure rod Uh and pigment (7.) ; v7., “‘ vit-

reous i tir " insepiddl of the clear peri. The nerve to the brain breaks up into

numerous Shere, which are distributed to the a ura Q)cell. x 153.

with the longitudinal muscle bands of the buds (Fig. 9 A, /.m.).

This mass is mesoderm, covered by ectoderm in the form of

epidermis. Both ectoderm and mesoderm of the buds are con-

wea tinuous with the same germ

à layers of the proliferating

...vt. region. Whether the prolifer-

ating mass comprises one or

more than one somite cannot

be positively stated from the

sections at hand, which are

ies ee zuo defective at this point.

an invagination of the epidermis (&A). Letter- — Ata very early stage the meso-

ing as in figure; Z,lens. x 267 i ‘

derm undergoes segmentation,

and the reproductive cells are discernible (Figs. 9, 9 B, 0’).

No buds provided with special natatory setze (Pubertdtsbor-

sten) such as usually occur in the stolons of syllids and in

the sexual buds of Sy//is ramosa! were observed.

1 M'Intosh. Challenger Reports, vol. xii, p. 201, Pl. XXXIV A, Fig. 9.

No. 424.] THE GENUS TRYPANOSYLLIS. 311

GENERAL CONSIDERATIONS.

From the standpoint of comparative morphology, the col-

lateral buds of Trypanosyllis may be regarded as structures

of the same order as the stolons of Autolytus or Myrianida,

but with the important difference that the buds in this case

are lateral outgrowths, whereas in stolonization they are the

result of linear growth and differentiation. As stolons, they

extend the organic axis of the parent, and the alimentary canal

passes through them all, to terminate at the anus located in

the pygidium of the oldest stolon.

At first thought it would seem as though the buds of

Trypanosyllis are more appropriately comparable to the sexual

zooids of Syllis ramosa. The superficial resemblance to the

latter is indeed striking. In both instances we have a lateral

outgrowth of the stock, to which it is attached at the head end

by a pedicle; in both, the zooids are the bearers of the sexual

products (which do not develop in the stock), and both have an

imperfectly developed prostomium bearing highly developed

eyes. There are, however, two important points of difference:

(1) the pedicle of the sexual zooid of Sy//is ramosa is segmented,

and the segments even bear rudimentary cirri!; (2) the sexual

zooids as well as the asexual branches of SS. ramosa contain a

branch of the alimentary canal, fully continuous with that of

the stock. In neither of these respects does the sexual bud

of T. gemmipara in the least resemble the sexual bud of

S. ramosa. The sexual zooids of the latter species are evi-

dently specialized, fructifying branches of a widely branching

asexual stock. They are equipped with large eyes and special

swimming organs (natatory setze), and break away from the

parent stock at maturity. In all these respects they differ,

it is true, from the purely vegetative branches, but these

differences are merely adaptive to the special functions of

insuring fertilization of the eggs and disseminating the species.

aether the somite or somites from which the sexual buds

of Trypanosyllis arise is to be regarded as homologous with

the others cannot be decided from the material at hand. No

1M’Intosh. Challenger Reports, vol. xii (1887), Pl. XXXIII, Fig. 11.

312 THE AMERICAN NATURALIST. . (VOL XXXVI.

somite unprovided with parapodia of the usual form was dis-

covered. A recent study of S. ramosa by Oka! has brought

out the interesting fact that the paired buds arise, not from

one of the original somites, but from a newly developed, inter-

calated somite, which differs from all the others in lacking para-

podia. Oka is doubtless right in regarding it as non-homologous

with the other somites,

While the stolons of Autolytus and of syllidians generally

are distinct and complete individuals provided not only with

ears ecto EM. «mus Ly

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d thins towards è blind extremities of "e diverticula.

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sexual, sensory, and locomotor organs, but with those subserv-

ing nutrition, circulation, and excretion, the sexual buds of

T. ingens, and even more ‘strikingly 7. gemmipara, exhibit a

higher degree of specialization in lacking functionally developed

vegetative organs. The zooid is therefore quite as incapable

of leading a prolonged independent existence as the famed

Palolo of the South Seas. It is no more than a living engine

for the dissemination of the genital products which it carries,

t Oka, Asajiro. Ueber die pierce bei ees ramosa, Zoilogical

Magazine, Tokyo, vol. vii (1895), p. 117.

No: 424.]: THE GENUS TRYPANOSELLLS, 313

and.that duty- must be accomplished solely by the expenditure

of.the stored-up energy which it has derived from the stock.

From the biological point of view the function of the zooids

is the same, whether they arise by linear or by collateral

budding. . In both cases . i

there is an alternation of P- ou Viste.

generations exactly com- e RE Cis

parable to that of the Nee

Hydromedusz. As regards AEN

SPee

S. ramosa the asexual SP eon mas

stock is practically.as ses- ioe | Mee

sile as the sponge in which Lj || mm. 5.

ibo: : cb qv e

it lives, and like. sessile cp 2 1

organisms of much lower. m | a c

grade it branches with e E =

great freedom and facility. -e e

It. is. not in the least sur- xo i er "y

ws co i €...

prising, therefore, to.find =p | 6 sol.

such.a form. producing by LD it SS ;

lateral budding free-swim- ce th ew

ming zooids, which. serve i d

as special disseminators of e iE =>

the.species and provide for BOIS

& sexual reproduction in. fre

e *

which epossotertilisapion 1 is m Q

assured. . 5t & H

- In the forms under con- 8 J

sideration the.case is SOm- pya; 17, — Ideal reconstruction of á mature male zooid

what different. Notwithe, | Eig, lieu din n aero vi

standing their large 5 size -`y The. sperm masses-are in every segment except the

sand: heavy. form, o differ- . . somites 2 arid $.' The aciculz are omitted except in

ent from the majority of - fie epe eh i arr agi a MM

syllidians, suggest that

they are somewhat sluggish in their habits, neither of them can

be considered as sessile. Iti is difficult. to see, however, that they

are any more inert than a multítude of annelids which have n

‘nothing’ aberrant in their mode of reproduction. ` Pumy

some more precise. meen, must be ‘sought. . ,

314 THE AMERICAN NATURALIST. [Vor. XXXVI.

The fact that Trypanosyllis is a genus of the Syllida must

not be overlooked. Of all the annelids, this family affords

the best and most numerous examples of asexual reproduction

by stolonization. The small size and comparatively simple

structure of most syllidians no doubt peculiarly adapt them to

this mode of multiplication. On account of its great advantage

in the dissemination of the species, this form of reproduction,

acquired by the presumably diminutive ancestors of the two

gigantic species of Trypanosyllis, might be retained in a grad-

ually modified form as by slow degrees the animal became larger

and more complex. By equally slow degrees the collateral type

of budding would replace the linear. The development in

both collaterally budding species of extensive intestinal diver-

ticula (Fig. 16) in every somite from the proventriculus to the

budding zone would obviously render inconvenient, if not

impossible, the production of large gonads in the stock. As

in many other syllidians the production of sexual cells would

be more and more relegated to the most posterior somites,

and to any linear or collateral buds that might develop in that

region.

The advantages of collateral over linear budding in greater

safety and compactness, as well as the possibility of securing

a more rapid development and a greater number of buds,

are obvious. Species of Myrianida produce stolons with the

greatest number of zooids, but the chain is not known to

have over thirty zooids at any one time? This is only 60 per

cent of the number of collateral buds present in the specimen

of Trypanosyllis gemmipara that has been studied. The 50

buds of the latter add practically nothing to its length, while

the 29 buds of the example of Myrianida figured by Malaquin

form a chain no less than 55 per cent of the entire length of

the worm. The liability to loss or injury to which such a

1 In 7. zebra, according to an observation recorded by Viguier in 1886 (Arch.

de Zool. Exp., sèr. 2, tome iv, p- 364, footnote), to which the author has kindly

drawn my attention, there occurs an apparently non-terminal stolonization which

perhaps is a t step towards collateral budding. It is not well, however, to push

the very far, as there is really little likeness between the two types of

gemimation as we know them at poe

2 See Malaquin, oc. cit., Pl. I. |

No. 424.] THE GENUS TRYPANOSVYLLIS. 315

lengthy chain would be exposed, particularly if very slender

and attached to a stock of vastly greater bulk and strength,

is evident.

The absence of organs of nutrition, the collateral mode of

origin, and the fact that the smaller and simpler members of

the genus still produce stolons point unmistakably to the high

degree of specialization attained by the sexual zooids of these

two species of Tryp yllis. Both in their structure and in

their mode of development they unquestionably represent the

most specialized mode of asexual reproduction, not only among

the Syllidae, but among all known annelids.

pit NE S^

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THE COURSE OF THE BLOOD FLOW IN

LUMBRICUS.

J.B. JOHNSTON AND SARAH W. JOHNSON.

ALTHOUGH this form is almost universally used as a type for

laboratory study, its circulation is still very imperfectly under-

stood. The present contribution aims to give a better under-

standing of the course taken by the blood, as determined by

observation and experiment upon large specimens of Lumbricus

terrestris. The following brief review of the anatomy of the

circulatory system, which is well understood, will serve as a

basis for the discussion of the blood flow.

A large dorsal vessel extends along the dorsal surface of the

intestine from the caudal end of the worm to the pharynx, over

which it ramifies. Strong pulsations of this vessel run in

more or less regular succession from the posterior to the

anterior end. In somites VII to XI, inclusive, five pairs of

circular vessels, the strongly pulsating %earts, encircle the

esophagus, connecting the dorsal vessel with the ventral.

The ventral is a longitudinal trunk lying beneath the intestine

and giving branches to the body wall and the nephridia. A

subneural and two lateral neural vessels constitute, by reason of

an irregular. network of anastomoses between them, a connected

system accompanying the nerve cord. This system is connected

through branches of the lateral neurals with, the body wall.

Connecting the dorsal and subneural vessels are the so-called

parietals. They occur, one pair in each segment beginning

with XII, close behind the septa and give off branches to the

body wall and the nephridia. The parietal in somite XII is

much larger than the others. In each somite two or more

unpaired ventro-intestinal vessels enter the wall of the intestine

from the ventral vessel, while two pairs of dorso-intestinals pass

from the dorsal vessel into the wall of the intestine.

os oe 1 From the Zoólogical propion of West Virginia University.

318 THE AMERICAN NATURALIST. [VoL. XXXVI.

In somite X a pair of /ateral vessels come from the dorsal

vessel and run forward on either side of the cesophagus to the

pharynx, giving branches to the cesophagus and body wall.

An anastomosing vessel arises from the lateral near its origin

in somite X and passes backward to join the large parietal in

somite XII. From this anastomosis, branches supply the

seminal vesicles, which are also supplied from the ventral ves-

sel. Connecting vessels between the laterals and subneurals

occur in several somites. |

The only points with regard to the course of the blood flow

upon which authors agree are that the blood flows forward in

the dorsal vessel to the hearts, downward in the hearts, and

backward in the ventral vessel. With regard to the further

course of the blood Perrier came to the following conclusions.

(as reported by Bourne and Harrington): the blood flows into

the dorsal vessel from the parietals and from it through the

dorso-intestinals ; the ventro-intestinals carry blood from the

intestine to the ventral vessel; from the ventral it is given to

the body wall and nephridia and is gathered up from these by

branches from the lateral neurals and parietals ; the flow in the

laterals is forward. :

Bourne (91) studied the blood flow in Megascolex principally

by holding the vessels with forceps and by cutting them to see

from which cut end the blood flowed. His conclusions are

nearly opposite to those of Perrier, namely, that the dorso-

intestinals empty into the dorsal and the parietals carry blood

away from it; the ventral gives blood to the intestine through

the ventro-intestinals; it also gives blood to the body wall and

nephridia, from which it is carried to the intestine (in Megas-

colex) by the intestino-tegumentaries. The lateral vessels

carry blood backward. ——— | :

Benham (91), working on Lumbricus, agrees with Perrier in

regard to the blood flow in all vessels.

Harrington (99) has given a full account of the anatomy of

the vascular system, with elaborate figures. His descriptions

and figures, however, do not always agree, e.g., the common

vessel branching into ventro-tegumentary and ventro-intestinal

described on page 142 is not shown ín the figures. Harrington

No. 424.] THE BLOOD FLOW IN LUMBRICUS. 319

DIAGRAMS TO ILLUSTRATE THE SCHEME OF CIRCULATION

GIVEN IN THIS PAPER.

A, cross section of body region ; &, cross section through hearts; C, sketch of

vessels in somites VII to XIV seen from the side. Arrows indicate the direction

and vessels supplying it; œ., oesophagus and vessels supplying it; æg., vessels

supplying cesophageal glands; 7., parietals; 5.7., subneural; s.v., seminal vesicles

and vessels supplying them; v., ventral vessel; v.i., ventro-intestinals; v.Z., ventro-

tegumentaries. The figures are unfortunately crowded, owing to the projection

on one plane of many vessels which lie at differentlevels. The lateral neurals are

not lettered in A and Z and are not shown in C.

320 THE AMERICAN’ NATURALIST.’ [VoL XXXVI.

was the first to describe the arrangement of vessels to the

integument, showing that the capillaries of the parietals

(** commissurals ") are directly continuous with those of the

véntro-tegumentaries, and that loops of these-capillaries lying

just beneath the. cuticle have the function of respiration. He

has shown also that in somites VIII to XIII inclusive the sub-

neural has a branch which supplies the nephridium and setiger-

ous glands and anastomoses with the lateral. The lateral has

connections with the dorsal in X and XII, as above described.

Anastomosing vessels continue backward through XIII and

XIV, making connections with the subneurals at each septum.

These points have been verified by the present writers, but

Harrington's statement that the true parietals (**commis-

surals") are absent in XIV and XV they cannot confirm.

True parietals are always found in all somites back of XII, and

occasional worms clearly show anastomosing vessels continuing

backward from the laterals, connecting with the parietals of

one or two somites.. The anastomoses between the laterals

and subneural have not always been found by the present

writers in all.the somites mentioned.

Harrington studied the course of blood flow iy watching

the pulsations of the vessels in small specimens of -Lumbricus

under a dissecting lens. He concludes that the blood flows

forward in the dorsal, downward in the hearts, backward in the

ventral, into the dorsal from the dorso-intestinals and out from

it into the parietals (* commissurals "), forward in the laterals,

and either forward or backward in the subneural.

The senior author has made observations on the course of

the blood flow in connection with class work for a number of

years. During this time the direction of flow in the subneural

has been demonstrated in many student dissections, the result

constantly agreeing with that stated below. The observations

reported here have been made on the largest worms obtain-

able, anesthetized and opened under physiological salt solution.

The course of the flow in each vessel has been determined by

the three methods of clamping, cutting the vessel, and watch-

ing the pulsations where these are visible. Injecting the

dorsal vessel was tried several times, but as yet no facts

No; 424.].. THE BLOOD FLOW: IN LUMBRICUS. 321

regarding the coutse-of the flow have been. made out by this

means, although the injection gives.a good demonstration of

the-anatomy of the Hpasiiatu fuo qus

DESCRIPTION oF x2 whe?

Many experiments. give no definite results,- owing to thd

condition of the worm or to violent disturbance of the circula-

tion. Great care is required in recognizing and interpreting

the results, especially in observing pulsations. It has been

found that when fairly clear and definite results are obtained,

the same result is given by the different methods.

I. The Dorsal Vessel. —'The fact that the blood flows for-

ward in the dorsal vessel up to the hearts is clearly shown by

the course of the pulsations, In front of the hearts the pulsa-

tions are not conspicuous, and special attention was given to

determining the course of flow. in this part of the dorsal.

Among the many worms dissected afew- were found in which

the pulsations of the dorsal were distinctly seen running for-

ward to the pharynx. The dorsal vessel. was clamped in front

of the hearts in.a number of worms, and in all cases the blood

collected behind the clamp and disappeared in front of it. The

blood flows forward throughout the whole length of the dorsal.

2. The Ventral Vessel. — The experiments noted below

and the general agreement of authors make it. certain that

the blood- flows backward in the ventral vessel back of the

hearts. The course of the flow. in front of the hearts was

shown by clamping the ventral vessel in the seventh somite.

In every case the blood collected behind the clamp and disap-

peared in front of it, indicating a forward flow.

3. The Subneural and Lateral Neurals. — The best results

are obtained. on these vessels by opening the worms from the

ventral side. When the subneural.vessel was cut the blood

flowed only- from the anterior cut end, Ze, backward. The

same result was obtained constantly. by pinching the subneural

with forceps: the blod collated: aen distinctly in front of the

forceps and di ^ These ——_ were

tried in all regions. of the worm. 1s

322 THE AMERICAN NATURALIST. [VoL. XXXVI.

4. The Parietals. — Clamping these vessels has given doubt-

ful results. The first experiments were made by pinching

them at about the middle of their length in a number of

worms. The blood seemed to collect between the dorsal ves-

sel and the forceps. More recently no results whatever have

been obtained from the parietal vessels by this method. This

is perhaps due to the fact that branches from the body wall

enter the parietals both above and below the forceps. In each

of eight worms a dozen or more parietal vessels were cut, and

in every case in which bleeding occurred the blood flowed from

the lower end, that is, toward the dorsal vessel.

Owing to the numerous connections of the parietal vessels,

it is necessary to use indirect methods to determine the course

of blood flow in them. In several sets of experiments the

attempt was made to determine the source of supply to the

parietals and their place of emptying, by clamping or cutting

other vessels. All three longitudinal trunks, dorsal, ventral,

and neural, were simultaneously clamped, with the following

results in each case. The dorsal vessel was filled on both sides

of the clamp. Although the pulsations were sometimes seri-

ously interfered with, its normal fullness was maintained.

This is to be explained, as will appear from what follows, by

the fact that in front of the clamps a complete circulation is

kept up and the fullness of the dorsal maintained by the dorso-

intestinals and parietals bot% emptying into it. On the theory

that the parietals and dorso-intestinals carry blood in opposite

directions, it must be expected that the dorsal would be empty

in front of the clamp in this experiment. If only one set of

circular vessels empty into the dorsal, its own’ pulsations

together with the other circular vessels must certainly drain

it of blood in front of a clamp. Behind the clamp it is to be

expected that the dorsal should be full; but the normal fullness

is not greatly exceeded, because the clamping of the ventral

and neurals cuts off the supply of blood to the posterior end

of the worm. It is also possible that the course of flow is

reversed in the parietals or dorso-intestinals or both, just

behind a clamp on‘the dorsal, because of the excessive pres-

sure developed by the pulsations of the latter vessel. In this

No. 424.] THE BLOOD FLOW IN LUMBRICUS. 323

experiment the ventral and neural vessels were filled in front

of, and empty behind, the clamps, indicating a backward flow

in both. The parietals were filled both in front and behind

the clamp. They were somewhat distended in front of the

clamps, owing to increased pressure from the subneural in

front of its clamp, while behind the clamps the parietals

continued to collect some blood from the body wall and

nephridia.

In four worms the ventro-intestinals in several segments

were cut in order to drain the intestine of blood in those seg-

ments, and the dorsal was afterward clamped at both ends of

the area in which the ventro-intestinals were cut. Between

the clamps the dorsal decreased and the parietals were nor-

mally full. In four other worms the same result was reached

when the dorsal was clamped in one place and the dorso-intes-

tinals were cut for several segments in front of the clamp.

Since the dorsal retained some blood and the parietals kept up

their normal fullness as long as the worm lived, while all other

supply to the dorsal was cut off, it is impossible to reach any

other conclusion from these experiments than that the parietals

. bring blood to the dorsal.

Pulsations of the parietal vessels are not always clearly visi-

ble. In four worms in which the pulsations of the parietals

were distinct and strong a large number of these vessels were

carefully studied. The contraction of the parietals occurs

opposite the front part of the pulse wave of the dorsal, z.e.,

while the dorsal is reaching its greatest fullness. The parie-

tals fill just after the contraction of the dorsal, z.e., opposite

the point of constriction of the dorsal behind the pulse wave.

The filling of the parietals is sudden and the contraction slow.

This, taken together with the order of the contractions, indi-

cates that the contraction of the parietals helps to swell the

dorsal just in front of the constriction which carries its pulse

wave along, while the rapid filling of the parietals is due to

the sudden closure of the parieto-dorsal openings by the con-

traction wave of the dorsal. The results of all the experi-

ments considered together strongly favor the conclusion that

the parietals carry blood to the dorsal.

324 THE AMERICAN NATURALIST. [Vor. XXXVI.

5. The Dorso-Intestinals. — When one clamp was put on the

dorsal vessel and anumber of parietals were cut just in front of

it, the dorsal vessel only partly emptied; but when the dorso-

intestinals in the same segments were cut the dorsal vessel

emptied entirely. This and the experiments on the parietals

described above, in which the cutting of the ventro-intestinals

in one case and of the dorso-intestinals in another resulted in

the decrease of the dorsal vessel in those segments, show that

the dorso-intestinals carry blood to the dorsal.

6. Lhe Lateral Vessels.— When the laterals are clamped

either at both their connections with the dorsal in somites X

and XII or in front of the connection in X, they remain full

of blood, sometimes much distended, in front of the clamps.

No vessels have given more positive and unmistakable results

than these.

CRITICISM AND CONCLUSIONS.

There is no difference of opinion in regard to the blood flow

in any part of the dorsal vessel. All authors agree that it

flows forward to the extreme anterior end of the worm.

Bourne alone suggests that the blood flows forward from the

hearts in the ventral vessel, since he considers that the strong

contractions of the hearts would force blood forward as well 2s

backward. Our experiments have confirmed this plausible

view, and it may be regarded as proved that the blood flows

forward in this part of the ventral.

Although previous workers have not agreed upon the func-

tions of the parietals and dorso-intestinals, they all agree in

assigning to these vessels opposite functions. The above

experiments on the parietals, although not wholly decisive,

give strong support to a different conclusion, namely, that

both parietals and dorso-intestinals carry blood to the dorsal.

It follows necessarily that the ventro-intestinals supply the

intestine with blood, as Bourne and Harrington contend,

According to Harrington’s interpretation all of the blood in

circulation would have to pass through the ventro-intestinals ;

in other words, all of the blood which is carried forward by the

dorsal and all that which passes downward from the dorsal in

No. 424.] THE BLOOD FLOW IN LUMBRICUS. 325

each somite through the parietals must be carried to the dorsal

by the ventro- and dorso-intestinals. It does not seem possible

for all of the blood to take this course, owing to the small size

of the intestinal vessels. It has been impossible for the pres-

ent writers to arrive at a definite conception of the blood flow

according to Harrington’s scheme. He represents the blood

as flowing in different directions in different parts of the same

vessel, and as changing its direction in the same vessel from

time to time. Our experiments give no confirmation whatever

to either of these conclusions, except that the direction is per-

manently reversed in the front end of the ventral vessel and in

the dorso-intestinals in front of the hearts as above described.

Temporary reversal of the flow in limited regions might easily

occur as a result of the animal's movements, and such phe-

nomena may have led Harrington to erroneous conclusions,

since he worked only by studying the pulsations through the

skin with a hand lens. His scheme of the circulation is full of

inconsistencies, due either to an attempt to avoid overloading

small vessels and consequent breakdown of the scheme, or to

the incomplete study of several genera, resulting in a patch-

work which is not true for Lumbricus at least. It is but

proper to say that these inconsistencies might have been

removed had the author lived to see his work through the

press.

Bourne states that the lateral vessels carry blood backward,

Harrington that they carry it forward. Our results positively

support Bourne's view and, together with the results on the

parietals, suggest a new interpretation of the morphology of

the lateral vessels. Bourne considers them to be the sub-

neural ends of the parietals in this region, dwt assigns to them

an opposite function. The following facts seem to show that

the lateral vessels are formed by anastomoses between succes-

sive parietals, which carry the blood from the head region to

the dorsal in somites X and XII, and that the lateral system

therefore represents the parietals in the head.

I. Absence of true parietals in front of somite XII.

2. Connections of the laterals with the subneural in several

somites (VIII-XII) which correspond perfectly to the ventral

326 THE AMERICAN NATURALIST. [Vor. XXXVI.

ends of parietals, Having branches to the nephridia as the

parietals do.

3. Connections of the laterals with the dorsal in somites X

and XII, corresponding to the dorsal ends of parietals.

4. Anastomoses between the parietals of somites XII, XIII,

and XIV, in apparent continuation of the laterals.

5. Presence of branches of laterals to the body wall along

the septa, corresponding to respiratory branches of the

parietals.

6. Presence of branches to the seminal vesicles, which are

also supplied by branches from the ventral. Here, as else-

where, branches of the parietals take up blood which is brought

from the ventral. The parietals are in the best position to

supply the vesicles which are developed from the septa.

The origin of the hearts and laterals and the position of the

laterals within the circular hearts are problems which require

more facts for their solution. An extended discussion of them

is beyond the province of this paper, but one or two sugges-

tions will be made. The suggestion of Harrington that the

hearts are modified parietals is negatived by the presence of

both parietals and hearts in somite X, and by the different

ventral connections of the parietals and hearts. It seems

more probable that the hearts are enlarged circular intestinal

vessels, serving to carry the blood brought forward by the dorsal.

The lateral vessels must have arisen as anastomoses between

successive parietals serving to return the blood from the head

(brought forward by both dorsal and ventral), to the dorsal

behind the hearts. The dorsal connections of the parietals in

front of the hearts have disappeared from disuse. The posi-

tion of these anastomoses within the circular hearts seems to

indicate that they, too, have been derived from the capillaries

of the cesophageal wall, although later than the hearts. This

suggestion would explain the branches of the lateral vessels to

the cesophagus and hearts, which otherwise are unexplained.

Conclusions. — The course of the circulation in each seg-

ment behind the hearts is as follows: the branches of the

lateral neurals and parietals collect blood from the body wall;

the parietals take- blood to the dorsal from the subneural, body.

No. 424. ] THE BLOOD FLOW IN LUMBRICUS. 327

wall, and nephridia; the dorso-intestinals give blood to the

dorsal from the intestine ; all of this blood goes forward in the

dorsal to the hearts; through them nearly all of it is given to

the ventral vessel, which sends it to the intestine through the

ventro-intestinals and to the body wall through the ventro-

tegumentaries. | e |

The head region is supplied with blood by the dorsal and

ventral vessels, and the neurals and laterals carry blood

from it.

The dorsal and ventral vessels carry mixed blood, the dorsal

receiving oxygenated blood from the parietals and non-oxygen-

ated from the dorso-intestinals. The neurals carry oxygenated

blood only. The dorsal and ventral vessels carry mixed blood

to the head region; the laterals carry backward mixed blood,

and the neurals oxygenated blood.

There is no support given by our experiments to the idea of

a more or less complete segmental circulation in Lumbricus. `

The circulation is strictly systematic, the blood making a com-

plete circuit from the hearts backward through a greater or

less number of somites, upward through the body wall, nephri-

dia, or intestinal wall, to the dorsal, and thence forward to

the hearts. This scheme of circulation is simple, and the

slight modifications of it, which occur in the head region only,

have received at least an attempt at explanation above. The

interesting and highly important question presents itself :

What happens when a worm's head is cut off and it lives

and regenerates a new head, or perchance a tail? "With the

removal of the hearts some substitute must be provided to

carry the blood from the dorsal to the ventral vessel. A series

of regeneration experiments have been begun to find the

answer to this question.

October 23, 1901.

328 THE AMERICAN NATURALIST.

LIST OF PAPERS CITED.

PERRIER (reported by Bourne and Harrington).

'91. BENHAM, W. B. The Nephridium of Lumbricus and its Blood Sup-

ply. Quart. Journ. Micr. Sci. Vol. xxxii, p. 293.

'91. BOURNE, A. C. On Megascolex cceruleus and a Theory of the Course

of the Blood in Earthworms. Quart. Journ. Micr. Sci. Vol. xxxii,

P. 49.

'99. HARRINGTON, N. R. The Calciferous Glands of the Earthworm,

with Appendix on the Circulation. Journ. of Morph. Vol. xv,

Supplement, p. 105.

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Organic Regulations. — Hans Driesch publishes, under the title

of Die organischen Regulationen, a pamphlet of 228 pages which

deals with a number of organic phenomena, some of which are better

known as adaptations, others as restitution of form and of function.

The subordinate title of the paper, ** Vorbereitungen zu einer Theorie

des Lebens,” indicates the ambitious aim of the author.

The contents are arranged under three headings: I, Descriptive

Part ; II, Theoretical Part ; III, Critique of Knowledge.

I. In the first part we find a description given of metabolic regula-

tions (adaptations), of morphological adjustments to the environment,

and of restitution or restorative regulative processes. An attempt is

made to sift out the essential features of these phenomena.

IL In the theoretical part the different kinds of regulations are

classified, the relation between regulations and stimuli is discussed,

and an analysis of form regulation is attempted. Under this same

heading a section is entitled * Thoughts on Respiration and Assimi-

lation." Here an hypothesis of the “real” meaning of the process

of oxidation in organisms is suggested, and a curious inquiry is made

into the fundamental but almost unknown field of assimilation.

Then follow two “proofs” of vitalism, or, as the author prefers to

call the latter, “ The Autonomy of the Life-Processes.”

The first proof of the autonomy of the life process is an extension

of the same argument previously employed by Driesch, and deals

with “The Differentiation of Harmonic Equipotential-Systems,” or,

more simply, with the formation of a new whole organism from a piece

of an egg or from a piece of an original organism, — the piece having

in both cases the potencies of the original whole in all its parts.

The second proof of vitalism is found in “The Genesis: and . Exist-

ence of Equipotential Systems with Complex Explicit. Potencies.”

The author believes “it is impossible that a complicated structure

(Tektonic) made up of many definitely arranged parts ( Specifitaten)

1 Driesch, Hans. Die organischen Regulationen : Vorbereitungen zu einer

Theorie des Lebens. Leipzig, W. Engelmann, 1901. 228 pp. Jone

329

330 THE AMERICAN NATURALIST. [Vor. XXXVI.

differentiated in three dimensions could divide itself by material

means into the elements of an equipotential system.” “If each

organism produced but one egg, then the materialistic conception of

heredity might be true, in which case there need not be of course a

regulative restitution; but since the organism produces many eggs,

the conception is impossible ; because, (1), in order that the develop-

mental phenomena may be deduced materialistically they must be

thought of under the assumption of a complicated primary machine,

but, (II), on account of the required division of the complex potencies

into the elements this assumption cannot be made."

Perhaps these statements may be clearer in the following quotation,

in which both the first and the second proofs of the autonomy of the

life process are considered together. “The preceding phenomena

can be interpreted only on the basis of machines of a complicated

sort — but such machines can, for other reasons, not exist. They

cannot exist, in the first place, because a machine does not remain

the same when we take away any of its parts, or when we rearrange

its parts; for machines do not possess the property of division." It

can scarcely be claimed by the author that the discovery that organ-

isms are not machines is new, but having admitted this, it does not

seem to follow that, therefore, there must be a vitalistic principle in

nature. There seems to us to be a great hiatus in Driesch's argu-

ment, and in any case his conclusions appear to be based rather on

our ignorance of what the organism really is than on a demonstration

that it is something that cannot be conceived under a causal-mechani-

cal point of view. ji

III. In the final section of the paper a number of philosophical

and metaphysical topics are discussed, and the author defines his

position in regard to the use and meaning of the terms “ mechanism ”

and “ causality.” bi ae i

ZOOLOGY.

Fauna of Switzerland. — The origin of the present fauna of

Switzerland from that of the glacial period has been the subject of

repeated investigations, but these studies were mainly restricted to

the higher forms of terrestrial life. A recent publication by Prof.

F. Zschokke' not only gives a condensed review of our knowledge

- lZschokke, F. Die Tierwelt der Schweiz im ihren Beziehungen zur Eiszeit.

Basel, 19or.

No. 424.] NOTES AND LITERATURE, 331

of the origin of the Helvetian fauna in general, but preéminently

applies the views gained by previous studies to the lower forms of

animals that inhabit the fresh-water lakes, streams, etc., of the country.

Professor Zschokke points out that the present terrestrial fauna

and the present flora belonging to the high mountain regions of the

Alps find a counterpart in the arctic regions, the identical species

being in many cases represented there. These two main ranges are

in some degree connected by intermediate, but isolated and discon-

tinuous, stations on the highest elevations of the intervening low

countries, or in localities which, owing to local conditions, possess a

particularly cool. climate (for instance, peat bogs). It is generally

agreed that this peculiar distribution is to be explained as a remnant

of the glacial period. During the latter the originally alpine and

arctic fauna and flora was widely distributed over the low countries

of central Europe, where it found congenial climatic conditions.

After the close of the glacial period, at the commencement of a

warmer climate, this glacial fauna and flora retreated in two direc-

tions : toward the north pole, and toward the high mountains to the

south; while in the intervening parts it remained only in a few scat-

tered colder localities, such as the highest summits of the mountains

of central Germany. The parts evacuated by this arctic element

were occupied by a new stock of life that immigrated chiefly from

the east (Siberia).

This is true for plants and terrestrial animals, such as mammals,

land mollusks, insects, reptiles; and in many cases we have direct

means of ascertaining and verifying this process, since we know

glacial and interglacial remains of the respective forms that have

been found in the low parts of central Europe.

While it thus seems positively assured, by paleontological facts,

that the origin of the alpine flora and terrestrial fauna of Switzerland

is to be traced back to glacial times, and lies in the vast country bor-

dering the ice sheet that covered north Europe to the south, we have

no such evidence as regards the fresh-water fauna. Nevertheless,

Professor Zschokke demonstrates that the distribution. of many fresh-

water animals of the alpine lakes and streams of Switzerland exhibits

the identical peculiar features that are known in terrestrial animals ;

they reappear in arctic fresh-water basins offering the same climatic

conditions, and are wanting in the intermediate tracts except for a

few scattered and exceptionally favorable localities. From this sim-

ilarity in distribution we are to infer that the origin of the distribu-

tion of these fresh-water forms was subject to the same laws.

332 THE AMERICAN NATURALIST. [VoL. XXXVI.

But there is an important difference of the fresh-water fauna from

the terrestrial. In the latter, the different faunas, the old glacial

and the newer postglacial, are, in the mountains, so to speak, on top

of one another, the postglacial fauna occupying the lower parts and

ascending the mountains only to a certain elevation, where it gives

room to the older fauna, which has taken to the summits. In the

fresh-water fauna, however, there is a much larger degree of inter-

mingling ; indeed, the postglacial immigrants have been able to reach

the most exposed and highest stations in the mountains, and are

found everywhere in close vicinity and association with the older

glacial elements. According to Zschokke, this fact is due, on the

one hand, to the character of the immigrants, which are cosmopolitan,

eurytherm forms of life, which may find congenial conditions of exist-

ence anywhere ; on the other hand, it is due to the peculiarities of

temperature in the fresh-water bodies, where the extremes are not

so much contrasted and lie closer together than those of the air

influencing the terrestrial animals.

The old glacial fauna of the glacial time did not only retreat in

these two directions mentioned, but there is a third refugium for

those that belong to the fresh water. In the depths of the great

lakes that fringe the northern foot of the Alps, similar conditions of

temperature prevail as in glacial times everywhere in fresh water.

These cold depths contain a fauna consisting of peculiar fishes: (Sal-

monidz) and crustaceans (the latter chiefly pelagic) that point dis-

tinctly to a northern origin. These forms apparently immigrated at

the close of the glacial period, when there was an abundance of

cold water derived from the receding ice. Later, after the abundant

supply of water had given out, these forms were cut off from the

communication with their northern homes, and their only chance to

survive was in these large and cold fresh-water lakes, which resemble

their original home at least in temperature.

Thus we may distinguish, among the fresh-water fauna of Switzer-

land as well as among the terrestrial fauna, two elements: (1) an

old glacial stock that lived during the’ glacial: period all over the

inhabitable parts of central Europe; and (2) a later immigration in

postglacial times.. The first one retreated out of the plains in two

directions : toward the north pole and toward the mountains. But

while for the terrestrial animals which fled to the'highlands no other

way was left than to go upward, for some of the fresh-water animals

a third asylum was open: the deep and cold fresh-water lakes on the

northerm:batder of thie jes <i» saw amot topo deat oegliio sio

No. 424.] NOTES AND LITERATURE. 333

North American Finches.'— This work, when complete, will of

course be the standard book of reference on North American birds.

It represents the matured views of our foremost ornithologist and is

remarkable for the clear and terse, yet ample, manner in which the

facts are presented. It includes not merely the birds of the United

States and Canada, but also those of Central America, Mexico, the

West Indies, and the Galapagos Islands. A full bibliography is

given for every species and subspecies; and there are numerous

measurements of specimens from different localities, indicating

slightly variant forms not designated by special names.

The present volume includes the sparrows and finches. Excluding

the Galapagos genera, and three species introduced from Europe, we

have 186 species, to which are added 165 subspecies, making 351

named forms in all. The species may be divided thus:

I. Monotypic species in monotypic genera . . + - + + Ib

2. Polytypic species in monotypic genera iva N

3. Monotypic species in polytypic genera. . =- - + + - 106.

4. 59-

Polytypic species in polytypic genera. . + + + + +

Total monotypic species, 119; polytypic species, 67.

A monotypic genus is one which has only a single species; a mono-

typic species is one having only one form, 2.¢., without subspecies.

The monotypic genera of the first group are distributed thus:

1. Great plains of North America: Rhynchophanes, Calamospiza, Centronyx.

2. Mountains of Western North America: Oreospiza.

3. Mexico and Central America: Plagiospiza, Pselliophorus, Pezopetes, Acan-

thidops, Rhodothraupis.

4. Cocos I., off Bay of Panama: Cocornis.

5. West Indies: Melanospiza (St. Lucia), Loxipasser (Jamaica), Loximitris

(Haiti).

It is significant and interesting that there are no monotypic genera

with monotypic species east of the great plains; or, excepting Oreo-

spiza, west of them to the Pacific. The conditions in the west, which

have produced so many distinct species and subspecies, have not yet

given us, in the groups under consideration, new generic types. In

other words, the evolution of the peculiar western genera antedates

the comparatively recent development of local specific forms. ‘This

agrees very well with the results obtained from the study of other

groups, eg., the flowering plants. We find numerous closely allied

1Ridgway, Robert. The Birds of North and Middle America, Part I, Buz.

No. 50. U.S. Nat. Mus. 1901. 715 pp. 20 pls- -` x

334 THE AMERICAN. NATURALIST. [Vor. XXXVI.

species belonging to old and widespread genera, pointing to a recent

period of plasticity of types, not going so far, however, as to affect

the genera. For the development of new genera, environmental con-

ditions of great stability and complexity, such as exist to-day in the

tropics, seem to be most favorable; and it may very well be that

most of the distinctive North American genera had their origin at a

time long previous to the glacial period, wlien such conditions more

nearly prevailed on this continent. The changeable conditions of

recent periods, while extremely favorable to the production of new

subspecies or even species, might from their very instability be inimical

to the development of new genera.

It is interesting to observe how much more plastic some species

are than others. Thus, in the genus Melospiza, M. Zimcoinii and

M. georgiana, though: very widespread, remain monotypic, while

M. cinerea has split up into no less than twenty-one subspecific

forms, many of very restricted distribution.

A very interesting discussion is given of the Galapagos genus

Geospiza. This includes twenty-seven forms, the extremes of which

are so distinct that they used to be assigned to different genera; yet

the gradation from one to the other is almost complete. Messrs.

Rothschild and Hartert proposed to regard a number of these forms

as subspecies of the others; but Mr. Ridgway urges that there is

really no alternative between regarding them all as forms of one

species and treating them all as specific units. Since the former

course is one that would commend itself to no one, the latter is

adopted, with the result that we have a series of “ species " far more

intimately related than the forms usually classed in this category.

We feel much as we might feel if suddenly confronted with the

whole series of extinct types connecting our very *good" genera

and species of the present day. Were the Galapagos less isolated,

and the competition with other groups of birds more severe, no

doubt several of the links would be conveniently missing by this

time, and we should not have occasion to dispute over the status

of the remainder. - CR ue miM i :

Many endemic forms are recorded from the islands of the Cali-

fornia coast and the West. Indies. These are in most cases regarded

as subspecies only ; though, in the inevitable absence of intermediate

series occupying the intervening territory (this being water), it would

seem more logical to treat them as distinct species. We seem to need

rm to express these allied insular forms ; but since their conti-

_nuity with the parent species is assuredly broken, I would rather write

ane ‘ae

VELIM LU

No. 424.] NOTES AND LITERATURE. ; 335

the names as binomials ; e.g., Pyrrhulagra ridgwayi for P. noctis ridg-

wayi Cory; Melospiza graminea for M. cinerea graminea (Townsend).

A very singular case is that of Æuetheia bicolor omissa, occupying

most of the West Indian Islands, but replaced in Jamaica, Haiti,

Barbados, and Grenada by Æ. b. marchii. Are we to suppose that

the latter once occupied the whole chain of islands but has given,

way to the former wherever it appeared upon the scene?

Two statements in the work seem to call for special comment.

One is as follows: * The necessity for beginning this work with the

highest instead of the lowest forms is to be regretted, and may be

explained by briefly stating that owing to inadequate facilities for

properly arranging the larger birds in the National Museum collec-

tion these are not available for study, and consequently it became

necessary either to begin with the smaller birds, already systemati-

cally arranged, or else postpone the work indefinitely."

That such a statement should be made regarding our National

Museum may well make us feel ashamed. Is this country too poor

to provide facilities for such a man as Mr. Ridgway, who returns to

it a thousandfold the small means it has placed at his disposal? Are

we so blind that we cannot see that scientific knowledge is more than

the equivalent of money ; is not merely convertible into that medium,

but is in itself far more nearly an end of national existence, since

it adds to the worth of the individual himself, and not merely to the

worth of that which is temporarily attached to him?

The other statement referred to is quite different. We are told:

“There are two essentially different kinds of ornithology: systematic,

or scientific, and popular. The former deals with the structure and

classification of birds, their synonymies and technical descriptions.

The latter treats of their habits, songs, nesting, and other facts per-

taining to their life-histories." The present writer has to confess

that this statement quite took his breath away. The study of living

birds, then, is “popular,” but the study of their mortal remains,

stuffed with cotton and provided with tags, or occasionally, perhaps,

immersed in alcohol, — this is “scientific.” It is hardly possible

that Mr. Ridgway could have intended his words to be taken literally.

No one could deny that Mr. Ridgway’s work on museum material is

in the highest degree scientific, and we may freely admit that nine-

tenths of what is written on the habits, nesting, etc., of birds is much

less so; but it surely does not follow that the story of life histories

is in the least degree less scientific, in itself, than any other branch

of ornithology.. One might as well say that the study of alcoholic

336 THE AMERICAN NATURALIST. [VoL. XXXVI.

brains is scientific (because the literature of that subject is so), but

the study of mental processes is popular, or non-scientific, for the

reason that we have a mass of trivial literature on psychology.

EDAC

Notes on Fishes. — In a recent visit to San Diego the writer saw

in the possession of an animal artist, Miss Annie Andrews, good

paintings of the threadfin, Polydactylus approximans, and the sea

bonito, Gymnosarda pelamis. The threadfin is common about Mazat-

lan, but had never been taken in the limits of the United States.

It was once described as Polynemus californiensis by Thominot,

from “California”; but that California which stretches from Rogue

River to Cape San Lucas is zoólogically very indefinite, comprising

three distinct marine faunas. The oceanic bonito is common at

Honolulu and in Japan, and was once before noticed by Eigenmann

at San Diego.

In the Scientific American for December 21 Mr. C. F. Holder

publishes a photograph of Zuvarus imperialis, a large and rare fish

of the Mediterranean, lately taken at Avalon on Santa Catalina

Island, off the coast of California. There is no question as to the

identity of the species with the genus Luvarus, and no specific

difference appears in Mr. Holder’s photograph, a copy of which the

writer has seen.

Mr. Holder also reports that he has seen two specimens of the

oarfish, Regalecus (russelli ?), taken in Avalon Bay. One of these, two

feet long, was examined by him while alive. « Its topknot," Mr.

Holder says, “was a vivid or scarlet mass of plumes. The dorsal

spines, which merged into a long dorsal fin, extended to the tail.

The color of the body was a brilliant silver sheen, splashed with

equally vivid black zebra-like stripes." Mr. Holder was unable to

obtain either specimen, the finders insisting on placing them on

a piece of board to be dried in the sun as « curios.” In this con-

dition the water soon evaporated, and practically nothing was left.

In Science for Dec. 13, r9or, Gill and Townsend give an account

of a large fish about five feet in length, dredged by the 4/batross at a

depth of 1050 fathoms off the Chonos Archipelago in Chile. By

some accident the huge specimen was cast overboard, and the

description is made from a photograph. The fish is of trachinoid

No. 424.] NOTES AND LITERATURE. 337

in the deep seas of the Antarctic, as well as the shore waters,

trachinoid fishes related to Percophis and Notothenia are numerous

and characteristic.

In the Proceedings of the New England Zoological Club, Mr. Samuel

Garman shows that the Japanese deep-water Chimzra described by

Professor Mitsikuri as Harriotta pacifica is the type of a new and

still more primitive genus, which he calls Rhinochimera. It differs

from Harriotta by the possession of teeth like the horny covers of

. the jaws on tortoises and birds, without the separate tritors found

in Chimera, Harriotta, and Callorhynchus. Garman divides the

group into three families, Chimzride, Callorhynchide, and Rhi-

nochimzridz, the latter including Harriotta. Garman makes the

important discovery that the frontal holder or cartilaginous hook

on the forehead of the male is present on the adult of Harriotta

and Rhinochimera as well as on Chimzra and Callorhynchus. As

with the ventral claspers, this is developed only when the animal is

sexually mature. Its presence is therefore a distinctive character

of all the living Chimera-like fishes, and it is found in no other

fishes. Mr. Garman’s paper, though brief and not illustrated, is a

most valuable addition to our knowledge of these fishes.

Under the head of * The Smallest Known Vertebrate," Dr. H. M.

Smith gives in Science for Jan. 3, 1902, an account of a diminutive

. goby only fifteen millimeters long when adult, found in Lake Buki

in southern Luzon, where, from its great numbers, it is an article of

food of considerable importance. It is named Mistichthys luzontus.

The genus is very close to the Japanese Euteniichthys, also very

diminutive, but it has larger and rougher scales than the latter.

In the Bulletin of the Museum of Paris for 1901, Dr. Pellegrin

notes a collection of fishes obtained by M. Diguet in the Lake of

Chapala and in Rio Lerma in Mexico. Most of the species have

been already noted by Jordan and Snyder. One new species, Xenen-

dum multipunctatum, is described from the pond called Agua Azul,

near Guadalajara. I cannot agree with Dr. Pellegrin that Ameiurus

dugesi of Lake Chapala is the same as Ameiurus catus, nor that the

little Gambusia infans is the same as G. affinis.

Under the head of “Les Poissons Vénéneux,” Dr. Jacques Pel-

legrin of the Museum at Paris publishes a valuable account of the

fishes of which the flesh is known or suspected to be poisonous.

It appears that in Tetraodon, Spheroides, Tropidichthys, Balistes,

Monacanthus, and other genera — mostly globefishes, filefishes, and

338 THE AMERICAN NATURALIST. [Vor. XXXVI.

trigger fishes — a. poisonous alkaloid is developed which causes a

very dangerous disease known in Cuba as ciguatera. This poison

attacks the nervous system, and through it other organs, often pro-

ducing death. Similar alkaloids of less virulence exist in sharks

and some other fishes. Dr. Pellegrin regards it as a kind of device

for the preservation of the species by the destruction of its enemies

with the death of the individual. The virulence of the poison is

less in fresh waters and grows less away from the. tropics.

Similar effects are sometimes produced by the flesh of species

otherwise innocuous which have been themselves poisoned by

poisonous mollusks, polyps, or fish.

Illness caused by decayed fish flesh or by undigested fish has no

relation to ciguatera. This Pellegrin calls *ichthyosisme," and it

may be produced by various species under conditions adverse to

assimilation.

In a monthly publication called /apan and America, for January,

1902, Dr. Jordan gives an account of the Salmonidz of Japan, this

paper being an epitome of one sent to Japan for publication. The

valid species known in Japanese waters are the Sake (Oncorhynchus

keta), the Ginmasu (Oncorhynchus kisutch), the Yesomasu (Oncorhyn-

chus masou or O. yessoénsis, accidentally omitted in this paper), the

Benimasu (Oncorhynchus nerka), the Yamabe or Kawamasu (Salmo

berryi), the Ito (Hucho blackistoni), the Iwana (Salvelinus pluvius),

the Malma (Salvelinus malma), the Amemasu (Salvelinus kundscha),

and the famous Ayu, one of the finest food fishes in all the world

(Plecoglossus altivelis). This dwarf salmon of.the rivers should by

all means be introduced into the clear streams of Maine, Quebec,

and California. Few finer food fishes exist anywhere. D.& 1.

Vascular System of Bdellostoma dombeyi. — C. M. Jackson!

has made the vascular system of Bdellostoma dombeyi the subject of

investigation. The author emphasizes the interest and importance

attaching to the Cyclostomata owing to their being the lowest of the

Craniata and possessing many structural features which must be

regarded as ancestral. The blood-vascular system is described in

detail, and certain points concerning its comparative anatomy are

discussed.

The heart is a simple tubular organ, situated in a pericardial

chamber which retains free communication with the peritoneal

1 Jackson, C. M. An Investigation of the Vascular System of Bdellostoma

oe of the Cin. Soc. of Nat. Hist., vol. xx, No. 1, pp. 13-48. 3 pls.

No. 424.] NOTES AND LITERATURE. 339

cavity. "There may be as many as fourteen pairs of functional

branchial vessels, all the efferent branchial arteries on each side

being connected by a longitudinal commissural vessel which con-

tinues forward as the common carotid of that side. "The dorsal

aorta extends forward into the head region. An inferior jugular

vein is present, and there is a persistent subintestinal vein which

does not join the portal system. These characters are among those

which the author regards as primitive. There is a portal heart whose

walls contain distinctly striated muscle fibers. "The contractility of -

the portal heart is included among the primitive characters.

The following features the author considers to have been second-

arily acquired. The venous system is asymmetrical. The branchial

vessels distribute “to gill slits instead of gill arches.” The portal

system receives blood from the anterior body region. The caudal

vein connects with the posterior cardinals. The portal heart has

well-developed valves. H.W. R.

On Hair in the Equide. 11 discusses the hair

of Equidz with special reference to peculiarities which, he believes,

are of taxonomic importance. The chief characters which he finds

to be of specific value are the shape and size, the extent of develop-

ment of the medulla, the relative thickness of cortex and medulla,

and the distribution and arrangement of pigment in the cortex. The

cuticle exhibits only sight modifications in the several members of

the group. :

Much greater variability in the character of the hairs was found in

Equus caballus than in the zebras. The hairs of zebra-horse hybrids

show surprisingly little variation, even though the dams were of

various breeds. The hybrid hairs resemble closely those of the

zebra sire.

No evidence was found, so far as hair was concerned, in support

of um telegony hypothesis. H. W. R.

Notes. — Wesley R. Coe describes the nemerteans collected by the

United po Fish Commission steamer Fish Hawk, at Porto Rico,

in the summer of 1899 (Bulletin United States Fish Commission, 1900,

Vol II, pp. 223-229). Eight species are represented among the

very few individuals collected. At least three of the species are new.

The collections of Professor Verrill, at Bermuda, and Ehrhardt, at

Barbados, exhibit the same marked scarcity of nemerteans. -

1 Proc. Royal Soc. Edinburgh, vol. xxiii, pp. 375-390- 6 pls.

340 THE AMERICAN NATURALIST. [Vor. XXXVI.

An isopod parasitic on the small hermit crabs found in the vicin-

ity of Woods Hole is described by Millett T. Thompson (Bulletin

United States Fish Commission, 1901, pp. 53-56, Pls. IX, X). The

female parasite is found on the abdomen of the host, the compara-

tively small male being attached to the marsupium of the female.

The isopod, probably representing a new genus allied to Phryxus

resupinatus Müller, has been named Stegophryxus hyptius, the generic

name being given in reference to the covering of the female, as it

lies upon the host, by its marsupium.

Prof. Carl H. Eigenmann reports on his investigations into the

history of the young squeteague of Buzzards Bay and Narragansett

Bay (Bulletin United States Fish Commission, 190t, pp. 45-51).

The young fish are pelagic in habit, subsisting entirely upon shrimp

and smaller fish. The report considers further the distribution of the

young squeteague, their rate of growth, and the changes accompany-

ing growth, in body proportions and coloration.

Captain Tuttle of the revenue cutter Bear has presented to the

Museum of Stanford University a pair of native boots which he

brought back from. a recent visit to the north, and which were-made

by a native on St. Lawrence Island in the Bering Sea from the skin

of a half-grown bull fur seal, taken in the winter of 1900 in the ice

adjacent to the island. St. Lawrence Island is 500 miles to the

northward of the Pribilof Islands, and it is interesting to note that

this is the first time the fur seal has been taken at s considerable

distance north of the Pribilof group.

BOTANY.

De Bary's Bacteria. — The wisdom of a third edition of de Bary's

Vorlesungen über Bakterien! so long after the death of its author

may well be questioned. New discovéries on every hand have

revolutionized old views, and to properly revise such a work would

require it to be doubled in size and practically rewritten. This

Dr. Migula has carefully refrained from doing. The title-page

reads, * Dritte Auflage durchgesehen und teilweise neu bearbeitet

von W. Migula," and in the preface apologies are offered for lack

1 De Bary, A. Wertesangen über Bakterien. "Leipsig, Wilhelm Engelmann,

1900.

No. 424.] NOTES AND LITERATURE. 341

of thoroughness on the ground of respect for the memory of the

author and for fear of disturbing the form or framework of the

book. Changes have been made only * wo sich eine absolut zwin-

gende Notwendigkeit ergab." De Bary, who loved thoroughness

so well, would never himself have consented to such a thing. It

is safe to say that the book will not please either the reviser or

the readers. It is at least questionable whether it is ever wise

to revise a book after an author's death, especially when very

extensive changes will be necessary to bring it up to date, and in

this case it is quite certain that Dr. Migula would have done much

better to have written an altogether new book. The first edition

of de Bary's Vorlesungen uber Bakterien, published in 1885, con-

tained 18 woodcuts.and 146 pages of text. The new edition con-

tains 41 figures and 186 pages of text. Three new chapters are

added dealing with sulphur bacteria, nitrogen bacteria, and the

question of immunity, toxins, antitoxins, toxalbumins, etc. The

book is about the size of Fischer's Vorlesungen and will find a

place beside it on the library shelf. It compares very favorably

with Fischer’s book, but, as in the case of the latter work, the least

well-made part of the compilation is that relating to plant diseases.

Four diseases only are noticed, and no contribution more modern

than Kramer's wet rot of the potato, published in 1892, finds men-

tion. In case of Wakker's hyacinth disease, de Bary's statement

that “succéssful infections and an exact following of the life history

of the bacterium are still wanting" is repeated. Under “pear

blight,” the long exploded statement that the organism produces

carbon dioxide, hydrogen, and butyric acid is once more repeated.

Under Kramer's “wet rot of the potato," it is thought to be safe

to hint that there may possibly be other bacteria capable of pro-

ducing similar phenomena. Of general statements, the following

sufficiently indicate the value of this résumé : ** The firm cell mem-

brane of the plant cell is an almost insurmountable obstacle to the

bacteria, especially as the very great majority of the bacteria are

not able to attack cellulose. It is true that in recent years espe-

cially there have been described very many so-called “ bacterioses,”

that is, diseases in plants caused by bacteria, but only a few will

stand searching criticism. . . . In the great majority, bacteria are

certainly not the cause. . . . Considering the uncertainty and inad-

missibility of most of the accounts relative to bacterioses, there is

no necessity of inquiring further about diseases which presumably

will sooner or later be stricken out of the list of the bacterioses.”

342 THE AMERICAN NATURALIST. (VOL XXXVI.

All of which means that to be a safe guide a man must actually

have done something in the particular field which he undertakes

to summarize. The book is well made and attractive in appearance.

E. F. S.

A New Elementary Text-Book. — A clear indication of the broad-

ened scope of botanical instruction is afforded by the discarding of

class books which only a decade or two ago were sufficient for the

purposes of the ordinary teacher. The books that have replaced

them are not only very different from their predecessors but from one

another. Some are questionably the equals in any respect of those that

are dropping out; others, though they have the merit of giving a

broadened view of the subject, overshoot the classes they are intended

for; still others are as one-sided on a new phase of the science as the

Miis ones were on another.

As long as there is individuality in teachers, universal satisfaction

is hardly to be expected from any text-book, and as questions of

temperance and cruelty have not yet intruded themselves into the

botany of secondary schools, it is probable that individuality in its

teachers may long resist the tendency to mechanical uniformity to

which many forces contribute with considerable persistency. With

any book, however much the teacher may desire to avoid giving the

impression that all desirable knowledge is contained between its

covers, it is difficult to prevent many pupils from at least tacitly hold-

ing this opinion, and good as many of the recent books are, it is to

be feared that their use is causing some of the wholesome everyday

knowledge of one's dooryard plants of a generation ago to be replaced

by a broader and doubtless more scientific, but unfortunately less

practically tangible knowledge of vegetation in the abstract.

To meet the undoubted need of a more comprehensive work than

Dr. Gray’ s Lessons without losing the advantage of its rational point

of view, excellent spirit, and good handling, Mr. Leavitt, at the

request of the botanical department of Harvard University, has based

on it a little book! which appears admirably adapted to the class

room, and which, by the introduction of a series of well-devised and

simple exercises, makes possible that most desirable of evolution E

the conversion. of the recitation room into a laboratory. GERA.

2 ‘Leavitt, R. G Outlines of Botany Jor the High School TF BEA and Class-

Room. Based. on | VS Lessons in cnet American Book Company. 272 pp»

383 figs.

No. 424.] NOTES AND LITERATURE. 343

Notes. — Mr. Gandoger has turned his discriminating eye on the

North American representatives of Astragalus and Oxytropis, and

in the number of the Bulletin de la Société Botanique de France of

January 5 he adds a considerable number to their named species,

varieties, and forms.

A Rhododendron related to Æ. punctatum, and from the Savannah

River, Ga., is described under the name Æ. cuthdertii, by Dr. Small

in Zorreya for January.

In Zorreya for January Mr. Howe extends the range of Arceutho-

bium pusillum to Nova Scotia and Newfoundland.

Three palms (Ptychosperma elegans, Archontophenix cunninghami,

and A. Alexandre), commonly cultivated under the name Seaforthia

elegans, are disentangled in the Gardeners’ Chronicle of January 11,

by Dammer, who illustrates the first and last named by reproductions

of photographs taken in the Buitenzorg garden.

The species of Porphyra of the Pacific coast of North America are

revised by Hus in Vol. II, No. 6, of the botanical section of Proceed-

ings of the California Academy of Sciences, issued January 4.

Professor Peck's * Report of the State Botanist" for 1900, sepa-

rately printed from the 54th Report of the New York State Museum,

contains plates illustrating a large number of edible agarics.

Bulletin No. 3 of the Lloyd Library is the first number of a

mycological series, and deals with the genera of Gastromycetes.

Mr. C. G. Lloyd is the author. Forty-nine figures, mostly half tones,

are used in illustration. y

Miss Vail gives a readable history of Van der Donck and his sev-

enteenth century New Amsterdam garden, in the Journal of the New

York Botanical Garden for December.

In ZZodora for December, Miss Day concludes her annotated list

of the herbaria of New England.

A short article on Lapham, whose herbarium is selene at the

University of Wisconsin, is contributed by x daughter to T he Wis-

consin Archeologist for January.

The Journal of Botany for December contains a good xerit of

]. G. Baker.

In a neatly gotten-up but rather crudely illustrated. booklet pub-

lished by William Briggs of Toronto, and entitled Sylvan Ontario,

Principal Muldrew of the Gravenhurst high school gives leaf keys to

344 THE AMERICAN NATURALIST. [VoL. XXXVI.

the recognized trees and shrubs of his district. It is pleasing to read

that the high school possesses an arboretum in which practically all

of the species are cultivated.

The relation of forests to snow is discussed by Carpenter in

Bulletin No. 55 of the Colorado Experiment Station, illustrated by a

series of photograms from excellent negatives.

In paper, presswork, and general appearance the third annual

report of the New York State College of Forestry, recently distributed,

is more like a product of the British colonies than of the United

States.

The value of growth rings in the so-called exogenous trunk, as an

index of age, is considered by Roger in the Revue Horticole of

January zr.

The distribution in Iowa of twenty native forest trees is discussed

by Professor Bessey in T'he Forester for December.

The big trees of California are the subject of a well-illustrated

article by R. T. Fisher in Zhe World’s Work for February, which

also contains illustrated articles on the cultivation of tobacco under

cloth in Connecticut and the growing of olives in California.

What the cedar of Lebanon may become in cultivation is shown

by photograms in the Gardeners’ Chronicle for January 4.

` PETROGRAPHY.

Tillman's ** Text-Book of Important Minerals and Rocks,” ' is

an attempt to present to the student a brief account of the character-

istics of the common minerals and rocks. The plan of the book is

good. It discusses the elements of geometrical crystallography in

24 pages, in which only the most fundamental forms of the six sys-

tems are described. The physical and chemical properties of

minerals are discussed in the next 5 pages. Then follow 65 pages

devoted to the description of about seventy-five of the commonest

mineral species, and about 4o pages in which the descriptive

material is embodied in tabular form. The final 22 pages treat of

the commonest rocks.

lTillman; S. E. A Text-book of Imp Minerals and Rocks. With tables for

See one "Ner Yorks, John Wiley & Sons, 900 186 pp.

No. 424. ] NOTES AND. LITERATURE. 345

For an elementary. treatise the book contains enough matter.

The author unquestionably did well when he decided to limit. his

descriptions to seventy-five species and make them fairly full rather

than to attempt to discuss a greater number and devote only a few

lines to each. _ There is nothing to criticise unfavorably in the chap-

ters.on crystallography and on the physical and chemical properties

of minerals. They are extremely brief, to be sure, but they are well

arranged. ‘Their topics are well chosen, and the statements in them

are accurate. The: tables are likewise good. . They contain no

peculiar features that need elucidation.

In the descriptive portion of the volume there is quite a little loose

writing, which is especially to be condemned in a beginner's book.

The order in which the minerals are taken up is that which seems to

appeal to most writers of elementary treatises, It is not scientific,

but it possesses the advantage of impressing on the student the

economic importance of the different compounds. The important

metals and their ores and compounds, other than the silicates, are

treated together. Then come the silicates. The grouping of the .

latter follows no general scheme. The order in which they are dis-

cussed *is mainly intended to emphasize and fix in mind their rela-

tionships and importance as rock-forming minerals." . There are

some errors in this portion of the book and many irregularities in the

headings of the paragraphs. Many chemical formulas are omitted

where they might serve a useful purpose. This is particularly true

in the case of the silicates, for which no composition formulas are

given. In a few cases not even is the composition referred to.

There is an unfortunate spacing of paragraphs, that undoubtedly

will cause confusion. After the heading *'Feldspathoid Group"

there is no spacing to indicate that this group does not include all

the minerals following it to the end of the book.

These faults, and others like them which might be mentioned, may

seem very slight and trivial. As a matter of fact, however, they are

serious in a text-book for beginners, since, though they may not teach

error, they certainly lead to misconceptions on the part of the

‘students. They may easily be corrected in a revision ; and a revised

edition, in which these corrections are made, would prove a very

satisfactory text-book. :

The second part of the volume is devoted to the common rocks.

It is so brief as to be of little value. The omission of a number of

the rock names mentioned would not have injured the book in the

least. Peperino, talus, detritus, eurite, ophiolite, itacolumite, and

346 THE AMERICAN NATURALIST. [VoL. XXXVI.

hydromica schists occupy the student’s attention when he might

better be learning something more about gabbro than that it is a

"coarse diabase.” The use of “syenitic” as an adjective synony-

mous with hornblendic is no longer to be tolerated in an elementary

text-book, in spite of “its wide popular use in this sense in this

country.” This wrong usage of the word will never be done away

with so long as it is taught in the text-books. 2:

There are some inconsistencies in the classification, but they are

not mischievous. There are also several statements concerning the

composition and origin of a few rocks that will not bear close

scrutiny.

Since there « are no very elementary. text-books dealing with rocks

published in English, the present volume has a place to fill. Itisa

welcome aspirant for the honor of initiating beginners into rock

study.. It would have been more welcome, however, had it.left

some of its useless baggage behind. We hope Professor Tillman

will revise the entire book, as there are many good points about it.

In its scope it certainly “fills a want.” The volume contains 38

crystallographic figures but otherwise is not illustrated. W.S. B.

The Differentiation at Magnet Cove. — Washington! has. made

new analyses. of some of the Magnet Cove plutonic rocks that lead to

interesting conclusions. He finds that many of the earlier analyses

are lacking in accuracy. The new analyses result in a change in the

former classification of some of the rocks and seem to render neces-

sary a few new rock names. From the analyses he calculates the

composition of some of the important qe to be as in the table on

opposite page.

The covite was formerly irdd asa onsite. Its qualitative

composition is close to that of theralite. The latter, however, con-

tains a large quantity of nepheline, while. the former contains a

comparatively . small quantity. On the other hand, shonkinite is

melanocratic, while covite is mesocratic (that is, it contains about

equal quantities of dark and light. constituents). . According to

Rosenbusch's definition covite is a typical shonkinite, but Rosen-

busch's shonkinite is an entirely different type from the prisia

shonkinite of Pirsson.

: The arkite. has been wrongfully designated a poni ti is most

nearly allied to missourite, but it differs from this rock in the same

way that covite differs from. shonkinite, że., it is a leucocratic

; Comieco, cU Fourn, of Geols, vol ix (1901); p. 607.-

No. 424.] NOTES AND LITERATURE, 347

aggregate of leucite and nepheline, with subordinate dark components.

The author defines it as a “holocrystalline, porphyritic, leucocratic

combination of leucite (or pseudo-leucite) and nephelite, with

pyroxene and garnet.” The jacupirangites of Brazil vary from rocks

rich in nepheline (true ijolites), through rocks composed predomi-

nantly of pyroxene with small and varying amounts of magnetite

and nepheline, to types rich in magnetite and with little or no nephe-

line. The Magnet Cove rock corresponds to the intermediate of the

Brazilian types.

I. Ik I1. IVi Xp FIA VER

Orthoclase.. . ..| 351. |.28.6 | 51-8.] 293 39

An s su 39.8 39.0 hee 22.8

Leucite . Ew ; 36.9

Nepheline. . . . 3-1 6.2 20.3 9.0 25.5 38.7 4.0

Cancrinite. . t3

Sodalite 5.2

Nosean Ls

irit 5.1 6.9 5.7 4 4.6

Hornblende 18.8 |Augite| 6.9 15.0

iopsi I1.0 . &6 9.0 10.8 ird 64.0

Biotite . . ‘ 7.9 5.0

Apatite. 5 1.0 3.0

phene . 2.7 5 3.1

Magnetite ror 2.5 8.7

Extra AlOs .. ——.—.L—42 Garnet 14.5 15.3

i6 Pulaskite, from Fourche Mt., Arka

II. Pulaskite sla Braddock's. uiid. 3 our M

III. Foyaite , Magnet

Covite, near Voc tal Magnet Cov:

- V,- Arkite (leucite-porphyry), near Diamond Jo Quarry.

VI. Ijolite, near Dr. Thornton's, Magnet Cove.

VII. Jac —— a C

oma idiom anticis the author discusses the dendi elatis

» the rocks with great fullness.. He represents the variations in

their composition by diagrams and with the aid of these calculates

the composition of the magma which, gave them birth by meth

that are somewhat novel, The peculiar differentiation of the mass is

- thought to be due to its small size.. The process was probably a “sort

of fractional crystallization, the magma being regarded. as a solution,

so that, in. accordance with the laws of cooling. solutions, the solvent

(the portion in excess) crystallizes out first around the borders on

RUE 1

s Journ. of Geol., y vol. ix (1901), P: 645.

348 THE AMERICAN NATURALIST. | [Von. XXXVI.

cooling of the mass.". The original body of magma was leucocratic,

so that the alumina. and alkalies with. silica enough for the forma-

tion of feldspar and. feldspathoids, crystallized first, and hence in the

periphery of the mass. -

Densities of Liquid and Solid Rock Magmas. — In view of the

fact that a recently proposed theory of volcanic action accounts for

the eruption of lava as the consequence of the expansion of liquid

magma in its passage to the solid state, a paper by Doelter,! in which

this author discusses the densities of liquid and solid magmas,

becomes of great interest. By means of a series of experiments, in

the course of which fragments of known density were allowed to float

or sink in molten magmas, Doelter obtained a series of results which

are embodied in the following table, the figures indicating specific

gravities :

NATURAL

SUBSTANCE. Morrzw. GLASSY. CRYSTALLINE.

Melanite 325 3.55-3.6 3-55-3-6 3-65-3-7

ugite . 33 2.92 2.92-2.95 3.2.—2.25

burgite 2.83 2.55—2.57 2.55-2.57 2.75-2.78

Lava (Ætna) 2.83 2.58-2.74 2.71-2.75 2.81-2.83

Lava (Vesuvius) 2.84 2.68-2.74 2.69-2.75 2.77-2.81

Nephelinite . 2.74 2.70-2.75 2.686 2.72-2.75

Leucitite. 2.83 2.60-2.68 2.68-2.72 2.75-2.79

The Laccolite of Shefford Mountain.?— Shefford Mountain is the

easternmost of the series of nine hills of igneous material that rise

above the St. Lawrence valley in the neighborhood of Montreal. It

is thought by Dresser to be an old laccolite in Lower Cambrian and

Trenton sediments. Its material consists of essexite, nordmarkite,

and pulaskite, the first two of which possess almost the typical

character of these rocks. Associated with these are dark-colored

dykes of a camptonitic. type, and others of a theralitic type, and light-

colored ones of trachyte and bostonite. The latter are the younger.

All the rocks are thought to be differentiated products of a single

magma. The primary magma, according to this view, had nearly the

composition of pulaskite. - Excluding the dykes the first differentiate

was tlie basic essexite, the second was the acid nordmarkite, and the

third the intermiediate pulaskite. The analyses of the essexite (I),

a Wekes Jahrb. of Univ., etc., Bd. ii (1901), p. 141.

? Amer. Geologist, vol. xxvii (Oct., 1901), p. 205.

No. 424.]

NOTES AND LITERATURE. 349

the nordmarkite (II), the pulaskite (III), and the mean between the

first two (IV) follow:

SiO, TiO, Al,O;

Sates gs 17-64

65.43 +16 16.96

59.96 .66 19.12

59.29 .84 17.30

Fe,03

FeO MnO

4.65 .46

1.53. | 49

1.73 -49

3.09 -43

CaO BaO MgO K,O Na,O P,O; CO, SO; Cl H,O Total

5.66 .13 2.94 3.10 5.00 .65 .39 .28 .07 1.10 — 99.86

1.36 ,22. 8:36. $98.02. - .06 04 .82— 99.84

4.24 .12-. 265° 49r 6:98 t4 08 .14 1.10 —100,17

3.531 .07 1.58 4.23 547 .34 +20 .17 96 — 99.85

PUBLICATIONS RECEIVED.

' (Regular exchanges are not included.)

AGLE, W. C. Eastern Peru and eng Seattle, Wash., Homer M. Hill de

lishing Company, 1902. 45 pp. Socents. — DAME, L. L., and Brooks, H. Han

Book of the Trees of New England. ad Ginn & Company. 8vo, xv + 196 pp.

87 pls. $1.35. — FABRE, J. H. Insect Life. Translated from the French by

the author of Mademoiselle Mori. New York, The Macmillan Company, 1901.

xii 4- 320 pp., 16 full-page illustrations. — GEGENBAUR, C. Vergleichende Anatomie

der Wirbelthiere mit Berücksichtigung der Wirbellosen. Bd. ii Leipzig, Wil-

helm Engelmann. viii + pps 355 figs. 20 marks; bound, 23 "=

— R. G. Catlins of Botany for the High Schóol Laboratory and Clas

R w York, American Book Company, 1901. 272 pp., 384 ledit

E .00. — ees W. H. D- Herbarium and Plant Description. Boston, Ginn &

mpany, 1901. Portfolio and 25 folders. 70 cents. — PACKARD, A. S.

dete his Life and Work. New York, Longmans, Green & Co., 1901. 8vo,

+ 451 pp., 10 pls. — PRATT, H. S. A Course in Invertebrate Zoology. Bos-

ton, Ginn & Company. xii + 210 pp. $1.35.

BRIGHAM, A. A. A Modern Dairy Barn. Bull. Rhode Island Agr. Exp. Sta.,

No. 8o. Pp. 17-107, numerous illustrations. — CoviLLE, F. V. Ribes aurum

and ribes lentum. Proc. Biol. Soc., Washington. Vol. xv, pp. 23-29. — DIXON,

R. B. Basketry Designs of the Indians of a sem California. Bull. Am. Mus.

Nat. Hist. Vol. xvii, pt. i, pp. 1-32. — ECKEL, E. C. Ihe Formation as the

asis for pieci Mappin A Weed n. id LS Vol pp. 708-71

EIGENMANN, C. H., and KEN Co Un ateral Coalo with Bilateral

Effect. ult N.s. Vol. xi; pP. Bolton breton C. H., and NORRIS,

A. A. Bergiaria. Comunicaciones del Museo Nacional de Buenos Altres.

Tome i, No. 8, p. 272. — EMBODY, G. C. Birds of Madison County, New York.

Generic Names of North American Skunks. Proc. Biol. Soc., Washington

Vol. xv, pp. 1-9. — JORDAN, D. S., and SINDO, M. A Review of the Pediculate

Fishes or Anglers of Japan. Proc. U.S. Nat. Mus. Vol. xxiv, pp. 361-381. —

Jorpan, D. S, and Sixbo, M. A Review of the Japanese Species of Surf

Fishes, or Embiotocide. Proc. U.S. Nat. Mus. Vol. xxiv, pp: 353-359- — JOR-

DAN, D. S., and SNYDER, J. O. A Review of the Discobolo vies of Japan.

Proc. U. S. Nat. Mus. Vol. xxiv, pp. 343-351. — KNIGHT. Ww. C., and SLOSSEN,

. E.E. Alkali Lakes and Deposits. Bull. Univ. of Nar Exp. Sta., No. 49. —

MEARNS, E. A. Description of a New Swallow from the Western United States.

. Proc. Biol. Sot., Washington. Vol. xv, pp. 31-32. — MERRIAM, C. H. Twenty

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Washington. Vol. xv, pp. 51-52. — NEEDHAM, J. G., and HART, The

Dragonflies of Illinois. Pt. i, Petaluridæ, Æschnidæ, and pie te ` Bui. fil.

tate Lab. o t. Hist. Vol. vi, art. i. September, 1901. — OBERHOLSER, H. C.

re of a ‘See of Humming Birds from eap and Colombia. Prec.

U.S. Nat. Mus. Vol. xxiv, pp. 309-342.— PALMER, W. and Riley, J. H.

t of Three New Birds from Cuba and ad. Kalis. Proc. Biol. Soc.,

hington. Vol. xv, pp. 33-34. — PEARSON, R. A. Market Milk: A Plan for

its Movement. Seventeenth Annual Keport of the Bureau of Animal Industry

(1900). Pp. 158-193, Pls. XXXIII-XLII.— PiNcHoT, G. Report of the For-

ester for r9or. From Annual Reports, Dept. of Agr. Washington, 1901. —

POLLARD, C. L. Nas J American Species of Chamecrista. Proc. Biol. Soc., Wash-

ington. Vol. xv, pp. 19-21.— SCHANS, W. Descriptions of New American But-

terflies. Proc. U.S. Nat. Mus. Vol. xxiv, pp. 383-460. — SHUFELDT, R. W.

Osteology of the Pinning, Annals of the Carnegie Museum. Vol. 1, pp. 295-

324. I901.— SiLLOWAY, P.M. Summer Birds of Flathead Lake: Bull. Univ. of

Montana, No. 3. 83 pp. 16 pls. — SiLLowav, P. M. Summer Birds of Flat-

head Lake. Bull, Univ. of Montana. Biological: Series No.1. 190r. — STANTON,

Vol. xxiv, pp. 301-307, Pls. XXV-XXVI. — SrEARNs, R. E. C. The

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271—299,

Pls. XIX-XXIV. era. L. A New mind i Sans fim Formosa.

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Anales del Museo Nacional de Montevideo. Tome iv, Ent. 22. 19or. — Dept.

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THE

AMERICAN NATURALIST

Vor. XXXVI. May, 1902. No. 425.

THE LAW OF ADAPTIVE RADIATION.

` HENRY FAIRFIELD OSBORN.

Oxe of the essential features of divergent evolution as

conceived in the branching system successively developed by

Lamarck, Darwin, Huxley, and Cope has béen termed by the

writer * adaptive radiation." This term seems to express most

clearly the idea of differentiation of habit in several directions

from a primitive type, as shown in the accompanying diagrams.

The law is a familiar one ; it results in the formation of analo-

gous radii in different groups of animals. The first compre-

hensive illustration of the law known to the writer is that

under the headings ** Homologous Groups " and * Heterology,"

in Cope's paper of 1868 on the “Origin of Genera," reprinted

in the Origin of the Fittest (pp. 95-106). This brilliant essay

is marred only by great confusion in the use of terms; but the

parallelisms in unrelated groups of amphibians and of mam-

mals such as marsupials and placentals, as first observed by

Owen, are clearly brought out.

In the present paper citations from earlier essays of my own

may be given bearing upon general adaptive radiation and the

independent production of analogous radii under the convergent

353

354 THE AMERICAN NATURALIST. [VoL. XXXVI.

principles of homoplasy, parallelism, and convergence, which,

as shown in the last number of the Naturalist, are by no

means synonymous terms or identical processes. The alto-

gether similar law of /oca/ adaptive radiation or incipient diver-

gence on a smaller scale in a single locality may now be more

clearly developed.

This idea of radiation becomes a means of interpretation,

and a way of imagining the relations of extinct and living

faunæ. As perceived by Cope, it applies both on a small

and on a vast scale.

I. GENERAL RADIATION.

In the * Rise of the Mammalia” ('93, pp. 30-33) the ancient

Mesozoic (Meseutheria) and modern Cenozoic (Ceneutheria)

differentiation of the placentals (Eutheria) was spoken of as

follows :

The Puerco is essentially an archaic fauna, and is to be regarded as the

climax of the first period of placental differentiation, a culmination of the

first attempts of nature to establish insectivorous, carnivorous, and her-

bivorous groups. These attempts began in the Cretaceous, and some of

the types thus produced died out in the Puerco, some in the Wasatch and

Bridger ; only a few flesh-eaters survive to the Miocene. It is most impor-

tant to grasp clearly the idea of this functional radiation in all directions

of this old Puerco fauna, resulting in forms like the modern insectivores,

rodents, bears, dogs and cats, monkeys, sloths, bunodont and selenodont

ungulates, and lophodont ungulates. This was an independent radiation

of placentals, like the Australian radiation of marsupials.

Some of the least specialized spurs of this radiation appear to have sur-

vived and become the centers of the second or mid-Tertiary radiation, from

which our modern fauna has evolved. Yet we have not in a single case

succeeded in tracing the direct connection. To sum up, we find on the

North American continent evidence of the rise and decline and disappear-

ance of monotremes and marsupials, and two great periods of placental

radiation,

ation, the ancient radiation beginning in the Mesozoic, reaching a

climax in the Puerco and unknown post-Puerco, and sending its spurs

into the higher Tertiary, and the modern radiation reaching its climax in

the Miocene, and sending down to us our existing types.

1 This statement has been modified by iibetquent discovery.

No. 425] LAW OF ADAPTIVE RADIATION. 355

In the * Origin of the Mammals ” ('99, p. 92) the idea was

developed as follows :

To guide our speculation in the unknown pre-Tertiary period, we may

gather certain positive principles from the known evolution of the Tertiary

Mammalia. First, we know that adaptive radiation, characteristic of all

vertebrates, and beautifully illustrated among Reptilia, is in a very high

degree distinctive of Mammalia, because of their superior plasticity.

The focal-types, or most primitive forms of the radiations, I-IV, «vere

certainly small, terrestrial, clawed, insectivorous or omnivorous forms.

It is noteworthy that in the evolution of each radiation, so far as we know

at present, land types and organs are invariably primitive, and water types

and organs are secondary, exactly as we find among the Reptilia. In fact,

we have not found a single instance in which a mammal or reptile series

is known to be transformed from a water into a land type; it is always the

reverse. There is certainly no evidence for a cetoid (Albrecht) stem of

the Mammals. Again, it is obvious that neither carnivorous nor herbivo-

rous types with highly specialized or reduced teeth and feet can be so

central as insectivorous and omnivorous types. In fact, the Insectivores

among Placentals, and Opossums among Marsupials, are the only animals

which have preserved the dental prototype close to that of the Promammal.

The radiations spoken of in this essay were:

I. Marsupial Radiation of Australia (Meteutheria).

Il. Tertiary Placental Radiation of the Northern Hemisphere, i.e.,

North America, Asia, and Europe (Ceneu eria). .

Ill. Tertiary Placental Radiation of South A merica (Ceneutheria).

IV. Cretaceous Placental Radiation of North America (Meseutheria).

V. Jurassic Radiation of Placentals and Marsupials.

In a subsequent paper, * The Geological and Faunal Rela-

tions of Europe and America during the Tertiary Period, and

the Theory of the Successive Invasions of an African Fauna "

(00), the subject was further developed as follows :

Now it is a well-known principle of zoólogical evolution that an isolated

region, if large and sufficiently varied in its topography, soil, climate, and

vegetation, will give rise to a diversified fauna according to the law of

adaptive radiation from primitive and central types. Branches will spring

off in all directions to take advantage of every possible opportunity of

securing food. The modifications which animals undergo in this adap-

tive radiation are largely of mechanical nature, they are limited in number

356 THE AMERICAN NATURALIST. [Vor. XXXVI.

and kind by hereditary, stirp, or germinal influences, and thus result in

the independent evolution of similar types in widely separated regions

under the Zaw of parallelism or homoplasy:

II. ADAPTIVE RADIATION of ORDERS AND FAMILIES AS BEAR-

ING ON GEOGRAPHICAL DISTRIBUTION.

This law causes the independent origin not only of similar genera but

of similar families and even of similar orders. Nature thus repeats herself

on a vast scale, but the similarity is never complete or exact. n

migrations are favored by over-population or geographical changes, a new

and severe test of fitness arises by the mingling and competition of the

parallel types.

Under the operation of these laws a most interesting generalization or

hypothesis can be made as to the three [zoólogical] realms : geographical

ded

> j

aI)

Fic. 1. — Ord f

during the Tertiary period. (From Osborn.)

isolation has been so continuous and prolonged that great orders of mam-

mals have been evolved . . . in each. Thus Arctogea, containing the

broadest and most highly diversified land area, appears hypothetically as

the center in which fourteen primitive and specialized orders radiated from

each other. In the southern portion of /Veogea at least four orders sprang

from primitive members of the above orders, and the Hystricomorph

rodents enjoyed their chief radiation. In Mofogea two orders were cut off

by the sea; one of them a rapidly declining type, the Monotremes, the

other, the Marsupials, enjoying a very highly diversified radiation. This

* At this time the distinction between homoplasy and parallelism was not

appreciated by the writer.

No. 425.] LAW OF ADAPTIVE RADIATION. 357

hypothesis is expressed in Fig. III (Fig. 1]. The other orders of mam-

mals, the Sirenia (probably a branch of the hoofed tribe), took the rivers

and coasts of America, Europe, and probably Africa as their radiating

center, while the Cetacea occupied the fourth or oceanic realm.

e mean to express by this hypothesis that REALMs [Fig. 2] were the

main centers of adaptive radiation of orders of mammals, but by no

ye ARCTIC

EURASIATIC NORTH TEMPERATE NORTH

(Palaearctic) (Hola €) ileyna

AWC TO CARD ;

CLE. — IND0-MALAYAN

Boy (Oriental)

N R

DaN Sin,

(Ethiopian) | . t

> A.P >

MALACASY ^ souT

(Ni

o p Eu ai

Fic. 2. — Division of the x into three realms and nine main geographical regions. The

tiene platform is raised to the 200 meter line showing the main Tertiary land connec-

tions. (From Osborn

means the exclusive areas of distribution, for during the periods of land

contact certain members of these orders found their way into adjacent

realms. Each realm, therefore, contains its pure autochthonous types an

its migrant or derived types. REGIONS, on the other hand, may be dis-

tinguished from realms as geographical and zoólogical areas, which have

been isolated from each other for shorter periods, either by climatic bar-

riers, as in the case of the Arctic or circumpolar region, or by great physi-

cal barriers, such as masses of water and of desert sands. In certain cases

these regions, such as Africa, appear to have been so large, distinct and

isolated, as to have become important centers of the radiation of certain

orders of mammals and almost attain the rank of realms, but regions in

general are chiefly and permanently distinguished by the adaptive radiation

of families of mammals.

In this paper Africa was treated hypothetically as a great

center of independent evolution and as the source of successive

northward migrations of animals. This hypothesis has recently

been confirmed by remarkable paleontological discoveries in

northern Africa. This adds to the above list of five radiations

a sixth, namely :

358 THE AMERICAN NATURALIST. [VoL. XXXVI.

VI. African Radiation of Placentals, chiefly Proboscidea ; Hyracoidea;

the families Antelopide, Giraffide, Hippopotamidea, etc.

X ) n

alll pO | i

Fic. 3.— Restoration of Antarctica, a hypothetical center of Tertiary adaptive radiation, by ele-

tion to the 304 ding line, showing old continental lines. (From Osborn

III. Tug Law of Locat ADAPTIVE RADIATION AS EXHIBITED

IN RELATED CONTEMPORANEOUS TYPES.

`- As seen in operation among the ungulates the competition

and range for food originates the lengthening of limb from

slow-footed into cursorial types, and the lengthening of teeth

from short-crowned (brachyodont) into long-crowned (hypso-

dont) types, and frequently the lengthening of skull from bra-

Chyceplialic into dolichocephalic types (Osborn, '02a).

The especial application to paleontology to be noted here is

: that as these types may have lived together or in proximity

and resorted to the same water courses for drink, their fossilized

No. 425.] LAW OF ADAPTIVE RADIATION. 359

remains are often found together. Yet if we examine anal-

ogous types living to-day, we see that they do not frequent

exactly the same feeding ranges nor do they subsist upon

exactly the same food; they thus do not compete. Good illus-

trations of this local adaptive radiation are seen in the distribu-

tion in Africa of the closely allied square-lipped Rhinoceros

simus with hypsodont. teeth, which lives upon grasses, and the

more pointed-lipped R. bicornis with brachyodont teeth, which

lives mainly upon shrubs. Both species belong to the same

phylum of rhinoceroses. Among fossil types closely related

to the above we.observe similar cases, such as the coexistence

in the lower Pleistocene of the hypsodont R. antiquitatis

(woolly rhinoceros) and the brachyodont A. hemitechus.

If carried farther than in the above instances, we may be

justified in placing these local adaptive radiations in separate

subfamilies, because in many cases they give rise to distinct

and long-persistent collateral phyla.

Examples of this kind are numerous among the ancient

Perissodactyla or ungulates related to the horses, tapirs, and

rhinoceroses, as seen below.

I. Stout-Foorep, HEAVY- IIl. SLENDER-FOOTED,

LIMBED PHYLUM LQieHT-LIMBE D,

CURSORIAL PHYLUM.

Families. Subfamilies. Subfamilies.

1. Palæotheriidæ Palzotheriine £8.) (Mid- Palaplotheriine (Middle

dle Eocene to Lower Eocene to Lower Oli-

Oligocene) gocene)

2. Titanotheriidze (Mid- Telmatototheriinz 77!

3. Hyracodontide

Paleosyopine Ø.

le Eocene to Upper

Eocene

Hyrachyine B. (Middle

(Middle Eocene to Up-

per Eocene)

Vin meu B. mmc

to Upper Eocene) Eoce

Lophiodontinz 7. (Mid-

dle to Upper Eocene) Colo

cene)

pper

4- Lophiodontidz Helaletine B. feud

don

tine ZB. crai

It will be observed at once that there is no inherent corre-

lation between brachyodontism and brachypody, or hypsodontism

and dolichopody, or elongation of the feet, as we might have

anticipated, although hypsodontism is gradually developed in

1 B. = brachyodont; H. = hypsodont.

360 THE AMERICAN NATURALIST. [Vor. XXXVI.

most long-footed series because subsistence upon grasses is

associated with such conditions of life as are afforded by exten-

sive open plains, long ranges for food, and rapid flight from

enemies. Again, as shown elsewhere, dolichocephaly and

dolichopody, brachycephaly and brachypody are frequently but

not invariably correlated. (See Osborn, '02a.)

The value of this law of Zoca/ adaptive radiation is especially

great as a means of interpretation of the frequent contempora-

neous existence or association of more primitive (brachyodont)

with more specialized (hypsodont) types. Among Tertiary

machzrodont cats it is seen in the contemporary long-limbed

Dinictis and short-limbed Hoplophoneus. In fact, the associa-

tion has been so often observed that if we find one phylum, we

may almost anticipate or predict the discovery of the other.

The law is made more clear by referring to the above table

and the following explanation of it: (1) As compared with

the Palzotheriinz, the Palaplotheriinz are so long-footed that

Huxley believed that they gave rise to the horse, and he actu-

ally placed Paloplotherium minus as the ancestor of the horse.

series. (2) The Telmatotheriinz are large animals also dis-

tinguished by very long, slender limbs; they independently

acquired horns, and the species 7. cornutum was believed by

Osborn to be ancestral to the titanotheres; but as a matter of

fact, as shown by Earle and Hatcher, this horn is a pure case

of parallelism, since the titanotheres probably sprang from the

short-skulled and relatively stout species, Pa/eosyops manteo-

ceras, a member of the Palcosyopinz. (3) The Triplopodinz

present the most extreme instance of light-limbed develop-

ment among the perissodactyl ungulates, since the limbs have

the proportions of some of the most slender and swift-

footed ruminant mammals, although these animals are found

in the same beds with the correlated subfamily Hyrachyinz.

(4) Among the Lophiodontidz, the Helaletinze bring out with

additional force the principle that this elongation of limb

occurred at a very early geological period; because the first

known member of this phylum appears way down in the Lower

Eocene, Wasatch, or Suessonien beds, in the genus Heptodon

. Cope, a remarkably light-limbed form, and it runs right through,

No. 425.] LAW OF ADAPTIVE RADIATION. 361

so far as we know, into the Oligocene genus Co/odon, the three-

toed type with extraordinarily elongate digits. So far as we

know, this light-limbed series is found both in America and

Europe, while the heavy-limbed Lophiodontinz are found only

in Europe.

Since the above was written the titanotheres have been

more carefully examined by the writer (Osborn, '02b), and, like

the rhinoceroses, they are found to subdivide into four con-

temporaneous phyla distinguished chiefly by dolichocephaly

and brachycephaly and by relatively long and short limbs,

thus affording another conspicuous illustration of this daw of

local adaptive radiation.

IV. RADIATION AND CORRELATION OF STRUCTURE.

In the careful consideration of adaptive radiation from cer-

tain stem types is to be found the true significance of Cuvier's

law of correlation as modified by the — to him — unknown prin-

ciple of evolution. Referring to the diagrams, Fig. 4, two

important principles are brought out: First, practically all the

adaptations known among mammals have arisen by combina-

tions of divergence independently pursued in the limbs and

teeth; for example, an herbivorous tooth type may combine

with a terrestrial, arboreal, or volant limb type, according as

the search for plant food is on the earth, in the trees, or in

the air. Although every imaginable combination (e.g., aquatic

limbs, myrmecophagous dentition) cannot be realized, yet these

combinations have been multiplied almost ad infinitum and

constitute the fatal defect of Cuvier's law as he conceived it.

As tested by a single case, the Eocene monkeys of the family

Notharctide acquired teeth exactly homoplastic with those of

Eocene horses, but the former were provided with arboreal,

the latter with terrestrial, limb types. Second, correlation of

limb and tooth structure in a given group is further condi-

tioned by the particular combination and degree of specializa-

tion of limbs and teeth which the radiation originates with.

For example, the primitive placentals combined tritubercular

insectivorous teeth with a generalized or probably terrestrial

362. THE AMERICAN NATURALIST. [Vov. XXXVI.

type of feet; the Australian marsupial radiation, on the other

hand, began with a dental type similar to that of the placentals,

LIMBS AND FEET

VOLANT

FOSSORIAL

RS ae

Short-limbed, nak

AMBU edad ie

impen yl, wisis

TERRESTRIAL

NATATORIAL

CURSORIAL

Amphibious Digitigrade

Aquatic ES

Unguligrade

TEETH

OMNIVOROUS ian

HERBIVOROUS ~ Shrub

Fruit

Root

MYRMECOPHAGOUS

ee Dentition reduced

Stem INSECTIVOROUS

Fic. 4 ^15. 1; i 3

Fish

CARNIVOROUS owes

E Bon te te

, (6) teeth among mammals.

but, as Dollo and Bensley have shown, the foot type was of

highly specialized arboreal character.

These two fundamental exceptions make clear why it is

impossible, as MY writers have observed, to reconstruct an

No.42 LAW OF ADAPTIVE RADIATION. 363

entire animal from either a claw ora tooth. Thus, while the

law of correlation is no less dominant than Cuvier supposed,

only a vestige is left of the mode of archetypal operation of

the law as he conceived it. It may be now restated as fol-

lows: Feet (correlated chiefly with limb and body structure)

and teeth (correlated chiefly with skull and neck structure)

diverge independently in adaptation respectively to securing

and eating food under different conditions. Each evolves

directly for its own mechanical functions or purposes, yet in

such a manner that each subserves the other.

Correlation is therefore not morphological, as Cuvier sup-

posed, but physiological, function always preceding structure.

It becomes closest where teeth and feet combine in the same

function as in the prehensile canines and claws of the Felidz,

and most diverse where the functions are most diverse, as in

the teeth and paddles of the Pinnipedia.

BIBLIOGRAPHY.

'68 Corr, E. D. On the Origin of Genera. Proc. Acad. Nat. Sci.

(October, 1868). Pp. 242 ef seg.

'93 Osporn, HENRY F. The Rise of the Mammalia in North America.

Proc. Amer. Assoc. Adv. Sci. Vol. xlii (1893), pp. 187-227.

.'99 Osporn, HENRY F. The ~ of Mammals. Amer. Journ. Sci.

Vol. vii (February, 1899), pp. 92-96.

'00 OSBORN, HENRY F. The uide and Faunal Relations of Europe

and America during the Tertiary Period, and the Theory of the

ste Invasions of an African Fauna. Science, N.S. Vol. xi,

o. 276 (April 13, 1900), pp- 561-574-

Bec Henry F. Dolichocephaly and Brachycephaly in the Lower

ammals. Bull. Amer. Mus. Nat. Hist. Vol. xvi, art. vii,

PP. 77—99.

Osporn, Henry F. The Four Phyla of Oligocene Titanotheres.

Bull. Amer. Mus. Nat. Hist. Vol. xvi, art. viii, pp. 91-109.

NEW AND LITTLE-KNOWN GUESTS OF THE

TEXAN LEGIONARY ANTS}!

CHARLES THOMAS BRUES.

Tue species included in the present paper are all Ecitophiles

which have been collected in the vicinity of Austin, Texas, by

Dr. Wm. M. Wheeler, Mr. A. L. Melander, and the author,

during the past two winters. The three species of Eciton

(E. schmitti Em. E. opacithorax Em., and E. cecum Latr.)

which occur here commonly seem to harbor a larger number

and a greater range of myrmecophiles than any other restricted

group of ants with which we are acquainted. Their nest mates

seem also to be more aberrant forms than those living with the

other ants, —a condition which may possibly be produced by

two unusual peculiarities of the Ecitons. Their eyes are very

small (even absent in the worker of E. cacum), and their power

of vision must be correspondingly poor ; then, too, their nomadic

life must prevent many myrmecophilous forms from living with

them, which would otherwise probably occur regularly. Eciton

schmitti and E. opacithorax change the sites of their nests quite

frequently and may often be seen marching in columns which

extend over long distances, and at such times their guests must

either travel along with them or be left to shift for themselves.

Eciton cecum is apparently always upon the move, but traveling

below the surface of the ground and not venturing out, at least

in the daylight.

Insects of three different orders are represented, Diptera,

Hymenoptera, and Coleoptera; the new Diptera which belong

to the family Phoride being among the most peculiar and remark-

able of described myrmecophiles.

1 Contributions from the Zoological Laboratory of the University of Texas, No. 25.

365

366 THE AMERICAN NATURALIST. [Vor. XXXVI.

COLEOPTERA.

CARABID.

1. Anillus affabilis sp. nov.

Length 2 mm. Rufotestaceous, darker anteriorly, elytra lighter than

head and thorax. Head smooth, shining ; frontal impressions deep but

not large. Antenna testaceous, scarcely longer than head and thorax,

slightly thickened towards apex. Palpi testaceous. Thorax distinctly

wider than long, widest near the anterior angles, narrowest at base. The

sides arcuate anteriorly and slightly sinuate posteriorly. Surface sparsely

but distinctly pubescent. Side margin very narrowly reflexed, distinctly

serrate, with four or five teeth near the slightly obtuse posterior angles.

Median line delicately but distinctly impressed. Posterior transverse

impression arcuate, the surface finely

rugose behind it. Elytra considerably

wider than thorax, distinctly serrate at

the broadly rounded humeral angles and

posteriorly to middle, where the teeth

become obsolete. Side margin regularly

arcuate, not at all angulated. A row of

large punctures near the lateral margin,

growing smaller and less distinct toward

the apex, form an ill-defined, broad lateral

stria. Dorsal striz almost obsolete, the

first three very faintly impressed, remain-

der indistinguishable. Elytra tra with three

long lateral setze, one at humeral angle,

one at posterior third, and one subapical.

Just anterior to the first long seta is à

shorter one. Elytra sparsely clothed

with erect pubescence. Legs pale tes-

Fic. 1. — Anillus affabilis sp. nov. taceous. 5

Described from three specimens collected at Austin, Texas.

Two were sifted from a nest of Eciton cavum Latr., and one

from a nest of Solenopsis geminata Fabr. While A. affabilis

may not be strictly myrmecophilous, it is evidently partial to ant

nests as a habitation.

This species most closely resembles A. explanatus Horn, from

which it differs in having broadly rounded humeral angles, no

angulation of the elytral margin, and by the three large elytral

No. 425.] THE TEXAN LEGIONARY ANTS. 367

The Californian A. debilis Lec. has been recorded from Texas

by Schwarz, but the species here described differs in having the

elytra serrate near the humeri. The original three-line descrip-

tion of A. debilis Lec! would not serve for identification, but

Horn? mentions that the elytra have the “ margin not serrate.”

STAPHYLINIDÆ.

2. Ecitonidia wheeleri Wasmann.’

A second specimen of this species was found April 20, 1901,

in a nest of Eciton schmitti Em., the same species with which

the type specimen was discovered. It is nearly as large as the

worker ants and quite similar in color and movements, so that it

is rather difficult to distinguish among a large number of ants.

It is undoubtedly mimetic of the ants, but whether its resem-

blance enables it to deceive the ants as to its identity would

seem very doubtful. The probability of its resemblance being

due to some outside influence of natural selection is heightened

by the fact that it must be exposed to insectivorous animals as

it marches along in the columns of the ants, which are often seen

in broad daylight and hence are exposed more than most species

of ants with their attendant myrmecophiles.

3. Ecitopora tenella Wasmann.*

The original types of this species were part of a collection of

numerous specimens from a nest of Eciton schmitti collected

during October at Austin. Since that time we have found

another species of this genus, the second to be recorded from

North America.

North American Species of Ecttopora.

Thorax one and one-half times as wide as long, strongly arcuate on the

sides, bns 6 ON o o ee ee tt laticollis sp. nov.

Thorax but slightly wider than long, almost straight on the sides, length

Tas mm SS S a tenella Wasm.

1 Trans. Amer. Phil. Soc., vol. x, N.S., P- 397- :

2 Trans. Amer. Ent. Soc., vol. xv, p. 26. * Loc. cit.

3 Neue Dorylinengáste, Zool. Jahrb., Abth. f. Systematik, Bd. xiv, Heft 3, P- 69.

368 THE AMERICAN NATURALIST. (VOL. XXXVI.

4. Ecitopora laticollis sp. nov.

Length 3 mm. Pale fuscous; head black, elytra almost testaceous,

thorax somewhat darker on the disk. First two abdominal segments with a

lighter band at apex, abdomen elsewhere fuscous, legs alutaceous. Antenna

fuscous, paler on three basal joints; beyond the third joint they are very

gradually thickened toward the tips, joints 4-10 being not quite twice as

wide as long. Eleventh joint conical, slightly longer than the two preced-

ing. Thorax one and one-half times as wide as head and one and one-half

times as wide as long ; widest at the middle, before which it is rather sharply

narrowed; behind it is not so strongly narrowed. Front margin of the

thorax somewhat concave on each side of the middle. Elytra not wider

than the thorax at their base. Abdomen at base as wide as the prothorax,

acuminate.

This species resembles Æ. tenella in general color and appear-

ance, but is larger, with darker head and much wider thorax.

Described from numerous specimens collected in a nest of

Eciton schmitti, February, 1901.

HisTERID E.

5. Ulkeus intricatus Horn.

Two specimens of this remarkable histerid were collected in a

nest of Eciton opacithorax Em., March 24, 1901. They agree

very well with Horn's original description, except that the frontal

carinze are divergent, not convergent above, and the network of

fine raised lines upon the discal portion of the thorax is very

faintly outlined in some places.

Horn makes no mention of the peculiar structure of the setae

with which the body is clothed. All the hairs upon the body

are provided with long barbs. Those upon the head have the

barbs shortest and scattered upon the central spine, but easily

distinguished from it. Those upon the remainder of the body

have no distinct central spine except at base, beyond which

they split up into about five or six sharp barbs. Upon the

dorsal surface of the body each forms a sort of stiff brush,

which is flattened so as to appear very slender when seen from

the side, but quite spreading when viewed from the front.

1 Trans. Amer. Ent. Soc., vol. xii (1885), pp. 142-143.

No. 425.] THE TEXAN LEGIONARY ANTS. 369

There can be no doubt that this histerid is a true myrme-

cophile of Eciton opacithorax Em., for Wasmann mentions it as

occurring in North Carolina near a nest of the same ant.!

6. Echinodes setiger Lec.

We have found this histerid in company with Eciton caecum,

although very rarely. It is not uncommonly seen, however,

in nests of Solenopsis geminata Fabr. and various species of

Pheidole.

HYMENOPTERA.

PROCTOTRUPID/E.

7. Ceraphron croceipes sp. nov.

Female. Length 1 mm. Black, shining, feebly punctured. Antenne

ten-jointed, black, scape ferruginous except at tip, where it is darker ;

pedicel pale at tip, constricted towards base, one and one-half times as

long as first flagellar joint; second joint as long as first but stouter, not

transverse; 3-5 equal in length, but gradually wider; 6 and 7 broadest,

subequal, quadrate; last joint as long as two preceding, and as wide at

base, acutely conically pointed at tip. Head finely punctured and white

pubescent. Frontal cavity deep, circular in outline, bare, and impunctured.

Groove in front of anterior ocellus punctiform. Mandibles dark yellow,

palpi pale. Collar black, with a lemon-yellow stripe on each side. Thorax

very delicately punctulate and white pubescent; mesonotum with a small

tooth at each anterior angle. Axille broadly meeting in front of the scutel-

lum, which is one-half longer than wide and fringed with stout golden

bristles on the sides, except near the apex, its sides strongly areolated ante-

riorly. Postscutellum rounded at tip, its sides sinuate and white pubescent.

Metathorax polished, its posterior angles produced and carinate. Meta-

pleure smooth, obliquely striolate behind, and bounded by two carine

above. Tegulz piceous, wings subhyaline, slightly infuscated toward apex.

Abdomen polished black, the second segment twice as long as the follow-

ing segments, coarsely striolate at base, with a tuft of white pubescence

at each anterior angle. Third and fourth segments about equal, fifth and

sixth shorter and equal. Legs, including cox, wholly deep yellow, except

the apical joint of all the tarsi, which is piceous. Posterior coxe with a

bunch of yellow hairs posteriorly.

Described from a female specimen collected in the galleries

of a nest of Eciton caecum Latr, at Austin, Texas, Feb. 3, 1901.

-1 Ein neuer Eciton-Gast aus Nord-Carolina, Deut. Ent. Zeit., Heft 2 (1897), p- 281.

379 THE AMERICAN NATURALIST. [VoL. XXXVI.

This species is quite closely allied to C. pedalis, flaviscapus,

glaber, and carinatus, but is distinct from all by the form of the

antenne, axilla, postscutellum, and maculation of collar.

Ecitonetes gen. nov.

Head globose, with a deep longitudinal frontal depression. Ocelli in

a triangle, small and close together. Eyes oval, one-half as long as head

and one-half as wide as long, coarsely faceted. Antenne inserted on pro-

jections near the base of the clypeus; ten-

jointed in female, scape stout, one and one-

half times as long as eye, pedicel one-half

longer than first flagellar joint, slender.

Antenna thickest at the seventh joint;

last joint about one-third as long as scape

and obtusely pointed at tip, equal in length

to pedicel. Maxillary palpi long, four-

jointed, last joint somewhat swollen; man-

dibles indistinctly bidentate. Thorax oval,

head, and without any furrows or grooves.

Scutellum absent, or at least not separated

from that part of the mesonotum which

extends over the place usually occupied by

ings Econ subapterus Ọ the scutellum. Metathorax produced into

posterior angles; much reduced, sloping on the sides and narrowed behind.

Wings very small, about as long as distance between posterior ocelli or

length of third antennal joint, with a strong vein along the costal margin

and three strong bristles on the apical front margin, each about as long as

the wing. Abdominal petiole very short, the abdomen squarely truncate

. at the base, where it is about as wide as the thorax. Abdomen oval, acumi-

nate, with six visible segments, the first of which is as long as the rest

together, second and third subequal, each about one-fourth as long as first.

— Legs rather short, stout; femora, especially posterior pair, swollen; anterior

and posterior tibial spurs well developed. First joint of all tarsi elongated

and longer than the three following ; claws simple.

8. Ecitonetes subapterus sp. nov.

. Length 2 mm. Light ferruginous, except the apical three joints of the

s antenne, — ee mack a sinuous dark band on the first abdominal seg-

rine

eT t LH > ' the narrow posterior Matai

No. 425.] THE TEXAN LEGIONARY ANTS. 371

of second and third segments, and an oval spot on the third segment, dark.

Eyes but little convex, finely hairy. Ocelli almost equidistant, small, and

rather close together. Basal half of antenna lighter than body, almost yel-

low. Thorax convex above, somewhat darker than the head. Abdomen

smooth, without striæ or grooves, convex above and rather sharply convex

below, so that the median line is quite sharply defined. Ovipositor black.

Described from one female specimen, collected in a nest of

Eciton cecum Latr., at Austin, Texas, during February, 1901.

While searching for the myrmecophilous Phoridz in the nests

of the large blind driver ant I chanced upon this small insect,

which, from its actions and color, simulated one of the very

smallest of the Eciton workers. Had it not been for its exceed-

ingly small size and more robust build it would undoubtedly have

passed unobserved

among the ants. On

examination with a

lens, however, it was

easily recognized as a

proctotrupid. I have

been unable to refer

it to any described

genus, although its

place in the Ceraphro-

nidze is quite certain. It seems most closely related to Lagynodes

Först., from which it differs by the ten-jointed antenne, shorter

scape, distinct ocelli, absence of groove on mesonotum, absence

of scutellum, and shorter legs. The dark spot upon the third

abdominal segment when strongly magnified is seen to consist of

a network of raised black lines, which are coarser and more

distinct posteriorly. This structure is interesting, as we know

that at the same relative position on the abdomen of other Eciton

myrmecophiles (Phoridz) we also find a spot where the integu-

ment is peculiarly modified. This suggests some possible way

that these blind ants may have of recognizing their habitual nest

mates by means of markings which they can perceive by touch.

Fic. 3. — Ecitonetes subapterus Q sp. nov.

372 THE AMERICAN NATURALIST. [VoL. XXXVI.

g. Telenomus texanus sp. nov.

Female. Length 1 mm. Black, shining, smooth, except thorax, which

is very finely punctured. Antenne piceous, lighter at base and apex of

scape and apex of pedicel. Mandibles yellowish at tips. Head but little

more than twice as wide as long antero-posteriorly, sparsely white pubescent.

Face shining, smooth, vertex distinctly reticulate with fine impressed lines,

reticulation extending down for some distance along the inner border of the

eye. Front ocellus in a depression, lateral ones contiguous with eye margin.

Eyes black, sparsely covered with short white hairs. Antennz eleven-jointed;

pedicel one and one-half times as long as first flagellar joint, which is nearly

twice as long as thick ; second and third thicker, but of same length as first ;

fourth small, transversely oval ; fifth larger, transverse ; three basal joints

of club nearly equal, subquadrate ; last joint conical. Thorax as wide as

head, more thickly and finely hairy, and not so shining. It is also reticulate,

but less noticeably than the head. Wings fusco-hyaline, marginal vein

three-fourths as long as stigmal, wing margins strongly fringed. Abdomen

very shining, slightly longer than thorax ; second segment about as wide

as long; the apical three segments. fringed with fine white bristles. Legs

and coxe deep yellow; hind femora infuscated on middle portion, tibia,

also, to a less degree. Tarsi paler, except last joint, which is black.

Described from a female specimen collected in a nest of

Eciton cecum Latr., near Austin, Texas, during March, 1901.

The insect was found among the ants at a distance of several

inches below the surface of the ground.

As all the other members of this genus are egg parasites it

is possible that this species may be parasitic upon some guest

of the Eciton, as is sometimes the case.

Telenomus texanus is most closely related to 7. geometre and

T. noctu, from either of which it is easily distinguished by the

form of the head, antennz, and marginal vein.

. Phaenopria acutiventris sp. nov.

Female. Length 1 mm. Black, shining, impunctured. Antennz rufo-

piceous, the three-jointed club piceous ; scape slender, arcuate ; pedicel

stout, oval, equal in length to first joint of club. First joint of flagellum one

and one-half times as long as the second, which is moniliform, third and

following joints increasing in size, moniliform except the last, which is oval

and as long as the two preceding taken together. Posterior margin of head

and anterior margin of collar white pubescent. Mesonotum considerably

‘narrowed in front, convex. Scutellum flat at base and strongly convex on

No. 425. THE TEXAN LEGIONARY ANTS.

373

the disk, without any pits or grooves. Wings hyaline, reaching considerably

beyond the tip of abdomen, strongly fringed. Metathorax pubescent, dark

rufous. Petiole and all the coxz and trochanters saturate yellow, the former

white pubescent. Femora and tibie strongly clavate, the swollen part pice-

ous, becoming concolorous with the coxz at the bases ; tarsi slender, fus-

cous, darker at the tips. Abdomen highly polished, as long as head and

thorax together, and conically pointed at the tip. Second segment reaching

to a little beyond the middle of abdomen; third and fourth segments very

short; fifth long and pointed, beset with a few stout white hairs.

Described from a female specimen collected in a nest of

Eciton cecum Latr., near Austin, Texas, during March, 1901.

I have placed this species in the genus Phzenopria because of

the form of the scutellum, although it resembles species of the

genus Tropidopria in the form of the abdomen.

PHORID/E.

Acontistoptera gen. nov.

Head very broad, more than one and one-half times as wide as thorax

at widest portion, widest at the front angles; half as long as wide ; seen

from above regularly arcuate in front, concave behind ; front angles sharp,

hind ones broadly rounded. Epistoma projecting slightly and visible from

above as a narrow band extending between the antennae. Eyes smaller than

‘in Ecitomyia, oval, their longest axis equal to diameter of large antennal

joint Upper surface of head with a pair of small median macrochætæ,

eight marginal ones on lateral and posterior edges, one over each eye, a

bunch at front angles, and a regular series of closely placed ones along the

front between the antennæ. Antennæ of the usual form, with an apical arista.

Palpi projecting, of the usual form. Head seen from the side higher than

long; front long, but slightly descending, rather acutely prolonged over the

deep antennal cavity. Mentum very large, bristly along the front margin,

and bearing several discal macrochete. Eyes very small, ‘contiguous with

front margin of the head below the middle of the antennal cavity ; omma-

tidia hemispherical, few in number. Sides of head with a few macrochete

below and behind the eye and a series of marginal ones from front edge to

middle of lower margin. oe

Thorax small; when seen from above, much narrower than the head and

owing to the narrowness of the dorsum. Dorsum with a pair of very long,

‘stout macrochztz at the anterior angles and a shorter pair directly posterior

| these. A pair of anterior, closely approximated, small discal macrochete ;

374 THE AMERICAN NATURALIST. [Vor. XXXVI.

two median larger and less approximate ones; and a pair of subapical

ones. Scutellum very small, with a pair of dorsal macrochete. Thorax,

seen from the side, but little larger than the head, strongly elevated in front;

coxz of the usual form.

Legs rather slender, espe-

cially the tarsi. Spurs of

four front tibize well devel-

oped. Wings about as long

as the width of the thorax,

arcuate, of nearly equal

bristles on its basal external

margin. Atabout the mid-

dle these suddenly enlarge

to form enormous macro-

chetz, which continue to

enlarge until at the tip they

are more than twice as long

as the wing and exceedingly

stout. There are about ten

of these large macrochete,

which are all strongly,

thickly; and almost scaly pubescent. Abdomen shaped as in Commoptera,

without any indications of segments or sclerites, except the one probably

corresponding to the fourth dorsal plate of Ecitomyia, which is small, with

- macrochætæ, and has a chitinous ring extending from its posterior edge,

which no doubt serves in connection with a gland like that of Ecitomyia.

Abdomen everywhere sparsely and regularly finely bristly.

Fic. 4. — A contistopt landeri Q sp. nov

11. Acontistoptera melanderi sp. nov.

"Length 1 mm.; of longest wing bristle, .4 mm. Light yellow, head

darker, fuscous in places, the occiput irregularly lighter, a small yellow

circular spot at the base of all the macrochzta. Antenne and palpi and

lower part of the head yeliosióh white. Thorax above but little lighter

than head. Wings almost white, their bristles black and very conspicuous.

. Abdomen, underside of thorax, and legs pale yellow. Legs finely black

hairy and more deeply colored distally.

Described from a single female specimen ! collected in a nest

of Eciton opacithorax Em., March 24, 1901, at Austin, Texas.

! After this article had gone to press di bars) fortius ast find ten more

eee: d Maher nest of the same ant on December 6 and 7; ep

AER.

No. 425.] THE TEXAN LEGIONARY ANTS. 375

On overturning a large stone, beneath which a small colony of

the ants had formed their nest, the fly in question was seen

darting about upon the surface of the ground in the exceedingly

nervous manner which characterizes the movement of Ecitomyia.

It appeared much larger than any specimens of Ecitomyia, but,

unfortunately, in capturing it the abdomen collapsed so that it

has been impossible to determine whether the large size was due

simply to a swollen condition of the abdomen or to something

attached to it (possibly one of the very large eggs or a larva).

This form is at once recognized by the marginal row of

enormous macrochzetz on the wings, which are also broader

than those of Ecito-

myia. The head is

exceedingly large and

wide in comparison

with the extremely

narrow thorax, which is

quite different from

that of any described

phorid, in that the dor-

sum is so narrow that

it allows the pleurz to

be visible from above

along their entire

length. The head is

very much flattened

and is remarkable for

the row of closely placed macrochztz along the anterior edge of

the front. Tue eyes are yat" the size of those of Ecitomyia and

have the same external structure. The mentum is large

and more dosely. continuous with the sides of the head than is

usually the case. The abdomen is wholly membranous, with

the exception of a single segment (the fourth ?), which lies directly

anterior to the abdominal gland instead of behind it as in Ecito-

myia; a rather remarkable difference were it not known that in

Fic. 5. — Acontistopt landeri Q sp. nov.

some cases nearly twice as large, swollen out and filled with a liquid in which can

be seen large developing eggs, some of which have the blastoderm completely

formed.

376 THE AMERICAN NATURALIST. | [Vor. XXXVI.

another Texan genus, Commoptera, the gland is situated in the

. middle of the segment and could thus give rise independently to

the condition of Ecitomyia or to that of Acontistoptera. The

legs of this form are long and quite slender, the front ones are

unfortunately broken off at the base of the femur in the only

specimen I possess.

Xanionotum gen. nov.

Head broad, one and one-half times as wide as the dorsum of the thorax

near the posterior angles; rounded triangular in shape, the anterior angles

not at all evident, posterior ones rounded ; twice as wide as long antero-

posteriorly, when seen from above, Eyes barely visible from above at the

extreme posterior angles of

N / the head. Four marginal

N ue PA macrochatz on middle of

Nue poa front, one on each side half-

AQ way toward the eye, another

24 just in front of eye, one at

F $ posterior angle, and a pair of

median posterior marginal

ones and two widely sepa-

rated discal ones. Antennæ

rather large, of the usual

form with apica arista,

attached at the lower part

of their unusually shallow

cavities. _ Palpi visible from

above for a distance equal to

diameter of second antennal

Fic. 6,—Xanionotum hystrix Q sp. nov. joint, rather slender, and with

e usual bristles. Epistoma

not visible from above. Ocelli absent. Eyes very small, situated slightly

below and in front of the center of the sides of the head. Cheeks bristly,

bearing a marginal row and an exceptionally strong macrocheta at the

lower posterior angle. Proboscis nearly as long as height of head, acumi-

nate, slightly geniculate at base. Thorax rounded, rather suddenly narrowed

behind, much narrower than the head and a little longer ; slightly wider than

long. It is very broadly rounded on the sides and at the anterior angles,

and not at all sinuate on the sides. Dorsum so wide as to almost entirely

conceal the pleura. Scutellum very small, without macrochætæ. Metathorax

visible behind it as a sclerite, which is but little larger than the scutellum.

Dorsum with a pair of strong macrochztz on the anterior angles, which are

No. 425.] THE TEXAN LEGIONARY ANTS. 377

discal ones. Thorax seen from the side considerably larger than the head,

the dorsum regularly arcuate. Coxæ as usual, the anterior ones rather short ;

legs slender, the tarsi long, spurs of four posterior tibiae well developed. Wings

nearly as long as width of thorax, clavate, about two-thirds as wide at base

as at apex, where they are rounded truncate. Wings very strongly bristly,

the macrochete longer than the wing, the longest being one and one-half

times as long, much more slender than those on the wings of Acontistoptera.

All the bristles are on the distal three-fourths of the outer margin, about

seven on the upper edge and three or four below. Abdomen of the usual

shape with the dorsal plate of only the fourth segment visible ; it is quad-

rate, somewhat narrowed in front and about as wide as diameter of second

antennal joint. The gland opening on the fourth segment small and with

the margin hardly at all thickened. The posterior margin of the four

anterior abdominal segments marked off by wide rows of enormous mac-

rochztz which extend more than halfway across the abdomen. Each row

contains about twenty bristles, those in the anterior rows being somewhat

the longest and nearly equaling the wing bristles in length and thickness.

Each bristle is not simple but composed of two distinct pieces, a short,

stout basal piece, apparently contiguous with the abdominal cuticle and

hollowed out into a spoon-shaped dorsal cavity at the apex into which the

bristle is articulated. Fifth, sixth, and seventh segments faintly indicated

by marginal bristles and by constrictions. Abdomen everywhere sparsely

short hairy. Sexual organs smaller than usual.

12. Xanionotum hystrix sp. nov.

Length 1.25 mm. Light yellow, almost white, the head much darker

rochaetze fuscous. Thorax tinged, darker above, especially in front. Legs

concolorous with the body, tarsi darker yellow, legs finely black hairy.

Described from a single female specimen ! collected at Austin,

Texas, March 24, 1901, in the same nest of Eciton opacithorax in

which the specimen of Acontistoptera melanderi was discovered.

Although the nest was carefully sifted it revealed no other

specimens of either form.

This form is undoubtedly the most remarkable phorid which

we have collected here, and although it was so closely associated

with the other new genus here described, there seems to be no

possibility of considering them as dimorphic forms of a single

species, as almost every part of the body is quite different in

1 We have since found another exactly similar specimen with the same ant,

December 6, 1901 - :

378 THE AMERICAN NATURALIST.

structure. It has been placed in a different genus on account of

the different shape and chzetotaxy of the head, its shorter thorax

with broader dorsum, abdomen with first dorsal plate present

and fourth wanting, and with the four rows of large spines.

We must of course expect to find a greater disparity between

these greatly degenerate forms than among more specialized

Diptera, but to include two such dissimilar species in the same

genus is hardly consistent

Re ME with the importance usu-

GAL ally laid upon structural

characters.

The head is very

strongly arcuate in front

and the lower margin is

not visible from above.

The antennz are larger

than in the other genus,

and the eyes slightly

smaller. The thorax is

not so unusual in shape,

although the pleurz are

slightly visible from above on the posterior part. The thorax in

both genera is, however, a closer approach to the typical dipteron

thorax than that of Ecitomyia. The rows of closely placed

abdominal spines are an unusual development, which give the

abdomen a most peculiar appearance. In this form the opening of

the abdominal gland is near the middle of the fourth segment.

Fic. 7.— Xanionotum hystrix Q sp. nov.

13. Ecitomyia wheeleri Brues.!

This form is an habitual nest mate in most of the Eciton caecum

nests which we have seen in this region, although the imagines

. become rare and perhaps disappear entirely upon the approach

.. of the summer heat and drought. :

—— The two specimens from a nest of Eciton schmitti Em. may

possibly have represented another closely allied species, but were

unfortunately sectioned before it was recognized that numerous

Species of myrmecophilous Phoridae occur in this region.

- DE -l American Naturalist, May, 1901.

ON THE STRUCTURE AND CLASSIFICATION OF

THE TREMATASPID.;E.!

WILLIAM PATTEN.

OF all the families usually included in the problematical and

heterogeneous group of animals called the “ ostracoderms,”’ none

is more interesting to the morphologist than the Tremataspidz,

as the little known about them shows they possessed a most

extraordinary structure, unlike in many respects that of any other

group of animals. While the character of the trunk scales, of

the orbits and other sensory openings, the minute structure of

the shell, and the presence of the newly discovered lateral-line

system clearly indicate the affinity of Tremataspis with Pteras-

pis, Cephalaspis, and Pterichthys, and through them with the

true vertebrates, other features, such as the general shape ‘of

the shield and its more superficial texture, which have long

been a source of perplexity to the paleontologist, clearly

point to the affinity of Tremataspis with arthropods like Limu-

lus, Apus, and the trilobites.

The importance of the Tremataspide to the morphologist

also lies in the fact that while the specimens are rare and more

. or less fragmentary, they are usually well preserved and give

fair promise that ultimately we shall be able to decipher in

detail the structure of all their hard parts. This knowledge

will certainly throw much light on the morphology of the whole

group of ostracoderms, and may afford decisive evidence of the

genetic relationship between the vertebrates and invertebrates.

en, therefore, through the generosity of the administra-

tion of Dartmouth College, I was granted a half-year's leave

of absence, I decided to make as thorough an investigation of

the ostracoderms as my time and means would allow, with the

special object of determining whether any evidence could be

1 This paper is an abstract of one about to be published in the Memoirs of the

Imperial Academy of Sciences of St. Petersburg.

379

380 THE AMERICAN NATURALIST. | [Vor. XXXVI.

found bearing out our assumption that they are an intermediate

group of animals related on one hand with the arthropods and

on the other with the vertebrates. My plan was to study all

the most important collections in Great Britain and the Conti-

nent and to purchase or collect material that might be used

for detailed study by sectioning or by other methods, as the

Fic, 1.— l i f th fada oe the hk

iin 39 di

specimens and casia in ipee hts d and the ime Academy, at St. Peters-

burg, and in the Dartmouth College Collect

d of Bs secon eso Bares

valuable type specimens permanently preserved in museums

could not be utilized in this manner.

It did not take long to discover that the following out of the

second part of my program, the collection of Thyestes and of

Tremataspis, was a most difficult task. So far as I know, every

fragment of these two genera has been taken from a shallow

pit about four feet deep and covering perhaps an area of three

or four hundred square yards, hidden in the heart of the remote

and otherwise little-known island of Osel in the Baltic Sea.

No. 425.] THE TREMATASPIDE. 381

The mysterious treasures of this classic spot have drawn to

its sides many famous scientific men from all quarters of the

globe. From time to time during the last forty years or more

many beautifully preserved eurypterids and an occasional Trema-

taspis have been taken from this insignificant pit in a pasture.

During the past twelve or thirteen years the spot has been

The d ] surf. f the head of Tremataspis.

FIG. 2.

worked more systematically by Mr. A. Simonson, who has col-

lected, with very few exceptions, all the material of Tremataspis

and Thyestes that has ever been found ; and yet, with the most

careful and painstaking work, and with considerable assistance

from common laborers, two or three, very rarely four, frag-

mentary heads of Tremataspis are all that reward the labors of

a wholé summer. I considered myself fortunate, therefore, in

being able to purchase nearly all of the fossils collected by Herr

Simonson during that season. The collection contained many

specimens of Thyestes, Cephalaspis, and Bunodes, together with

four heads of Tremataspis, some of which were in exceptionally

382 THE AMERICAN NATURALIST. [Vor. XXXVI.

good condition; but, unfortunately, none of the last-named

genus showed the presence of the plates in the oral region.

All of these specimens have been added to the collections in

Butterfield Museum of Dartmouth College.

The four heads of Tremataspis (which have been cleaned

with great care) have enabled me to make out several new

details concerning the sensory openings on the dorsal surface;

they were the first to show the system of lateral-line pits,

although these pits were afterwards seen, on most of the

St. Petersburg specimens. But they were of special value in

that they enabled me to work out the structure of the anterior

margin of the dorsal shield and to discover three new openings

Fic. 3. — Head of Tremataspis seen from the side.

in the series of so-called gill openings described by Rohon.

These parts in the Petersburg material were absent or had been

destroyed by. rough handling.

The reconstruction of the oral region was made from a

study of the single specimen and its cast that have already

been described by Rohon. The original fossil had lost many

details through repeated handling and the apparently incautious

attempts to clean out the matrix between the edges of the

plates. The original mold, however, was in nearly perfect con-

dition, and by taking several impressions of it in dentist’s wax

a beautiful reproduction of the original untouched fossil was

obtained, from which were worked out all the details in the

arrangement of the oral plates shown in the restoration.

These casts and the enlarged model are now — in

Butterfield Museum at Dartmouth College.

Our observations on the structure of Troinatipis have

brought out the following principal facts:

The Lateral-Line Organs of Tremataspis consist of a series

of shallow groove-like dots and dashes, arranged in linear

series on the dorsal surface of the shield. We distinguish a

No. 425. THE TREMATASPIDA. 8

393

circumorbital, marginal, anterior transverse, and a posterior

dorsal line. The first two lines appear to correspond with the

circumorbital, and the trunk line of Pterichthys. The circum-

orbital line is represented in Tolypaspis by a V-shaped ridge.

The Sensory Openings of the Dorsal Shield. — There are four

centrally placed openings and two pairs of marginal ones on

the dorsal shield of Tremataspis. Interpret these openings

provisionally as follows : the posterior central opening is that

of an olfactory organ; the three anterior central ones belong

to the median eye; the anterior lateral openings contain the

lateral eyes, and the posterior lateral pair, a segmental sense

organ comparable with the dorsal organ of embryo Limuli.

The olfactory opening and both pairs of lateral openings in

all well-preserved specimens possess sharply scalloped margins

- and a reticulated bony floor. Transverse sections show that the

floor is an extension of the inner layers of the shell.

In transverse sections through the frontal depression the

median slit is seen to be a true perforation of the shell, its

infolded margins forming a short flattened tube.

The median orbits are separate, nearly spherical chambers,

enclosed in a network of bony tissue formed by ingrowths of

the inner layers of the shell The narrow median canal that

appears to connect the two orbits is closed by a deep-lying

bony floor, also formed from the inner layers of the shell.

The /ateral eyes were small and subordinate in function to

the median ones. They probably occupied the anterior pair

of marginal openings, the rounded incisions on the edge -of the

openings possibly indicating the presence of compound eyes,

consisting of a few large ommatidia. The lateral eye orbits

agree with those of Limulus in being shut off from the interior

of the head by a bony network arising from the inner layer of

the shell.

The posterior marginal openings agree in position with the

so-called dorsal organs of Limulus, a pair of segmental sense

organs serially homologous with the lateral eyes and lying in

larval Limuli opposite the fourth pair of thoracic appendages.

The Median Eye.— The three anterior median openings I

have compared with the triocular median eye of Limulus, Apus,

384 THE AMERICAN NATURALIST. | [Vor. XXXVI.

trilobites, Merostomata, and other arthropods, but not with

the three frontal ocelli of adult insects. In Limulus, where

the structure and development of this organ is best known, the

retinas of the median ocelli arise from two pairs of segmental

sense organs, that during the closing in of the brain migrate

from the margins of the cephalic lobes to the roof of the fore-

brain vesicle. Here the ocelli come to lie at the blind end of a

long tubular outgrowth of the brain roof. The distal end of the

tube then divides into two vesicles, lying in the median line, one

in front of the other. The united retinas of one pair of ocelli

form one of the vesicles and, at a considerably later period, lie

in a degenerate condition, deeply buried beneath a median

tubercle on the dorsal surface of the head. The other two reti-

nas lie close together in the second vesicle, beneath two median

lenses. These two terminal vesicles are found in a more or less

modified form in many Crustacea and without doubt in the trilo-

bites and Merostomata also, since the arrangement of their sur-

. face lenses is, in some case, precisely the same as in Limulus.

The median eye of Tremataspis, like that of Limulus, probably

consists of a complex group of three ocelli derived from the

incomplete fusion of two pairs. They were, no doubt, true

cerebral eyes, lying at the end of a tubular outgrowth of the

brain. The distal end of this tube was probably bifurcate, the

anterior vesicle containing one pair of ocelli lying beneath, or in,

the median pit and the posterior vesicle lying in thepaired median

orbits. The anterior and posterior vesicles of Limulus and

Tremataspis are represented in true vertebrates by either the

vesicular ends of two separate outgrowths from the brain roof,

one behind the other, or by two terminal vesicles, one in front

of the other, arising from a common tubular outgrowth.

According to this view, the visual organs of vertebrates are

derived from three pairs of segmental sense organs, originally

situated near the margins of the cephalic lobes. The median

eyes, which were the most anterior, were the first to be con-

verted into cerebral eyes of the vertebrate type. This change

took place in the arthropods, the various steps in the process

being clearly seen in insects, crustaceans, and arachnids.

The transfer of the lateral eyes to the cerebral vesicles and

No. 425.] THE TREMATASPID.E. 385

their consequent inversion took place much later, probably in

the intermediate type of animals to which the Tremataspidze

belong. It was during this period that the median eyes

reached their highest development and the lateral eyes degen-

erated or, in some cases, disappeared completely from the sur-

face. We have no means of knowing whether their temporary

decline in functional importance was the cause, or the result,

of their transformation into eyes of the cerebral type.

The postorbital opening probably contained the forerunner of

the vertebrate olfactory organ. I have identified it with the

frontal organ of Limulus, Branchipus, Apus, and others. This

organ in the arthropods presents extraordinary variation in its

position, but can always be identified by the peculiar histological

structure of the terminal organ and its nerve, and by the origin

of the nerve in the brain. In Limulus, no doubt, the organ was

originally a visual organ serially homologous with the lateral

and median eyes. In the adult it lies on the ventral side in

front of the chelicerz. It gradually loses the histological

characters of a visual organ and finally presents many points of

resemblance to the olfactory organ of a vertebrate, especially

in the structure and relations of its nerves. The hypostomeal

. eyes of trilobites are very probably homologous with the olfactory

organs of Limulus. In Branchipus the same organ has moved

from its original position on the ventral surface to a point on

the dorsal surface almost as far back as the median ocellus. In

Apus the two organs have moved still farther back and have

united behind the median ocellus to form an unpaired organ in

precisely the same location as the post-orbital opening of

Tremataspis.

The Oral Plates have been worked out anew from the single

fossil and its cast that was used by Rohon. My description

differs from his in several very important respects.

According to my interpretation of this important fossil there

are on each side nine large oral plates arranged in four rows.

Some of the marginal plates are provided with one or more

rounded incisions lying opposite corresponding incisions of the

ventral and dorsal shields. The marginal plate of the fourth

row is probably a compound plate. There is a row of four or

386 THE AMERICAN NATURALIST. | [Vor. XXXVI.

five small plates on each side, just behind the anterior margin

of the dorsal shield.

A small triangular median plate lies in the anterior oral

region. Its anterior margin seems to be articulated to the

hinge-like process on the anterior median margin of the dorsal

shield. The outer surface of the plate is smooth and provided

with a low, keel-like ridge that gives it the appearance of the

rostrum of an arthropod. Its apex lies considerably below the

surrounding surface in a median depression that probably leads

into a small circular oral cavity like that of an arachnid. There

is no trace of a large transverse oral opening, like that described

by Rohon, between the anterior plates and the anterior ventral

margin of the dorsal shield.

The anterior margin of the dorsal shield is deflected sharply

downward to form a low wall in front of the oral region. A

small keel lies in the middle line on the posterior surface of the

wall, with its rounded edge directed backwards. Two rounded

toothlike projections of the rim, directed ventrally, lie on either

side. The lateral margins of the dorsal shield are folded sharply

toward the ventral median line and present three large rounded

incisions that must be regarded as a forward extension of the

series of six incisions of the ventral shield described by Schmidt

and Rohon.

The Appendages. — The most anterior incision is the largest

and is clearly the same as that so well seen on the margin of

the dorsal shield in Tolypaspis, Cyathaspis, and Pteraspis, and

which has been regarded as an opening for the lateral eye.

Lindstróm's important discovery of an appendage in Cyathas-

pis, my own discovery of fragments of the appendages in Trema-

taspis, and a renewed examination of the pteraspids in the

British Museum indicate that in these four genera the large

anterior marginal incision served for the attachment of an oar-

like appendage similar to that in Pterichthys and Bothriolepis.

The remaining openings, which are unquestionably serially

homologous with the first, must have served for the attachment

of other appendages of a similar nature. They decreased in

size from before backwards, and were pn too delicate to be

well poonvel) in a fossil condition,

No. 425.] THE TREMATASPIDA. . 387

The Entapophyses and the Trabecule. —1t is doubtful whether

the endolymphatic ducts of Rohon are actual perforations of the

shell. In sections they are seen to be deep tubular infoldings,

probably closed at the inner end. They appear to be comparable

with those infoldings, or entapophyses, on the dorsal shield of

Limulus which serve for the attachment of dorso-ventral muscles.

This interpretation is strengthened by the fact that serial sec-

tions show the presence of another pair of bony ingrowths, just

behind and in line with the so-called endolymphatic ducts. They

are long deep plates that unquestionably serve for the attach-

ment of muscles, since the frayed-out ossified tendons are still

attached to their inner ends. These entapophyses are of great

importance, as they indicate a similar arrangement of muscles

» and appendages to that seen in Limulus. They probably served

for the attachment of dorso-ventral muscles, arising from the

dorsal surface of the cartilaginous cranium and from the various

pairs of cephalic appendages.

The presence of a similar set of muscles in Pteraspis and

Cyathaspis is indicated by radiating impressions or lobes on the

inner surface of the shell, similar to the radiating muscle

markings on the dorsal wall of Limulus.

Transverse sections of the whole head show that the shell is

specially thickened in the region of the crista occipitalis, and

that the margin of the shield is considerably strengthened by

a network of bony trabecule uniting the edges of the dorsal

and ventral walls. The marginal trabecule are very similar

to those I have described in Limulus. The marginal cells of

Eukeraspis are probably produced by a special arrangement

of these trabecule. |

A loose network of bony trabeculz is also developed round

the median and lateral openings of the dorsal shield and along

the incisions of the ventral wall where the appendages are

attached. With the above exceptions, the inner surfaces of

the dorsal and ventral shields are smooth.

The minute structure of the shell of Tremataspis and other

members of the ostracoderms can be best explained, as we have

pointed out elsewhere, on the assumption that it is a modification

of a three-layered dermal skeleton similar to that of Limulus.

388 THE AMERICAN NATURALIST. [Vor. XXXVI.

The Resemblance between the Tremataspide and the Arthro-

pods.—The following summary of the principal characteristics

of the Tremataspidz shows how strongly they resemble the

arthropods, and how surprisingly the resemblance has been

strengthened at certain points by important discoveries in

unexpected directions.

The Tremataspidz resemble the arthropods in (a) their gen-

eral arthropod appearance, (4) in the minute structure of the

shell, (c) in the structure and arrangement of the openings for

the eyes and olfactory organs, (7) in the character and arrange-

ment of the oral plates, (e) in the absence of vertebrate jaws

and mouth and in the presence of a small centrally placed oral

opening similar to that of an arachnid, (/) in the presence of

numerous pairs of jointed appendages.

The presence of segmented appendages in Tremataspis and

related forms is indicated by the following evidence: (1) the

presence of a pair of oarlike jointed appendages, unlike those

of any true vertebrate, in Pterichthys, Bothriolepis, Cyathaspis,

Pteraspis, Tolypaspis, and Tremataspis; (2) the presence of a

fringe of jointed and movable appendages (25-30 pairs) along

the ventral margin of thetrunk of Cephalaspis ; (3) the presence

of a pair of crushing mandibles like those of an arthropod in

the head of Cephalaspis; (4) the presence in Pterichthys,

Bothriolepis, and Tremataspis of oral plates that appear to be

movable laterally, like the jaws of an arthropod ; (5) the presence

in Tremataspis of a series of eight other pairs of openings like

the ones to which the large swimming appendages are attached ;

(6) the presence in Tremataspis of two pairs of entapophyses

which, like those in Limulus, apparently serve for the attach-

ment of muscles moving several pairs of appendages; (7) the

presence in Cyathaspis and Pteraspis of from four to six radiat-

ing grooves on the inner surface of the dorsal shield, which, like

similar markings in Limulus, indicate the points of attachment

of dorso-ventral muscles moving several pairs of appendages.

The concurrent testimony, from so many different and

independent sources, to the fundamental similarity between

the Tremataspidz and the arthropods shows very clearly that

the resemblance between them is due, neither to incidental

No. 425.] THE TREMATASPIDE. 389

parallelism, nor to mimicry, but to genetic relationship and

community of origin. It could not have been a remote relation-

ship, signifying merely a common origin of both types from some

annelid ancestor, because the resemblance consists in the com-

mon possession of highly specialized structures characteristic

of the fully established arthropod and vertebrate types.

The genetic relationship, therefore, between the Trematas-

pida and the arthropods can mean nothing less than the

derivation, through changes in structure and function, of one

type from the other.

But we must not too hastily conclude that the Tremataspidz

are true arthropods, and by transferring them from one group

to the other avoid the real problem at issue. We cannot so

easily ignore the profound significance their structure has for

the origin of vertebrates. Their vertebrate affinities are too

obvious to be denied.

It is quite out of the question to separate the Tremataspidae

very far, either from the Cephalaspide on the one hand, as is

conclusively shown by the similarity in the structure of their

orbits, or from the Pteraspide and Pterichydz on the other,

as is shown by the appendages and the lateral-line markings,

or from either, as is shown by the microscopic structure of the

shell. The whole group must be kept together. But, as it is

quite impossible to include in the arthropods a group of animals,

nearly all of which have osseous dermal skeletons, and some

of which have such characteristically vertebrate bodies and fins

as the Cephalaspide and Pterichydz, or to include in the ver-

tebrates animals having many pairs of jointed appendages, it

becomes necessary to create for them a new class, one that

shall occupy a position between the true vertebrates and arth-

ropods, and unite these two great groups into one compact

phylum. 2

I propose for this new class the name Peltacephalata, and

include in it forms like Pterichthys, Cephalaspis, Pteraspis,

Tremataspis, and related genera. The class may be provi-

sionally characterized as follows:

The Peltacephalata were arthropod-like animals, moving

about through the soft mud on the bottom of shallow waters,

390 THE AMERICAN NATURALIST. [VoL. XXXVI.

in the typical arthropod position. In many cases most of the

body was probably concealed, leaving only the prominent median

eyes exposed. The presence of paired oarlike appendages indi-

cates the power of free swimming, but the more or less rigid and

clumsy appendages and heavily armored body could have pro-

duced little more than brief, spasmodic excursions, like those

of adult Limuli and eurypterids, or jerky, intermittent flights

through the water, like those of a copepod. And, just as in

these examples the shape of the body and the position of the

appendages in reference to the center of gravity compel the

free-swimming individual to reverse the usual position of dorsal

and ventral surfaces, so in the Peltacephalata the prevalence of

the same conditions must have forced them, after leaving the

bottom, to turn over and swim with the neural side uppermost,

in the true vertebrate position. The swimming movements

were probably aided in some cases by numerous small appendages

on the head and trunk. Fishlike caudal fins and tail were

used in swimming and in reversing the position of the dorsal

and ventral surfaces.

It was not till this new method of locomotion had completely

replaced the old that the eyes left the hamal surface (their

position in most adult arthropods) and returned to the neural

surface of the body (their position in embryo arthropods and

their permanent position in vertebrates).

The exoskeleton was a true dermal armor of ectodermic origin,

intermediate between the type presented by Limulus and that

of the more modern vertebrates. It consisted of three prin-

cipal layers, the middle one containing large, more or less

regular spaces or cancella. The matrix was strongly laminated

and penetrated by numerous dentine-like tubules, or pore canals,

and contained either unipolar or multipolar osseous lacuna.

The trunk was covered with rhomboidal scales or with seg-

mentally arranged ringlike plates. The presence of a system

of superficial sense organs is indicated by numerous pitlike

markings arranged in linear series.

A flattened cartilaginous cranium was present, but notochord

and vertebral arches were absent or rudimentary. Median and

lateral eyes were enclosed in bony orbits, sometimes protected

No. 425.] THE TREMATASPIDA. 391

by hard convex coverings continuous with the outer layers of

the shell. The median eye was large, complex, and important

functionally. It consisted of two pairs of ocelli, one pair com-

pletely united in the median line and the other nearly so. The

lateral eyes were reduced in size and in functional importance.

The nasal pit was unpaired and, in some cases, situated behind

the median eye. The mouth was small, circular, and situated

near the center of a group of oral plates. No upper and lower

jaws were present.

The head of the Peltacephalata may be regarded as a

modification of the cephalothorax of an arthropod ancestor,

consisting of three principal groups of segments, namely: the

precesophageal, including all the parts derived from the cephalic

lobes; the true thoracic segments; and the highly modified

vagus segments, formed by the forward migration and their

complete union with the thorax of from two to four abdominal

segments. The brain, like that of vertebrates, probably con-

sisted of three groups of neuromeres derived from these three

sources. |

The Peltacephalata have their nearest relatives among the

known invertebrates in the Trilobita and Merostomata, hav-

ing retained to a considerable extent the general shape of the

body, the structure of the head, and the mode of life character-

istic of these arthropods.

Sufficient data are as yet unavailable for a permanent arrange-

ment of the Peltacephalata into orders and families, but some

modifications of the old arrangement may be made to advantage.

The old subdivisions into Osteostraci and Heterostraci, pro-

posed by Lankester, should be abandoned, as they do not mark

natural divisions. The discovery of heavily armored oarlike

appendages in Cyathaspis and Tremataspis, and their probable

presence in Pteraspis and Tolypaspis, unite these genera more

closely with one another and with the Pterichyde than ever

before. It is therefore inadvisable to isolate the pteraspidian

section merely on the absence of multipolar bone cells, unless

the Pterichthydz are united with the remaining families

under the heading Osteostraci. But such an arrangement

would not sufficiently emphasize the resemblance between

392 THE AMERICAN NATURALIST. [VoL. XXXVI.

the oarlike appendages of Cyathaspis, Tremataspis, and

Pterichthys, and the difference between these appendages and

those of Cephalaspis.

Moreover, the Pteraspidee approach the Pterichydz more

closely than do the Tremataspidz in the division of the cephalic

buckler into separate plates, and in its separation into a true

cephalic or rostral portion bearing the median eyes, and a

thoracic one to which the oarlike appendages are attached. On

the other hand, Tolypaspis, which must be placed close to

Pteraspis and Cyathaspis on account of the minute structure

of the shield, shows no trace of a subdivision of its dorsal shield

into separate plates.

It seems to me, therefore, that we must recognize four

subdivisions of the Peltacephalata of about equal value, viz.,

the Pteraspida, Tremataspide, Pterichydz, and Cephalaspidz.

The Cephalaspidz occupy a somewhat isolated position on

account of the very peculiar shape of the head and the position

and character of the appendages, although on the other hand,

as shown by the connecting form Thyestes, a close relationship

between Cephalaspis and Tremataspis is indicated by the resem-

blance between their median, lateral, and postorbital openings.

The Pterichyda and Tremataspide are bound together more

closely than before, owing to their possession of oarlike append-

ages, large centrally placed orbits, and to the arrangement of

the so-called lateral-line organs and oral plates.

The Syncephalata. — For the great phylum of the animal.

kingdom formed by the union of the vertebrates and arthro-

pods I propose the name Syncephalata. The delimitation

of the Syncephalata can be only roughly determined, especially

at the lower end of the phylum. The main stalk consists of

the Arachnida (including the Trilobita, Merostomata), the Pelta-

cephalata, and the Vertebrata. The point of divergence from

the main stalk of such groups as the Insecta, Crustacea, and

the simplified and aberrant forms, like the Ternicata, Amphioxus,

Balanoglossus, and others, are of minor importance and do not

concern us here.

The justification of the term Syncephalata lies in the fact

_ that in this vast series of segmented animals the concentration

No. 425.] THE TREMATASPIDE. 393

and specialization of the anterior body segments into a head

region is definitely begun and completed. It is only when this

group is viewed as a whole that we see these momentous

structural advances in their true perspective, and can follow

the endlessly varied theme that leads steadily and consistently

onward toward the completion of the most complex organic

structure that has ever been produced, the vertebrate head.

DARTMOUTH COLLEGE,

January, 1902.

oneal

a m

DR SE, Bar

A joe i

VARIATION IN THE POSITION OF THE

ADDUCTOR MUSCLES OF ANA-

DONTA GRANDIS SAY.

ELLIOT R. DOWNING.

TunoucH the kindness of Mr. Frank C. Baker, curator of

the Chicago Academy of Science, I have had the privilege of

examining and measuring a number of exceptionally perfect

shells of Amadonta grandis Say. These shells were taken

in August, 1897, from “ South Pond,” Lincoln Park, Chicago,

where they were closely associated as a colony. As the pond

was thoroughly cleaned four years previously, the age of the

oldest shells is definitely known. Not only do the annual

limits of growth show, but internally the muscle scars and the

paths of migration are very distinct (see figures).

During part of the year shell growth occurs rapidly and

muscle migration is also rapid ; again, growth nearly ceases,

and the narrow dark band is deposited. During this slow

growth the muscle is about stationary, and then the very

distinct scar is formed.

The questions which I wished to settle were :

. Does the muscle in its migration move at a rate propor-

ose to shell growth, so that the muscle | retains a fixed

relation to the shell’s proportions?

2. If not, how does it vary?

I chose for this study the ratio,

distance from the umbo to the muscle

distance from the umbo to the shell margin

In all the shells the separation of the anterior adductor from

the anterior retractor is marked by a very distinct ridge, making

a fine line which lies in the path of migration. The distance

measured along this line on the curve of the shell, from the umbo

395

396 THE AMERICAN NATURALIST. | [Vor. XXXVI.

to the point where the outer boundary of the anterior adductor

scar cuts the line, made the first term of the ratio (see xi, x,

etc., Figs. rand 3). The second term was the distance from the

umbo along this line produced to its intersection with the shell

Fic. 1. Pic, 2.

EI. Fic. 4

Fic, 1. — Interior of a four-year-old shell, impressi hasized by ink lines.

Fic, 2. — Exterior of a four-year-old shell.

Fic, 3. — Interior of a two-year-old shell.

Fig. 4. — Exterior of a two-year-old shell,

4;, 42, anterior adductor scars; 7, 3 posterior adductor scars; vr, x3, etc., points on

ridge between anterior adductor and anteri rior tarator, fixed by its Taleo weg the

a ductor’s outer margin ; 71, Y2, etc., limit of posterior muscle

on at which adducto: in i th

ptt OT E ng to the ridge.

margin. A similar ridge is formed at the anterior margin of

the posterior adductor. The first term of the ratio for the

posterior adductor is the distance from the umbo to the point

where this line is tangent to the adductor scar (71, yz, etc.

Figs. 1 and 3). The second term is the distance measured

No. 425.] ANADONTA GRANDIS SAY. 397

along this line produced, from the umbo to its point of inter-

section with the shell margin. The proportions then are:

umbo to anterior adductor umbo to posterior adductor

: prod E

umbo to anterior margin umbo to posterior margin

The umbo in this species comes to a sharp point, which was

in no case eroded. It afforded a definite point from which to

measure. The measurements were made along the outer sur-

face of the shell. The shells were thin and quite transparent,

so that lines and points marked on the inside were readily

traced by a soft pencil on the outside by holding the shell to

a strong light (Fig. 4). Most of the shells were four years old,

a few three, and some two. (The exteriors of two- and four-

year-old shells are given in the figures) Measurements were

made and the ratios calculated for two hundred and seventy

anterior adductor scars, and for the same number of posterior

adductor scars.

In order to plot a curve for these ratios, the following classes

were taken for the anterior adductor :

CLASSES. FREQUENCIES.

585 — .604

605 — .624 8

625 — .644 27

645 — -664 38

665 — .684 9o

685 — .704 64

705 - -724 27

725 — -744 13

745 - -764 ee

270

The classes for the ratios of the posterior adductor are:

CLASSES. FREQUENCIES.

575 — -594

595 —-614 8

615 — -634 25

635 — -654 71

655 — -674 96

675 — -694 45

695 — -714 19

415 —.734 4

335 — -754 a

398 THE AMERICAN NATURALIST. [VoL. XXXVI.

Different limits for the classes were selected in the seriation

so as to give in each case the most nearly normal distribution

of frequencies.

Laying off the frequency polygon by the method of rec-

tangles gives Figs. 5 and 6, both curves of Type 4. The essen-

tial data for these curves are:

ANTERIOR, POSTERIOR,

A .6783 6613

Vi .0033 —.0037

Us —.00000 138 —.00000297

B, 00005 .O4I

B, 74.47 1388.8

E —142.94 -2771.47

a (skewness) —.00028 -.022

g (index of variability)

.0287 +.00083 .0254 4.00073

Discussion of results.

I. The distance of the adductor muscle from the umbo, meas-

ured along the line of migration, varies considerably zn its relation

to the distance to the shell margin, measured along the same line

produced. The extreme ratios are .591 and .754. The meas-

49 mm.

mm.

‘urements in the latter case are To produce the former

proportion we should have to have 37.3 mm. instead of 49 mm.,

— a change of 11.7 mm., or nearly x of the total measure-

ments.

2. While the range of variation then is comparatively large,

yet the frequencies are well concentrated at or near the mode.

In 93 per cent of the cases the position of the muscle would not

vary 3 mm., in either direction from the mean position.

3. Roughly speaking, we may say that the muscle is situated

two-thirds of the way from the umbo to the margin. Accurately,

the mean ratios are .6757 for the anterior muscle and .6608

4. It was thought at the outset of the investigation that the

results might indicate a difference in the rate of migration of

the adductor muscles, one approaching the shell margin at a

more rapid rate than the other. The negative skewness of both

No. 425.] ANADONTA GRANDIS SAY. 399

curves, however, would indicate a tendency toward smaller

ratios ;. that is, a decrease in the relative distances from the umbo

to the muscle impression. This may mean that the muscles

migrate constantly less rapidly than the margins grow. Since the

skewness is least in the anterior adductor curve, the anterior

adductor tends to move toward the margin at a more rapid

Fic. 6.

Fic. 5.

885 .605 .615 .635 .655 .675 .695 .725 .745 163 -575 -595 615 -635 655 .675 695 -715 735 -755 ;

1 8 37 489 Uny € 1.8 s y 95 «4 mw 4 t

Fic. 5. — Frequency olye for anterior adductor ratios.

Fic. 6. — Frequency polygons for pos:erior adductor ratios-

rate than the posterior, although there is a tendency in both

to lag behind the rate of growth of the shell margin. The

greater variability of the anterior muscle position would

indicate that it is the muscle most concerned in the evolu-

tionary process. However, variability and skewness are both

so slight that the above conclusions must be tentative until

400 . THE AMERICAN NATURALIST. ;

|. ^ . further evidence can be adduced. We seem to have in this

: form a species that is stable, rather than one that is in-

process of rapid evolution.

I am indebted to Dr. C. B. Davenport for valuable sugges-

|. tions and assistance in carrying on this is, iic. aniey

EN . HULL BIOLOGICAL LABORATORY,

CAGO UNIVERSITY

: NOTES AND LITERATURE.

ZOOLOGY.

Temperature of Insects. — Professor Bachmetjew's! paper is one

of those rare publications which is full of interest not only to the

specialist in entomology but to biologists in general. The Russian

author, with the equipment of the trained physicist, approaches a

subject that has often been studied before, and after treating it in an

exhaustive manner reaches new and important results, which would

carry conviction in their very simplicity, even if they were not sub-

stantiated step by step by detailed tables of observations. The work

of all previous investigators in determining the vital temperature of

insects is briefly and critically reviewed as a preface to each of the

main sections of the work.

In order to determine the temperatures, the insect was spitted

through the thorax on a thermoelectric needle consisting of fused

manganin and steel wires connected with a galvanometer. A detailed

account of the somewhat complicated apparatus and the method of

using it are given in an appendix (pp.:138-142). A number of

different insects, mostly larger moths, butterflies, and beetles, both

pupal and imaginal, were used in the experiments.

The first part of the.work is devoted to a consideration of the body

temperature of insects. In his earlier experiments, Bachmetjew came

to the conclusion that the temperature of the insect body varies within

very considerable limits, apparently without any serious consequences

to the life of the animal. He found, moreover, that in resting insects

the temperature is the same or very nearly the same as that of the

surrounding air, Subsequent experiments, however, led him to con-

clude that this is true only under ordinary conditions of moisture,

temperature, etc., since these factors, when abnormal, have a very

pronounced effect on the body temperature. Under normal condi-

tions, when the temperature of the atmosphere is raised, the tempera-

ture of the insect, though rising, lags at first more and more behind

1 Bachmetjew, P. Temperaturverhiltnisse bei Insekten. Experimentelle ento-

mologische Studien vom physikalisch-chemischen Standpunkt aus. Bd. i, pp. 4-

160. Leipzig, Wilhelm Engelmann, 1901.

401

402 THE AMERICAN NATURALIST. [Vor. XXXVI.

that of the atmosphere, and only begins to approach atmospheric

temperature just before partial heat paralysis of the wing muscles

sets in. . After death the temperature of the insect and the air are

the same. But when the air is very damp the body temperature of

the insect is higher than that of the air. This is explained as due to

evaporation of the body fluids and to respiration, the former having

a tendency to diminish, the latter to raise, the temperature of the

insect. Bachmetjew predicts that the study of the dependence of the

body temperature on that of the air, under different conditions of

moisture, will ultimately enable us to determine the metabolism of

these animals, and hence their vitality at different temperatures.

The influence of the activity of the insect on its body temperature,

long since noted by Newport (1837) and others, is exhaustively

studied by Bachmetjew. While a moth is moving its wings its body

temperature keeps rising, but falls suddenly with the cessation of this

movement. The insect was studied under three conditions : first, at

the ordinary temperature of the room ; second, at a higher tempera-

ture in the thermostat ; third, at a lower temperature in a cold-air

bath. Experiments conducted at room temperature show that the moth

(Sphingid) is incapable of raising its own temperature higher than

38.5" C. by means of muscular movement. Fluttering of the wings

does not produce as high a temperature as “humming.” At about

38° C. the insect often suddenly changed from humming to fluttering,

or rested completely. Bachmetjew interprets this change as due to

partial heat paralysis of the muscles. It is a transitory phenomenon,

which disappears with the sinking of the temperature during rest to

that of the surrounding atmosphere. The temperature at which the

wing muscles are paralyzed (in Deilephila euphorbiz) increases with

an increase of the temperature (at ordinary moisture) and reaches

45.5^ C., after which the moth loses the power of humming. Com-

plete, #2, no longer transitory, heat paralysis of these muscles super-

venes at a body temperature of 49.77 C. In somewhat moister air

this result does not set in till 53° C. is reached. Ina single experi-

ment on Deilephila at low atmospheric temperature the muscle

paralysis also appeared, but at a lower temperature. At a body tem-

perature of — 0.5° C. all movements ceased, fluttering began at 12? C.,

and humming not till 20° C. had been reached. It would seem,

therefore, that the temperature of partial paralysis of the wing mus-

cles is directly proportional to the body temperature of the insect, as

is also the case for higher temperatures. According to Bachmetjew,

these effects of partial paralysis play a great róle in the production

No. 425.] NOTES AND LITERATURE. 403

of color aberrations in butterflies. He also suggests that further study

of these effects may explain why so many moths are nocturnal, while

the butterflies are diurnal. Some experiments on the influence of

respiration showed that Deilephila at 29.4° C. atmospheric tempera-

ture could raise its body temperature through at least 3° C. by

means of breathing alone.

The second and more important portion of Bachmetjew’s paper

deals with the vital extremes of temperature. It is divided into two

sections, one dealing with the maximum, the other with the minimum

temperature. The vital maximum is the highest temperature at which

an insect is able to live. Experiments on Saturnia pyri showed that

the insect becomes very restless at a temperature of about 39° C.

and dies when the body reaches a temperature of 46^ C. This is

also very near the lethal temperature for plants (Sachs and Schultze).

This lethal temperature, however, depends on a number of factors.

In general, it may be said that if the insect at high temperature first,

has not been exhausted, Z^, has been artificially fed; second, is not

desiccated, że., is in a sufficiently moist atmosphere; and third, pre-

sents the same conductivity to heat and the same body size for a

given species, — its life will depend only on the coagulation or non-

coagulation of its body fluids. Hence, the vital maximum is only

another expression for the coagulation point of the body fluids. And

if one knew the amount of water in the insect's albumins, especially

of those albumins essential to life, the question of the vital maximum

would resolve itself merely into a determination of the amount of

water.

Bachmetjew's study of the vital minimum, Ze, the lowest tempera-

ture at which an insect can live, brought out some startling results.

He found from experiments on a great number of insects that

different species died at very different temperatures. But his most

interesting results refer to the critical point, which is the temperature

to which the fluids of the insect may be undercooled before they

begin to congeal and then suddenly rise in temperature till the nor-

mal congealing point is reached. Bachmetjew points out the resem-

blance of this phenomenon to the well-known undercooling of water,

. which can be cooled to — 25° C. without freezing, but at once rises

to o? C. to freeze. Bachmetjew discovered the undercooling of the

body fluids of insects by accident in an experiment on Saturnia

pyri 9. The insect was cooled to — 9.4^ C., whereupon within a

minute's time the temperature bounded up to — 1.4° C., the normal

congealing point of the body fluids, and then remained constant for

404 THE AMERICAN NATURALIST. [Vor. XXXVI.

eleven minutes. In this case the critical point is —9.4° C., and

— 1.4? C. is the normal congealing point. This insect revived within

an hour after the experiment, and laid eggs on the following day.

From this Bachmetjew concludes that the mere congealing of the

body fluids is not lethal. In a second experiment a moth of the same

species showed a critical point of — 11.6? C. (4.25 P.M.), whereupon

the temperature rose at once to — r.1? C. The insect was kept in

the cold, the temperature of its body again sinking to — 15.67. At

6.15 P.M. it was removed to the temperature of the room, but could

not be revived. It follows that the insect dies if its body is still

further cooled after the rebound (the limits being not necessarily

higher than — 2.5? C. nor lower than — 15.6? C.), or, as a general rule,

it may be stated that the insect dies if its temperature be again

reduced to about the point from which it rebounded. Further exper-

imentation on this interesting subject led to the following general

conclusions here briefly transcribed. The extreme degrees of under-

cooling of the fluids differ in different insects, and these extremes

occur at nearly the same rate of cooling. The critical point, so far

as its absolute minimum is concerned, is greater in pupæ than in

imaginal moths and butterflies, whereas the maximum differs in pupe

and imagines. Owing to lack of material, the behavior of the larve

could not be determined. With respect to sex, the degree of under-

cooling of the fluids in normal specimens is lower in the males than

in the females. This is also the case after brief fasting; after pro-

tracted fasting, however, the degree of undercooling is lower in the

males, but finally becomes the same in both sexes. On the other

hand, the normal congealing point of the fluids is lower in the females

than in the males. After fasting, it is the same in both sexes ; but

after protracted fasting, the relation is again reversed. Further

investigation of this question showed that the insect juices have a

lower critical point when the insect is fasting, but so far as its abso-

lute magnitude is concerned, it diminishes on continued. starvation.

Repetition of undercooling gave the following results: On freezing

a second time strong Lepidoptera exhibit a much greater degree of

undercooling than on the first freezing ; on being frozen a third time

the fluids show almost no undercooling. This is also shown by weak

Lepidoptera on the second cooling. Bachmetjew also studied the

influence of the fluid coefficient on undercooling. If M be taken as

the total weight of the living insect, and Pits weight after drying on

a water bath for a long time at a temperature of 115° C., 4 — P

No. 425.] NOTES AND LITERATURE. 405

would represent the weight of the fluids which leave the body on

evaporation at this temperature. The relation

is known as the fluid coefficient and signifies the percentage of fluids

in a unit of weight of the living insect body. Experiment shows that

the smaller the fluid coefficient the lower lies the normal congealing

point of the fluids. The critical point is also influenced by the fluid

coefficient, but this influence cannot be stated in general terms till

the composition of the fluids has been further studied. Time also

influences the critical point. If the temperature (7) to which the

insect is undercooled coincides at the same rate of cooling with the

critical point (.&,), the juices at once begin to congeal (z.e., time = a);

but if ¢ does not coincide with Ay, the congealing of the fluids is

delayed in proportion to this difference (A, — /).

Undoubtedly Bachmetjew's results are of a far-reaching character

and will ultimately form the basis for important work along theo-

retical lines in physiology, and for practical applications of great

moment (in economic entomology, e.g. !), for they throw light on the

geographical and climatic distribution of organisms, the resistance of

animals and plants to cold and heat, and the problems of anabiosis.

That Bachmetjew himself is very sanguine concerning the results

that may ultimately flow from his work is apparent when he says.

“ Es eröffnet sich somit ein ganz neues Gebiet für die Forscher, und

wer weiss, ob die Zeit nicht nahe ist, wo man den màrchenhaften

hundertjáhrigen Schlaf auch bei Menschen künstlich hervorrufen

könnte! Die Insekten wenigstens bieten die Möglichkeit dazu.”

. W. M. W.

An Important Paper on Phoride.— Theodor Becker, of Liegnitz,

Prussia, has recently published a work! of roo pages, with five

plates, on the family Phoridz, which deserves notice among zoolo-

gists in general because it is one of the finest pieces of systematic

work that has been published on the Diptera.

The family Phoridz includes only small species, generally from

two to four millimeters in length, which do not offer to the observer

with a hand lens a satisfactory series of specific characters. The

genus Phora was early described and generally recognized: from its

1 Abhandlungen der k. k. zool-botan. Gesellschaft in Wien, Bd. i, Heft 1, 1901.

406 THE AMERICAN NATURALIST. [Vor. XXXVI.

peculiar venation; this led to the description of a large number of

species by the earlier dipterists, among them Meigen, Macquart,

Zetterstedt, Bohemann, Haliday, Rondani, and Egger, the types of

which ultimately found lodgment in various public museums of

Europe. Most, if not all, of these describers based their species

chiefly on size and color, having but little comprehension of the

real specific distinctions. When they came to identifying each

other’s descriptions, confusion was worse confounded, and down to

the present time it has been impossible to get the family, which is

mostly comprised in the single genus, into intelligible shape.

Mr. Becker secured for study the material contained in twelve

public museums of Europe, including all the types now in existence

of the earlier descriptions, with a few exceptions. He not only

made a thorough study of all these collections, but he has pub-

lished a full report in this work on the named species and types

in each, thus putting the old species in a perfectly clear light and

preventing future disputes over alleged types and misnamed species

in these collections.

In addition to this material and that in his own collection,

Mr. Becker was able to study the collections of some eight dipterists

of the present generation, so that his work may fairly be termed

exhaustive.

Sixty-five European species of the genus Phora are described,

of which twenty-two are new. The remainder of the family, as

represented in Europe, consists of Trineura, three species; Coni-

cera, two species; Gymnophora and Metopina, one each.

he arrangement of the paper is admirable, and includes the fol-

lowing sections: table of genera of the world; structure and char-

acters of the genus Phora; analytical table of species; description of

species; enumeration of the named species in each of the type col-

lections examined, with the proper status of each specimen; a brief

division on biology; other European genera and species; extra-

European genera, with their species; index of European species,

including synonyms; list of accepted European species; list of

extra-European described species in the family, with references;

explanation of plates, and table of contents. This will give an

idea of the completeness of the work.

The characters used are largely those of the bristles, the “ chzeto-

taxy" of recent writers; without the use of these bristles it would

be impossible to write an intelligible description of many of the

species. There is no family of flies in which it is more essential.

No. 425. NOTES AND LITERATURE. Oo

407

One might wish to see a more extended notice of the biology

of the Phoridz, but as long as we have not yet nearly reached the

point where we can extend our classification to the larval stages, it

is not improper to allow the natural history of the species to form a

separate subject.

This paper will be of great advantage to American workers, as

it will enable them to ascertain how far our species are identical

with the European, Becker's descriptions being so exact that it will

be possible to determine without comparison of specimens, I should

judge.

To take up a family that is in a state of chaos and transform it

into order and beauty by a single publication is a great achieve-

ment, and one not accomplished without long and arduous study.

Mr. Becker has produced a monumental work, easily the greatest

he has yet attempted, and one which may well be taken as a model

by younger entomologists. J. M. A.

Habits of Insects. — An English rendering of the first volume of

Fabre's delightful Souvenirs entomologiques. Etudes sur l'instinct et les

maurs des insectes, though with an exceptionable title and an over-

burdened title-page,! is to be heartily welcomed. It makes access-

ible to a larger circle some of the early work of a keen inquirer into

the faculties of insects. Well and favorably known since 1879,

Fabre's observations have instigated similar and successful work

elsewhere, and it is only from the philosophical side, Fabre being a

rigid opponent to any form of evolution, that his writings are open to

hostile criticism.

The volume under notice begins with an account of the habits and

life history of Scarabeus sacer, and is devoted almost wholly to the

higher Hymenopteta, though incidental observations concerning other

insects are given.

With due allowance for the many difficulties, the translation is

fairly well done. Editorially the volume cannot be considered as

altogether satisfactory; the supervision of an entomologist should

have precluded the translation of gri//on indifferently as “ cicada,”

“cricket,” or “grasshopper,” the almost universal use of “feet”

1 Fabre, J. H. Zmsect Life. Souvenirs of a Naturalist. Translated from the

French by the author of Mademoiselle Mori. With a preface by David Sharp.

Edited by F. Merrifield. With illustrations by M. Prendergast Parker. London,

Macmillan & Co.; New York, The Macmillan Company, 1901. xii + 320 pp»

408 THE AMERICAN NATURALIST. [Vou. XXXVI.

(pattes) for “legs,” and the frequent confusion between genus and

family.

Footnotes recording similar work or opposite conclusions would

have been valuable, and the lack of an index is especially regret-

table. SH

BOTANY.

The Rhodomelaceæ. — Originally planned as one of the series

of monographs of the marine organisms of the bay of Naples,

the author! of this work has extended its scope until now it

covers the entire family of the Rhodomelaceæ, as represented in

all waters. Of its large quarto pages rog are given to the gen-

eral part, covering the anatomical development of the stem, the

morphology of the vegetative organs, and the reproductive organs ;

588 pages are given to the special part, with detailed studies of all

the species found in the Neapolitan region, and of all other species

authentic specimens of which were accessible to the author; 248

species are elaborately treated in this part. The third part, “ Sys-

tematic Results,” 34 pages, includes notes on phylogeny, on the

relation of the Rhodomelacez to other families, and a synoptical

view, practically a key to the genera of the Rhodomelacez, giving

under each genus the names, with descriptions, of the species

described in the second part, and of such other species as the

author had reason to consider sufficiently studied to leave no doubt

of their position under his arrangement. The large genera Laurencia

and Polysiphonia are excepted from this full treatment, only a por-

tion of the species being mentioned, about which the many other

species can be grouped; even with this reduction, 320 species are

given in this third part.

A monograph of this character, from the hands of the one person

competent for the task, is an important event, and the care and

thoroughness with which it is done are remarkable. The author

undertook ihe task in 1878, and some of the plates were printed

in 1885; after all, this long stretch of time seems none too much

for the enormous amount of work involved. What the future may

! Falkenberg, R. Flora und Fauna des Golfes von Neapel. 26. Monographie.

Die Rhodomelaceen. Herausgegeben von t der Zoologischen Station zu Neapel.

I9OI. xvi + 754 pp., 24 pls.

No. 425.] NOTES AND LITERATURE. 409

bring, no one can tell, but it is difficult to imagine anything that

will affect, except in details, so logical and well-grounded a classi-

fication as this. :

In 1889 Schmitz published his Systematische Uebersicht der bisher

bekannten Gattungen der Florideen, and in 1897 “ The Rhodophycez,"

in Engler and Prantl, Die Watiirlichen Pflanzenfamilien, was from the

manuscript left by Schmitz at his death; in both of these the Rho-

domelacez were based on Falkenberg's studies, but in both changes

were made, with Falkenberg's consent, from his original plan, to con-

form with Schmitz's general system. In the present work no such

change was needed.

The nomenclature of the present work, as compared with J. G.

Agardh's, the former standard, shows considerable change. Few

new species have been described, and consolidation of existing

species has, apparently, at least equaled their division; but the

larger genera have been split up, so that the number of new bino-

mials is quite considerable. The increased number of genera seems

the result of a logical employment of certain definite characters

throughout the family, — monopodial or sympodial growth, radial or

dorsiventral character of the frond, endogenous or exogenous char-

acter of the regularly distributed branches, presence or absence of

“leaves” in addition to the branches, number of pericentral cells,

their persistence unchanged or ultimate division, etc. The sexual

organs are practically uniform throughout the family, and where

the arrangement of the tetraspores appears to offer distinctive char-

acters, it is probably due to the structure of the branches in which

they are formed. In conformity with the general algological prac-

tice, no attempt has been made to substitute dead and forgotten

generic names for long-established ones of later date, but in some

cases the older names are given as synonyms; this last may be

quite a convenience for persons wishing to attach their names to

new binomials, though not familiar with the plants in question.

Many American forms are studied and figured, representatives

occurring of the new genera Brongniartella, Bryocladia, Dasyopsis,

Falkenbergia, Herposiphonia, Heterosiphonia, Lophocladia, Lopho-

siphonia, Ophidocladus, Pterosiphonia and Wrightiella; genera,

that is, that may be considered as new, for although most of them

appear in Schmitz's papers previously mentioned, they now for the

first time are given with full characters and list of species included.

Rhodomela floccosa of our northwest coast is transferred to Odon-

thalia, a very satisfactory place for the luxuriant, pinnately branched

410 THE AMERICAN NATURALIST. [Vor. XXXVI.

plant of the northern Pacific; but there are other forms passing

under this name of quite different habit, for which a place must be

found somewhere else. Polysiphonia bipinnata Post. and Rupr. is

referred to Pterosiphonia, but with exclusion of Polysiphonia califor-

. nica Harv., hitherto generally regarded as a synonym. ‘This trans-

fer being based on original material is undoubtedly decisive, but,

as in the case of Rhodomela floccosa, we are left with a long series

of forms, some of which will not go into Pterosiphonia. It may

be that they can be included in Polysiphonia californica, but more

study is needed. Chondria baileyana Harv: and C. sedifolia Harv. are

restored to specific rank, but in actual collecting it is not easy to

draw the line between the former and what is called C. senuissima on

the northeast coast, and between the latter and C. dasyphylla.

In regard to the Baltic forms of Polysiphonia violacea (Roth)

Grev., which Reinke considered identical with Z. harveyi Bailey

and P. o/neyi Harv. of the American coast, denying autonymy to

these two species, the author considers Reinke’s identification an

error, Harvey’s types being amply distinct from the Baltic forms.

This is a relief to American algologists, who were about ready, if

P. violacea and P. harveyi were united, to accept one name for all

four-tubed Polysiphonias whatever. The union under Rhodomela

pesos (Woodw.) Ag. of such various forms as A. /ycopodioides (L.)

, R. virgata Kjellm., and A. rochei Harv. is possible only by

endi an extreme range in habit and mode of fruiting. It still

seems as if A. subfusca, taken in so broad a sense, must be an

aggregate, to be divided sooner or later.

A work as thoroughgoing as the present, and starting from the

foundations, must continually reach conclusions differing from those

of previous writers, and the author states these divergences and

contradictions with great frankness; not ill-naturedly, but sometimes

apparently with a little impatience with errors which could have

been avoided by a little more careful observation. Practically

every one who has written on or referred to the Rhodomelacee

comes in for correction sooner or later, most of all the late Pro-

fessor Agardh, both as the most conspicuous writer and because

. in spite of, perhaps in consequence of, his remarkable intuitional

perception of systematic relations among the alga, he was never a

careful and punctilious student of the development of their struc-

ture, and it is upon the development, rather than on the mature

structure, that the classification of the present work is based. As

Teei with the elaborate synonymy of Bornet and Flahault’s

No. 425.] ' NOTES AND LITERATURE. 411

monograph of the heterocysted Nostochacez, the synonymy here is

quite meager; those who are interested in the ganzen Ballast veral-

teter Namen aus jener Zeit are referred to Agardh's, Kutzing’s, and

Harvey’s works. References to standard plates are abundant, but

there are practically no references to published exsiccata, which is

often unfortunate ; a plate can tell only what the artist saw, or even

only what he chose to represent; the plant itself, if in proper con-

dition, is ready to answer questions that never occurred to the artist

or the author. The plates in the present work give an instance of

this: when a figure is intended to show the position of certain cells,

it shows that with the utmost distinctness, but usually nothing more ;

no indications of thickness of cell walls, character of chromatophores,

etc.; where some other character is under consideration, that is given

the prominence, to the exclusion or subordination of all others. The

plates, however, tell admirably what they undertake to tell; if we

were told also where we could find the plant itself, nothing more

could be asked. But some remarks by the author as to specimens

which belong to several distinct species and appear as autograph

authentic specimens of a single species may account for a reluctance

to give exsiccatze numbers.

The work is written in a clear and comprehensible style, and now

and then contains a graphic expression, which, if not necessary for

scientific value, certainly does not detract from it. It would seem

that any student, with even a fair knowledge of German, could use

the work readily. The ordinary American student, however, will

be more likely to borrow the copy of some well-to-do friend than to

own one, the price being 120 marks.

Californian Nitophylla.'— In the historical sketch with which

this paper opens, the first reference to Californian Nitophylla is given

as *W. H. Harvey, 1858, Pt. II, p. 104, Suppl, p. 128." This is

misleading, as Part II of the Nereis was published in 1853 ; Part III,

with the supplement, in 1858. Thus the history of Nitophyllum in

California dates back five years earlier than given by the author.

The latest reference is in 1898, when J. G. Agardh published

Vol. III, Part III, of the Egzerzsis (noticed in the American Natu-

ralist for June, 1899), giving fourteen species for the west coast of

America. Mr. Nott’s careful study of living and dried material,

from all parts of the coast, shows that six of these must be

1 Nott, Charles Palmer. Nitophylla of California, Description and Distribu-

tion, Proc. Cal. Acad. Sci., Ser. 3, Botany, vol. ii (1901). 62 pp- 9 pls.

412 THE AMERICAN NATURALIST. [Vor. XXXVI.

considered merely forms of the other eight. A new species,

N. corallinarum, is added, together with the New Zealand species,

N. harveyanum, making the final list ten.

The genus has a world-wide distribution and includes many hand-

some species, with large, delicate, more or less veined membranes of

various shades of red. The Californian species are not inferior to

others in size and beauty, and the present paper gives a careful

study of them, with ample description, full synonymy and references.

The considerable variation occurring in some species is noted as

accounting for the new species proposed by Agardh; unless one had

a large series of forms of JV. ruprechtianum, for instance, it would be

perfectly natural to describe extreme forms as distinct species.

The plates illustrating this paper are by a photolithographic process,

and, while showing fairly well the habit of the broader membraned

species, are less satisfactory with the other species. Dependence in

determining must be had on the text and the references to exsiccate.

Alaskan Algæ.— This paper! gives quite an addition to our

knowledge of the marine and fresh-water flora of our northwest

coast, nearly half of the marine and more than half the fresh-water

species being new to Alaska. The really northern flora of the west

coast begins at Puget Sound, practically all the species from this

point north being distinctively northern in character, though some

of them extend south to central California. Of the red and brown

alez, 55 are circumpolar or found in the north Atlantic, 49 peculiar

to the Pacific. In the table, p. 394, these 49 species are in a column

headed “ Peculiar to the Pacific Coast of North America." As this

column includes such Asiatic species as Cystophyllum lepidium and

Odonthalia kamtschatica, and such south Pacific species as Macro-

cystis pyrifera, the heading is very misleading. If it were intended

to say that they were not found on any coast of North America

except the Pacific, it would be true, but that is not the natural

meaning of the words. Nine new species are described and fig-

ured by Mr. Saunders, Streblonema pacificum, S. irregulare, S. minutis-

simum, Dermocarpa fucicola, Homeostroma lobatum, Mvyelophycus

intestinalis, Coilodesme linearis, Mesogloia simplex, and Alaria fra-

gilis; also Pleurophycus gardneri Setchell and Saunders ; a few less

familiar forms already described are figured, a plate is given of a

s Saunders, De Alton. Papers from the Harriman Alaska Expedition. XXV.

The Algæ. Proc. Wash. Acad. Sci, vol. iti (Nov. 15, 1901), pp. 391-486,

-o Pls. XLITI-LXIL.

No. 425.] NOTES AND LITERATURE. 413

Liebmannia (?) but without specific name, and some 60 species of

desmids are figured on Plates XLIII and XLIV.

Nereocystis Priapus (Gmelin) Saunders takes the place of JV. Luet-

&eanus Mert, universally accepted since its publication in 1829.

While it is probable that Gmelin's plate of Ulva priapus, published

in 1768, represents a fragment of a frond of this species, no one

before Mr. Saunders has proposed the change, though Gmelin's

name has often been mentioned as a possible synonym.

Some curious phrases occur: p. 426, Alaria lanceolata is said to

be “easily recognized by the tufts of long cryptostomata "; p. 434,

Jridea membranacea J. Ag., “To this species Dr. Farlow has very

questionably referred," etc. The context shows that * questionably ”

is used in the sense of ** doubtfully," and the word is used in apparently

the same sense (p. 440) in regard to G/orosipAonia californica. On

p. 438 there are notes on the “ perithecia " of two species of Odon-

thalia. There is a curious tendency to give specific names the femi-

nine ending in all genera whose names end ina. In the table, p. 394.

this is consistently carried out, — Streblonema pacifica, Homeostroma

undulata, etc. In the descriptive text, later, some are changed to

neuter, others continue feminine. Possibly these are only instances

of the misprints which abound in the names throughout the paper,

due in part to scanty time allowed for correction of proof, and per-

haps to volunteered corrections by sóme other than the author. See

P- 414, Cladophora arctica for C. arcta.

The plates are clearly drawn and printed, the descriptions of new

species are fairly complete, and, as a whole, the paper is a valuable

addition to our knowledge of the plants of our northwestern

possessions.

Agardh's Alga.'—— In the notice of Part III of this work, in the

American Naturalist for June, 1899, attention was called to the long

time, the greater part of the nineteenth century, of Agardh's work in

this field, and the hope was expressed that the paper under consider-

ation might not be the last. One more part has been issued, but

only half of the proof had been read by the author at the time

of his death. In this part there is a rearrangement of the genus ©

Gracilaria, notes on some other genera, and an article, “On the.

Principles of Classification to be adopted for the Florideæ.” This

he considered as of great importance, as a final statement of his

1 Species, Genera et Ordines Algarum. Auctore Jacobo Georgio Agardh.

Vol. iii, pars iv (Lund, 1901), pp. 149.

414 THE AMERICAN NATURALIST.

views, and something of a protest against the Schmitzian principles.

At eighty years of age one does not readily give up the principles

upon which one’s life work has been based, but every system, when

it has done its work, must give place to something else, at least until

our knowledge has advanced far beyond its present state.

But the Agardhian system has been very useful in its day, and

with the death of its author we lose one of the most conspicuous

figures in the botanical field. Born in 181 3, the son of C. A. Agardh,

the foremost algologist of his time, his publications range from 1836

to rgor, the first part of the work whose last part we have just

noticed appearing in 1848. His main characteristics were his quick

grasp of a situation, however complicated ; his unerring instinct for

really important characters; his prompt recognition of true affinities.

With this type of mind, the toilsome, plodding investigation required

by modern conditions was not to be expected ; but it was the type of

mind needed to bring order out of the chaos of conflicting schemes

prevailing at the time, and to arrange the great numbers of new

forms coming to light in all parts of the world.

In person, as in mind, Agardh might be classed with the Norse

giants. Tall, well formed, athletic, dignified, serenely confident of

his position, he was a benevolent, gracious potentate of botany.

Very liberal in the distribution of specimens, he yet, like all royal

personages, held some in favor and some in disfavor, and more than

one private student in America has received from him a finer set of

his algz than can be found in Berlin, or some other great botanical

centers. The name Agardhia having been used in honor of his

father, Agardhiella, a genus of red algz with one handsome species

on our Atlantic coast and one on the Pacific coast commemorates

him, while many species bear his name. The red algae were his

special field, and in spite of all of the modern advances it will be

long before his works cease to be the place to which one will naturally

first turn when studying these plants.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS, AND DEATHS.

EDUCATIONAL GIFTS.

Allegheny College, $200,000, from various donors.

erst College, $15,000, from an anonymous donor; $1000, from

arren F. Draper.

Athol (Mass.) Public Library, $15,000, from Andrew Carnegie.

Atlantic, Iowa, $12,500, from Andrew Carnegie, for a public library.

Baraboo, Wis., $12,000, from Andrew Carnegie, for a public library.

Barnard College, $250,000, from John D. Rockefeller; $250,000, from

other sources.

Beatrice, Neb., $20,000, from Andrew Carnegie, for a public library.

Benton Harbor, Mich., $15,000, from Andrew Carnegie, for a public library.

Blue Island, Ill., $15,000, from Andrew Carnegie, for a public library.

Boston Public Library, $100,000, from the estate of the late Nathan

Haskell Dole.

Bozeman, Mont:, $15,000, from Andrew Carnegie, for a public library.

Brazil, Ind., $20,000, from Andrew Carnegie, for a public library.

Brown University, a conditional gift of $75,000, from John D. Rockefeller;

$100,000 and the residuary estate (which may amount to $500,000)

the will of Geo. L. Littlefield.

Buchtel College, $20,000, from several donors.

Carroll College (Wis.), $20,000, from Ralph Vorhees.

edar Falls, Iowa, $15,000, from Andrew Carnegie, for a public library.

Charlotte, Mich., $10,000, from Andrew Carnegie, for a public library.

Chicago Heights, Ill., $10,000, from Andrew Carnegie, for a public library.

Clark University, $100,000, by the will of Jacob Wheelock.

Columbia University, $3000, from Adolph Lewissohn; $100,000, from an

anonymous donor; $50,000, by the will of Mrs. Lila Currier; $11,000,

from an anonymous donor.

Cooper Union (N.Y.), $300,000 each, from Andrew Carnegie and the ,

descendants of Peter Cooper; an anonymous gift of $250,000.

Denison, Iowa, $10,000, from Andrew Carnegie, for a public library.

Denver Public Library, $200,000, from Andrew Carnegie.

Fulton, N.Y., $15,000, from Andrew Carnegie, for a public library.

Greensburg, Md., $15,000, from Andrew Carnegie, for a public library.

Hamilton College, $50,000, from the alumni.

Hampton, Iowa, $10,000, from Andrew Carnegie, for a public library.

415

416 THE AMERICAN NATURALIST. (VoL. XXXVI.

Harvard Medical School, $1,000,000, from John D. Rockefeller; $250,000,

from Mrs. C. P. Huntington; $100,000, from James Stillman; $471,225,

from others.

Harvard University, $450,000, by the will of George Smith; $100,000,

by the will of Robert C. Billings; $100,000, by the will of Jacob

wW |

elock.

Haverford College, $50,000, from Mrs. Charles Roberts.

Iowa City, Iowa, $25,000, from Andrew Carnegie, for a public library.

Las Vegas, N.M., $10,000, from Andrew Carnegie, for a public library.

Lehigh University, $5000, from Warren A. Wilbur

Little Falls, Minn., $10,000, fróm Andrew Carnegie, for a public library.

Littleton, N.H., $15,000, from Andrew Carnegie, for a public library.

ondon, Ohio, $10,000, from Andrew Carnegie, for a public library.

Maquoketa, Iowa, $10,000, from Andrew Carnegie, for a public library.

Massachusetts Institute of Technology, $100,000, by the will of Robert €

Billings.

Melrose (Mass.) Public Library, $25,000, from Andrew Carnegie.

Mt. Clemens, Mich., $15,000, from Andrew Carnegie, for a public library.

Nakoma, Ind., $20,000, from Andrew Carnegie, for a public library.

New Albany (Ind.) Public Library, $35,000, from Andrew Carnegie.

New Brunswick (N. J.) Public Library, $50,000, from Andrew Carnegie.

Newton, Kan., $10,000, from Andrew Carnegie, for a public library.

New York Botanical Garden, $5000, from Mrs. George Whitfield Collett.

Oskaloosa, Iowa, $20,000, from Andrew Carnegie, for a public library.

Paris, Ill., $18,000, from Andrew Carnegie, for a public library.

Redfield, S.D., $10,000, from Andrew Carnegie, for a public library.

Reno, Nev pe 5,000, from Andrew Carnegie, for a public library.

haieta Academy of Medicine, $5000, from Charles T. Ham, for medical

researc

Teacher’s Collega, Columbia University, $250,000, from an anonymous

donor. :

Tipton, Ind., $10,000, from Andrew Carnegie, for a public library.

University of Chicago, $1,250,000, from John D. Rockefeller.

University of Pennsylvania, $2500, from William Ivins; $2500, from

James Hay; $5000, from Ralph C. Stewart; land valued at $12,000;

from Gen. Isaac C. Wistar; anonymous gifts of $15,000.

f University of Wooster (Ohio), a conditional gift of $100,000, from Dr.

D. K. Pearson; $5000, from the directors of the Pennsylvania

Railroad ; $100,000, from Andrew Carnegie; $50,000, from .

— .. Severance.

Vassar College, a conditional gift of $200,000, from John D. Rockefeller.

Washington (Mo.) University, $25,000, by the will of Geo. E. Leighton;

. $20,000, by the will of William E. Huse.

breuis cer and Lee University, Bist from John D. Rockefeller;

n E SP PEN

No. 425.] GIFTS, APPOINTMENTS, RETIREMENTS. 417

Waukesha (Wis.) Public Library, $15,000, from Andrew Carnegie.

Waynesburg (Pa.) College, $36,000, from various donors.

William Jewell College; a conditional gift of $25,000, from John D. Rocke-

feller.

San Bernardino, Cal., $15,000, from Andrew Carnegie, for a public library.

Santa Rosa, Cal., $20,000, from Andrew Carnegie, for a public library.

Saratoga (N.Y.) Public Library, $20,000, from Andrew Carnegie.

Stevens Institute of Technology, $5000, from Alexander C. Humphreys.

Syracuse University, $100,000, from John D. Rockefeller.

Yale University, $5000, froni the class of 1876; $100,000, by the will of

rs. Lila Curger.

Yankton (S.D.) Public Library, $10,000, from Andrew Carnegie.

APPOINTMENTS.

Dr. A. F. Adams, assistant in histology in the University of Toronto. —

Dr. Leon Asher, professor of physiology in the university at Bern. —

Prof. Saverio Belli of Turin, professor of botany in the university at

Cagliari. — Dr. Max Belowsky, custodian of the Mineralogical Petrological

Institute in Berlin. — Dr. Max Bleibtreu, professor of physiology in the

University at Bonn. — Dr. Georg Bóhn, honorary professor of geology in

the university at Freiburg i. B. — Dr. G. Brandes of Halle, scientific

director of the zoólogical gardens there. — Dr. H. C. Bumpus, director of

the American Museum of Natural History. — Dr. Wesley R. Coe, assistant

professor of comparative anatomy in Yale University. — Dr. John M.

Coulter, professor of botany in the Manila Normal School, Philippines. —

Dr. Otto Drasch, professor of histology and embryology in the university

at Graz. — Théophile Durand, director of the botanical gardens at Brussels.

— M. H. Embrer, assistant in biology in the University of Toronto.— _

Dr. A. Ernst, docent for botany in the university at Zurich. — Dr. Ferdi-

nand Filarsky, custodian of the botanical section of the Hungarian National

Budapest. — Dr. Alexander Fleroff, docent for botany in the

university at Moscow. — C. M. Fraser, assistant in zoólogy in the Univer-

sity of Toronto. — Dr. Ph. Glangeaud, adjunct professor of mineralogy

in the university at Clermont, France. — Dr. Caswell Grave, director of

the U.S. Fish Commission Station at Beaufort, N.C. — Prof. J. W.

Gregory, acting head of the geological survey of Victoria. — Dr. Grinchant,

adjunct professor of mineralogy in the university at Caen, France. — Dr.

A. C. Haddon, advisory curator of the Horniman Museum at Forest Hill

near London. — Dr. R. W. Hall, instructor in biology at Lehigh University.

— Dr. Fr. C. C. Hansen, professor of anatomy in the university at Copen-

hagen. — C. Willard Hayes, geologist in charge of geology on the U.S.

Geological Survey. — Dr. E. Johann Gerhard Holm, professor and director

of the paleontological section of the Royal Natural History Museum in.

418 THE AMERICAN NATURALIST. [VoL. XXXVI.

Stockholm. — Dr. R. E. Hooper, assistant in histology in the University

of Toronto. — Dr. Jaroslav J. Jahn, professor of geology and mineralogy in

the Brünn Technical School. — Dr. Max Keernicke, docent for botany

in the university at Bonn. — Dr. Richard Kolkwitz, botanist in the royal

water-testing establishment in Berlin. — Dr. H. B. Kümmel, state geologist

of New Jersey. — E. A. Macallum, assistant in biology in the University of

Toronto. — Dr. W. J. Macallum, assistant in histology in the University of

Toronto. — Dr. S. Magocsy-Dietz, professor of vegetable morphology and

physiology in the university at Budapest. — Dr. R. Martin, professor of

anatomy in the university at Giessen. — Dr. Franz Wilhelm Negr, docent

for botany in the university at Munich. — Dr. Paul Parnuntier, associate

professor of botany in the university at Besancon. — Dr. Paulke, docent for

geology in the university at Freiburg i. B. — Dr. W. H. Piersol, instructor

in biology and histology in the University of Toronto. — Dr. F. Ptieninger,

docent for geology and paleontology in the university at Tübingen. —

Dr. Ernst, Freiherr Stromer von Reichenbach, docent for paleontology and

geology in the university at Munich. — Dr. Adalar Richter, professor of

botany in the university at Klausenberg. — W. M. Smallwood, associate

professor of zoólogy in Syracuse University. — Dr. Srdinko, docent for

histology and embryology in the Bohemian University at Prag. — Dr.

Studniczka, docent for zoólogy and comparative anatomy in the Brünn

Technical School. — R. B. Thompson, assistant in botany in the University

of Toronto. — Dr. Wilhelm Trabut, professor extraordinary of mineralogy

in the university at Vienna. — Dr. Karl von Tubeuf, head of the biological

division of the Imperial Health Department of Germany. — Dr. M

Wadsworth, geologist for the Pennsylvania State Board of Agriculture. —

enry L. Ward, custodian of the Public Museum at Milwaukee. — Dr.

Eugen Warming, director of the Geological Survey of Denmark. — Dr.

S. H. Westman, assistant in histology in the University of Toronto.

RETIRED.

Dr. Chr. Aurivillius from the charge of the entomological collections in

the Royal Swedish Museum. — Professor W. H. Brewer from the chair of

agriculture at Yale after thirty-seven years of service. — Dr. E. Bugnion

from the chair of anatomy in the university at Lausanne. — Francois Crépin

.. from the directorship of the botanical gardens at Brussels. — Miss Susan

M. Hallowell from the chair of botany at e arg College. — Professor

R Sadebeck from the directorship of the Botanical Museum at Hamburg.

— Professor E. B. Tylor from the keepership of the University Museum,

Oxford.

No. 425.] GIFTS, APPOINTMENTS, RETIREMENTS. 419

DEATHS.

Mr. Alfred W. Bennett, the well-known English botanist, January 23,

aged 69. — Giuseppe Camillo Giordano, professor of natural history in the

Technical Institute at Naples, November 17. — Professor Alpheus Hyatt,

at Cambridge, January 15, aged 63. — Professor Axel Key, anatomist and

bacteriologist, at Stockholm, December 27. — Dr. Johannes Christoph

Klinge, head botanist of the St. Petersburg Botanical Gardens, aged 51. —

J. H. Krelage, botanist (Liliacez), in Belgium, December 1. — Professor

Ivan Muschketoff, geologist of the St. Petersburg Mining Institute, Janu-

ary 25. — C. L. A. de Nicéville, state entomologist of India, at Calcutta,

December 3, of malarial fever. — Charles Roberts, British surgeon and

naturalist, January 8. — Dr. E. Selenka, professor of zoólogy in the univer-

sity at Erlangen, January 20, aged 6o. — Flaminio Bandi de Selvi, ento-

mologist (Coleoptera), at Turin. — James P. Shipman, local geologist at

Nottingham, England, November 21, aged 53. — Dr. Charles Stuart, an

English naturalist. — T. T. T. Thorell, arachnologist, at Helsingborg,

Sweden, December 23, aged 71. — P. C. Truman, entomologist, at Voca,

So. Dakota, October 27.

(No. 424 was mailed April 24.)

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| VoL. XXXVI, No. 42627

THE

MERICAN |

NATURALIS

A MONTHLY JOURNAL qp e

DEVOTED TO THE NATURAL SCIENCES =

IN THEIR WIDEST SENSE EK EE

= =e CONTENTS

E -The Life Hi: ES y f Ul 1s hy li Sar Seka :

; II. The Nemertean Parasites of Crabs . . .

< — dp ona iaa] Origin of the Cnieted apogei

prs Pueri ed” ing et €

The American Naturalist.

ASSOCIATE EDITORS:

merican Museum

> A. ALLEN, PH.D of Natural History, New York.

ANDR

| ambridge.

; » University of Michigan, Ann Arbor.

IN Š SEE SD., U.S. Department Agriculture, Washington.

HARD STEJNEGER, Smit, Washington.

chao raska, me

v Na List is an illustrated monthly magazine

cand will aim to present to its jas the atop

| 1eral

IHE

AMERICAN NATURALIST

Vor. XXXVI. June, 1902. No. 426.

THE LIFE HISTORY OF ULULA HYALINA

LATREILLE.

J. F. McCLENDON.

WnirE collecting insects in Galveston, Texas, during June,

1900, I found specimens of U/u/a hyalina Latreille, and at the

suggestion of Dr. Wheeler kept them alive for the purpose of

obtaining their eggs and following their larval and pupal devel-

opment. The insects would fly against the sides of the large

glass jar in which they were kept, and finally killed themselves ;

but two females deposited their eggs before they died. After

many of the eggs and young larvae were destroyed by various

accidents, I finally succeeded in raising two larvae. One of these

I preserved when full-grown, the other after it had pupated.

Later I obtained several imagoes from different localities and

one full-grown larva from Austin, Texas. After reading the

few notes that have been published on the life history of this

interesting insect, I concluded that a more thorough treatment

of the subject would not be out of place.

The first notes on the life history of the Ascalaphide, the

family to which our insect belongs, were published in 1826

1 Contributions from the Zoólogical Laboratory of the University of Texas, No. 27.

422 THE AMERICAN NATURALIST. [Vor. XXXVI.

by Guilding, who is perhaps best known to zodlogists as the

discoverer of Peripatus. These notes were on the life history

of Ulula macleayanus Guilding,! which is synonymous with

Ulula hyalina, or a variety of it, occurring in the island of

St. Vincent, West Indies, where Guilding obtained his specimens.

His description is meager and, to some extent, erroneous ; but

the arrangement of the repagula has not been observed subse-

quently, to my knowledge. Hagen published a paper? in 1873,

L] LER

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Fic. 1. — Ulula hyalina Latr, Male.

Latr

which contained short descriptions of sixteen species of Asca-

laphidæ. He repeated Guilding’s descriptions and also described

the larva of Ulula senex Burm., ignoring the fact that it was

synonymous with Ulula hyalina and, consequently, also with

U. macleayanus. Westwood wrote a review in 1888 of the

previous papers on the life history of the Ascalaphidz? and

added a short life history of a Ceylonese species, perhaps

Ascalaphus insimulans Walk., accompanied by figures, but

i n Genus Ascalaphus, Transactions Linnean Society, vol. xv, p. 509.

Larven von Ascalaphus, Stettiner entomologische Zeitung, Jahrg. xxxiv

* Notes on the Life History of Various Species of the Neuropterous Genus

Ascalaphus, Transactions Entomological Society, London, 1888, pp. 1-12, Pls. I, II.

No. 426.] ULULA HYALINA LATREILLE. 423

failed, as did Guilding and Hagen, to work out the mouth parts

thoroughly. A good description of the ant-lion’s mouth parts,

which are very similar to those of the ascalaphid larva, may be

found in a paper by Redtenbacher!as early as 1884. Hagen

and Westwood do not appear to have been familiar with this

paper.

Ulula hyalina is distributed over the southern half of the

United States, Mexico, and the West Indies, but is compara-

tively rare in many of the places where it is found. The insect

(Fig. 1) when at rest remains motionless

on some small branch or stalk, head down,

with wings and antennz closely applied

to the branch, and abdomen erected and

often bent so as to resemble a short

brown twig or dried branch. On being

approached, the insect moves to the oppo-

site side of the branch, and, on being

further disturbed, flies to another branch

and alights with head up, then quickly

turns and assumes its characteristic atti-

tude. I found specimens on stalks of

green sedge near the beach at Galveston,

Texas. The insect contrasted strongly

with the green stalk, but there were,

near the tops of the stalks, brown Ni. uM Feu

seeds which resembled the insect and iie, resting on a stalk of

made it hard to find. gun mp.

Guilding says the eggs (Fig. 3) are placed in double series

of 64 to 75, near the end of a branch, and are fenced off by

little rods, which he called *repagula" (Fig. 3), placed on end

and arranged in circles around the branch below the eggs,

thus preventing the approach of insects and the wandering

abroad of young larvæ until they can climb over the repagula

and have likewise acquired strength enough to resist ants and

other insect enemies. I observed one of my specimens from

Galveston deposit its eggs and repagula; but it was too weak

1 Uebersicht der Myrmeleoniden-Larven, Den£schriften der Kaiserlichen Aka-

demie der Wissenschaften, Bd. xlviii, Taf. VII. Wien, 1884.

424 THE AMERICAN NATURALIST. [Vor. XXXVI.

from confinement to remain on the stalk on which it rested

and fell to the ground, so that I could not tell how it would

have arranged the eggs under normal circumstances. These

repagula I find to be in all probability abortive eggs, since

dissection shows that some of the tubules of the ovary pro-

duce eggs, and others repagula.

The eggs hatch after nine to ten days. The young larva

remains quiet a day or two, after which it seeks the ground.

The larva, while growing in size, always

retains the form shown in Fig. 4, except that

after hatching and after each moult the head

is proportionally larger. It hides in some

slight depression or under the edge of a

stone, with its body covered with sand and

its mandibles widely extended so as to touch

the fringe of hairs on each side of the head.

Its brown color simulates the surroundings.

Its body is hidden by the covering of sand,

and the head is somewhat concealed by its

f peculiar covering of hairs (Figs. 4, 5, 12), so

? 1 that small insects may crawl, unawares, too

yf near the extended mandibles, In this case

ii the larva thrusts out its head and snaps the

Fic.3.— Ulula hyalina, Mandibles together, pinioning the victim on

cade. the curved points. It then proceeds to suck

the arrangement of the Out the juices of its prey like an ant-lion. In

ber itii the latter this is accomplished, according

gear peut ad to Redtenbacher, by the expansion of the

~ pharynx, the juices passing through the

duct formed by the mandible and maxilla fitting together

(Fig. 6). The wound is kept open by the maxilla working like

a piston in the groove of the mandible. The palpi at the same

time move back and forth slightly. The labium and ligula

are folded back into the mouth and adhere together so as to

close the orifice in front. I have observed that the Ulula

larva soon kills its victim, and at intervals opens its mandibles

slightly, until one of them comes out, and then sticks it into a

new place. This is continued until the skin is sucked dry,

No. 426.] ULULA HYALINA LATREILLE. 425

when the larva throws it aside and assumes its characteristic

attitude and awaits another victim. On being disturbed the

larva crawls away and seeks some other retreat. It often

changes its hiding place at night, probably on account of

scarcity of food. It always walks forward, contrary to the

habit of antlions. The larval life lasts about sixty-two days,

during which time the larva moults twice. It moults a third

time inside of the cocoon, when it changes to the pupa.

As the habits of the Ulula larva are somewhat peculiar, it

becomes of interest to compare them with the habits of the not

very remotely related ant-lions (Myrme-

leonida). According to Redtenbacher,

the Myrmeleonidz (Formicaleo, Acan-

thaclisis, and perhaps Palpares) lie

quietly during the day like U/w/a

hyalina, except that the body, instead

of being simply covered with sand, is

buried slightly beneath the surface.

They can walk backward as well as

forward. At night they wander about

in search of prey. Myrmeczlurus can

walk forward as well as backward, but

digs a pit like the ordinary ant-lion.

These latter, however, never walk for-

ward. The digging of a pit by the

ant-lion may be but a step removed

from the habit of Palpares in burying

its body. The ant-lion also has the Fic. 4.— Utula hyalina. Full-

instinct of wandering at night in pnm

search of a more favorable situation when food is scarce.

When the Ulula larva is full-grown it seeks some hidden place

at night in which to pupate. Having found such a place, it spins

a web, covering it with sand and such other small objects as

may be at hand. It then gets inside the web and begins spin-.

ning a cocoon. The next day it remains quiet, and at night

continues the work. I had an opportunity of observing several

stages of the process, as my specimen spun its cocoon against

the side of a glass jar partly filled with sand. The side next

426 THE AMERICAN NATURALIST. [Vor. XXXVI.

to the glass could be seen through until after the third night,

so that it must have required more than three days to con-

struct the cocoon.

The mandibles (Fig. 15, zz) of the pupa are toothed on their

inner edges for the purpose of enabling it to bite a hole in the

cocoon when the metamorphosis has been completed. I did

not have an opportunity to observe the escape of the imago.

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. Figs. 5-1 t. — Ulula Ayalina. Larva. Fig. s, three small setæ seen in optical section and a

f ng one, surface view; Fig. 6, the head of a larval skin of the first moult,

ith

E m U

c, clypeus; 7 labrum; a, mandible; »', maxilla;

2 seb ?; »'", mentum ; 2, labial palpus ; 4, point of contact of mentum and clypeus ;

, lobe of th n; y, lobe of the gena ; o, ocular peduncle

Below I have added descri

ptions of the egg, repagulum, larva,

pupa, and cocoon. i

Egg and Repagulum.

. The repagula are abortive eggs. Some tubules of the ovary bear eggs,

According to Guilding the eggs are placed in a double alternating series

of 64 to 75, near the extremity of the branch, and the repagula (barriers)

are placed in circles around the branch, below the eggs (Fig. 3).

No. 426] ULULA HYALINA LATREILLE. 427

Larva (Fig. 4).

Length 13 mm. Head cordate, broad behind, tapering anteriorly, swollen

beneath, thicker behind than in front, fuscous varied with black, covered

with hair except on mid-ventral line, deeply emarginated on posterior border

above, leaving an angular projection in center of emargination. Anterior

border fringed with serrate

beset at the end with serrated

hairs, two of which are very uti.

large and inclined backward. B

Eyes black, six on upper sur- 4 iva

face of ocular peduncle, five 4

forming an incomplete circle Wow

around the sixth, one on under 4

surface of ocular peduncle near

the posterior outer margin. F16. 12——U7u/a hyalina. menaa rana NEG 0e

ular peduncle ; 2, antenn:

Antenna (Fig. 12, a) 1% times

as long as ocular peduncle, basal segment very large and broad at base,

second segment much smaller, third still smaller and proportionately much

Shorter, and followed by a piece not definitely segmented and still more

slender; the succeeding nine segments are of nearly equal size, the terminal

segment is as long as the three preceding and ends in a tuft of small hairs ;

there is a lobe above the base of the antenna

ending in a fringe of five hairs, — three stout

serrated ones alternating with two slender smooth

ones which terminate in stellate enlargements.

Clypeus (Fig. 9) narrow behind, continued in front

as a pair of semicircular lobes over bases of man-

Ü Fe. 13.— U?wla dibles, middle of anterior border continued down-

t” Agalina, Meo ward and reaching the mentum. Labrum (Fig.

larva: x, mesothorax; 5, an` 10, 2) small, infolded into the mouth. Mandible

ticular membrane; c, coxa;#, — long, swollen at base, straight for three-fourths its

z", tms me length, then curving inward; curved portion free

from hairs, inner edge set with three teeth, middle

one largest, posterior one smallest, space between anterior and middle one

smaller than that between middle and posterior one, mandible grooved on

ventralside. Maxilla swollen at base, lying in. groove of mandible, with which

it forms a duct leading to the mouth, edge near tip. Mentum

(Fig. 6, m”) nearly square, anterior border produced in the middle so as to

428 THE AMERICAN NATURALIST. [Vor. XXXVI.

reach the clypeus, lateral margins produced into a pair of triangular lobes

(Fig. 11, x) inclined forward and devoid of hairs, each divided by a suture

into two triangles; ligula infolded into mouth, adhering to the labrum;

basal segment of labial palpus very large, flattened, fourth segment as long

as second and third combined. On each side of the mentum arises a lobe

of the gena (Fig. 11, y) which projects over the base of the mandible,

deeply emarginated on anterior border. The hairs (Fig. 5) on the head,

as well as on the body, are of peculiar structure: the base of a hair is con-

stricted so as to close the cavity within, and the integument is raised around

the constricted portion; the hairs are of two kinds, smooth and serrated ;

some of the serrated hairs are enlarged at the tip.

Thorax flat, much broader behind than in front, luteo-fuscous mottled

hairs, all of which are fuscous or black;

prothorax freely articulated with meso-

thorax, much narrower than head, much

broader than long; metathorax broader

_ than head, bearing above three ellipsoid

swellings, each of which has the margin

depressed below the general level; one lies

near the anterior border, the other two near

the lateral borders and are connected by

a deep groove; two pairs of lateral lobes,

anterior pair short, conical, inclined for-

ward, terminating in a few black setae which

are packed together so as to appear like

the end of the lobe, second pair very long,

inclined forward at base, then bent slightly

backward, fringed with large, serrated hairs; there is a pair of spiracles below

ar t 1 TA 1 J aL +h,

£141 x

4 d

Fic. 14. — Ulula hyalina. Pupa,

p air of lobes; | ,

a pair of ellipsoid swellings above, similar to those on mesothorax, a pair of

lateral lobes, shorter than pair on mesothorax, inclined forward, fringed

with large hairs; a pair of spiracles below, near bases of lateral lobes ; legs

luteo-fuscous with fuscous hair, anterior pair small, each succeeding pair

larger, coxa very long (see Fig. 13),! trochanter small and almost rigidly

attached to the femur, tibia slender, tarsus of a single small joint, ungues

black, much curved.

Abdomen broad and thin, dorso-ventrally compressed, pointed behind,

lateral margins very convex, luteo-fuscous mottled with fuscous, covered

with hairs which vary from fuscous to black, flattened and wrinkled below,

. ! Redtenbacher (/oc. cit., Fig. 116) gives a different interpretation to the joints

of the leg of the antion. He calls coxa what I believe to be a much-developed

articular membrane (Fig. 13, y), and what I have called the trochanter he regards

as part of the femur. The articulation between the trochanter and femur is not

well developed and allows very little movement, but it cannot be overlooked. I

may add that my i ion i ed

pretation is based on comparison with the imago.

No. 426.] ULULA HYALINA LATREILLE. 429

somewhat convex above, divided into nine segments ; tergites, except last

two, separated along middle, so as to disclose the articular membranes, —

each tergite, except the last two, with a cross groove nearly reaching the

lateral borders; each segment bears a pair of lateral lobes fringed with

fuscous hairs; each segment also bears a pair of spiracles below, near the

bases of the lateral lobes; last segment conical, truncated. There is a

circle of black curved spines around anus.

Pupa (Fig. 14).

Length r2 mm., diameter in cocoon 6 mm., breadth of abdomen 4 mm.

The pupa resembles the imago in general, but is much smaller and com-

paratively shorter.

Head short, and compressed against thorax; eyes fuscous, with a deep

vertical fold at right angles to the sulcus; antenne comparatively short,

Fic. 15. — Ulula hyalina. Mouth of pupa from below: Z, labrum; , mandible ;

Ż, maxilla; 2’, maxillary palpus ; 7, labium; ’, labial palpus.

curved backward over the head, not knobbed, white; face light yellow,

clypeus not distinct from labrum; mandibles (Fig. 15) stout, armed each

with nine to we instet t inner edge of mandible rufo-fuscous, teeth black ;

gular region lla bilobate, maxillary palpus of three segments ;

labium a er: labial palpus slender, smaller at base than at

tip; mouth parts and vertex clothed with fine white hair.

Thorax short, cream-colored varied with ferruginous ; prothorax com-

pressed against back of head; wing sacs small, white; legs slender, white.

bdomen short, curved under so that the anus reaches the mouth, cream-

colored, varied with ferruginous, covered with fine white hair, a narrow

mid-dorsal groove ; each segment except the last two with a pair of spiracles.

Cocoon spherical, diameter 7 mm., made of silk.

UNIVERSITY OF TEXAS, AUSTIN,

ovember 20, 1901.

THE NEMERTEAN PARASITES OF CRABS.

"WESLEY KR. COE.

YALE UNIVERSITY.

In 1844 Kolliker found a small species of nemertean among

the egg masses carried about on the abdominal hairs of a ** small

crab" at Messina. This worm he named Nemertes carcino-

philos! and gave a fairly good description of its anatomy (45).

Fifteen years later van Beneden found a similar species among

the eggs of the green crab (Carcinus manas) on the coast of

Belgium and, having overlooked Kólliker's descriptions, rede-

scribed and figured it as Polia involuta (61). The worms were

said to be small, slender, yellowish or rosy in color, with two

ocelli and a very short proboscis, armed with central stylet only.

They lived in delicate mucous sheaths among the crab's ova,

and often two individuals — a male and a female — lay side by

side, or with their bodies somewhat folded, in the same tube.

Van Beneden decided that the worm was not a true parasite,

but rather that it merely occupied the egg masses of the crab

as convenient places for building its tube and depositing its

ova, as well as a place well protected and furnished with food.

The females were 2—3 cm. in length, although the males meas-

ured scarcely 1 cm. Some of the younger individuals were

only 2 mm. long. There are figures showing the form and

color of the living worms, several of anatomical details, of

the eggs in various stages of development, and of the early

embryos before and after the molting of the larval skin.

These figures are generally accurate, as is also the interesting

account of the development. The large nerves passing forward

from the dorsal ganglia are described as excretory canals open-

ing on the lateral margins of the head, although the author

states that they seem to be a continuation of the ganglia. The

1 Spelled cartinophilos.

431

432 THE AMERICAN NATURALIST. [Vor. XXXVI.

ciliated canals leading from the cerebral sense organs were in

other species of nemerteans looked upon as excretory canals.

The figures of the proboscis with its single central stylet and

large glandular masses on each side are quite characteristic.

McIntosh (73) gives a good description of the general anat-

omy of this species and an interesting account of its behavior

in confinement, as well as the method of deposition of its ova.

He also briefly describes the segmentation of the egg and

gives several figures of the developing embryos. There is a

good colored drawing of the living worm and figures illustrat-

ing the anatomy of the anterior portion of the body and of

the proboscis, all of which are described in detail.

In 1874 Dieck (74) found at Messina (where Kólliker's speci-

mens of N. carcinophilos were obtained) a number of nemer-

teans among the egg masses of Galathea strigosa. But Dieck's

description does not apply to any of the metanemerteans, and

the species was named Cephalothrix galatheg. The species

agrees fully with Kolliker's JV. carcinophilos in size, form, and

color of body, in not having the head demarcated from body,

in having two comma-shaped eyes in front of the ganglia, and

in the absence of cerebral sense organs and cephalic furrows.

In all these respects the description exactly corresponds with

that of JV. carcinophilos, but Dieck describes certain other

anatomical peculiarities which separate the two forms widely,

and certain others which have been found in no other nemer-

tean. The proboscis is described as being without stylets,

and the mouth is said to lie behind the brain. Dieck fur-

ther describes remarkable appendages on the head which are

believed to aid the worm in retaining its position on the crab.

These *fingerfórmige Greif oder Haftorgane . . . sind so con-

tractil, dass sie nur bei starker Ausdehnung deutlich ins

Auge fallen " (74, p. 502) ; but he shows no indication of them

in any of his figures. It seems possible that they may have

been formed by a too severe pressure on the glass covering

the worms, and thereby rupturing the integument. Dieck

thinks he may have seen similar appendages on the posterior

end of the single male which he found, but he could not be

certain whether he saw them or not.

No. 426.] MWEMERTEAN PARASITES OF CRABS. 433

He also describes the ovarian pouches as opening ventrally,

and states that their external openings are provided with con-

tractile lids which serve to close the openings, except when

the ova are being extruded. Both Dieck and van Beneden

speak of the minute white specks seen on the bodies of the

females with ripe ova, while Dieck’s observations on the

appearance of the egg strings, the eggs themselves and their

early development, as well as the appearance and peculiarities

of the embryos, answer equally well for Kolliker’s species ; and,

finally, his statement that the worms, after having devoured

the eggs of the crab, find their way to the gills, where they

live as ectoparasites, is in perfect accord with the observations

on carcinophila described below, although his interpretation of

the life history of the worms seems to be in error.

While it would be unwarranted to say that Dieck’s descrip-

tions of the anatomy of his Cephalothrix galathee are incorrect

in so far as they do not agree with the structures found in

Kolliker's species, yet it would seem most remarkable if

there were to be found in the same locality two species of

nemerteans belonging to entirely different orders which agree

so perfectly in color, size, external appearance ; in possessing

the same peculiarities in regard to the ocelli, structure of

body walls, mucous glands, and other features; in baving the

same peculiar habits due to a parasitic life; in laying eggs

which have the same appearance and mode of cleavage, and

which in their development give rise to perfectly similar

embryos. Likewise remarkable among the nemerteans,

although it is known in a few forms, is Dieck's observation

that while most of the eggs are laid before cleavage, yet ferti-

lization and early cleavage sometimes take place within the

body of the parent. McIntosh states that this is also true in

N. carcinophila. It should be noted in this connection that

Dieck found but a single male, which was 2 cm. in length,

although the females were abundant and of much larger size,

some of them being as much as 7 cm. in length. He con-

sidered it probable that the sperm from a single male entered

the ovaries of all of the five or six females which might live on

the same crab.

434 THE AMERICAN NATURALIST. [Vou. XXXVI.

Whether the species described by Kölliker, van Beneden,

and McIntosh will prove to be widely different, closely related,

or identical with that studied by Dieck, future investigations

must decide. Suffice it to add that Dieck was quite unac-

quainted with the descriptions of any of the other authors

mentioned, and believed that he was describing a crab para-

site for the first time. He shows by his descriptions that he

knew very little of nemertean anatomy, and this fact might

easily account for the strange anatomical peculiarities which

he found the worms to possess. Careful comparison of his

figures with those of van Beneden (61) and McIntosh (73)

reveals such slight differences that we may well consider

them as belonging to the same species; that they represent

worms of different orders seems incredible.

Joubin (93) has found Kolliker’s species abundant on Car-

cinus manas at Roscoff, on the northern coast of France, and

Giard (88), on the coast bordering the Bay of Biscay, on the

same species of crab. Giard (p. 496) found these worms on

almost every crab examined which carried eggs, and has

observed the same or a very similar species on Xantho

Jforidus. This he calls Polia xanthophila, but gives no dis-

tinguishing peculiarities except that it is smaller in size.

On the Challenger Expedition Willemoes-Suhm (74) found

a small species of nemertean on the body of a species of crab

(Nautzlograpsus minutus) which lives on the gulf weed in the

“Sargasso Sea,” between Bermuda and the Azores. These

little worms were only about 1-2 mm. long, but none of them

were sexually mature. The suggestion was made that the

nemertean inhabits the crab only when young, and that it lives

freely on the gulf weed when mature, This form agrees with

the other species of Carcinonemertes in possessing two large

ocelli, although there is said to be a pair of accessory ocelli of

extremely minute size near the proboscis. These are repre-

sented in the figure (74). It is intimated also that the mouth

lies behind the brain. The color of the worms was brownish.

The stylet lies just back of the brain, and the proboscis is

~ Very short. The figure shows indications of two pouches of

_ accessory stylets, although these are not mentioned in the

No. 426.] WEMERTEAN PARASITES OF CRABS. 435

descriptions. Some of the worms are said to be found on —

various parts of the crab’s body, being most abundant on the

abdomen, but the gills are not mentioned. An account of

this species is given also in the WVarrative of the Cruise of the

Challenger" (Vol. I, p. 169) and is reproduced in Hubrecht’s

report on the nemerteans (87).

A further instance of a nemertean parasitic on Crustacea is

mentioned by Quoy and Gaimard (33), who describe from

Amboina a small species, apparently belonging to the genus

Tetrastemma, under the name of Borlasia quadripunctata.

They state that although it usually lives in the sea, yet it

is also found at times living in the barnacle (Anatifa).

So far as I know, the first observations of a nemertean para-

site of the crab in this country are those by Prof. J. P. McMur-

rich, who writes me that he found the worms quite numerous

among the egg masses attached to the abdominal hairs of a

“lady crab" (Platyonichus ocellatus) July 6, 1889. These

worms laid several strings of eggs in the manner characteristic

of the species. The eggs developed normally, as described by

van Beneden ('61), McIntosh (73), and Dieck (74).

I first found this nemertean on the gills of the lady crab at

North Dennis, Mass., situated on Massachusetts Bay, Aug. 6,

1898, and have since found it abundantly in other regions

south of Cape Cod.

The gills of a single crab often harbor as many as forty to

sixty of these worms, which are of small size and sexually imma-

ture when found in this position, becoming full-grown and sexu-

ally mature only when living among the crab's ova. As found

on the gills the worms vary in color from pale ocher to salmon,

depending largely on the coloration of the intestinal canal.

They are of all sizes up to about 15 mm. in length when

extended and very slender — almost threadlike. The body is

of about the same diameter throughout and is often folded once

or twice on itself as it lies between the gill plates of the crab,

imbedded in considerable mucus. The crab's gills are some-

times undoubtedly injured by the abundance of the worms, so

that I am almost sure that the latter feed upon the blood in the

gills. This was also Dieck’s conclusion in regard to his

436 THE AMERICAN NATURALIST: [Vor. XXXVI.

Cephalothrix galathee, where he believed that he found the crab's

blood corpuscles within the intestinal canal of the worms.

In many of the crabs on which I found only a small number

of the parasites no injury to the gills was apparent, the ability

on the part of the crab to repair its injured tissues being

sufficient to prevent the gills being destroyed. When the

worms were abundant, however, some of the gill plates were

blackened, torn, and degenerated.

A large number of lady crabs obtained from Woods Hole

and Martha’s Vineyard, Mass., in July and August, 1900, were

examined, and the parasitic nemerteans found in the gills in

about one-tenth of the number. When present on one side of

the body they were almost always found on the other side also.

I found them only on female crabs, and did not find them in

the gills when the crabs were carrying eggs. The worms

cling tenaciously to the gills when these are removed from the

crab. A gill cut from a crab and placed in a dish of sea water

is not deserted by its parasites for several days, — or not until

it decomposes. When forcibly removed from the gills the

worms live several weeks in sea water, crawling sluggishly

about, and often collecting in masses with their bodies placed

lengthwise or folded sharply so that the anterior portion of the

body lies parallel and in contact with the posterior portion,

exactly as when living in the gills. Masses of them often col-

lect on the surface of the water where it comes in contact with

the sides of the vessel.

They secrete a very sticky mucous covering and when touched

with a needle or drawn into a pipette attach themselves to it

with exasperating tenacity. After living several weeks without

_ food the worms become much smaller than when first collected.

. Several dozen spider crabs (Libinia canaliculata), some of

which carried eggs, a number of blue crabs (Callinectes hasta-

tus) with eggs, and many rock crabs (Cancer amenus) and green

crabs (Carcinus manas) without eggs were examined without

finding any nemerteans. I quite expected to find them on the

No. 426.] NEMERTEAN PARASITES OF CRABS. 437

body, and a portion of them will be found among the gill plates

after twelve to thirty-six hours, but many more remain clinging

to the crab’s legs. Their behavior to lady crabs, however, —

males as well as females, — is quite different, and they quickly

find their way to the gills. The few rock, spider, and blue crabs

with which I experimented in this way did not prove attractive

to the worms, although occasionally one of them would enter the

gills. Far more of them remained `;

attached to the basal joints of the

legs for several days.

Sexually mature worms may be

found early in July (and probably

also in June) on the lady crabs

carrying eggs. But the crabs with

eggs are shy or else frequent deeper

water, so that I did not find them

nearly so abundant in the localities

noted as were the males and smaller

females without eggs.

All my observations on the p. — Carcinonemertes epialti. Ante-

spegies seeni to indicate Idt ihe e podm < botret ione vem sa

worms spend their whole existence matic. »,rhynchodzum; o, ocellus; xv,

on the crab, for I have found them oreli and other organs of head ; s central

commissure

i stylet; őr, brain; dc, dorsal

I Dru 2 ages of development of brain; d. cesophagus; P, pate

from the egg to the sexually mature chamber o iet i, intestine; Zn,

lateral nerve.

worm. I have not, however, fol-

lowed their history during the winter months, but suspect

that this time is occupied as a period of slower growth.

_ The worms apparently occupy nearly a year in attaining

sexual maturity, and their life history is briefly as follows:

Eggs laid in mucous tubes among the egg masses of the crab

in June and July ; cleavage regular and nearly equal, with the

formation of free-swimming ciliated blastula which develops

into a ciliated embryo provided with ventrally placed mouth, a

pair of ocelli, and an anterior and a posterior flagellum, or tuft

of much longer, consolidated cilia. The embryo leaves the egg

membrane in this condition and usually remains in the mucous

tube or among the egg masses of the host, but may swim freely

438 THE AMERICAN NATURALIST. (VoL. XXXVI.

in the water. The larval integument with its cilia and flagella

is apparently shed, as described by van Beneden (61) and

Dieck (74). At this time the embryos assume the form of the

adult and crawl about instead of swimming. The integument

of the young worms now becomes covered with cilia, as in the

adult. After remaining for a time among the egg masses of

‘Siti Pee

prit. ut

"n -£

! 1

Fic. 2. DU agone, section through anterior portion of body of C. eren Trees. diagram-

tic, and the proboscis is represented a di too large. cg. kan esi

Ex t var ": me, £c, anterior, middle, and posterior icc n

tively; 47, brain; z, intestine. x 150.

the host, or perhaps until her eggs have hatched, they wander

about on her body, eventually reaching the gills. They are

found in this position in July or August, and later, and here

they probably remain until the crab produces another batch of

eggs the following season. At this time they migrate again to

_ the egg masses, where they become sexually mature. Those

No. 426.] NWEMERTEAN. PARASITES OF CRABS. 439

embryos which swim away and which do not chance to find

another suitable crab probably perish. The observations of

the European writers mentioned above

are mainly in accord with the account

as here given.

The mature worms often become

25 mm. or more in length, are generally

bright reddish orange in color, but some

are reddish ocher and others brick red.

Their anatomical details agree closely

with such descriptions of the European

species as have been given, although

these descriptions refer mainly to the

external features. McIntosh’s colored

drawing (Pl. I, Fig. 5) of the worm, his

figures of the anterior portion of the

body and of the proboscis, as well as

his detailed description of these parts,

agree in most respects with the New

England form, except that I find the

posterior, glandular portion of the pro-

boscis (Figs. 2—4, 6) much shorter than

McIntosh represents. Yet it seems

highly probable that such differences as

appear to exist are largely accidental,

and that the New England form is

specifically identical with Kolliker's

Nemertes carcinophilos. Joubin (93) eant dose ds Mid

also gives a colored figure of the worm moved from the worm. ac, mc,

H x : č i fe, anterior, posterior, and middle

as it lies folded in its mucous sheath ‘Gambers respectively; z, gland

i cells; e, canal connecting anterior

TE the egg decus of the crab. and middle chambers; fs, rem-

In internal organization the worms © nants of proboscis sheath attached

: A a o terio: ber; t connec-

agree closely with a second species which- ripen da vd eir due

I found abundantly on the gills of another berisimbedóed ; 4, “basis of central

vow x 300. ;

species of crab (Epialtus productus) at

Monterey, California, and which is described in detail below:

. Both the Atlantic and the Pacific forms show such wide

deviations from all other species of the genus Eunemertes, in

440 THE AMERICAN NATURALIST. [Vor. XXXVI.

which Kolliker’s form has been placed by Joubin, Biirger, and

others, that the establishment of a new genus is imperative.

Carcinonemertes gen. nov.

Parasitic nemerteans living on various species of Crustacea.

Body small, slender, often filiform, rounded, and of about the

same diameter throughout; head without distinct lateral

grooves, not demarcated from body. Body not usually coiled

or much twisted, but often folded sharply so that the anterior

portion of body lies parallel and in contact with the posterior

portion. Mouth and proboscis open together; cesophagus

extremely short, opening broadly into the intestine through

a large muscular chamber situated immediately behind the

brain (Fig. 6); intestine broad, with short lateral pouches

which are but little developed in posterior portion of body.

Proboscis sheath without muscular walls, consisting merely

of a thin membrane closely applied to the small proboscis.

Proboscis but little developed, very small in size, and extremely

short, without lateral pouches of reserve stylets, but armed with

central stylet and basis only. Central stylet minute, usually

one-third to one-half as long as basis, which is small and slender.

Stylet region of proboscis can be withdrawn but little behind

brain; consequently anterior chamber is very short, without

distinct muscular layers, without distinct nerves, and without

a thickened glandular epithelium such as occurs in almost all

other nemerteans. Stylet apparatus imbedded in a strong

muscular enlargement provided with numerous large glands

(Figs. 2-4). Chamber immediately behind stylet, small but

muscular, and with a lining of flattened epithelium, while

the posterior proboscidial cavity is very short, often almost

spherical, highly glandular, connected closely with the rudi-

ments of the proboscis sheath and imbedded in the connective

tissue which lies internal to the body musculature.

Cerebral sense organs probably wanting. Ocelli 2 (occa-

sionally fragmented into 4).

Cephalic glands massively developed ; a remarkable develop-

ment of submuscular glands extends throughout the whole

No. 426.] MNWEMERTEAN PARASITES OF CRABS. 44I

length of the body, usually forming a distinct layer internal to

the muscular walls of the body, and often thicker than all the

other layers of the body wall combined.

Body musculature consists of a thin, oblique or circular mus-

cular layer and a somewhat thicker, but yet weak, longitudinal

layer internal to the former.

Brain and lateral nerves as in other metanemerteans.

Usually oviparous, though fertilization often takes place

internally, and sometimes a portion of the ova of an individual

Fic. 4.— Carcinonemertes epialti. Horizontal section of proboscis in its natural position,

showing the posterior chamber lying at right angles to the general axis of the proboscis.

Reference letters as in Fig.3. x 300

may be retained in the body until the development of free-

swimming embryos. Development without complicated meta-

morphosis, although the layer of ciliated cells originally covering

the embryo is shed as development proceeds.

Carcinonemertes carcinophila (Kölliker).

Body slender, commonly 6-15 mm. long when found on gills,

20-70 mm. long when sexually mature; color yellowish orange,

442 THE AMERICAN NATURALIST. [VoL. XXXVI.

pale reddish, rose pink (McIntosh), or bright brick red; posterior

proboscis chamber very small, rounded; in ordinary states

of contraction central stylet lies immediately behind brain.

Basis of central stylet slender, about .025—.03 mm. in length

by .006—.008 in average diameter. Central stylet about .008—

012 mm. long, or between one-third and one-half as long as

basis. In general anatomical features the

species closely resembles C. epialti, which

is described in detail below.

Parasitic on the gills of various species

of crabs when young, migrating to the egg

UU BU masses of the crab at the approach of

ey voril ii toria sexual maturity, the young returning to the

TUM um gills after a short period of development.

Distribution: Mediterranean Sea, Bay of Biscay, English

Channel, on Carcinus manas; both north and south of Cape

Cod, Mass., on Platyonichus ocellatus; France, Bay of Biscay

(the same or a closely related species) on Xantho fforidus;

Mediterranean Sea (possibly the same species), on Galathea

strigosa.

Carcinonemertes epialti sp. nov.

This is a much smaller and less slender species than the above

when sexually mature, and differs from it in regard to the size

of the posterior chamber of proboscis, in the stylet apparatus,

and in many other anatomical details, although the differences,

as will be described below, are not very considerable.

In general appearance, in color, arrangement of ocelli, cesoph-

agus, intestine, and brain the two species are very similar.

C. epialti also lives when sexually mature among the egg masses

of a crab — in this case Epialtus productus, the common kelp

crab of the California coast.

Upwards of one hundred of these little worms were found

among the eggs of a single crab at Monterey, Cal., Sept. 3,

1901. In practically all, the sexual products were nearly mature,

but no eggs were laid in confinement. The worms lived only

a few days in a dish of sea water and appeared less hardy than

the species on the Atlantic coast. I was unable to determine

No. 426.) NEMERTEAN PARASITES OF CRABS. 443

whether the worms pass their early life on the gills of the crab,

as does C. carcinophila, but suspect that this may be the case.

The species may be described in detail as follows: Body

small, rounded, slender, of the same diameter throughout ; sex-

ually mature individuals about 4-6 mm. in length and less than

half a millimeter in diameter; head not demarcated from body ;

lateral grooves and cerebral sense organs very inconspicuous

or wanting.

Color, bright orange, sometimes inclining more to reddish

and sometimes to yellowish. Head a little paler, for the color

is largely due to the intestinal lobes which extend forward to

the brain.

A pair of ocelli of irregular outline, but sometimes crescent

shaped, lie about halfway between the tip of the snout and

the brain (Fig. 1). Sometimes the ocelli are irregularly frag-

mented, and the pigment is aps in four irregular masses.

REL) nem

27 EL.

Carp = EQ

N a Sr

2 oy ee

= Orr c

—— SAY ee Ce LIE)

/ LE IU

n 4X ams mud 4 d y 7

T eG te

E QAS A AS A t SS

-A 4 NI SS ee Xy: ASS

[ ERIT! EN ol Ne S

s o REL "n

ER ga ie TUNE

Fic. 6. — C. epialti. Oblique section through anterior portion of body. 7, opening of rhyncho-

dzum; cg, cephalic glands; ac, dor anteri or and posterior proboscis cha mbers; sg, sub-

muscular glands; vc, ventral co ssure.of brain; e, cesop ic, rudimen

intestinal caecum; 7, intestine.

Proboscis sheath greatly reduced, extending but little poste-

riorly to the brain, where it becomes united with the posterior

chamber of the proboscis (Fig. 3). The sheath consists merely

of few fibers of connective tissue supporting a very thin, flat-

tened epithelium, and can be seen only in favorable preparations.

444 THE AMERICAN NATURALIST. [Vor. XXXVI.

Proboscis very minute and short, extending scarcely more

than its own diameter posteriorly to the brain (Figs 1,°2;Gp

Rhynchodzum (Fig. 6, 7) slender; cesophagus separates from

proboscis just in front of brain (Fig. 6). Anterior chamber of

WZ,

————

Fic. 7.— C. epialti. Transverse section of body immediately back of brain. The posterior

chamber of the’ proboscis (c) is firmly imbedded in the surrounding connective tissue.

TL 1s f th y " * itat (i ) a cesophagus lined with cilia;

cm, Im, circular and longitudinal layers of muscles; sg, submuscular glands; Zx, lateral

nerve; zz, integument. x 200.

proboscis (Figs. 2, 3, ac) very small, not as long as the diameter

of a brain lobe, lined with thin, scarcely glandular, epithelium.

Stylet region swollen (Figs. 2-4) and provided with large

and abundant gland cells (g) which open both into the anterior

chamber and into the narrow canal connecting this with the

cavity behind the stylet region.

Basis of central stylet slender, about three to five times as

long as broad (Figs. 3-5), measuring about .027—.033 mm.

in length and .005—.008 mm. in diameter. Basis slightly larger

posteriorly than at attachment of stylet, often somewhat asym-

metrical, as shown in Fig. s. Stylet rather slender, a little

less than half as long as basis, measuring .012—015 mm. in

length. Basis imbedded among the gland cells and surround-

ing muscles in a single layer of columnar cells with oval nuclei

No. 426.] NWEMERTEAN PARASITES OF CRABS. 445

(Figs. 3, 4) at right angles to its longitudinal diameter. There

is no trace of accessory stylets.

The usual small, oval middle chamber lies directly behind

the stylet region and connects with the anterior chamber by a

canal (Figs. 3, 4) which passes close beside the basis of the

central stylet and which, though narrow, is broader than in

many other metanemerteans. The middle chamber, behind

the stylet, is highly muscular, lined with flattened epithelium,

and is often filled with fluid containing an abundance of gran-

ules resembling hardened secretions (Fig. 4). These, I think,

originate in the posterior chamber as described below.

The proboscis now bends sharply on itself in ordinary states

of contraction and ends in an oval chamber with small lumen

and very massive glandular walls (Figs. 2-4, 6). The cells

lining this chamber are highly columnar, irregularly arranged

QOL CD Ie

eee

AON

Fic. 8. — Transverse section of body of C. epialti, showing the thick layer of submuscular glands

(se) and the ovaries (ov) with large ova. The intestine (Z) is reduced to a narrow canal,

Other reference letters as in Fig. 7. x 200.

in several layers, and are thickly packed with secretions which

have great affinity for ordinary stains. This posterior cham-

ber is closely imbedded in the surrounding connective tissue

(Figs. 4, 7), and this appears to be connected with the muscular

446 THE AMERICAN NATURALIST. [Vor. XXXVI.

walls of the cesophagus. Its movements are doubtless to a great

extent dependent on the contractions of the cesophagus, which,

as described below, is converted into a sort of muscular pharynx.

The stylet can hardly be moved much beyond the external

opening of the rhynchodzeum, and from a study of its structure

alone it is hard to conceive how it can be moved for even this

short distance, imbedded as it is among the other tissues. By

crushing and many kinds of stimuli I have seldom been able to

cause the worms to move the stylet region to any extent either

forward or backward. It nearly always remained in the vicin-

ity of the brain, as shown in the figures. It is my opinion

that the proboscis can be everted only far enough to bring the

stylet a little beyond the opening of the rhynchodzeum on the

tip of the snout, as figured by van Beneden (61), and that

the cesophageal muscles aid in this movement. At the tip of

the snout the stylet can puncture the tissues and blood vessels

of the crab’s gills. With the rhynchodaeum of the worm widely

opened and closely applied to the point of puncture, the blood

and nutritive fluids exuding from the wound can be drawn

directly into the rhynchodzeum and thence into the cesophagus

by the contraction of the muscular walls of the latter.

The cesophagus, which leaves the rhynchodzum just in front

of the brain (Fig. 6), passes beneath the ventral commissure as

a narrow tube lined with rather flat cells, as in other genera.

Just back of the brain, however, it becomes enormously enlarged

with high, columnar, ciliated epithelium, richly provided with

gland cells. This portion of the cesophagus is highly muscular

and somewhat barrel shaped (F ig. 6), projecting a little way

backward into the broad intestine which immediately follows

posteriorly. Its posterior portion is therefore surrounded by

the intestine, as shown in Figs. 6, 7. Its opening into the

intestine is wide and has thickened lips. The backward and

forward motion of this barrel-shaped portion of the cesophagus

in all probability aids in the eversion of the proboscis, as well

as acts as a suction pump to draw in the nutritive fluids from

the crab's gills.

The intestinal lobes surrounding the end of the cesophagus

(Figs. 6, 7) indicate rudiments of the intestinal caeca found in

No.426.] WEMERTEAN PARASITES OF CRABS. 447

other genera. The intestinal canal is broad, with short lateral

pouches which become very much reduced towards the pos-

terior end of the body.

The nervous system shows few deviations from that in related

genera. The brain is fairly well developed as shown in Fig. 1.

From the dorsal lobes a pair of large nerves (Fig. I, zv) pass

anteriorly to the eyes and anterior portions of the head. These

are easily seen in living worms. I found no indications of

cerebral sense organs either when the specimens were stained

Fic. 9. — C. epialti. Transverse section of body showing the large number of spermaries (£)

and their distribution throughout the body. Reference letters as in Fig.7. x 200.

in toto or when examined in sections. I also failed in my

attempts to locate the efferent nephridial ducts.

Throughout the head the tissues are crowded with the cephalic

glands. Those situated more anteriorly open mainly on the tip

of the snout (Fig. 6, cg), but farther back they open directly

outwards on all sides of the body. Back of the brain they pass

gradually into the submuscular glands which extend as a dis-

tinct layer throughout the length of the body. The glandular

cells composing this layer open directly outward to the sur-

face of the body (Figs. 6, 7, 8, sg) and are situated on the

whole circumference of the body immediately internal to the

448 THE AMERICAN NATURALIST. | (VoL. XXXVI.

longitudinal muscular layer. The glandular layer is in most

regions so massively developed that it exceeds in thickness all

the other layers of the body wall combined. The secretions of

these glands furnish the sticky mucus by means of which the

worms cling so tenaciously to the crab or to other objects.

The outer epithelium is as in other genera, and is richly pro-

vided with glands.

The muscular layers of the body wall consist of a thin,

external circular or oblique layer of muscles and an internal

longitudinal layer (Figs. 7, 8), somewhat thicker than the former,

but yet thinner than in most related genera. The lateral nerves

occupy the usual places internal to the longitudinal muscular

layer. In this species, however, they lie internal also to the

thick layer of submuscular glands (Figs. 6, 8, 9, 7»), and there-

fore nearer the center of the body than in other genera where

these glands are not so highly developed.

There is very little body parenchyma, the intestine filling

most of the space internal to the glandular layer, except at the

time when the genital products are developing. The pouches

of genital products become enormously developed and encroach

greatly upon the intestinal canal at the time of sexual maturity

(Figs. 8, 9). The genital pouches extend much farther forward

than in almost any other nemertean, reaching very nearly to the

brain. The ovaries (Fig. 8, ov) are regularly paired, with a

single large pouch containing usually from 12 to 30 ova between

each pair of intestinal lobes. The spermaries, on the other

hand, are far more numerous, surrounding the intestinal canal

on all sides. As many as fifteen or more separate spermaries

(Fig. 9, /) are sometimes found in a single transverse section.

As in most parasitic animals the abundance of sexual products

is greatly in excess of that in related non-parasitic forms.

This is also well illustrated in Bergendal’s recent descrip-

tion (00) of Gononemertes, a nemertean parasitic in the tuni-

cate Phallusia. There is a resemblance also in other ana-

tomical features, — in the excessive development of the cephalic

glands, in the short posterior chamber of the proboscis, and in

the slight development of the intestinal cæca. Of the two

genera, Carcinonemertes appears to be far more degenerate than

No. 426.] WEMERTEAN PARASITES OF CRABS. 449

Gononemertes, although it still retains ocelli and has a central

stylet in the proboscis.

Summary. — The above observations seem to lead to the

following general conclusions: (1) that the nemerteans inhabit-

ing various species of crabs are distributed throughout the

North Atlantic and into the Pacific Ocean, (2) that the New

England form is identical with the long-known European

species, (3) that several European forms thought to be widely

different are either closely related or identical, (4) that a// the

species recorded show great similarity of structure, and may be

closely related, (5) that the worms are true parasites and are

not found except on the body of their host, spending practi-

cally their whole existence on the crab's body — in the gills

when young, on the egg masses when mature, (6) that in dif-

ferent geographical regions the same species of worm may

infest different species of crabs, (7) that the worms crawl about

on the bodies of the crabs and are thus easily transferred from

one host to another, (8) that by means of the free-swimming

embryos the species may be distributed widely, although the

young usually remain among the egg masses until they are

past the free-swimming stage.

LIST OF REFERENCES.

BENEDEN, P. J. VAN.

1861. Faune littorale de Belgique, Turbellariés. Mém. Acad. Roy.

Belgique. Tome xxxii, pp. 18-23, Pl. III, Figs. 1-30.

BERGENDAL, D.

1900. Ueberein Paar sehr eigenthümliche nordische Nemertinen. Zool.

Anzeiger. Bd. xxiii, p. 321.

BÜRGER, O.

1895. Die Nemertinen. Flora und Fauna des Golfes von Neapel.

`- Bd. xxii

DIECK, G.

1874. Beiträge zur Entwicklungsgeschichte der Nemertinen. Jenaische

Zeitschr. f. Naturwiss. Bd. viii, pp. 500-520, Taf. XX, XXI

Le laboratoire de Wimereux en 1888.

450 THE AMERICAN NATURALIST.

GIARD, A.

1888. Bull. Scientifique de la France et de la Belgique. Tome xx,

p- 496.

HuBRECHT, A. A. W.

1887. Nemerteans. Vol. xix. Challenger Reports. P. 25.

JousBINn, L.

1893. Les Némertiens. Faune Française. Pp. 212-213, Pl. III,

Fig. 81. Paris.

KOLLIKER, A.

1845. Ueber drei neue Gattungen von Wiirmern. Verh. Schweiz. Nat.

Gery Chur. Pp. 89-93.

McINTosH, W. C.

1873. Monograph of British Annelids. Pt.i, Nemerteans. Ray Society,

London.

Quoy, j. R: C., and GAIMArRD, J. P.

1833. Voyage de PAstrolabe. | Zoologie. Vol. iv, p. 291; Atlas,

PL ; Figs. 20, 21. ° 1826-1829. — Paris.

WILLEMOES-SUHM, R. v.

1874. Land Nemertean from Bermuda. Ann. Mag. Nat. Hist.

Vol. xiii, p. 411, Pl. XVII, Fig. 4

ON THE ASEXUAL ORIGIN OF THE CILIATED

SPONGE LARVA.

H. V. WILSON.

IN 1891! and in 1894? I published certain observations which

led me to believe that in monaxonid sponges free-swimming

larvae identical in structure with those developing from eggs

are in some species produced asexually. For one such species,

Esperella fibrexilis Wilson, the observations referred to are

in brief as follows: (1) Parenchymal cells well supplied

with yolk granules are found collected together in abundant

small groups of varying size. Such groups have no definite

shape and may contain few cells or many, and the compo-

nent cells may lie together very loosely or be packed pretty

closely? The microscopic pictures indicate that the groups

owe their origin to wandering cells that have collected together.

(2) Rounded masses of similar cells are found, the cells tightly

packed and polygonal in outline. To such masses, which are

surrounded by a follicle composed of a single layer of flattened

cells, I have applied the term “ gemmule.” The smaller gem-

mules are regarded by me as arising from the amorphous masses

of parenchymal cells above mentioned, and the microscopic pic-

tures further indicate that the gemmules grow both by cell

divisions and by fusion with one another. The gemmules vary

greatly in size. The smallest show zz section only a few cells,

the largest several hundred cells,* the diameter of the largest

gemmules bearing to that of the smallest about the ratio of

tentoone. With increasing size of the gemmule, the component

1 Notes on the Development of Some Sponges, Journ. of Morph., vol. v, No. 3.

? Observations on the Gemmule and Egg Development of Marine Sponges,

Journ. of Morph., vol. ix, No. 3.

3 Journ. of Morph., vol. ix, No. 3, Pl. XIV, Fig. 8; Pl. XV, Figs. 15-15.

t Itid, Pl. XV, Fig. 9, 5 cells; Fig. 16, 18 cells; Fig. 11, 37 cells; Fig. 9,

about 6o cells; Fig. 21, about 100 dte: Fig. 18, several hundred cells.

451

452 THE AMERICAN NATURALIST. '[Vor. XXXVI.

cells are found to be more tightly packed, the fine yolk in

the cell bodies is more abundant, and the nuclei are smaller.

Hence, to demonstrate cell outlines in the largest (mature)

gemmules! very thin sections are necessary. (3) Mature

gemmules are found in which the compact mass of cells is

broken up into a few large masses, on the surface of which

are semi-detached single cells, and between and round which

lie scattered a few similar free cells. The shape and position

of the partially free cells indicate that they are separating as

individuals from the subsidiary masses into which the gemmule

has broken. Gemmules in this stage are larger than in the

mature undivided condition, the spaces between and round the

subsidiary masses presumably being filled with absorbed fluid.

(4) In other gemmules the subsidiary masses (always irregular

in shape and of varying size) are numerous and small, and the

free cells are abundant. (5) In yet another stage the constitu-

ent cells of the gemmule are found to be completely separated

from one another, save that small multinucleate masses continue

to be present. The cells of the outermost layer are flattened

and form an investing membrane for the inner mass, the cells

and multinucleate masses of which are connected together by

delicate processes? (6) Transitional stages show that the outer

layer of cells becomes the surface epithelium of the larva, which

is ciliated except at the posterior pole, while the inner mass

becomes histologically differentiated into the several kinds of.

cells forming the parenchyma of the larva. The larva itself is

a typical monaxonid larva, essentially identical in structure with

those that are known to develop from segmenting eggs.

Thus an unbroken series of stages is found leading from a

small mass composed of a few parenchymal cells to a ciliated

larva. The various stages in this gemmule development,

including the resulting larvze, are scattered in great abundance

throughout the sponge body, and their presence is associated

with degeneration in the structure of the parent sponge.

Nowhere in this continuous series of stages is an egg, either

undivided or segmenting, interpolated. Very small ova, many

! Journ. of Morph., vol. ix, No. 3, Pl. XV, Fig. 18.

* Ibid., Pl. XVI, Fig. 21. 3 Ibid., Pl. XVI, Fig. 22.

No. 426.] THE CILIATED SPONGE LARVA. 453

times less the diameter of the mature gemmule, are however

present, although not common, in the parent parenchyma.

My conclusion that a larva, identical in structure with the

typical egg larva of the group, is produced asexually in the

manner just indicated has not seemed to certain reviewers

(Maas,! Minchin?) altogether worthy of credence. And it may

be that subsequent researches will show that such a conclusion

is not necessitated by the facts. What I may point out here,

however, is that the conclusion rests upon a series of micro-

scopic pictures, the arrangement of which is left to the intelli-

gence of the observer, and that this is precisely the kind of

evidence on which rests our belief that in other sponges an egg

development occurs. If, to be sure, the microscopic pictures

(images) have been portrayed by me in an inaccurate fashion,

then there is no need to consider the logic involved in their

arrangement. This is in reality the point of view from which

Maas has criticised my account, although he professes to find

in my figures evidence for the correctness of his interpretations

of them. Maas, in short, very freely and confidently expresses

(98, p. 370 ; '96, pp. 233—234) the belief that the bodies I figure

and describe as gemmules are merely eggs surrounded by masses

of nourishing cells ; and that the dissolution of the mature

gemmule into subsidiary masses and ultimately into separate

cells is only the segmentation of the egg. This process of dis-

solution specially awakens his incredulity: * Wie man es sich

vorstellen soll, und welche Kräfte es bewirken sollen, dass ein

Aggregat von Zellen, eine Gemmula, sich in einzelne Stücke

weiter und weiter spaltet, darnach will ich hier nicht fragen,

sondern nur darnach, warum Wilson in dem ganzen Vorgang

nicht eine Furchung, in dem Auseinanderbrechen der Massen

in kleine und kleinere Stücke nicht eine Zellteilung gesehen

hat!?'" (96,p.234). The mechanics of the process to me seem

unworthy of so much wonder. Absorption of water by the

1 Die Embryonal-Entwicklung und Metamorphose der Cornacuspongien, Zai.

Jahrb., Abth. f. Anat. und Ontog., Bd. vii, 1893; Erledigte und strittige Fragen

der Schwammentwicklung, Biol. Centralbl., Bd. xvi, Nr. 6, 1 96.

2 The Position of Sponges in the Animal Kingdom, Science Progress, vol. i,

N.S., July, 1897.

454 THE AMERICAN NATURALIST.. [VoL. XXXVI.

mass, coupled with the passage of individual cells from a resting

condition, in which mutual pressure gives them a polyhedral

shape, into an active condition would appear to be the essential

features of the process and are phenomena not altogether out-

side our experience. Maas has obviously been led astray in his

criticism by that well-known condition of mind in which things

that are not familiar seem impossible. In a more open and

receptive mood he would probably recognize the fundamental

differences between the figures of gemmules shown in Pls. XV,

XVI of my '94 paper and the figures of Fiedler! depicting the

growth of the ovum in Spongilla, and would not attempt to

explain the former as variations of the latter; more especially

when in the same paper (Pls. XXIII, XXIV) I myself repre-

sent stages in the growth and segmentation of sponge ova (for

Tedanione and Hircinia), the mode of growth in one species

(Hircinia acuta) closely resembling that of Spongilla as described

by Fiedler.

Since the publication of my results in 1894 there have

appeared but two communications in which this question is

touched upon, on a basis of actual observation. Vosmaer and

Pekelharing? found in Esperella egagropila certain reproduc-

tive bodies of a problematical nature. Not having at the time

a sufficiently complete series of stages in the development of

these bodies, they do “not enter into the question whether

Wilson's statements about Esperella fibrexilis are applicable to

our sponge." They confirm my observation that the formation

of these * gemmules'' is associated with degeneration in the

structure of the parent sponge, which they find “entirely

degenerates and finally dies off.” The other communication

is from Professor Ijima of Tokyo. This author in his recent *

beautiful and much admired work on the systematic zoólogy

and histology of the Hexactinellida? reports observations in

the main similar to mine, and is disposed to draw the same

,. Ueber Ei und Spermabildung bei Spongilla fluviatilis, Zeitschr. f. swiss

2 OF tions on Sponges, Verhandelingen der Koninklijke Akademie van

Wetenschappen te Amsterdam, ı 1898.

Studies on the Hexactinellida, Contribution 1, Journ. Coll. Sci. Imp. Univ.,

y Pu NE vol. gin Igor.

No. 426.] THE CILIATED SPONGE LARVA. 455

conclusion with respect to the origin of the larva. Ijima’s

observations are as follows :

In Euplectella marshalli undifferentiated cells, styled ** archze-

ocytes," are found in abundance scattered through the sponge

body. Such cells occur “solitarily or in irregular groups of

two, three, and so on, up to tens and in certain positions even

to hundreds or thousands." The growth of the archzocyte

groups ** may take place not only by cell division of their cells,

but also by fusion of originally separate groups." The smaller

groups are flat, the larger ones form solid and compact masses,

the shape of which is at first rather irregular. '* With continued

increase in the number of the cells and consequently in the

size of the mass, the latter assumes a roundish, oval or broadly

lobose shape, measuring up to 100 m or more across." Such

masses are common in all large individuals. Ijima believes,

on the strength of his own and earlier observations, that

such congeries of archzeocytes are of general occurrence in

the Hexactinellida.

In certain archzeocyte congeries the cells undergo a histologi-

cal change, the cell body developing spherules and increasing in

size. The transformed cells are known as **thesocytes." The

spherules are not firm solids but are of a soft, perhaps even

fluid, nature. Ijima concurs in the opinion of F. E. Schulze

that the formation of spherules is a metabolic process analogous

to the formation of glycogen in liver cells, and that the spherule

substance is to be looked on as reserve nutriment. The sub-

‘stance of the spherules is probably of an albuminous nature.

Ijima's description of the spherules strongly suggests that the

bodies are comparable with the well-known G/anzkérper in the

` rhizopod Pelomyxa (see Amer. Nat., Vol. XXXIV, No. 403).

All the archzocyte congeries are not transformed into theso-

cyte masses. Of those which are not so transformed, some

possibly represent an early stage in spermatogenesis, but Ijima

believes that “a good part of the primitive archzocyte congeries

are directly and actively concerned in the formation of certain

reproductive bodies, asexual or sexual but other than sperma-

tozoa.” The further history of such reproductive masses in

Euplectella marshalli was, however, not followed. But Ijima

456 THE AMERICAN NATURALIST. [Vou. XXXVI.

regards it as probable that similar masses constitute the anlages

of the superficial buds known to occur in some hexactinellids

(Rhabdocalyptus, Lophocalyx).

In two other species what appears to be the complete intra-

parental history of the archaeocyte congeries was made out.

But before describing Ijima’s observations on these forms I

wish to digress and point out how much commoner archzocyte

_masses seem to be in the Hexactinellida than ova and sperm.

Ijima recalls the fact that very little is known concerning

the ova and spermatozoa of these sponges. Schulze (80, '87)

describes sperm masses and ova (0.3 mm. in diameter) as pres-

ent in more or less abundance, and usually in the same indi-

vidual, in Euplectella aspergillum. Schulze was never able to

discover any distinct segmentation stages. Ijima himself says:

* [t is somewhat surprising to me that, although the different

hexactinellid species histologically studied by me are not few

in number, yet scarcely a single case in which either of the

sexual products was indisputably developed came under my

observation." As regards the male elements Ijima says he

is quite in the dark, that he has never seen spermatozoa or

indisputable sperm masses, nor indeed does he feel disposed

to say that he has seen ova. In one specimen of Euplectella

marshalli a number of cells were observed, the larger ones

measuring IO-I5 4, sometimes 23 p, in diameter. Such cells

are connected by transitional stages with, and are undoubtedly

derived from, archzocytes. Ijima thinks these cells are young

stages in the development of bodies similar to those described

by Schulze as ova in Euplectella aspergillum, but does not

commit himself to the view that they are eggs. In another

sponge, a specimen of Leucopsacus, comparatively large spher-

ical or ovoid cells, 20-40 p in diameter, were found in some

abundance. Such cells were not connected by transitional

stages with archzocytes, and the describer again does not find

the evidence for their ovum nature absolutely convincing.

The fact that Ijima found no indubitable sexual elements is

the more surprising since in the case of Euplectella marshalli

he made special search for them, examining numerous speci-

pm collected xus the four seasons of the year. The rarity

No. 426.] THE CILIATED SPONGE LARVA. 457

of these elements suggests that the sexual breeding season of

the Hexactinellida, zf zt occurs annually, is one of very short

duration.

In the two species above alluded to, Leucopsacus orthodocus

and Vitrollula fertile, Ijima has discovered not only archzeocyte

masses but also larva, the only hexactinellidan larva as yet

observed. Reserving for a later contribution his detailed

observations on these bodies, Ijima contents himself for the

present with a sketch of their structure and development.

The larvae are spindle shaped, thicker at one end, and consist

of a superficial flagellated layer covering an internal mass of

cells. The hexactinellidan larva is thus fundamentally similar

to that of monaxonid sponges. In an earlier stage the embryo

is spherical in shape, with no discernible flagellation, and con-

sists of a compact mass of small cells, those of the superficial

layer showing an epithelium-like arrangement but not differing

in general appearance from the internal cells. “Ata still earlier

stage there is found a simple cellular mass agreeing in all

respects with the latter" (the stage just described) “except in

having no distinct epithelial covering. And that simple cel-

lular mass is in all appearance nothing else than an advanced

stage of what I have called the archzocyte congeries, of which

there exists a series of different sizes, leading down uninter-

ruptedly to the little groups of cells so commonly found on the

chambers. At all events, there is nothing else than these

compact groups of small cells to which the origin of the develop-

ing embryo can be traced back with any degree of probability.”

Ijima appreciates the remarkable character of the development

just sketched and, before definitely formulating an opinion as

to the asexual origin of the larva, wishes to continue his obser-

vations on a more plentiful supply of material than he has

hitherto had. His position is best stated in his own words :

“To repeat, to me it seems certain that the embryo in a very

early stage of its development consists of a small assemblage

of uniform-looking cells, which differ in no distinguishable

feature from the archeocytes. If the resulting body were

something comparable to a bud or gemmula, I would probably

have felt no hesitation in concluding that the cells were really

458 THE AMERICAN NATURALIST. [VoL. XXXVI.

archzeocytes, and that we had here to do with a case of asexual

reproduction. But, free-swimming larvze, essentially similar to

those developed from ova in other sponges, being at issue, the

question whether true ova are not somehow complicated in the

cell mass whence the larva arises seems to claim to be brought

on the tapis, all the more, since our knowledge of the hexac-

tinellidan ovum is far from being satisfactory.” And again,

p. 188, after mentioning my observations and conclusion, “I

conceive the mode of origin and growth of the archexocyte

congeries in the Hexactinellida to be just the same, and it

seems to me not impossible that in the hexactinellid larve

which I have seen we have simply a new case of the ‘gem-

mule larva’ or bud embryo." After referring to Maas's view

that my conclusion rests upon a mistaken interpretation of a

process of oógenesis, Ijima goes on to say with regard to his

own observations, p. 189: ‘So far as concerns the archzocyte

congeries of the Hexactinellida, I can confidently state that

among the constituent cells in any stage of its growth, there

exists not one which, on account of its size or of other external

peculiarities, can be recognized as an egg. If it be that so

many cells are aggregated for the sake of the nutrition of a

developing ovum, this ovum is to be expected to deviate more

or less morphologically from the rest as it approaches maturity ;

however, no sign of such a differentiation is noticeable. Further,

all the cells in a congeries, large or small, are tolerably uniformly

and compactly packed together, so as to directly touch one

another; and where they are somewhat loosely arranged, there

is not a trace of any substance between them. So that I am

decidedly. against the assumption that some of them are, at

any stage of the growth of the congeries, engulfed among

certain others as pabulum. If, after all that, a portion or all

of the cells in a congeries giving rise to an embryo are still to

be looked at in the light of blastomeres that have arisen by

segmentation from a single egg cell, one is driven to the

assumption that the original ovum is, like the blastomeres

themselves, as small-bodied as, and indistinguishable from, an

archzeocyte. This would be very remarkable in an ovum; and

. moreover, under that supposition, it becomes imperative to deny

No. 426.] THE CILIATED SPONGE LARVA. 459

egg nature to the large ovum-like cells described by Schulze

and by myself from Euplectella.” Such an hypothetical mother

cell of an embryo as Ijima suggests in the above excerpt would

be something so different from an ovum that to class it as such,

it seems to me, would be to confuse our ideas concerning repro-

ductive cells. It would rather be analogous to a spore. How-

ever, both Ijima's observations and my own on the character

of the smaller groups of archzeocytes and their growth by fusion

render unwarranted the assumption that any such cell exists,

that is, as a typical and necessary condition in the development

of an embryo. That a single cell may occasionally and inci-

dentally give rise to a gemmule and so to an embryo is, of

course, a possibility. And thinking over the possibility of such

a case tends to clear up our ideas as to the nature of the typical

reproductive archzeocyte which acts in coóperation with others

to form a new individual. If the hypothetical, exceptional

archeocyte, which independently produces an embryo, is to

be looked on as equivalent to a spore, the same view must be

entertained regarding the typical coóperative archzocyte. In

the one case we have a single spore producing the individual,

while in the other case several spores unite, as, for instance, in

the production of a myxomycete plasmodium (sporangium).

Into the deeper-lying question as to how fundamental is the

difference between an ovum and a spore there is no need to

go. That there are, except in some of the plants, certain great

and obvious differences no one will deny.

From a theoretical standpoint further investigations of this

type of development in the sponges are much to be desired,

and Professor Ijima’s contribution will scarcely fail to call

them out.

UNIVERSITY OF NoRTH CAROLINA,

December 20, 1901.

AGGREGATED COLONIES IN MADREPORARIAN

CORALS.

J. E. DUERDEN.

ProF. G. von Kocu,! in the eighth of his well-known

« Kleinere Mittheilungen über Anthozoen," describes what

he terms * Aggregated Colonies” as occurring in the simple

Mediterranean coral, Balanophyllia verrucaria. The designa-

tion *aggregated" is employed by the author for such colonies

as have been formed through the secondary fusion of individ-

uals which were originally distinct, thus distinguishing them

from most other Anthozoan colonies which are produced by

the budding or fission of a single individual.

Von Koch collected around the small island of Vivara in the

Mediterranean Sea a number of Balanophyllia which, instead of

retaining the simple habit usual for this genus, consisted of two

or more polyps. The corallites were fused with one another

and possessed a common base, and many of the septa of adja-

cent calices were continuous. The tentacular system of each

individual polyp was distinct, a mouth occurred in the center

of each oral disk, but the column wall passed uninterruptedly

from one polyp to another. DUM :

An examination of serial transverse sections of the corallum

of two such colonies revealed that the corallites were inde-

pendent towards the base, but united in a common secondary

skeletal formation. In the case of a colony constituted of two

individuals, one much larger than the other, von Koch con-

cludes that the union had arisen through a larva settling near

an older polyp; after the first skeletal rudiments were formed

their coralla became fused through secondary deposition of

calcareous matter, the larger polyp contributing most.

1 Koch, G. von. Kleinere Mittheilungen über Anthozoen. VIII. Aggregirte

Kolonien von Balanophyllia verrucaria Aut. Morph. Jahrb., vol. xviii.

461 :

462 THE AMERICAN NATURALIST [VoL. XXXVI.

Sections of another colony, composed of four individuals,

showed that the four corallites remained at approximately the

same proportional size throughout, and the writer assumes that

the mass was formed by four larvz which settled close to one

another and practically at the same time, each constructing its

own corallum, and contributing an equal share to the formation

of the common investing skeleton.

The late Prof. H. de Lacaze-Duthiers! has recently given an

account of somewhat similar aggregations of the simple coral

Caryophyllia, obtained from Port-Vendres. On many of the

larger corallites smaller examples were growing, in such a way

as to leave little doubt that the * bouquet" was a result of the

adherence of Caryophyllia larvae to the corallum of a polyp

already established. Occasionally the individual members of a

colony are polyps of practically equal size, as if originating

from larvz which settled simultaneously and in close proximity.

In most of the examples described and figured by Lacaze-

Duthiers the corallites retain their individuality, without the

formation of a common secondary skeletal mass, but in one

specimen (/oc. cit., Pl. XV, Fig. 12) two calices appear to have

fused at their margin.

Prof. C. Schuchert has drawn my attention to the colonial

condition generally exhibited by the rugose coral, Strepte-

lasma (Paleophyllum) divaricans (Nicholson)? Representatives

of the genus Streptelasma are usually simple, but numerous

specimens of the above species in the collections of the United

States National Museum, collected from near the top of the

Lower Silurian of Ohio, constitute small colonies of two to six

individuals. The colonies were at first supposed to have been

formed “by lateral gemmation, or rarely by fission,” but a

close examination shows that such an explanation is very

improbable. Each member retains more independence of form

than is usual in colonies produced by gemmation or fission.

In many instances two or more corallites are found adhering to

some foreign object, such as a shell of Rhynchotrema capax or

x Lacaze-Duthiers, H. de. Les Caryophyllies de Port-Vendres, Arch. de Zoól.

Exp. et Gen., Ser. 3, vol. vii, ¥:

.* Geol. Surv. Ohio, Pal. 11, c p. 220, Pl. XXII, Figs. 10, 10 4.

No. 426. MADREPORARIAN CORALS. 6

493

a monticuliporoid bryozoan. The corallites of the different

colonies vary greatly in size, but the members of any colony

are practically equal, showing them to be derived from polyps

of a similar age; there is no suggestion of a larger parent

corallite and smaller buds. They seem best regarded as aggre-

gated colonies.

The common West Indian coral, Siderastrea radians (Pallas),

in the course of its development, has recently provided me

with actual instances of colony formation by the fusion of

originally free, distinct larve. The process of growth has

been followed from the time of fixation of the free-swimming

individuals as far as the production of the early skeleton.

Small, unfixed, adult colonies of S. radiaus occur in plenty

in the shallow waters of Kingston harbor, Jamaica, and are

easily kept alive in aquaria in the laboratory. The physiologi-

cal activity of numbers of unicellular algz, symbiotic within

the endodermal tissues, renders unnecessary any artificial means

of aération of the water. Colonies are occasionally met with in

which the internal cavity of many of the polyps is crowded with

larva. These are shot out from time to time and are thus

free to establish new colonies in turn. As a rule, only one or

two larvz are extruded at a time, but upon disturbance of a

fully charged colony a score or so would appear together. In

one colony the intermittent liberation of fresh individuals was

continued for about a month.

Immediately on being set free within the water the larvae

were able to swim about, and revolved around their principal

axis, being provided with a uniform layer of cilia. They

measured two to three millimeters in length and were mostly

pear-shaped, the narrow aboral end being anterior, or forward,

in swimming. The broad oral end was posterior and dark

colored, the latter condition owing to the presence of zoóxan-

thellze (yellow cells) within the outer ectoderm. In a few larve

the forward aboral end was the broader, as appears to be more

usually the case in actinian and madreporarian larvae when

first extruded. The majority were opaque, without any out-

ward indication of internal mesenteries, but occasionally a

distended specimen was met with in which the walls were

464 THE AMERICAN NATURALIST. [VoL. XXXVI.

nearly transparent and revealed eight internal mesenterial

divisions.!

During the first day the Siderastrzea larvae remained near

the surface of the water or gathered around the sides of the

vessels in which the colonies were kept. Afterwards they

traversed the water as a whole, though some preferred the

neighborhood of the bottom of the vessel From the first

day many would sink to the bottom and there lie motionless

for a time, after which the swimming movements might

recommence.

By the evening of the second day a few individuals had

fixed themselves permanently, either to the sides or bottom of

the vessels, or to the foreign objects partly incrusted by the

parent colonies. At first the larvae would

adhere by means of the actual tip of the

narrow extremity, but this would soon

flatten out and form a broad base, a small

rounded oral aperture appearing at the

free extremity. Whether or not any

individual larva would settle seemed very

uncertain, for out of several hundreds set

free comparatively few became permanently fixed. If fixation

were not accomplished within the first few days it seemed to

be impossible afterwards. In one instance about a score of larvae

were isolated, and nearly all kept alive for a period of twenty

days, without any of the specimens settling, although various

suitable objects were placed in the vessels.

iile many larva fixed themselves isolated from others, a

decided disposition to settle simultaneously in small groups

was evident. Thus in Fig. 1, where three larvae are repre-

sented, all in the first stage of fixation, their narrow apices are

so close as to be nearly touching. It is clear that when they

1 The larve of West Indian corals are usually set free at a stage at which

three or more pairs of mesenteries are already developed, the stomodzeum fully

formed but non-functional, and the interior wholly, or in part, occupied by a

lated tissue breaks. dein and i is seen to be extruded at intervals through the oral

aperture.

No. 426.] MADREPORARIAN CORALS. 465

flatten out by this extremity their walls will be in close con-

tact, and a colony of three young polyps will result.

Such a colony, adherent to a pebble, and formed of seven

individuals, is represented in Fig. 2. The drawing was made

two or three days after fixation of the larva, when the adult

polypal form was beginning to be assumed, and the tentacles

were just appearing as rounded prominences. All the polyps

are at practically the same stage of development, proving that

they settled synchronously, or nearly so. They are irregularly

arranged with regard to one another, and in their effort to fully

expand the walls press upon one another, and produce distortion

of the proximal part of the column.

To the under surface of another small pebble thirty-eight

larvae attached themselves in groups of two, three, or more.

One of these groups contained a dozen or so young polyps, all

touching, the mutual pressure producing an angulation of the

normally circular base. Another group of thirty-two became

adherent during a single night to the surface of a small glass

dish. In this case nearly all the members, upon flattening,

were touching to a greater or less degree.

Fixation having taken place, the larva, now to be regarded

as polyps, seemed remarkably vigorous, and underwent the

next stages of development with comparative rapidity. In

466 THE AMERICAN NATURALIST. [Vor. XXXVI.

sharp contrast with this the unfixed larve remained all the

time at practically the same stage as when first extruded.

The glass vessel to which some of the polyps became adher-

.ent was broken into small fragments, with the polyps still upon

them. These could then be transferred from one aquarium

to another, or examined in small dishes as transparent living

objects under the microscope. In this way the growth of the

various organs was followed day by day. Some specimens

were kept alive for a period of seventeen weeks, the develop-

ment of the tentacles and septa being followed as far as the

third cycle. For the present purpose the normal development

will be succinctly reviewed, so as to compare the modifications

introduced as a result of the primary grouping of many of the

polyps in colonies.

On fixation the tissues of the polyps became transparent and

the six pairs of primary mesenteries were seen to be already

present, four pairs attached to the stomodaeum and two pairs

incomplete. About, a month elapsed before the members of

the second cycle began to appear, being first represented by a

pair in each of the dorsal exocceles, then later by a pair in each

of the middle exocceles, and finally by a pair in each ventral

exoccelic chamber.

The first tentacles commenced to make their appearance two

or three days after settling, as six rounded outgrowths over

alternate mesenterial chambers, and early showed an opaque,

white, knob-like apex distinct from a short, more transparent

stem. The six prominences appeared simultaneously, con-

stituting a first cycle, and were equal in size and distance

apart, and in communication with the exoccelic chambers.

About four weeks passed before another cycle of six tentacles

appeared, internal to the first, and situated over the entocoelic

chambers. The further tentacular development was somewhat

complicated and need not be here considered.

Three or four days after fixation the skeleton was first

bserved in the form of six radiating septal upgrowths, prac-

tiodlly equal in size. At the same time a narrow peripheral

calcareous ring was formed, its outer surface uncovered by the

soft polypal tissues, and therefore to be regarded as an epitheca.

No. 426.] MADREPORARIAN CORALS. 467

The six septa were free from one another and from the epitheca,

and situated at equal distances apart within the primary ento-

coelic chambers, thus alternating with the cycle of six exoccelic

tentacles first to arise.

A day or two after the formation of the first cycle of ento-

septa the six exoccelic septa began to make their appearance,

in some cases simultaneously, but successively in others, in

bilateral pairs from the dorsal to the ventral aspect of the polyp.

Under ordinary transmitted light no basal skeletal formation

could be detected, but with polarized light the presence of

crystalline particles could be demonstrated. In polyps macer-

ated at this period a very thin basal disk was recognizable,

Fic. 3.

formed of an aggregation of irregular granules, and continued

at its edge as the upturned epitheca, the six or more radial

septa arising from it vertically some distance within.

A wide range of variation was exhibited by the different

polyps as to the rate of growth of the various organs, especially

in the later stages. The individuals which were larger on fixa-

tion soon outstripped the others; scarcely any succumbed,

and it was possible to feed them on fragments of worms and

mollusks.

Such a regular, symmetrical development was characteristic

only of the isolated polyps, free to expand equally on all sides.

The polyps in groups were crowded to such a degree as to

necessitate arranging themselves at all angles with regard to

one another and the incrusted object. Sometimes part of the

base of one polyp would be reclining against part of another ;

but none of the individuals seemed to be overpowered by such

468 THE AMERICAN NATURALIST. | [Vor. XXXVI.

close proximity. As a whole, the growth of the colonial polyps

was much less forward than that of the isolated specimens.

All were normal as regards the occurrence of the six pairs of

primary mesenteries—four complete and two incomplete; the

tentacles also were represented by a primary cycle of six exo-

ceelic representatives and a secondary inner cycle of six

entoceelic members. Variations and irregularities, however,

were introduced in connection with the formation of the skele-

ton. The epitheca was invariably produced at the free basal

edge of the column wall, and under such conditions as those

represented in Fig. 2, it is clear that its outline would be irreg-

ular, and that the epitheca of one polyp would be in contact

with that of the contiguous polyps to the extent the polypal

walls were adherent. Likewise the basal disks of two polyps

reclining against one another would be formed at a sharp angle.

The septa of the colonial polyps exhibited from the begin-

ning many irregularities and imperfections. Such a series is

indicated in Fig. 3. The drawing was taken from the under

surface of a living colony of seventeen polyps, adherent to

a fragment of glass, and viewed by transmitted light, so that

only the basal skeletal structures are exhibited. The polyps

are seen to vary greatly in size; the outline is angulated where

they are in contact with others, and rounded on the free

border. The two larger polyps at each extremity of the colony

partly overlap one another at an angle, though this is not very

obvious from their flat basal expansions alone. The thickened

margin to each polyp represents the epitheca, that of two con-

tiguous polyps being fused along the line of contact. The

basal disk is yet so thin as to be practically transparent and is

indicated by the dotted areas.

The radiating dark bands, as a rule simple, but sometimes

forked, represent the adherent surface of the septa; the latter

on account of their vertical thickness are non-transparent like

the epitheca. In no polyp are the two orders of six septa fully

or regularly developed in the manner already described for

isolated polyps. Any number of septa from four to twelve are

represented, as a rule with a suggestion of alternating large

and small members.

No. 426.] MADREPORARIAN CORALS. 469

The development of the colonial polyps was not followed far

beyond the stage represented in Fig. 3. Growth was very

slow compared with that of the isolated individuals, and most

were preserved for further study. There was no indication,

however, but that under favorable natural conditions the

aggregations would have grown into fully mature colonies. In

the later stages the primary irregularities due to crowding

would probably have been outgrown, and the groups would

then present all the characters of normal colonies arising by

budding.

In one feature the polyps constituting an ordinary colony of

S. radians differ from those in a colony formed of aggregated,

primarily independent units. Polyps arising by gemmation

are, as a rule, only incompletely cut off from the parent; their

internal cavities are from the beginning in communication with

those of the adult polyps, so that the nutrient fluid can circu-

late from one individual to another. Such would not be the

case at first with aggregated colonies, and at the very early

stage at which the observations on Siderastrea were made I

could not assure myself that intercommunication was estab-

lished. The intervening external epitheca would for some

time interfere with such a possibility, but no doubt this struc-

ture, along with the basal plate, would be left behind in the

upward growth of all except the marginal polyps. For, except

a most rudimentary formation at the margin, the epitheca is

absent from mature colonies.

It does not seem probable that the phenomenon of aggre-

gated colonies is a prominent feature of coral growth. Still,

it is necessary that its possibility should be recognized, for

wherever it occurs the resulting colony will probably differ

somewhat from one produced by normal budding or fission,

while when occurring in usually simple corals its non-recogni-

tion might lead to disastrous results.

All students of both fossil and recent corals are familiar with

the fact that very often simple coralla are found adhering to the

dead portions of colonies, undoubtedly the young of these.

Among West Indian corals the occurrence of young polyps

under such conditions is very frequent in Manicina areolata

470 . THE AMERICAN NATURALIST. | [Vor. XXXVI.

(Linn.) and Favia fragum (Esper). A colony of Manicina will

sometimes possess five or six adhering young polyps, the dif-

ferent members of which exhibit one, two, three or more oral

apertures, and in other ways are regularly developed. Their

structural details prove that they have originated not by fission

or budding, but from larvze which have settled, probably imme-

diately or shortly after extrusion, in close association with the

parent colony. Colonies of Manicina and Favia are often

found charged with larve, and their extrusion has been fre-

quently observed.

Much variation exists as to the activity displayed by the

larvae of these and other coral species immediately on extru-

sion. In some cases the larva are quite vigorous, and able to

swim about from the beginning; and under natural conditions

these would no doubt continue their activities for a long time,

and might be drifted great distances before settling. Others

again sink immediately on liberation from the interior of the

parent, and falling upon the nearest object may settle there.

In most instances the nearest object will be the dead lower

part of the parent colony. It is thus easy to see how, under

ordinary natural conditions, such aggregations as those found

by von Koch and Lacaze-Duthiers may have arisen. The

grouping of the larva in S. radians appeared to be less fortui-

tous than this, for practically all the specimens were able to

swim about immediately on being set free. It may be that

the tendency towards aggregation above described was in some

way influenced by the cramped artificial conditions of the small

aquaria. ;

The possibility of extruded larve settling in numbers imme-

diately around the parent, instead of drifting widely apart, is a

phenomenon likewise to be looked for in the field study of the

closely allied Actiniaria. On the coral reefs in West Indian

waters occur large irregular patches, often several feet across,

of one or other of the anemones, Actinotryx sancti- Thome

Duch. & Mich. and Ricordea florida Duch. & Mich. The

polyps are usually adherent to some dead coral slab, and in

a single patch there may be hundreds of individuals. Both

species exhibit asexual reproduction by fission, and in any

No. 426.] MADREPORARIAN CORALS. 471

group are to be found specimens with from one to three or

more oral apertures on a single disk. One may assume that

all the individuals constituting a patch have arisen asexually

from one or a few primary polyps. But such does not appear

to be the case. Aggregations have been observed in which

one large example would be surrounded by a number of others,

all smaller but practically uniform in size. Both species are

found to extrude larvae very freely, and there seems no doubt

that the patches are in the main formed of individuals derived

from such larvz as settled near the parent. A certain num-

ber of fission polyps would also be expected. Externally it

would be practically impossible to distinguish between larval

polyps and fission polyps, but the internal mesenteries afford

distinctive characters as to the one or the other form.

Similar extensive patches of the large discosomid, Stoichactis

helianthus (Ellis) are not infrequent. Their occurrence is

probably to be explained in the same manner as above, that is,

as due almost entirely to the aggregation of distinct larve.

Asexual reproduction seems to be very rare in this species.

Polyps of the northern species of Metridium usually occur

in groups. From the researches of G. H. Parker! and others,

asexual reproduction seems to play a considerable part in the

crowding habit of these, but the possibility of aggregating

larvae also should not be overlooked, even though M. margi-

natum extrudes unfertilized eggs.

JoHNs HOPKINS UNIVERSITY,

BALTIMORE, Mp.

l Parker, G. H. Longitudinal Fission in Metridium marginatum Milne-

Edwards, Ju. Mus. Comp. Zoöl. Harvard, vol. xxxv, 1899.

UTAH CHILOPODS OF THE GEOPHILID#.,

RALPH V. CHAMBERLIN.

In this paper are presented descriptive accounts of six new

species of chilopods belonging to the genera Geophilus and

Linotzenia, four to the former and two to the latter. The

material upon which the descriptions are based was collected

for the most part during the spring and summer seasons of

1901. By way of showing the position of the new forms in

their genera analytical keys are given for the species here first

described and also those known previously from the western

United States.

KEY TO WESTERN SPECIES OF GEOPHILUS.

a,. Frontal plate discrete; last ventral plate wide.

ġ Anal pores present; nei pores numerous; pairs of legs

53-55 (9); length 50-53 m : i glaber Bollman

Č, Anal pores absent; pleural "Tora two; pairs of legs 47-53;

length 37-47 m . . cephalicus Wood

a,. Frontal plate not disce! ; dnd aid present

6,. Last ventral plate relatively narr

cı Claw of anal legs obsolete ; pen pores ten or eleven ; basal

plate free; pairs of legs 73 (3); length 39 mm

fomir Meinert

c, Claw of anal legs long; pleural pores eighteen or twenty;

basal plate partly covered; pairs of legs 73-75; length

49-55 mm neatotus sp. nov.

€. Claw of Sem he shot or eS ; pleural pores nine ;

pairs of legs 45-49; length 20-31 mm. xenoporus Sp. nov.

c, Pleural pores thirty or more; basal plate partly covered;

pairs of legs 64-67 ( 9); length 36 mm. cal eer Bollman

6,. Last ventral plate relatively wide.

c Claw of anal legs obsolete; basal plate thrice wider than

long; pleural pores partly covered; coxz of prehensorial feet

armed ; length 29 mm. (?) atopus sp. nov.

€, Claw d anal legs short ; land R i times wider than

long ; pleural pores free ; coxz of prehensorial feet unarmed ;

length 40-43 mm. : - i . glyptus sp. nov.

473

474 THE AMERICAN NATURALIST. (VoL. XXXVI.

Geophilus nealotus sp. nov.

Description. — Rather robust, very gradually attenuated anteriorly, more

strongly posteriorly ; prosternum and head with a few long hairs or sub-

glabrous, the latter partly granular or mostly smooth, polished, the body

nearly glabrous, smooth and polished; head light chestnut or brown, body

dorsally light brown to yellow, ventral plates and legs yellowish, antenne

the same or darker.

Antenne rather short (4.4—4.6 mm.); the four basal joints sparsely

pilose with long hairs, the others closely clothed with finer short hairs ;

joints all long, the ultimate much shorter than the two preceding taken

together

Cephalic plate much longer than wide (5 :3.8 nearly) ; posterior border

truncate ; impressed with two lateral and a single median sulcus, the latter

within a more or less evident longitudinal furrow, which is wider and

shallower anteriorly ; covering the anterior border of the basal plate ; basal

plate large, subequal to the first dorsal scutum, 2.5 times wider than long.

Claws of prehensorial feet when closed reaching to the distal portion of

the first antennal joint; each claw with a small obtuse tooth at base ; coxa

armed with an inconspicuous obtuse tooth ; sternum wider than long, more

than twice the outer height of the coxa, anterior border widely sinuate,

unarmed.

Dorsal scuta with a single median and two lateral longitudinal sulci,

which become obscure on a few of the first middle plates but are again

distinctly impressed caudad ; often a few of the last middle scuta with a

transversely oval or oblong depressed and darker colored area marked off

anteriorly and posteriorly by a crescentic impression; anterior prascuta

very short, gradually increasing in size to the first posterior segments, which,

are moderate, then more rapidly decreasing to end of body.

nterior spiracle very large, vertically oval or suboval, a few following

of the same shape, others round; decreasing in size from the first to the

eae! caudad becoming very sm

scuta with a median longitudinal sulcus, which may be faint or

Kon absent on a number of the middle plates, its place there being taken

sb two more or less distinct lateral sulci ; a median suboval or somewhat

r poriferous area on posterior portion of each plate ; with some-

times but one, but more commonly with two, transverse furrows, these on

each side often broken up into several lines diverging toward the middle

"t the plate.

Legs sparsely pilose ; first pair distinctly shorter and more slender than

those. ag; anterior pairs stouter and shorter than posterior pairs;

anal lees ied longer than the penultimate, Me swollen, each pro-

— with a rather long claw.

osterior pleura inflated, with 18-21 mirins. pores on each side,

few innermost covered by the last ventral ins last ventral plate

No. 426.] UTAH CHILOPODS OF THE GEOPHILIDZ. 475

relatively rather narrow, its sides very gently curved convexly (9) or

concavely ( 2), converging posteriorly ; anal pores moderately large.

Pairs of legs of male seventy-three, of female seventy-five. Length

of male 55 mm., width 1.6 mm.; length of female 49 mm., width 1.5 mm.

Adolescens. — A a, 30.5 mm. long measures .7 mm. in width ;

length of antennae 2.6 mm. ; pleural pores very small, eight on each side ;

pairs of legs the normal oum (75) ; coloration as in adults.

Habitat.— Under stones in the gullies of the hillsides north of Salt

Lake City.

The types were collected March 28, 1901.

Geophilus xenoporus sp. nov.

Description. — Moderately robust, distinctly attenuated caudad and

cephalad ; head and prehensorial feet sparsely pilose with long or moder-

ately long hairs, cephalic plate punctate, especially at sides, with moderately

coarse puncte, the prosternum and coxa of prehensorial feet sparsely

minutely punctate, dorsal and ventral plates nearly glabrous, smooth or a

little roughened ; head and prehensorial feet with prosternum light brown,

body, legs, and antennz yellow.

ntennz rather short (d 1.602 mm. 9 2.4-2.5 mm.); first seven

or eight joints sparsely pilose, the others, especially the final ones, densely

clothed with short straight hairs; articles all long, the ultimate subequal

to the two preceding taken together.

Cephalic plate much longer than wide (7:5 nearly), narrowed poste-

riorly; posterior border somewhat rounded, covering the ba sal plate ante-

riorly ; with two rather long lateral sulci and a brief but sharply impressed

median line immediately caudad to the frontal suture; basal plate rather

large, width about three times the middle length, the plate being more

exposed laterally.

Claws of prehensorial feet when closed reaching or a litle surpassing

the distal end of the first antennal article; claw armed at base with a long

subacute tooth; inner side of coxa long, armed similarly to claw but tooth

somewhat stouter; prosternum nearly equal in length and breadth, less

than twice the height of the coxa, the anterior border narrowly and acutely

incised, unarmed.

Dorsal scuta bisulcate, the sulci distinct throughout; anterior prascuta

short, the middle long or very long, the posterior moderate

First spiracle mtg oval, the others circular, not inch decreasing

in size caudad, all

Ventral Pati cens a median sulcus which is deep, especially cephalad.

Legs sparsely pilose ; the first pairs shorter and much more slender than

those eres: ; the anterior pairs stouter than the posterior but not much

differing in length ; anal legs of male much swollen, those of the female more

slender, with a ca or indistinct claw in the male, longer in the female.

476 THE AMERICAN NATURALIST. [VoL. XXXVI.

Pleure of last segment not much inflated, with about nine small pores

on each side, these arranged along a suboval line on ventral and lateral

surfaces ; last ventral plate narrow, the sides straight and parallel; anal

pores small.

Pairs of legs of male 45, of female 47—49. Length of male 19-21 mm.,

width .5—.6 mm. ; length of female 30-31 mm., width .9—1 mm.

Adolescens. — Specimens 13 mm. long measure .4 mm. in width ; length

of antenna 1.2 mm.; pleural pores four or five on each side; color pale

yellow.

Habitat. — Canyons near Salt Lake City, mostly among decaying leaves

and in vegetable mo

The types were sdti in City Creek Canyon, April 1, 1901.

Geophilus atopus sp. nov.

Description. — Rather slender, very gradually attenuated cephalad,

more strongly caudad, depth of the body in the middle region as great as

e width; head sparsely pilose, smooth, the basal plate and exposed dorsal

partió ót prosternum more densely covered with hair, the dorsal scuta .

mostly very sparsely clothed with short hairs, not at all roughened ; cephalad

light brown, head a little darker at sides, caudad yellowish, antennze and

legs concolorous with adjacent parts of bod

Antenne short (2 mm.) ; the first three or four joints sparsely pilose with

moderate straight hairs, the others more closely clothed with short hairs

which increase in density to ultimate joint; ultimate article nearly equal in

length to the two preceding taken together.

Cephalic plate much longer than wide (5:4 nearly); anterior and pos-

terior margins truncate; with two distinct lateral sulci and a fine discon-

tinuous median line; frontal border of basal plate concealed, exposed

portion thrice wider than long.

Claws of the prehensorial feet when closed reaching to the middle of the

first antennal article; claw armed at base with a pale obtuse tooth ; coxa

armed similarly to claw; prosternum wider than long (9:8 approximately),

less than twice the height of the coxa, anterior margin with only a shallow

sinus, unarmed.

Dorsal scuta with two rather faint lateral sulci and a single median

sulcus, these more strongly impressed caudad ; first dorsal præscuta short,

irregularly increasing in length to the pne region, where they are long

excepting the last few, which are v

The first anterior spiracle subround, those following round, gradually

decreasing in size caudad, finally becoming small or very small.

Ventral scuta faintly sulcate, the anterior ones with a median depressed

area Leve) on the plate; ventral pores large, arranged over and about

the ressed area, extending forwards and backwards along the median

s= add iiaae to ~ sides, farther caudad the pores mostly restri

No. 426.] UTAH CHILOPODS OF THE GEOPHILID.£. 477

a transverse band ; posterior border of the anterior scuta produced back-

wards from the sides to the middle, there touching or slightly overlapping

the succeeding plate, exposing the praescutum of the latter only at the sides ;

posterior borders of plates after the twenty-second straight.

Legs, excepting ultimate, pilose with moderately long straight hairs ;

first pair more slender than others, not much shorter ; anterior pairs much

stouter than the posterior, not much shorter; anallegs much swollen, claw

obsolete, clothed more densely and, more often, with shorter hairs than

pairs preceding.

Pleurz of last segment not much inflated, with about twelve pores of

several sizes, these arranged subseriately, partly covered by the last ventral

plate; last ventral plate wide, much wider than long (1.85 : 1), sides con-

vexly curved, converging posteriorly.

Pairs of legs of male 69. Length of male 29 mm., width .7 mm.

Habitat. — The type specimen, a male, was found buried in damp saw-

dust under a piece of timber, Salt Lake City.

The type was taken June 3, 1900.

Geophilus glyptus sp. nov.

Description. — Body robust, caudad strongly, cephalad but little attenu-

ated; head pilose at sides, more especially posteriorly, basal plate and

exposed dorsal portions of prosternum more densely pilose, prosternum

ventrally subglabrous, first dorsal plates sparsely provided with short hairs,

the plates caudad becoming subglabrous, ventral plates glabrous ; head

and prosternum with the prehensorial feet chestnut, body above and below

amber brown, or in middle portions yellowish, legs and antenne concolorous

with the body. i

Antenne short (3-3.4 mm.), stout, attenuated from base distally ; first

four or five joints sparsely provided with long hairs, the others with shorter

hairs, which decrease in length and increase in density toward ultimate

article ; articles long, the ultimate shorter than the two preceding taken

together.

Cephalic plate a little longer than wide (9: 8.2 nearly) ; posterior mar-

gin truncate, anterior margin extending forward from the sides to the

middle, sides rounded, impressed with two lateral and a single median

sulcus; basal plate partly covered, exposed portion over four times wider

than long.

Claws of prehensorial feet not much curved, when closed reaching to or

a little beyond the distal end of the first antennal article; each claw armed

at base with a moderately small obtuse tooth ; inner side of coxa short or

very short, unarmed ; prosternum wider than long (not quite 9:8), over

twice the outer height of the coxa; anterior margin widely sinuate.

Dorsal scuta impressed with two lateral and a single deep median sulcus ;

anterior prescuta very short, middle long or very long, posterior short.

478 THE AMERICAN NATURALIST. [Vow. XXXVI.

First two anterior spiracles very large, round, those following abruptly

smaller, also round, gradually decreasing in size caudad, in the middle and

posterior segments very small.

Ventral plates with a median sulcus which is strongly impressed cephalad,

becoming rather faint caudad.

Legs all pilose; the first pair much smaller than those succeeding; ante-

rior pairs much stouter and also longer than the posterior ; anal legs long,

moderately to strongly swollen, armed with a short or very short claw.

Last pleure moderately inflated, with about twelve pores which are sub-

seriate and free, or sometimes a few covered by the last ventral plate ;

last ventral plate moderately wide ( d) or narrower (9) ; anal pores present, `

moderate in size.

Pairs of legs of male 67, of female 69. Length of male 43 mm., width

1.3 mm. ; length of female 40 mm., width 1.2 mm.

vidodpeieni- — Specimens 30 mm. long measure .8 mm. in width; pro-

portionately wider cephalad than adult; length of antenna 2.6 mm.;

pleural pores seven or eight on each side ; coloration nearly as in adults.

Habitat. — Along the streams in the canyons near Salt Lake City.

The types were collected in Neff's canyon, Salt Lake County, June 30,

1900, by my brother, Mr. Seth C. Chamberlin.

KEY TO WESTERN SPECIES OF LINOTJENIA.

4,. Pairs of legs 71-81 ; individuals large.

ó,. Pairs of legs 71 ; length 60 mm., head not punctate or spotted.

parviceps (Wood)

5, Pairs of legs 81; length 140 mm., head minutely spotted with white.

epileptica (Wood)

. Pairs of legs 37-47; individuals small. (Frontal — discrete, anal

pores present, last ventral plate wide or moderate.)

6,. Anal legs shorter than the penultimate ; tem pores 5-7, moder-

ate; posterior border of cephalic plate straight; pairs of legs

39-41 ; length 13.5-16.7 mm miuropus sp. nov.

6,. Anal legs longer than the EEE rend pores 7-11, small ;

posterior border of m = xd pairs of legs 43-47;

length 20-21 mm ; micropus sp. nov.

Linotenia miuropus sp. nov.

Description. — Robust, gradually attenuated caudad, more strongly

cephalad ; head and entire body moderately or subdensely clothed with

short hairs, smooth ; ferruginous, ventral plates and legs paler.

Antennz short (maximum length 1.2 mm.); moderately hirsute, decreas-

ing in density distally, ultimate article subglabrous, hairs short or very

short ; first articles of medium length, shorter distally, the ultimate longer

than the two nme together.

No. 426.] UTAH CHILOPODS OF THE GEOPHILID.E. 479

Cephalic plate wider than long ; posterior border covered by the basal

plate, substraight ; impressed with a median sulcus ; frontal plate discrete ;

basal plate in male less than thrice wider than long (2.8:1 nearly), in

female more than four times wider (4.35 : 1).

Dorsal scuta not sulcate; anterior przsscuta short, middle and posterior

moderately long.

Ventral plates with a distinct median sulcus.

Spiracles all round, the first rather small, those succeeding scarcely

decreasing in size, the posterior being not much or not at all smaller than

the anterior.

Legs sparsely hirsute with moderately long hairs; first pair distinctly

smaller than others ; the anterior and posterior pairs subequal; anal legs

of male shorter than the penultimate, much stouter, densely clothed with

short hairs, armed with a slender claw, anal legs of female slender, not at

all swollen, similarly hirsute to other legs.

Pleurz of last segment with 5-7 moderate pores on each side; last

ventral plate relatively intermediate in width, wider than long (6.7 : 5.7

nearly), sides curved, converging posteriorly ; anal pores moderate.

Pairs of legs of male 37-39, of female 41. Length of average male

16.7 mm., width .9 mm. ; length of female 13.5 mm., width .65 mm.

Habitat.— About a dozen specimens were found in Logan canyon,

Cache county, near the mouth, and three were taken among the pines on

the divide between Daniell's Creek and Hobble Creek canyons, Utah county.

The types were collected May 25, 1901.

Linotenia micropus sp. nov.

Description. — Robust, cephalad strongly, caudad more gradually atten-

uated ; head and body above and below pilose, more densely in female

than in male, last ventral segments more densely pilose than others, proster-

num nearly glabrous ( 2) or sparsely hirsute (9); ferruginous throughout.

^. Antenne very short (1 mm.), stout; moderately pilose ; articles mostly

short, the ultimate longer than the two preceding taken together.

Cephalic plate as a whole subequal in width and length, exposed portion

wider than long in the ratio of 6: 5.4; posterior border produced backwards

from sides to middle, subtriangular, overlapped by the basal plate ; basal -

plate thrice wider than long.

Claws of prehensorial feet when closed not reaching the front margin

of head by a considerable space; claw armed at base with a long stout

tooth, coxa unarmed; prosternum much wider than long (7: 4.2 approxi-

mately), two and one-half times longer than the coxa, anterior margin widely

sinuate, unarmed; inner margin of coxa extremely short or entirely

concealed.

Dorsal scuta not sulcate; anterior praescuta short, longer in male than

_in female, the middle and posterior moderately long.

480 THE AMERICAN NATURALIST.

Ventral plates with a single median sulcus which caudad becomes indis-

tinct or obsolete, with a transverse furrow or depression; excepting the

first few, with a rather large median poriferous area on the anterior por-

tion of plate, this often, especially caudad, prolonged backwards in the

form of a narrow tongue; ventral pores also arranged on each side of the

sulcus in front of the posterior border.

Anterior spiracle intermediate in size, those succeeding very gradually

decreasing, caudad becoming small.

Legs pilose with mostly rather long hairs; first pair distinctly reduced,

the second pair intermediate; anterior and posterior pairs not much differing

in size; anal legs of male strongly swollen, densely clothed with short to

very short fine hairs, longer than the penultimate, anal legs of female not

at all swollen, similarly pilose to other legs, armed with a short pale claw.

Pleurz of last segment with 7-11 small obliquely seriate pores on each

side; last ventral plate wide, sides curved, strongly converging posteriorly,

anterior border produced forwards from sides to middle line, making the

mee: tocan: anal pores present.

rs of legs of male 43-47, of female 47. Length of average male

21 mm., width 1 mm. ; length of female 20 mm., width 1.2 mm.

Adolescens. — Specimens 12.5—13.5 mm. long measure .4—6 mm. in

width; length of antenne .6-.7 mm.; pleural pores 4-6 on each side ;

color yellowish or light brown.

Fetus.— One individual examined is still coiled up within the egg, the

yellowish outer membrane, or shell, of which, however, is evenly split down

one side from end to end, the thin transparent inner membrane being still

intact; in this condition the egg is spherical and measures 1 mm. in

diameter.

An individual 4 mm. long is free excepting posterior segments, these

still covered by the membranes; legs short and rather stumpy, closely

crowded ; antennz bent back under the head along the not yet fully

ed mouth parts.

pecimens 4.5-7.5 mm. in length still spirally coiled; width .4 mm.

and under ; antennae extended, as usual relatively long, clavate, joints very

short excepting the ultimate, the sy as long as the three or four pre-

ceding taken together ; color brow

Habitat.— In the quaking aspen ugar pine woods along the head waters

of Manti and Pole canyons, San Pete County, and about the glacial lakes

at the head of Little Cottonwood canyon, Salt Lake County.

All the specimens of the fetus stage, ten in number, were found together

with the body of the — coiled about them, the nest being a pit in the

underside of a log.

The types were cni in Pole canyon, July 15, 1901.

LATTER-DAY Saints’ COLLEGE, SALT LAKE CITY,

December 17, 1901.

ON THE COLOR VARIATIONS OF THE COMMON

GARTER SNAKE.

EDWIN C. ECKEL.

I. EUTJENIA SIRTALIS IN THE ADIRONDACK REGION OF

New YORK.

IN an earlier paper! I have noted that Cope's treatment of

the color forms of Eutenia sirtalis, at least so far as the sub-

species occurring in the northeastern United States are con-

cerned, can hardly be regarded as satisfactory. My acquaintance

with the extreme western and southern forms of the species is

entirely too limited to permit the expression of an opinion on

the closeness with which his subspecific grouping, in those

areas, approximates to the facts; but it seems probable that

the same conditions prevail there as here.

In a catalogue? (now in press) of the reptiles and batrachians

of New York, prepared by Dr. F. C. Paulmier and myself at

the request of Dr. F. J. H. Merrill, director of the New York

State Museum, no revision of Cope's grouping was attempted.

All.the subspecies described from this state by Cope and

G. M. Allen were included, as it seemed undesirable, in a paper

designed for popular use, either to omit these forms without

presenting reasons for such action, or to discuss these reasons

in the necessary detail. ;

For this reason the results of the collection work which has

been accomplished during the past year, in critical localities,

are presented in this paper.

My thanks are due to Dr. F. C. Paulmier, who has kindly

placed the resources of the zoólogical department of the State

Museum at my disposal, and to Mr. Raymond L. Ditmars, who

1 Snakes of New York, Amer. Nat., February, 1901

? Paulmier, F. C., and Eckel, E. C. Catalogue of the Reptiles and Batrachians

of New York, Bulletin New York State Museum.

481

482 THE AMERICAN NATURALIST. [Vor. XXXVI.

has given me many facts regarding the distribution of the

various color forms of the common garter snake.

It is to be regretted that the results so far obtained are

chiefly negative, tending to cast doubt upon the tenability of

previously described subspecies, without pointing out the direc-

tion in which we are to look for more significant variations. A

large series of living specimens, collected in such widely sepa-

rated areas as the Adirondacks, the Catskills, the Hudson

Highlands, Long Island, the shore of Lake Ontario, and the

counties of the “southern tier," might indicate the presence

of certain variations which could be correlated with differences

of habitat, or are associated with such other aids to polytypic

evolution as would justify us in considering them as subspecies.

It is hardly necessary to say that, for the purposes of this

study, living specimens are absolutely essential. Colors alter

so rapidly on immersion in alcohol that alcoholic specimens are

worthless so far as color variations are concerned.

1. Subspecies previously described from the Adirondacks.

The subspecies of Eutenia sirtalis which have been noted as

occurring north of Pennsylvania are as follows: Æ. s. graminea

(Cope), E. s. ordinata (Linn.), E. s. sirtalis (Linn.), E. s. obscura

(Cope), and Z. s. pallidula (Allen). Of these, all save Æ. s.

graminea have been found in the Adirondack region. In the

present paper I purpose discussing the effect, upon the tena-

bility of the subspecies Æ. s. obscura and E. s. pallidula, of the

examination of material collected in New York state during the

past field season. In a future paper I hope to discuss the

remaining subspecies. The characters of Æ. s. obscura and £E. s.

pallidula, as given by Cope and Allen respectively, are as

follows :

Eutenia sirtalis obscura (Cope).

Cope: in his-last discussion! of this subspecies, describes it

as a form i in which the spots have entirely (or almost entirely)

1 Cope, E. D. Crocodilians, Lizards, and Snakes of North America, Af.

Nat. Mus. (1898), p. 1074.

No. 426.] THE COMMON GARTER SNAKE. | 483

disappeared by fusion, leaving the stripes intact. The colora-

tion above is therefore deep brown or almost black, with three

yellowish stripes, of which the laterals are less distinct than

the dorsal. The gastrosteges are grayish green, with the usual

black spots near the ends.

In this subspecies he includes one specimen from Mitchell's

Bay, Ontario ; three from Lac qui Parle, Minnesota ; and five

from Westport, Essex County, New York. These last were

collected by Baird and described! by him as Eutenia sirtalis.

The specimens assigned to the subspecies are, it is true, all

from northern localities, but no definite statement is made as

to the range of the form ; and it is probable that Cope did not

intend to associate this coloration with any particular area or

life zone. A proposed subspecies, based upon specimens from

widely scattered localities (the intervening areas being occupied

by other subspecies), is, in general, to be regarded with sus-

picion, unless positive evidence, other than that afforded by

similarity of coloration, can be adduced in favor of its subspecific

value. Positive evidence of this character is rarely obtainable,

particularly in the case of reptiles. In the case now under

discussion, I believe that it can be shown that obscura inter-

grades completely with more typical forms of Eutenia sirtalis ;

that its color characters cannot be correlated with any particular

climatic conditions ; and that therefore individuals exhibiting

these color characters cannot be regarded as subspecifically

distinct from those showing the coloration of typical Eutenta

szrtalis.

Eutenia sirtalis pallidula (Allen).

In 1899 G. M. Allen described? the subspecies pallidula

from specimens taken near Intervale, N. H., giving its geo-

graphic distribution (p. 64) as “from the White Mountains

of New Hampshire and the Adirondacks of New York north-

ward into New Brunswick and Nova Scotia, and possibly

farther." Later in the same paper (p. 65) he states that

1 Baird, S. F.,and Girard, C. Catalogue of North American Reptiles E P

2 Notes on the Reptiles and Amphibians of Intervale, N. H., Proc.

Nat. Hist. (1899), p. 64 et seg.

484 THE AMERICAN NATURALIST. | [Vov. XXXVI.

pallidula is the characteristic form of the Canadian zone, and

restricts “the name Thamnophis sirtalis Linn. to the brighter-

colored form found in the Transition and Austral zones of the

. east." This restriction necessitates a redescription of Eutenza

sirtalis sirtalis, which he accordingly gives. His summary

of the differences between the two subspecies is as follows:

“ Thamnophis sirtalis pallidula needs comparison with no other

of the sirtalis group except T. sirtalis proper, from which it

differs in the obscurity of the dorsal stripe, which is grayish,

not yellow; the ground color, which is olive brown, not black

or blackish ; in the chestnut color below the lateral stripe,

where sirtalis is olive; in the lighter color of the belly, espe-

cially in the younger examples; and in the interlinear spots as

previously described” (Z.e., the spots of pallidula were described

as being composed of chestnut scales, with black edges and

interspaces, and those of sirtalis proper as being composed of

scales usually entirely black, occasionally faintly reddish in the

middle).

As can be seen, this is an attempt to correlate a certain

variation with a given “life zone"; and, if justified by the facts,

would be a distinct advance upon the subspecific treatment of

Cope. For this reason the question of its validity requires a

more detailed discussion, for even a slight variation — if obvi-

ously arising from definite causes — is worthy of consideration.

As will be seen later, however, I cannot admit that pallidula

fulfills these necessary conditions. Not only do the characters

assigned to it occur frequently in specimens outside of the

Canadian zone, but all specimens from that zone do not exhibit

these characters.

2. Specimens collected during 1901 in New York State.

Specimens from Moody, Franklin County, N.Y.

1. Dorsal stripe very faint, but still visible on close examina-

tion as a dull yellow-brown line. Lateral stripes fairly well

shown; greenish yellow. Sides below lateral stripes brownish

green. Gastrosteges greenish; black spot on anterior edges,

near sides, often communicating by a thin black band along

No. 426.] THE COMMON GARTER SNAKE. 485

anterior edge of the scutum with the lowest row of spots.

Color, above lateral stripes, olive green. Interspaces (between

scales) light greenish. Spots present, but not very marked.

Scales composing spots chestnut, with black edges and inter-

spaces.

Second upper labial on right side divided; left side normal.

First temporal on both sides of head divided, its anterior

portion appearing as a small separate triangular plate.

2. Dorsal stripe fairly distinct, lateral stripes less so. Sides

below lateral stripes dark green. Gastrosteges greenish. Color

above lateral stripes olive green; interspaces light greenish.

Spots present, composed of chestnut scales, with black edges

and interspaces.

3. Dorsal stripes almost invisible; lateral stripes fairly well

shown. Sides below lateral stripes brownish. Gastrosteges

greenish. Color above lateral stripes olive brown; interspaces

light greenish. Spots very indistinct (owing to brownish back-

ground), composed of chestnut scales, with black edges and

interspaces.

4. Dorsal stripe visible but not distinct. Lateral stripes

visible ; yellowish. Sides below lateral stripes light brown.

Gastrosteges greenish. Color above lateral stripes olive brown ;

interspaces light greenish. Spots of chestnut scales, with black

edges and interspaces.

5. Dorsal stripe almost invisible, only showing on close

examination. Indications of lateral stripes on second and third

scale rows; yellowish. Sides below lateral stripes light brown.

Gastrosteges slate gray, tinged brownish. Color above lateral

stripes olive brown ; interspaces light greenish. Two rows of

spots between lateral and dorsal stripes ; the scales, in general,

being black (a few, however, are chestnut, with black edges),

with black interspaces. Another series of spots, similar in

color, along the first and second scale rows.

6. Dorsal stripe visible throughout; yellowish. Lateral

stripes faint. Sides below lateral stripes light brown. Gastros-

teges slate gray. Other colors as in No. 5.

7. Dorsal stripe visible. Lateral stripes very faint. Sides

below lateral stripes brown. Gastrosteges grayish, tinged with

486 THE AMERICAN NATURALIST. [Vor. XXXVI.

brown. Color above lateral stripes olive brown. Interspaces

light greenish. Spots of chestnut scales, with black edges

and interspaces. Preocular divided on right side; left side

normal.

8. Dorsal stripe very faint; lateral stripes faint. Color

above lateral stripes brown, tinged olive. Interspaces light

greenish. Four series of spots between lateral stripes. Scales

of the spots chestnut, with black edges and interspaces.

The four series connect across back, forming bars or network.

Top of head deep green. Chin and throat orange. Rostral

yellow. Lower labials white, tinged yellowish on upper

edges. Upper labials yellow to yellowish green, black edges

anteriorly. Sides below lateral stripes, together with exterior

portions of gastrosteges, brown. Remaining portions of gas-

trosteges olive brown. Lower side of tail deep orange.

9. Dorsal stripe pure and rather deep yellow near head,

becoming browner yellow and somewhat less distinct poste-

riorly. Lateral stripes yellow, rather distinct. Color above

lateral stripes olive brown ; interspaces light greenish. Spots

present, not very distinct ; composed of chestnut scales, with

black edges and interspaces. Sides below stripes brown;

gastrosteges lighter brown.

10. Dorsal stripe yellowish brown, distinct throughout.

Lateral stripes brighter and somewhat more yellowish than the

dorsal. Color above lateral stripes brown, tinged olive. Inter-

spaces light greenish. Spots composed of chestnut scales,

black borders and interspaces. Sides below stripes, as well as

exterior edges of gastrosteges, a soft light brown. Remaining

portions of gastrosteges greenish gray.

To the descriptions of these specimens from Moody, Franklin

County, N.Y., I will add for comparison descriptions of a few

other living specimens of Eutenia sirtalis examined this year:

11. Sconondoa Creck, one-half mile east of Oneida, N.Y.

Dorsal stripe not apparent. Lateral stripes faint, but visible ;

yellowish brown. Sides below lateral stripes brown. Gastros-

teges greenish. Ground color above lateral stripes dark olive

brown. Spots present, but not apparent against the dark

background.

No. 426.] THE COMMON GARTER SNAKE. 487

12. Same locality. Dorsal and lateral stripes distinct, yel-

low. Sides below lateral stripes, and gastrosteges, greenish.

Spots distinct. Ground color above lateral stripes olive brown.

13. Rensselaer, Rensselaer County, N.Y. Dorsal stripe con-

spicuous throughout; clear yellow. Lateral stripes fairly

distinct. Sides below lateral stripes brownish green; gastros-

teges dark greenish. Ground color above lateral stripes very

dark brown, almost black. The location of the spots, on this

dark background, is only indicated by the blackness of their

interspaces, the normal interspaces being light greenish.

14. Rochester, N.Y. Dorsal stripe fairly conspicuous. Lat-

eral stripes indistinct. Ground color above lateral stripes soft

olive. Spots of chestnut scales, with black edges and inter-

spaces.

15. Ausable Forks, Essex County, N.Y. Dorsal stripe dull

yellow brown, but fairly distinct. Lateral stripes faint. Sides

below lateral stripes brown. Gastrosteges brownish green.

Ground color above lateral stripes dark greenish brown. Spots

indistinct.

16. Same locality. Dorsal stripe very faint; dull dark

brown. Lateral stripes visible; yellowish brown. Sides below

lateral stripes dark brown. Gastrosteges grayish green. Above

lateral stripes dark olive brown. Spots indistinct because of

darkness of ground color; composed of scales only a little

redder than the ground color, with black interspaces.

17. Same locality. Dorsal stripe rather faint, dull brown.

Lateral stripes very faint. Sides below lateral stripes brown-

ish green. Gastrosteges greenish. Ground color above lateral

stripes greenish olive. Spots small and indistinct ; composed

of chestnut scales, with black edges and interspaces. Eight

superior labials on each side resulting from the division of the

(normal) second upper labial. Lower anterior portion of pre-

frontal plates on each side divided, so as to give, in effect, two

loreals, one above the other.

- The following table gives certain data concerning the locali-

ties from which the specimens in the above list were obtained.

In the cases of Moody, Rochester, Oneida, and Rensselaer the

elevations given are those of the exact collecting localities.

488 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Concerning the other localities I have no exact data as to the

collecting range, and have therefore given the geographic

position and elevation of the villages.

DATA CONCERNING LOCALITIES DISCUSSED.

LocArriTYy. jupe ied | LATITUDE. | LONGITUDE.

1 | Moody, Franklin County, N. Y. ..... 1600” 44° 12° 74 39.

zo dntervale, NA 0... 1i enm 540° 44? 157 71° 25°

3 | Ausable Forks, Essex County, N.Y... . 600° 44° 26° 21" AT.

4 | Westport, Essex County, N.Y....... SUI 44° 11* 73" 29

5 | Rochester, Monroe County, N.Y. ... . 280° 43? 12" 77° 30

6 | Oneida, Oneida County, N.Y.......- 440° 43? o6 75° 3

7 | Rensselaer, Rensselaer County, N.Y. . . 60’ 42° 38’ 73° 44

The specimens from Franklin County, N.Y., were collected

by me during August, 1901, at Moody, a small settlement on

the east shore of Tupper Lake. All the specimens from that

locality described above (Nos. 1-10) were obtained at an eleva-

tion of about 1600 feet A.T. Though the ophidian fauna of

the Adirondacks appears to be scanty, so far as species are

concerned, the few species occurring within the region are

represented by numerous individuals. The specimens above

described were not selected because of color, but are simply

those which reached the museum alive and in good condition.

The locality is, of course, well within the Canadian life zone,

and the specimens can therefore be compared directly with

those described by Allen from Intervale, N.H.

As can be seen from the descriptions given (Nos. 1-10), the

specimens obtained from this one small area furnished fairly

good examples of three of the four subspecies credited to the

Adirondacks, together with specimens intermediate between

each pair of the three. If we omit the specific and higher

characters, there is hardly one point in which the ten specimens

agree. The dorsal stripe is, in most, somewhat indistinct ; but

in one specimen it is quite bright throughout, and in others

barely visible. In color it varies from dull brown to yellow —

never gray or grayish. The spots are usually of chestnut

scales, with black edges and interspaces; but the chestnut

No. 426.] THE COMMON GARTER SNAKE. 489

areas may be very small and only faintly reddish. The distinct-

ness of the spots varies with the ground color. In one speci-

men the sides below the lateral stripes are dark green; in

another, brownish green; in the remaining eight specimens,

various shades of brown. In two specimens various characters

have aided each other in such a way as to give a fairly uniform

reddish brown coloration above the lateral stripes, thus even

going a step beyond the condition shown by Cope's Eutenta

sirtalis obscura. The gastrosteges vary in color from greenish

gray to slate gray or even brown.

The general darkness of color is much accentuated by the

age, not necessarily of the individual, but of the skin which

the specimen is wearing at the time of examination. Six of the

specimens brought from the Adirondacks have shed their skins

since their arrival at the State Museum. Four of these had

been examined with some care previous to this time, and

reéxamination after shedding developed the fact that, while the

markings are unchanged, the colors are sufficiently brightened

by shedding to produce an impression of much lighter tint.

To sum up these statements, I find that ten specimens of

the common garter snake obtained (by an unmethodical selec-

tion) from an area less than two acres in extent, located well

within the Canadian life zone, represent, according to the

criteria of various authors, the following subspecies:

NuMBER OF SPECIMENS REPRESENTED.

According to E. s. sirtalis. E. s. obscura. E. s. pallidula.

SM o ord no ee IO SE one

GU c ee V 8 2 ER

ADADn 22 E dl rs -e di

The specimens k (Nos. 15, 16, and 17) collected by Mr. Van

Allen Lyman near Ausable Forks, Essex County, N.Y., which

is also within the Canadian life zone, exhibit similar variations.

1 Three of h ten specimens show charactors side with neither of Allen's

descriptions ; epted by him as pallidula,

De accep

though few of them exhibit all the characteristics of that form.

490 THE AMERICAN NATURALIST.

The thirteen specimens so far discussed prove that the

Canadian life zone contains garter snakes not exhibiting the

characters of pallidula. An examination of the descriptions

of specimens 11, 12, 13, and 14 (all of which are from points

in the Transition or Upper Austral zone) will show that speci-

mens exhibiting the characters of pallidula are not confined to

the Canadian zone. It would seem impossible, therefore, to

agree with Mr. Allen in assuming any relation between the

characters of pallidula and the climate of the Canadian life

zone. Asa matter of fact, specimens exhibiting these charac-

ters may be found in almost any part of New York state. I

believe that at the present time Mr. Ditmars has, in the New

York Zoólogical Garden, living specimens of this type captured

in New York City.

If, however, no relation can be established between the col-

oration of the form pallidula and any particular habitat,

pallidula can no longer be regarded as a definite variation from

the type, due to definite climatic conditions. It then falls to

the same level as the form oscura, and apparently no reason

can be given for assigning subspecific rank to either.

New York STATE MUSEUM,

December 5, 1901.

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

E MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L MARK, Direcror, No. 133

NOTES ON THE DISPERSAL OF SAGARTIA

LUCLE VERRILL.

G. H. PARKER.

SrupENTS of the New England sea anemones have for some

years past been acquainted with a small but striking species of

Sagartia that inhabits, often in great numbers, the higher tidal

pools and similar situations on our coast. This sea anemone is

about a quarter of an inch in height, slightly less in diameter,

and of a dark green color; it is usually striped vertically with

a few orange lines and has a cluster of some forty-eight whitish

tentacles. It was described nearly four years ago under the

name Sagartia lucie by Verrill (98), who first observed it in

1892 and annually after that for six seasons near New Haven.

On the authority of W. R. Coe, Verrill reports it common at

Woods Hole in 1898. He suggests that it may have been

introduced into the region about New Haven on oysters that

are annually brought from the south in large quantities and

planted in the waters of Long Island Sound. He notes its

rapid increase in numbers. That it is multiplying and spread-

ing considerably is indicated by the following records of earliest

occurrences, which, though probably open to revision, give a

rather definite idea of the direction and rate of dispersal.

When not otherwise stated, these records are based on the

observations of the writer.

New Haven, Conn. Verrill, who for a long time has been

especially acquainted with the marine invertebrates of this

region, states that he did not observe this species between

1865 and 1890. His attention was first called to it in 1892

by Miss L. L. Verrill, who found it in tidal pools at Outer

491

492 THE AMERICAN NATURALIST. [VoL. XXXVI.

Island, Conn. It was then much less abundant than in 1898,

when it was described.

Newport, RJ. This anemone was not found in the region

about the Newport Marine Laboratory in 1887, nor in 189o,

though in both seasons the actinians of the Newport region

were extensively collected. In the summer of 1895 it was the

most abundant species in the cove next the laboratory.

Woods Hole, Mass. S. lucie was not observed by Verrill in

his very exhaustive study of the fauna of Woods Hole between

1871 and 1887. In my own collecting at that place in 1889

it was not obtained. It was reported from Woods Hole in

1898 by W. E. Coe, and, in a letter to me from T. H. Morgan,

the same year is given as the earliest at which Professor Morgan

is certain of its occurrence in that locality. It is now abundant

on the wharves about Woods Hole, and especially on the eel-

grass in the Eel Pond.

Nahant, Mass. This species was not found at Nahant

between 1882 and 1898, though collections of invertebrates

from this locality were made usually several times annually in

the years between 1882 and 1898. In the autumn of 1899 à

few specimens of S. /uciæ were found in a small high tidal

pool on Little Nahant. During the summer of 19o1 Albert

P. Morse, without knowledge of the previous occurrence of

this species at Nahant, collected specimens of it in that locality.

It is now very abundant in most tidal pools on Little Nahant.

Salem, Mass. It was collected at Salem Neck, near Fort

Pickering, and near the Salem end of the Beverly Bridge in the

summer of 1901 by Albert P. Morse.

Salem is the most northerly locality for which I have any

record of the occurrence of this species, and, since it is not

included in the Preliminary Catalogue of the Marine Inverte-

brates of Casco Bay, Me., prepared a year ago by Kingsley

(01) it may be that Salem is not far from its present most

northerly limit.

The five localities already mentioned fall into the following

series when arranged according to the sequence of earliest

occurrences : New Haven (1892), Newport (1895), Woods Hole

eai Nahant (1899), and Salem (1901). This series shows,

No. 426.] SAGARTIA LUCIA VERRILL. 493

I think, beyond a doubt that the species has migrated eastward

from New Haven and northward to Salem, having covered the

distance between these two extremes probably within ten years.

As this actinian is remarkably hardy, thriving well in high tidal

pools where the water is often temporarily much diluted and

very variable in temperature, it is unlikely that the change

from the warmer waters south of Cape Cod to the colder ones

north of that point will noticeably retard its dispersal. Like the

introduced periwinkle, Littorina littorea (see Bumpus, '98), this

species will probably gain an extended distribution both north

and south of Cape Cod, though in this instance the invasion

comes from the south instead of from the north, as with the

periwinkle. It is hoped that these notes will call the attention

of observers to S. /uciæ, so that definite information may be

obtained as to its present distribution, both north and south,

and likewise evidence of its further dispersal.

LIST OF REFERENCES.

Bumpus, H. C.

'98. The Variations and Mutations of the ‘Introduced Littorina. Zod.

Bull. Vol. i, pp. 247-260.

KINGSLEY, J. S.

‘Ol. Preliminary Catalogue of the Marine Invertebrates of Casco Bay,

Me. Proceedings of the Portland Society of Natural History.

Vol. ii, pp. 159-183.

VERRILL, A. E.

'98. Descriptions of New American Actinians, with Critical Notes on

ther Species, I. Amer. Journ. of Sci. Ser. 4, Vol vi,

PP- 493-499.

NOTES AND LITERATURE.

BIOGRAPHY.

Lamarck’s Life and Work.' — It would be difficult to single out

from the naturalists of the past one more appropriate for biographical

treatment at the present time than Lamarck. The renewed interest

taken in his work in the last two decades, and the appearance of

such books as Darwin's Life and Letters and the recent biography

of Huxley suggest some extended treatment of the greatest of the

precursors of the Darwinian movement. Unfortunately, the material

for such a study is meager in the extreme, and Dr. Packard has been

obliged to search assiduously for the relatively few facts he can

record of the life of the great French naturalist. The biographical

portion of the work is contained in the first sixty-five pages. Then

follows a general estimate of Lamarck by contemporary and later

biologists. Lamarck's wide range of activities is next reviewed in

chapters dealing with his work in meteorology and physical science,

in geology, in invertebrate paleontology, in general physiology, in

botany, and in zodlogy. This is followed by a historical treatment

of the theory of evolution leading up to Lamarck’s contribution to

it. The last chapter is on Neolamarckism, and the volume is con-

cluded by a bibliography and a short index. There are ten full-

page illustrations, four of which are portraits of Lamarck.

The biographical chapters, which really form the introduction to

the body of the work, give one the impression of uncritical methods.

Dr. Packard was unable to find in Paris any exact statement of

Lamarck’s birthplace, and he undertook the very worthy task of

seeking it out for himself. His belief, however, that the house that

he was shown at Bazentin was the place in question rests upon

the unsupported statement of the village schoolmaster, who, we are

told, left his duties in the schoolroom to point out the ancient struc-

ture. Possibly Dr. Packard may be correct, but the whole account

has much more the air of French politeness than of scientific

acumen. Still more unsettling are the statements concerning

1 Packard, A. S. Lamarck, the Founder of Evolution, his Life and Work.

New York, Longmans, Green & Co., 1901. xii + 451 pp» IO pts-

495

496 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Lamarck’s death. This occurred, according to Dr. Packard, on

Dec. 28, 1829 (p. 56), the obsequies were celebrated on the Sunday

preceding December 23, the burial took place December 30 (p. 57);

and the first assembly of professors after his death was on

December 22 (p. 62). From this confusion of dates one may seek

refuge in a biographical dictionary to find that Lamarck died

December 18 and was buried probably December 20. It is really

unfortunate that in the one extended account of Lamarck’s life in

English the date of his death should not be accurately recorded.

Although the biographical introduction is somewhat disappoint:

ing, the body of the work more than makes good this defect.

Dr. Packard is to be congratulated on having hit upon the happy

idea of allowing Lamarck to expound his own views, and a large part

of the 300 pages that follow the biography is made up of translations

of well-selected passages from the works of Lamarck. Nowhere

else in English is one likely to find so true an exposition of the

Philosophie zoologiqgue as in the chapter devoted to this subject,

and the reading of this portion of the work is to be recommended

to every student of zoólogy that is not already familiar with

Lamarck's original publications.

The concluding chapter deals with Neolamarckism, a movement

with which Dr. Packard has had much to do and about which he

can consequently speak with authority. The impression made by

this chapter is, however, one of extreme vagueness. What Neola-

marckism is, is nowhere made very clear. Darwin’s views and

Lamarck's are contrasted, and much stress is laid on the importance

of environment as a factor in the production of variations; but the

idea is not even suggested that all such material may be only grist for _

the Darwinian mill. Moreover, a strange inconsistency runs through

much of this chapter, for Dr. Packard seems to think that by

pointing out the weaknesses of Darwinism, he is strengthening

Lamarckism. This leads him to give numerous quotations from

the works of eminent biologists who were keen enough to expose

some of the weak points in natural selection, but who did not per-

ceive that by so doing they were ranking themselves Neolamarckians.

The last quotations in criticism of natural selection are taken from

discit ; had a few been drawn from Darwin the proof would have

been complete. The real question, however, is not whether

ee ae is true and Darwinism untrue, — for these theories are

|. not necessarily incompatible, — but what are the factors of evolu-

. tion; and on this point Dr. Packard has not much to say, though in

No. 426.] NOTES AND LITERATURE. 497

the body of the work Lamarck tells us many things worthy of careful

perusal and thought. While this last chapter, like the biographical

introduction, leaves much to be desired, Dr. Packard’s volume is so

replete with good translations of well-chosen passages from Lamarck

that one cannot hesitate in pronouncing it the most complete and

truthful statement of Lamarck’s views that has thus far appeared in

English.

ZOOLOGY.

Benham on Flatworms. — The present volume,’ which is Part

IV of Lankester’s Treatise, is the third of this useful series to make

its appearance, and covers, in Chapters XIV to XXI inclusive, the

Turbellaria, Temnocephaloidea, Trematoda, Cestoidea, Appendices

to the Platyhelmia, and Nemertini. Each chapter opens with a

synopsis of the classification adopted for the group, and this is

followed by an admirable historical summary, which, though brief,

is notably clear, complete, and well balanced. The next section

deals with the general characters of the group under consideration,

and contains an analysis and discussion of each subdivision in order,

closing with a list of the chief works on the group. The “ideal”

platyhelminth, exploited in the opening chapter, is rather too gener-

ally used to meet the approval of present-day zoólogists, even though

it affords an easy standard of comparison for the beginner. It is a

great pleasure, however, to see some important theoretical explana-

tions presented in a general text, and that in a clear and attractive

manner which serves to make the mass of detail comprehensible.

But it may be seriously questioned whether anything is gained by

raising to the rank of phylum every group which after intensive

study appears to be sharply set off from its nearest of kin.

For Turbellaria the author has used the classification of Lang

and von Graff, and has made a happy selection of figures to illus-

trate the points under consideration. The short chapter on Temno-

cephaloidea follows Haswell's work very closely and gives the best

presentation of this little-known group accessible in any text-book.

Among the Trematoda, Monticelli’s classification, as modified by

1 Treatise on Zoölogy. Edited by E. Ray Lankester. Part IV, The Platyhel-

mia, Mesozoa, and Nemertini. By W. Blaxland Benham, D.Sc., M.A. London,

Adams and Black; New York, The Macmillan Company. 204 pp., 114 text-figs.

498 THE AMERICAN NATURALIST. [Vor. XXXVI.

Braun, has been adopted in the main, although such minor changes:

as those in the arrangement of the families will hardly be regarded

by most helminthologists as an improvement. A few minor errors

have crept into the chapter, as the representation (Fig. IX) of the

lateral nerve as the chief longitudinal stem, and the denomination of

the basement membrane as an internal limiting cuticular membrane.

The classification used for the Cestoidea is a combination of those

used by Braun and Lang,to the former of which it is certainly

inferior. The use of Cestoidea merozoa instead of Cestoidea polyzoa

Lang is a noteworthy improvement as avoiding the confusion and

incorrect inference in the latter term; but why not a single term

rather than the cumbersome double form? The arrangement of the

families is open to the criticism that no grouping can be considered

more than temporary which is based on so artificial a feature as the

number of suckers. Both text and literature are deficient in respect

to the older work of Cohn and Lühe, and one is struck by the entire

omission of references to the very extensive and important work of

Stiles, although one cut was taken from Stiles and Hassall.

The next chapter, which is referred to on the title-page and in

the preface as dealing with Mesozoa, distinctly repudiates that term

and considers as separate appendices to the Platyhelmia the Rhom-

bozoa, Orthonectida, Trichoplax, and Salinella. An extended note

by the editor of the series adds here a valuable statement of recent

results of importance which would have been welcome in other

. places also.

The final chapter, dealing with Nemertini, follows the work of

Bürger closely. Two pages of addenda and corrigenda have been

added by Mr. Punett without doing more in the opinion of the

reviewer than emphasizing the unfortunate delay in the appearance

of the book. Scant justice is done some authors, and as striking a

paper as that by Woodworth on Planktonemertes is entirely passed

Over.

There are some features which appear rather clumsy in American

eyes, such as the use of Roman numerals to designate the figures,

and the habit of beginning to number the figures anew in each

chapter. One may justly say also that the figures are inferior to

such as are used in this country to-day, and far behind those found

in continental texts. As instances of the use of mediocre figures

for forms of which good representations are easily accessible may be

cited those of Bothriocephalus (p- 112) and Echinococcus (p. 1 31),

while that of Dipylidium (p. 1 35) is little more than a caricature.

No. 426.] NOTES AND LITERATURE. 499

Much of the poor effect of the figures may be attributed to the

author’s inclination to use ideal or diagrammatic representations.

It is difficult to criticise fairly a work which, as the editor of the

series says in a prefatory note, was written in 1897 and was entirely

in proof in 1898. The epoch-making work of Looss on Trematoda

and of Cohn and Liihe on Cestoidea, as well as many shorter con-

tributions, have added so largely to our knowledge of these groups

that what was thoroughly good in 1898 would now be strikingly out

of date. One cannot help wondering how the work could have been

held so long unpublished. In spite of these disadvantages, it will

stand as a clear and well-balanced presentation of the subject,

admirably arranged and suggestive in treatment.

Henry B. WARD.

Structure and Metamorphosis of Actinotrocha. — In the Journa?

of the College of Science of the Imperial University of Tokyo, Japan,

Ikeda! gives an account of studies on the development of Phoronis,

from the unsegmented egg through the metamorphosis. For the

earliest stages, the author was able to obtain living material in

quantity, of the species Phoronis ijimai, while the structure of the

larva was studied in four species (or forms) of Actinotrocha taken

in the region of the Misaki Marine Biological Station.

The author gives some interesting details as to the life history of

Phoronis, showing that the adults probably die every year, the colo-

nies being completely replaced by the metamorphosing larve. He

criticises the present classification of the species of Phoronis, holding

that several of the so-called species are not really distinct.

A detailed account is given of the cleavage, gastrulation, formation

and structure of the larva, and the transformation into the adult.

Ikeda's results on some important or disputed points are as follows :

The formation of the mesoblast begins at the beginning of gastrula-

tion, by the irregular pushing of some of the entoblast cells into the

cavity of the blastula. Mesoblast formation continues in the same

manner from the two “anterior diverticula” of the archenteron, and

from a ventral groove leading backward from the blastopore. The

mesoblast cells are at first irregularly scattered, and only later arrange

themselves to form the boundaries of the body cavities. The latter

^ are thus not formed from enteric diverticula, as has sometimes been

1 Ikeda, Iwaji. Observations on the Development, Structure, and Metamor-

phosis of Actinotrocha, Journ. Coll. Sci. Imp. Univ. Tokyo, Japan, vol. xiii,

Pp- 507-592, Pls. XXV-XXX. i

500 THE AMERICAN NATURALIST. [VoL. XXXVI.

described. The posterior so-called “anal pit” is said by Ikeda to

be the Anlage of the nephridia, and to have no connection with the

so-called “primitive streak." In many details Ikeda's results differ

from those of Masterman, — especially in many of the points which

the latter emphasizes as indicating affinities with the Diplochorda.

Thus, Ikeda considers as artefacts, due to reagents, the oral and

pharyngeal grooves which Masterman had compared with gill slits,

the *neuropore," the *subneural gland," and certain parts of the

vascular system described by Masterman. He was further unable

to find the * proboscis pores " of Masterman, the “trunk nephridia,"

the ventral blood vessel, the dorsal mesentery, the collar nerve ring,

the ventral nerve commissure, or the perianal nerve ring. The

impression is thus given that his results are largely opposed to those

of Masterman; the latter, however, points out in a recent review’

of Ikeda's paper that in regard to the fundamental structure of

Actinotrocha the points of agreement are more important than those

of disagreement; indeed, he claims Ikeda's work as a corroboration

of his own.

Ikeda has given especially valuable observations on the relation

of the body cavities and vascular systems of the adult to those of

the larva, — a complex matter, on which little had been done, and

which perhaps requires still further elucidation. The vascular

system of the larva is very simple as compared with that of the

adult, not forming a closed system at all; the transformation of the

one into the other in the fifteen or twenty minutes occupied by

the metamorphosis is therefore a complicated matter. In the same

way the body cavities of the larva and adult by no means correspond.

Ikeda states that the collar cavity of the Actinotrocha is largely

transformed into the “ring vessel" of the adult, — the collar cavity

of the latter being largely a new formation. HSI

A General Course in Insect Anatomy.? — This is a new edition of

a little book that is important both of itself, and because it represents

the fundamental laboratory course in the chief center of entomologi-

cal instruction in America. The book has been evolved along with

the laboratory which it represents, and each new edition marks

Progress in the knowledge of even those subjects which are dealt

with in the most elementary instruction.

1 Quart. Journ. Micr. aps vol. ve (1902), pp. 485-492.

? Comstock, J. H., Kellogg, V. L. The Elements of Insect Anatomy: An

Outline for the Use of uae, in i Msi! Laboratories. Ithaca, Comstock

poner: Co., 1901. 145 pp., 11 figs.

No. 426.] NOTES AND LITERATURE. 501

The changes to be noted in this edition are not very extensive.

External anatomy is studied in grasshopper and beetle; internal

anatomy, in the larvae of Corydalis and Holorusia; there are two

chapters devoted to comparative anatomy of mouth parts and wings;

there is a brief opening chapter on terminology, and another con-

cluding one on methods of insect histology. The growth of eleven

pages is chiefly due to the (new) chapter on the anatomy of the larva

of the giant crane fly (Holorusia) by Professor Kellogg. This chapter,

proposed as an alterhative to the one on Corydalis, is a very desira-

ble addition whether Corydalis be obtainable or not. Such state-

ments as this, *The internal anatomy of all insects is exceedingly

similar," continue to be repeated in the latest text-books of zoólogy ;

but it would seem that even the average “pure morphologist," for

whom one grasshopper constitutes an entomological summer, should

eventually learn.their absurdity.

In the old chapters there are new paragraphs here and there. In

the study of even such well-worn subjects as the skeleton of the grass-

hopper the discovery of new sclerites still goes on. Thus, in the last

‘edition were noted for the first time sternum and sternedlum, and in

this one we note such new parts as żrochantin of the mandible and

antennary sclerite, etc.

For a simple, straightforward, condensed guide to the laboratory

study of elementary insect anatomy, there is no such book elsewherc.

j. GN,

The Breeding Habits of Cancer magister. — In 1884 Prof. J.

Brown Goode wrote! concerning Cancer magister: “Nothing is

known regarding the spawning season and habits of this species.

'The occurrence of a female with spawn in the San Francisco market

has not yet been recorded by any naturalist.” This last statement

is still true, there being no scientific record, so far as I can learn, of

the capture of this crab while it was carrying eggs. This is rather

remarkable, since Cancer magister is the largest of the edible crabs

of the Pacific coast of the United States, and extends from Sitka on

the north, as far as Magdalena Bay, Lower California, to the south.

In San Francisco Bay and vicinity they are common, and thousands

are annually brought to the markets, where they hold as important

a position as does "e in. the commerce of the. eastern

seaboard.

Y The Fisheries and € Industries of the United States. Section 1, Natural

Animals. 1884.

. -History of Useful Aqua

502 THE AMERICAN NATURALIST. [VoL. XXXVI.

On the 29th of December, 1900, two females carrying eggs were

Mr. J. B. Babcock of the California State Fish Commission. These

were procured in the market and had been caught in San Francisco

Bay. The eggs of one of these were in various stages of gastru-

lation, while the others were older, showing slight pigmentation of

the eyes. During the following summer, while aboard a fishing

steamer off the California coast, I was shown six females “in berry.”

In one of these the embryos were half-developed zoéas, and the

young of another were only slightly more advanced, while the

remaining carried larva almost ready to burst their membranes.

The captain of the vessel stated that he had taken egg-bearing

females in every month of the year, and that the “yellow eggs”

(young stages) occurred from the first of October until some time

in January. This statement is supported by the discovery of seven

other females taken off Moss Landing in Monterey Bay during

January, 1902. All these carried eggs, which were in various

stages, ranging from late gastrulation to half-developed zoéas.

It thus appears that the breeding season of Cancer magister com-

mences in the late summer or during the autumn and usually

continues until some time in the following summer. H. H.

Pratt's Invertebrate Zoölogy.! — This new book by Professor

Pratt of Haverford College consists of a series of directions and

descriptions for the guidance of students in their laboratory work

in connection with comparative anatomical study of invertebrated

animals. The types for study are selected from the following groups

and in the order named: insects, myriapods, Crustacea, annelids,

platyhelminths, Bryozoa, mollusks, ascidians, echinoderms, Hydrozoa,

sponges, Protozoa. Several types from each of the important phyla

are treated, and thus within groups the practical study is made truly

comparative. This is important, for such comparisons. as untrained

students can make for themselves are necessarily limited in the

usual series of types with a single representative of each important

phylum. Another valuable feature of these practical directions is

that each lesson is complete in itself, so that the sequence of types

may be varied at the will of the teacher. This is also of advantage

in selecting the exercises of which the book contains more than

ordinarily can be accomplished in a single year's course. The

author suggests the possibility of beginning with the last lessons on

1 Pratt, H. S. Znvertebrate Zoology. Boston, Ginn & Company, 1902. 210 pP-

No. 426.] NOTES AND LITERATURE. 503

the unicellular forms and working upward to the higher animals,

but he prefers the arthropods for the introductory work because they

are so convenient for teaching the fundamental principles of

comparative anatomy.

Apparently there is no special reason for the position of the lesson

on the ascidians between those on the mollusks and echinoderms.

Probably most teachers will prefer to place this type at the end of

the course on invertebrates, when there is a natural transition to

the study of the vertebrates.

In the plan of the laboratory directions there is a general resem-

blance to the well-known guides by Huxley and Martin, and Marshall

and Hurst; but there is a great and important difference in that

there is not placed before the student a description so complete as

to limit the laboratory work to that of merely examining the natural

objects in order to verify the printed statements. On the contrary,

we find a judicious combination of description of the difficult and

time-wasting points with practical directions for suggested problems

which are not beyond the student's ability and time for investiga-

tion. Such an arrangement gives a desirable mean between two

common extreme methods of laboratory teaching, — the one aiming

to inform the student through verification of quite complete descrip-

tions, the other attempting to stimulate originality and investigation

by leaving him largely dependent upon the natural materials, with a

minimum of direction. A combination of the two methods ina labo-

ratory manual will be welcomed by all teachers who believe that the

aim of zoólogical instruction is not only to furnish students informa-

tion concerning the science, butalso to give them practice in acquiring

some knowledge through their own investigations.

In addition to the practical directions, the book contains a useful

appendix on classification of animals and one with short definitions

of the principal groups. An excellent glossary of zoólogical terms

serves also to indicate where they occur in the lessons ; and there

is a general index.

In all essential respects this new laboratory manual appears to be

excellent, and it deserves the attention of all teachers who use

invertebrate types in their courses of zoólogy. u AER

Notes on Birds. — A very good local list of the birds of north-

western Montana is presented in he Summer Birds of Flathead

Lake, by P. M. Silloway, issued as a bulletin of the University of

Montana. It includes an annotated list of 128 species noted between

504 THE AMERICAN NATURALIST.

June.s.and August 29, and notes on the nests and eggs of about

twenty-four common species. ‘There are sixteen excellent plates

made from photographs of the nests and eggs. The list will be of

particular value to western ornithologists, but it is also of interest to

those in the East who care to learn something of the western repre-

sentatives of some of our eastern forms.

"The Proceedings of the Nebraska Ornithologists’ Union at its Second

Annual Meeting, held in Omaha, Jan. 12, 1901; contains not only

a very creditable number of interesting papers, but gives evidence

of an energy and ability among the leaders of the union that

augur well for the future of ornithology in the state. The papers

presented cover an unusually wide range ; they include an historical

account of thé formation of the union ; ; a list of the breeding birds

of the state ; accounts of the nesting of Bell's vireo ; accounts of

young birds in captivity and in the nest ; a description of a peculiar

disease of birds' feet, and an account of internal parasites of

Nebraska birds; and papers dealing with methods of teaching

bird study in schools and of keeping migration records. A short

paper on the blue jay falls so far below the level of general excel-

lence as to emphasize the serious tone of the other papers. The

paper on Nebraska records gives assurance that the interesting

problems of distribution which the position of the state presents

will be worked out with care and patience.

The Smithsonian lastiution publishes a reprint of E Oberbelsen s

* Catalogue of a Collection of Humming Birds from Ecuador and

Colombia," from the Proceedings of the United States National Museum.

"The catalogue contains descriptions of one new species and three

new subspecies, besides many of the collectors' field notes.

CORRESPONDENCE.

To the Editor of the American Naturalist :

Sin: In his article in the May Naturalist on the “ Structure and

Classification of the Tremataspidzs," Dr. William Patten returns again

to the question of the origin of the vertebrates from a supposed

arthropod ancestor. It will be remembered that in two earlier

papers! he compared the sutures and other markings on the head

shield of trilobites with those on the dermal armor of Pterichthys and

Bothriolepis ; and we are confident that paleontologists will dissent

as strongly from his latest as they did from his earlier conclusions

that ostracoderms and arthropods are genetically related. The

view expressed by him about eight years ago, that the three-layered

dermal skeleton of ostracoderms is a modification of that in arthro-

pods similar to Limulus, also lacks confirmation from other sources.

His present procedure, however, is of the boldest; for on the start-

ling assumption that ostracoderms were animals having many pairs

of jointed appendages, — on merely a suspicion that this was true, —

he finds it necessary to “create for them a new class, one that shall

occupy a position between the true vertebrates and arthropods, and

unite these two great groups into one compact phylum." ;

His account of the habits, development, and even some of the

morphological characters of these * arthropod-like animals" is of

such extremely speculative nature that few can hope to read with

him the unwritten records of the past. For if it be not by intui-

tion, how else can one know that ostracoderms originally progressed

*through the soft mud on the bottom of shallow water with the

usual position of dorsal and ventral sides reversed” ; that after leav-

ing the bottom they righted themselves into the true vertebrate posi-

tion; and that this acrobatic performance was accompanied by a

migration of the eyes from the haemal to the neural surface of

the body? Or in what antediluvian aquarium, we wonder, was

the locomotion of Eurypterus observed to be “by brief, spasmodic

excursions"? And what authority have we for supposing that the

“ hypostomeal eyes of trilobites” (Lindström) were in the slightest

! Quart. Journ. Mier. Sci., vol. xxxi (1890), pp- 3597365; Anat. Anzeiger, vol. ix

(1894), pp- 429-438.

505

506 THE. AMERICAN NATURALIST. | (Vor. XXXVI.

sense visual organs, or, in fact, anything but mere muscular

impressions ?

But these are trite criticisms in comparison with what might be

said respecting Patten’s remarkable declaration that ostracoderms

possessed many pairs of jointed appendages. To refute this is to

nullify the author’s sole new claim that these creatures are “ genet-

ically related to arthropods, or that, through changes in structure and

function, one type has been derived from the other"; for all other

of the well-known resemblances may be explained as due to mimicry,

or to incidental parallelism dependent on environment.

We have already characterized this assertion of Patten's as a sur-

mise; we will go further, and say that it is not only an unproved

hypothesis, but one that is absolutely unsupported by any specimens

that have yet been described. If examples of Cephalaspis have

been found which display “a fringe of 25 to 3o pairs of jointed and

movable appendages along the ventral margin of the trunk," the

present reviewer for one will confess that he has never seen nor

heard of them. As for * Lindstróm's important discovery of an

appendage in Cyathaspis," this detached and unjointed fragment is

probably to be interpreted as a spine or cornu. But Professor

Patten does not stop here: he points to a series of marginal openings

in the shield of Pteraspis, Cyathaspis, and Tremataspis, commonly

regarded as branchial, and declares that they “must” have served for

the attachment of appendages. He surmises that in Tremataspis

the latter ** decreased in size from before backwards, and were possibly

too delicate to be well preserved in a fossil condition." Fearful lest

our rude touch should annihilate these frail conjectural organs, we

will simply refer those interested to Dr. Traquair's comparison of

Tremataspis with Birkenia, in which the branchial openings (com-

monly so-called) are disposed relatively as in the skate and shark.’

A comparison, also, of the anterior ventral plates of Tremataspis

with the very similar ones of Drepanaspis, in our opinion would

have been far more instructive than our esteemed friend's attempt

to homologize certain of them with the jaws of an arthropod.

"The position of the mouth in Tremataspis we shall believe, until

the contrary is proved, to have been correctly determined by Rohon,

who places it immediately behind the rim of the head-shield, as in

other ostracoderms where the oral plates are satisfactorily known.

Patten, however, basing his conclusions on the solitary example

: studied by Rohon, challenges the latter's interpretation, and would

1 Trans. Roy. Soc. Edinb., vol. xxxix (1899), pt. iii, p. 859.

No. 426.] CORRESPONDENCE. 507

have us suppose that one of the plates of the ventral armor functioned

as oral. The author’s employment of such generic and family mis-

appellations as “ Tolypaspis " and “ Pterichthyda ” respectively, is

contrary to all recognized principles of nomenclature.

That part of Professor Patten’s article which embodies a redescrip-

tion of the dorsal and ventral shields of Tremataspis, based on four

or five unusually perfect specimens from the Isle of Oesel, possesses

great merit, and paleontologists should be truly grateful to him for

having increased our knowledge of this primitive chordate in several

important respects. The more extensive memoir on the same sub-

ject which is promised will be awaited with great interest; and

further enlightenment with regard to the gill openings and position

of the mouth in ostracoderms will be particularly welcome.

C. R. EASTMAN.

CAMBRIDGE, MASS.

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BALDWIN, J. M. Fragments in Philosophy and Science, being Collected

Essays and oe m York, Scribners, dues xii, 389 pp. 8vo. $2.50.

AMPBELL, D. H. A University Text Book of Botany. New York, Mac-

millan, 1902. xv, 579 pp., 8vo, 15 pls., 493 tex is $4.00. — CosTANTIN, M.

L'hérédité acquise. Les conséquences horticoles, e er et médicales. Scien-

tia: Biologie, No. 12. Paris, C. Naud, 1901. 86 pp. 8vo. 2 francs. — DANNE-

MANN, F. rundriss einer Geschichte der ilius ssenschaften zugleich eine

Entführung in dei Studium der grundlegenden inibi mo s Litteratur.

Bd. i, zweite spon Leipzig, Engelmann, 1902. xiv, 422 pp., 8vo, 57 figs

9 marks. — FRY . E. Grammar School Geography, with Supplinwat by

W. M. Davis. cea Ginn & Company, 1902. viii, 195 pp.; viii, 30 pp., 4to. —

Honce, C. F. Nature Study and Life. Boston, Ginn & Company, 1902. xvi,

514pp»8vo, ro6figs. $1.75. — MACDOUGAL, D. T. Elementary Plant Physiology.

New York, Longmans, Green & Co., 1902. 138 pp., 8vo, 108 figs. — SACHA-

ROFF, N. Das Eisen als das thätige Tess dus Enzyme und der lebendigen

Substanz. Jena, Fischer, 1902. 83 pp., 8vo, 2 pls. 2.50 marks

ALLEN, J. A. A Further Note on the Generic Names g^ ue Mephitinz.

Proc. Biol. Soc. Wash. Vol. xv, pp. 59-66. — ALLEN, G. M. e Mammals of

Margarita Island, Venezuela. Proc. Biol. Soc. Wash. va ZW pp oru

BAKER, F.C. The Mollusca of the Chicago A The Gast da. Bull. Chic.

Acad. Sci. No. 5, Pt. ii, pp. 136-418, Pls. Xxviti- XXXVI. — BANKS, N. Papers

from the Hopkins Stanford iocum at youre VII. Entomological Results,

(6) Arachnida. Proc. Wash. Acad. Sci. Vol. iv, pp. 49-80, Pls. I-IT I. — COCKER-

s m A. A New Currant from Arizona. Proc. Biol. Soc. Wash. Vol. xv,

— Dat, W. H. Illustrations. dod Descriptions of New, Unfigured, or

impact TEE Shells, chiefly American, in the U. S. National Museum. Proc.

U.S. Nat. Mus. Vol. xxiv, pp. 499-566, Pls. XXVII-XXXIX. — ECKEL, E. C., and

PAULMIER, F.C. Catalogue of New York pets and Batrachians. Bull. N.Y.

State Mus. No. 51, pp. 355-414. 1 pl, 24 text-figs. — EIGENMANN, C. H., and

KENNEDY, C. H. The Leptocephalus of the cu Eel and Other American

Leptocephali. li ul. U.S. Fish Com. for igor. pp. 81-92. 14 figs. — EVER

met B. w. List of o Fishes known to occur in the Great Lakes or

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River. Rep. U.S. Fish Com. for rgor. pp. 209-240. — EVERMANN, B. W., an

. GorpsBoRovcH, E. L, A. Report on Fishes collected in Mexico and Central

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Jorpan, D. S., and SNYDER, J. O. A Review of the Zalnoid Fishes and Related

Forms found in the Waters of Japan. Proc. U. S. Nat. Mus. Vol. xxiv, pp. 595-

662. 10 figs. — JoRDAN, D. S., and SNYDER, J. O. A Review of the Hair

— of Japan. Proc. U.S. Nat. Mus. Vol. xxiv, pp. 567-593- 5 figs. — Jo.

N, D. S, and SNYDER, J. O. A Review of the Trachinoid Fishes and their

Serco Allies found in the Waters of Japan. Proc. U. S. Nat. Mus. Vol xxiv. —

p. 461-497. 7 figs. — KENDALL, W. Notes on the Silversides of the Genus

Manila of the East Coast of the United States, with Descriptions of Two New

Subspecies. Rep. U.S. Fish Com. for 1901. pp. 241-267. 6figs.— Lyon, M. W., Jr.

Description of a New Phyllostome Put t Rol the Isthmus of Panama. Proc. Biol.

Soc. Was

oc. Vol. xv, pp. 83-84. — Merriam, C. H. Tw o New € from the

Alaska Pask. Proc. Biol. Soc. Wash. Vol. xv, pp- 77-79- — MERRIAM, C. H.

A New Bobcat (Lynx Uinta) from the Rocky Mountains. Proc. Biol. pus Wash.

Vol. xv xv., pp. 71-72. — MERRIAM, C.H. Two New Shrews of the Sorex Tenellus

e Bio

p. 67-69. — MILLER, G. S, Jr. A New Rabbit from Southern Texas. Proc.

Biol. Soc. Wash. Vol. xv, pp. 81-82. — OSBORN, H.,and BALL E.D. A Review

of the North American Species of Athysanus (Jassidz). Bull. Ohio State Univ.

Ser. 6, No. 14, pp. 231-256, Pls. X VI-X VII. — RICHMOND, C. W. List of Generic

Names proposed for Birds during the Years 1890 to 1900 Inclusive, to which are

added Names omitted by Waterhouse in his * Index Generum Avium." Proc.

U.S. Nat. Mus. Vol. xxiv, pp. 663-729. — RIGGS, E. S. The Dinosaur Beds of

the Grand River Valley of Colorado. Field Columb. Mus. Publ. No. 60, Geol. Ser.

Vol. i, No. 9, pp. 267-274, Pls. XXXIV-XX XIX. — Rices, E. S. The Fore Leg and

Pectoral Girdle of Morosaurus, with a Note on the Genus Camarosaurus. Field

Columb. Mus. Publ. No. 63, Geol. Ser. Vol. i, No. 10, pp. 27 5-281, Pls. XC-XCII. —

SurrH, H.M. Description of a New Species of Blenny from Japan. Bull. U.S.

Fish Com. for rgor. pp.93-94. 1 fig.— Situ, H. M. Notes on the Tagging of

Four Thousand Adult Cod at Woods Hole, Massachusetts. ef. U.S. Fish

Com. for rgor. pp. 193-208. — SMirH, H. M. Note on the Scotch Methods of

mene Haddocks. s U.S. Fish Com. for 1901. pp. 269-271. — VAUGHN,

. W. Some Recent Changes in the Nomenclature of West Indian Corals.

Proc. Biol. Soc. Wash. Vol. xv, pp- 53-58. — WASHBURN, F. L. Oregon Water

Supplies. Examination of Several Samples of Drinking Water. The Process

described. Rep. Oregon State gendi, fer rgor. Appendix B. 10 pps 3 pls.—

510 THE AMERICAN NATURALIST.

Witson, H. V. The Sponges collected in Porto Rico in 1899 by the Fish Com-

mission Steamer Fish Hawk. Bull. U.S. Fish Com. for 1900. Vol. ii, pp. 375-

411. ;

Bulletin jn Hopkins Hospital. Vol. xiii, Nos. 133, 134. April and May.

— Entomological Society of Ontario. 8 Ann. Rep. for rgor. Toronto, 1902.

128 S 56 figs. — Higher Science. March. — Journal of Geography. Vol. i,

- 2-3. February and March. — Naturae A e Jahrg. xxiii, r9or. Berlin,

Vdeltéider, Igor. 498vo. 4 marks. — Rp of the s Science

Association of Staten posts Vol. viii, Nos. 12, 13. Sami and M

evista Chilena de Historia Natural. Año v, N OS. II, I2. — Revista de Pies

Habana. Afio ii, No.2. Febr ruary, 1902.

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— I AGaseolAbnormality n Cate Paws .

" II. Th M ce sa E See ee VAR, di rie io mm

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THE

AMERICAN NATURALIST

Vor. XXXVI. July, 1902. No. 427.

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L. MARK, DIRECTOR. No. 134.

A CASE OF ABNORMALITY IN CATS' PAWS.

FREELAND HOWE, Jn.

THIs paper contains an account of facts learned by the

study of the walking pads, the muscular, vascular, nervous,

and skeletal systems of the manus of a polydactyle and syn-

dactyle cat. The cat furnishing the material for this study

Was one of a strain of polydactyle cats living in Cambridge,

Massachusetts, and descended from a polydactyle cat which

lived at the astronomical observatory of Harvard University.

All four legs of the cat dissected were preserved in alcohol

and were kindly given me for study by Dr. C. B. Davenport,

under whose direction the work was done. To him and to

Prof. E. L. Mark I wish to extend my thanks for kind advice

and criticism.

Each fore paw had six toes; the toes of the hind paws were

fused in pairs almost to the ends of the claws, as is shown in

Figs. 1 and 2.

SII

512 THE AMERICAN NATURALIST. [Vor. XXXVI.

The syndactyle hind paws have not been dissected, but a

drawing of the palmar and dorsal surfaces is shown in Figs.

I and 2 respectively. Walking pads and distal phalanges

both indicate that the four digits are fused in pairs.

I wished to determine, if possible, which toe of each fore

paw is the extra one, and what is the nature of the evidence.

I have been unable to learn of any account of a study on

material precisely similar to this. The collection of facts in

Bateson's “Materials for the Study of Variation” covers the

ground worked over by previous authors, and this, with Poul-

ton's papers, includes an account of all the published facts on

the subject, as far as known to me. Previous study has been

only on the walking pads and

on the skeleton. Bateson

described principally skeletal

structures, and Poulton studied

the walking pads. They have

drawn their conclusions as to

the true nature of polydactyl-

ism from facts learned from

these two organs only. It is,

of course, desirable to have as

Fio. 2. — Dorsal aspect of hind paw of abnormal complete a knowledge as pos-

cat, showing claws fused in pairs.

sible of the anatomy of the

paw, as a basis for any conclusions to be drawn. I find from my

dissections that the evidence furnished by the walking pads and

skeletal system is borne out by the other organs mentioned.

The general appearance of the polydactyle paw studied, apart

from the increased number of digits, is quite different from

that of a normal fore paw. The radial digit in the polydac-

tyle paw extended much nearer to the end of the paw than

the pollex normally does, the digits being more nearly of the

same length than in the normal paw. A comparison of the

relative positions of the ungual and middle phalanges during

the retraction of the former will be made when we come to

the study of the bones.

The walking pads of the polydactyle paw (Fig. 4) differ from

those of the normal paw (Fig. 3) in that the phalango-metacarpal

Fu: Fic. 2.

Fic. 1. — Palmar aspect of hind paw of abnormal

cat, showing fused pads.

No. 427.] ABNORMALITY IN CATS’ PAWS. S43

pad (pulv.phix-mt'carp.) of the polydactyle is five-lobed

instead of three-lobed as normally, and the lobes in the

polydactyle paw are more clearly marked off from one another

than in the normal paw. The parts of the five-lobed pad

which appear to correspond to the phalango-metacarpal pad

of the normal paw are the three external lobes which are

more closely associated with one another than with the two

internal lobes; while the two internal lobes are more closely

applied to each other than to the three external ones. That

Fic. 3.— Palmar aspect of right fore paw of normal cat.

Fic. 4. — Palmar aspect of right fore paw of polydactyle cat. A, radial digit; B, second

digit, etc. ; AE qeedbs II, index; ZZZ, medius; ZV, annulus; V, minimus; J£x.5P 7. dg.,

flexor sublimis digitorum; 7v. 9A/x., phalangal pads; pulv.phix.-mt’carp., phalango-

metacarpal pad; fuly.fis., pisiform pad.

five lobes instead of three existed in the polydactyle paw is

probably due to the fact that one lobe was added by the for-

mation of an extra digit, and that the growth of the radial

digit, which occurred to so unusual an extent that it func-

tioned more like a walking digit than the pollex normally

does, excited the growth of a phalango-metacarpal pad on

this digit also. In both the normal and the polydactyle paw

there is a walking pad on the distal end of each middle pha-

lanx (pulv.phix.) and on the pisiform bone (pulv.pis.).

514 THE AMERICAN NATURALIST [Vor. XXXVI.

MUSCLES.

As in the normal fore paw, the muscles extensor carpi radi-

alis longior and extensor carpi radialis brevior (Figs. 5, 6,

ext.carp.r.lg. and ext.carp.r.brv.) are inserted on the proximal

dorsal surface of the second and third (counting from the

radial side) metacarpal bones respectively.

Considering the evidence of these two muscles alone, we

should infer that in the polydactyle paw the radial side is nor-

mally formed and that the extra digit is external to (on the

- -|- Li ext.com.dg. --2-- |

-H -|-|--- ---extmindg. ---- T

-RE----- eiicasp tinm. o

ho hr,

ext.carp.r bro

DIR uem a extcazp.rdg.-----

| IV

A JT

Fic. 5. Fic. 6

Fic. s.— Dorsal aspect of left manus of normal cat, showing dissection of muscles. ext.carf.r.

. extensor carpi ialis brevior; ext.carp.rig., extensor carpi radialis longior;

igitorum ; ex.

radial

ext.carp.uin., extensor carpi ulnaris; ext.com.dg., extensor communis

min.dg., extensor minimi digiti.

Fic. 6. — Dorsal aspect of left manus of polydactyle cat, showing dissections of same muscles

ulnar side of) the third one, for in the polydactyle paw there

are three digits external to the insertion of the extensor carpt

radialis brevior, whereas in the normal paw there are only

No. 427.] ABNORMALITY IN CATS’ PAWS. 515

two. Such an inference, however, is not borne out by the

evidence of other muscles and tissues.

Muscles extensor communis digitorum and extensor minimi

digiti (Figs. 5, 6, ext.com.dg. and ext.min.dg.) in both normal

Fic. 8.

Fic. 7. l asp f right f normal cat, showing di tion of deep muscles.

ext.ix., extensor indicis; ex£.»f'carf. poll., extensor metacarpi pollicis.

Fic. 8.— Dorsal f righ f polydactyle cat, showi deep muscles as in

Fig. 7- (See text for description of ext.ix.r and ext.iz.2.)

and polydactyle paw extend to the proximal dorsal surface of

the middle phalanx of each of the four external digits. The

distribution of these muscles points to the inference that the

516 THE AMERICAN NATURALIST. [Vor. XXXVI

four external digits of the polydactyle paw correspond to

the external four in the normal cat, and that the extra digit

occurs on the radial side of the four external digits, an infer-

ence which, with some modifications, seems to be confirmed

by other facts.

Muscle extensor carpi ulnaris (Figs. 5, 6, ext.carp.uln.) in

both normal and polydactyle paw is inserted on the ulnar

metacarpal.

^. The indicator (Fig. 7, ext.2x.) is somewhat variable in respect

to its insertion, even in normal specimens, sometimes supplying

the pollex, index, and medius, sometimes only the pollex and

index. In the polydactyle paw a peculiar arrangement exists,

in that there are two muscles in place of one. That which,

from relative position and insertion, seems to correspond to

the normal one (Fig. 8, exzzx.7) is distributed to the second

(5) and third (C) digits.

In addition to this muscle there is under it, and distinct

from it, another muscle (ext.2x.2), which passes in the same

general direction to the two internal digits (A and B, Fig. 8).

This muscle has a more distal origin than does the one sup-

plying digits B and C; it originates from the dorsal border

of the ulna and passes directly over the muscle extensor meta-

carpi pollicis (ext.mt' carp poll). Comparisons of the two indi-

cators with each other and with the normal muscle seem to

point to a readjustment to meet a new condition of the manus.

In both the normal and the polydactyle manus the radial

metacarpal furnishes insertion for the extensor metacarpi polli-

cts (Figs. 7, 8, ext.mt carp poll).

Muscle flexor carpi radialis in both the normal and the poly-

dactyle paw has its tendons inserted on the proximal ends of

the palmar surface of the second (counting from the radial

side) metacarpal.

he ulnar part of the flexor sublimis digitorum in both

normal and polydactyle paws has tendons extending to the

first and second digits, counting from the z/zar side (Figs.

3, 4, Jix.sb Ldg., while the tendons from the radial part of

the muscle extend in the normal paw to the four, and in the

polydactyle paw to the five, digits nearest to the radial side.

: No. 427.] ABNORMALITY IN CATS’ PAWS. 517

The fact that in the polydactyle paw the union between the

tendons to the digits marked /V and V (Fig. 4) extends fur-

ther distally than in the normal paw (Fig. 3) is apparently

less important than the fact that in the polydactyle paw the

radial part of the muscle has five tendons while the normal

paw has only four.

Muscle flexor profundus digitorum (Figs. 9, 10, fix. profnd.dg.)

has in the normal paw five, and in the hexadactyle paw six,

£ 1 2 E : A +i f deen muscles

Fic. 9. — Palmar aspect of left

Fic. 10.— Palmar aspect of left manus of polydactyle cat, showing mus g. 9

A, radial digit of polydactyle manus; 7, that of normal manus; posuer dg., flexor

profundus digitorum; Zrsór.1-4, uet Pa 1-4.

tendons distributed one to each digit. There is no evidence

here as to which digit is the extra one.

Muscle flexor carpi ulnaris is inserted on the pisiform bone

in both normal and polydactyle paws.

There are four /umbricales in both normal and polydactyle

paws (Figs. 9, 10, Zór. 7-4). In both cases these are inserted

one each on the radial side of the proximal phalanx of each

of the four external digits. In the polydactyle paw there is

no lumbrical superficial to the radial border of the distal part

of the flexor profundus digitorum.

5 18 THE AMERICAN NATURALIST. [VoL. XXXVI,

Broop VESSELS.

The arrangement of the veins on the dorsal surface of the

normal and the polydactyle fore paw is shown in Figs. 11 and 12.

The only points that can be taken as homologous for deter-

mining the corresponding veins in the normal and polydactyle

paw are the most distal point of the loop formed by the anas-

tomosis of the ulnar (v.7/7.) with the radial (v.7.) vein. This

point in both normal and polydactyle paws seems to be between

digits marked /// and

IF. Starting from this

are three branches, and

in the polydactyle paw

four branches, which

contribute to the radial

vein. These facts

suggest the conclusion

that the extra digit of

the polydactyle paw is

on the radial side; but

owing to the uncer-

Fic. rr.

Fic. 11. 1 aspect of lef t howing tainty of the exact

shea veins. v.udn., ulnar vein; v.r., radial vein. position of this point

ie Wem S src Pelion left manus of polydactyle cat, of reference, the evi

dence from the veins

is of less value than that from some of the other organs.

In studying the arteries the only fact which throws any light

on the problem is the relative size of the digital branches from

the palmar arch (Figs. 13, 14). The branch which supplies

the radial digit is small in both the normal and the polydactyle

paw. In the normal paw the branch to the index is as large as

the branches to each of the other three digits; but in the poly-

dactyle paw the branches to digits 4 + B and B + C (Fig. 14),

while about equal to each other in size, are much smaller than

those to the external digits.

No. 427.] ABNORMALITY IN CATS’ PAWS. 519

NERVES.

The radial nerve after becoming subcutaneous follows the

courseiand distribution of the dorsal veins, which are shown

in Figs. r1 and 12. The median nerve (Figs. 15, 16, n.n.)

besides supplying the muscle flexor communis digitorum is dis-

tributed to four consecutive digits, beginning with the radial

side, in the normal and to five consecutive digits in the poly-

dactyle paw. In the normal manus the z/zar nerve divides,

just below the olecranon, into an inner (ventral, 7.2/z.v.) and

an outer (dorsal, z.zZz.d.)

branch (Fig. 15). The

outer (dorsal) branch

passes to the outer side of

digit V and also sends dor-

sally a branch to the inter-

nal side of the same digit

and to the external side of

digit 7V. In both normal

and polydactyle manus the

internal (ventral) branch

of the ulnar (Figs. 15, 16,

n.uln.v.) supplies the three

external digits. Fic. 13. — Palmar aspect of left manus of normal cat.

art.r., radial artery.

For a point of reference

in comparing the nerves we

may takea small branch which passes from the uzarto the median

nerve. This unites with that branch of the median which in

the normal manus (Fig. 15) passes to the ulnar side of digit 7/7

and to the radial side of digit ZV. Zn the hexadactyle manus

(Fig. 16) the branch of the median nerve which is joined by the

ulnar nerve likewise passes to the ulnar side of digit III and to

the radial side of digit IV. There is, then, in the abnormal

manus radially to the nerve of reference one more digit than

exists in the normal manus. This makes it apparent that the

modification producing polydactylism has occurred on the

radial side of the manus.

Fic. 14. — Palmar aspect of left manus of polydactyle cat.

520 THE AMERICAN NATURALIST. [VoL. XXXVI.

SKELETON.

The ulnar, radial, and carpal bones are practically the same

in number, relative size, shape, proportion, etc., in the normal

and abnormal manus. There are in the polydactyle paw six

metacarpals, a pollex of two phalanges, and five

digits, each with three phalanges. All the bones

of the normal paw (Fig. 17) are larger and stouter -

than those of the polydactyle paw (Fig. 18)

Bones of a normal paw were prepared for the

ES? purpose of making comparisons with the abnor-

mal skeleton in respect to weight, relative pro-

portions, etc. In making these comparisons,

allowances are made for the general differences

mentioned above.

The radial sesamoid of the carpus of the six-

toed cat (ses., Fig. 18) is fused to the radial side

of the scapho-lunar

(scph-lun.) and furnishes

the place of articulation

for the metacarpal of the

pollex. In the polydac-

tyle manus the four

external metacarpals

(mt carp.) have the same

articulations with the

distal row of carpals as

in the normal manus. In

the normal manus the

Fic. 15. — Palmar ured of right manus of normal cat. Pp ollex articulates with

n.m., median nerve; z.u/n.d., dorsal ot una the trapezium (trz.)

nerve; sz.uln.v., ventral ban of ulnar n E A

Wii. 5. Palmar aspect of Hight manta of pei xm a, While in the abnormal

| manus (Fig. 18) the

metacarpal of digit B eee with the trapezium. Metacarpal

of digit 4 (Fig. 1 18) arti ly stated, with the radial

sesamoid (ses.), sh i is fused w the sapbolonar (scph-lun.).

The metacarpal bones in both manus are similar, except that

there is in the abnormal manus no “groove” for the radial

Fic. 15. Fic. 16,

No. 427.] ABNORMALITY IN CATS’ PAWS. 521

artery. The metacarpals, as well as the other bones of the

pollices, vary considerably in length and thickness, and in this

respect will be con-

sidered more closely

later.

There is no par-

ticularly noticeable

variation in the proxi-

mal series of pha-

langes (PAZx.przx.).

These bones have the

least distinctive char-

acters, differing from

one another chiefly in

size,

The series of mid-

dle phalanges (phéx.

m.) furnishes very

interesting condi-

tions. In both the

normal and abnormal

manus the three ex-

ternal (ulnar) middle

phalanges are carved

away on the ulnar

side to allow for the

retraction of the cor-

responding ungual

phalanges (phlx.ung.).

The middle phalanx

Fic. 17- . Fic. 18.

Fic. 17. — Dorsal aspect of skeleton of left manus of normal cat.

of the index of the Fic. 18. — Dorsal aspect of left manus of hexadactyle cat. A, radial

normal manus is like- digit of — ma) I, that of voies manus ; d

wise carved away on pals; xz., uncinate; os mag., os magnum; fALr.»., middle

ges; flr.

the ulnar side, but in phalanges; pæ pisiform; r, radius; scph-lun., scapho-lunar ;

thicábhnormal fanus — 7" sesamoid ; £rz., trapezium; £rzd., trapezoid ; w/m., ulna.

the digit (C, Fig. 18) next to the three ulnar digits is carved

away on neither side and is therefore “ indifferent." The middle

phalanx of digit B (Fig. 18) has no counterpart in the normal

522 THE AMERICAN NATURALIST. [VoL. XXXVI.

manus, except that it is somewhat like a pollex, which, when

retractable, as occasionally happens, is carved away on the

radial side.

The ungual phalanges are too nearly alike in the normal and

the polydactyle paw to be of service as far as our purpose is

concerned.

GENERAL CONSIDERATIONS.

The facts learned from the dissection of the polydactyle

manus are in part contradictory; that is, some facts suggest

that the extra digit occurs on the radial side of the paw,

whereas others indicate an extra digit on the ulnar side; but,

on the whole, the balance of the evidence points to the exist-

ence of the extra digit on the radial side of the three ulnar digits.

The evidence which the bones furnish seems to be the most

satisfactory, and is borne out in the majority of cases by the

other tissues. The middle phalanges of digits ///, /V, and

V (Figs. 17, 18) are carved away on the ulnar side, as nor-

mally. The middle phalanx of digit C (Fig. 18) is an indif-

ferent digit, Z.e., carved away on neither side. In this it more

resembles a pollex than a digit. The middle phalanx of

digit B (Fig. 18) is carved away on the radial side, in this

respect resembling a pollex more than a digit. Digits ///, /V,

and V in both manus are distinctly similar. Regarding the

articulation of the metacarpals with the carpals, it is seen that

the five external digits of the abnormal manus have the same

articulation as do the five digits of the normal manus. The

pollex of the abnormal manus has the articulation abnormal, in

that it is with the radial sesamoid, which in this case is fused

with the scapho-lunar. Here the five digits nearest the ulnar

side are normal

Considering the three ulnar digits of the abnormal manus to

correspond to normal digits, one is naturally led to inquire what

modifications the manus has undergone that there should be

three digits instead of two on the radial side of the three ulnar

normal ones. Further evidence from the bones is interesting

in respect to this query.

No. 427.] ABNORMALITY IN CATS’ PAWS. 523

A comparison of the indices and the weights of the indi-

vidual bones of both the normal and abnormal manus is shown

in the accompanying table. For the purpose of more accurate

comparison of the two sets of bones, I have figured the per-

centage which each bone bears to the total weight of all the

bones of the manus to which it belongs (including ulna and

radius). The indices of the bones were obtained by finding

the ratio between the minimum thickness and the maximum

length of each bone. The actual points of measurement are

shown by the dots in Fig. 17, ZZZ. Inspection of the bones

first leads us to see that digit A (Fig. 18) is much longer than

the normal pollex and presents in the undissected manus more

the character of a finger than of a pollex. The bones, however,

are distinctly those of a pollex, since there are two phalanges

only; but they are longer and more slender. In this respect

they resemble a finger. The bones of digit B (Fig. 18) are more

like a pollex than are those of the digit which normally adjoins

the pollex. One can also see that digit C (Fig. 18) is more

like a pollex than is the digit which is normally fourth from the

ulnar side. What further facts are there to bear out the idea

that where there are normally two digits, there abnormally occur

three, each of which partakes somewhat of the general charac-

ters of the others ?

It is a fact that where normally two digits are found, namely,

a pollex and an index, there are found in this special case

three, and that the material which would normally form two

digits has so distributed itself that each of the three digits

which actually occurs partakes of the nature of the other two.

Is the abnormal pollex two-thirds pollex and one-third index ;

the digit next the pollex, one-half index and one-half pollex ;

and the digit C (Fig. 18) two-thirds index and one-third pollex?

There seems to be some relation of this sort.

If we compare the indices and the relative weights of the

two manus, we come to a like conclusion (see accompanying

table). For example, comparing the percentages of total

weights in the abnormal manus, metacarpal A (Fig. 18) is more

like the other metacarpals of its manus than is metacarpal 7 of

the normal manus like the other metacarpals of its manus;

524 THE AMERICAN NATURALIST. [VoL. XXXVI,

NORMAL.

GRAMS PERCENTAGE | MILLIMETERS | PERCENTAGE

WEIGHT. WEIGHT. MEASUREMENT. INDEX.

PS a att tact ao 3.467 333 45 X 1030 0437

Ea EO Noo uis 2.679 257 52 x 878 0592

Mapai a .699 .067 — =>

Metacarpals

E 135 O12 32 * riz .286

II 423 .040 28 x 284 .0986

III 525 054 34 x 321 0997

IV 445 043 30 X 300 100

V 348 033 28 x 247 113

Totals of metacarpals . 1.876 .182 — as

Prox Y Tv

‘ f 0092 a2 x 52 402

ME ga ee sa ae .157 .O1 58 22 X 136 235

BRL. bg ec un .188 0187 33 X 1$5 .203

IE QE IUE LIN -161 0154 4o X 547 .204

Mornok RA oy RU 47 .0122 20 X PES 245

oe of proximal Er

ges. 220 :0713 — emp

Middl phalanges

1. Te 097 .0093 23 X 94 244

III 108 0103 23 X 116 200

IV 094 -0090 23 X 108 213

v 078 0075 28 x 82 341

ora e middle dini

: 37 0361 — aes

Pig phalanges

f. .150 0144 "4 X 104 333

WW. iw. ui. 116 OIE 30 X 100 300

MEI. QU ay 12$ .0120 28 x 100 |. .280

IN uc d uv ust .102 .0098 28 x 101 .278

Na iir y .o089 .0086 28 x 93 -301

Totals of distal pee

ge. 582 -0559 "€ e

CM 1M all bones en 11.030 — — =

No. 427.] ABNORMALITY IN CATS’ PAWS. 525

POLYDACTYLE.

RAMS PERCENTAGE | MILLIMETERS | PERCENTAGE

WEIGHT. WEIGHT. MEASUREMENT. | INDEX.

Ina . 2.829 330 40 X IOI5 0394

Radius . 2.188 255 42 x 850 o

Carpets í .529 061 —

Metacarpals

A. .097 OII 20 X 157 .127

B. .261 .030 27 X 231 113

C: 295 .034 26 X 260 .100

EH .292 034 26 x 278 .094

IX. .291 .034 4 X 260 .o91

V. im .02 at X 414 118

Totals of metacarpals L473 170 xm mU

Proximal phalanges

NES uou 054 0063 I9 X 103 .184

B. 123 0144 31 X 120 .258

Ga 123 0144 30 X 126 .238

Hi. 133 o166 28 x 140 .200

IV. 123 0144 27. X 132 .204

ae .090 OIOS 28 x 101 .277

Totals of proximal o

langes i .646 0766 — E

T phalanges

2. 073 .008 5 25 x 85 -294

c 071 .008 3 23 x 83 277

HE .078 .00Q1I 18 X IOI .178

IV. .073 .0085 20 X 96 .208

Le 057 0066 | 27 X 72 375

ane of middle we

lan .352 .0410 — —

Distal phalanges

A. .092 0107 24 X 79 304

B. 110 0128 25 X 99 .252

Cc. 105 0123 28 X 102 .274

ntf. .096 0112 28 x 99 .282

IV. .o8o .0093 26 X 94 .276

Yoo .069 .0080 27 x 89 315

Totals of distal dou

ges -552 0643 en ue

Total of all bones 8.569 — se ue

520 THE AMERICAN NATURALIST.

likewise, the metacarpal of digit B is more like metacarpal A

than the metacarpal of digit // of the normal manus is like its

pollex. Again, the metacarpal of digit C is more like its pollex

metacarpal A than is the metacarpal of digit ZZZ of the normal

manus like its pollex.

The same fact is also borne out by the comparison of the

indices, which are mathematical expressions of the forms of the

bones and can, therefore, be combine in the same way as

the percentage weights.

That the two normal radial digits have given place to three,

each of which partakes somewhat of the nature of the others,

is shown by a consideration of the tissue systems, and in no

way does this explanation meet with a decided contradiction.

The evidence of the dorsal veins is negative and not

contradictory.

I know of no case of polydactylism similar to the one here

described. One which is somewhat similar is described by

Windle (Journ. of Anat. Vol. XXVI, 1891, p. 100), who in

conclusion says: * The musculature shows that where there is

an additional digit on the radial side, it and the digit next to

the index both partake of the nature of thumbs and may be

looked — as the first and second digits of a hexadactylous

manus.'

In the case described in this paper, there is no reversion,

and the anatomy of the polydactyle paw, as here worked out,

has no bearing on either the * pre-pollex" or the “ post-mini-

mus" theories. The abnormality seems to result purely from

a readjustment of parts. The only definite statement which

can be made in regard to this case is that where normally two

digits occur, three have here appeared, and that each of these

three partakes more of the nature of theothers than one of the

two normal digits does of the other.

ZOGLOGICAL LABORATORY,

ARVARD UNIVERSITY.

THE GASTRULATION OF THE EGG OF BUFO

LENTIGINOSUS.

HELEN DEAN KING.

AN examination of the literature dealing with the early

development of the amphibian egg shows many conflicting

observations and theories regarding the origin of the blasto-

pore and the manner of formation of the mesoderm and

notochord. It is evident, therefore, that more forms must be

studied and the observations in some cases carefully repeated

before any general conclusions regarding the origin of these

structures can be drawn for the entire group.

The amphibian eggs that have been most studied are those

of Rana, Triton, Axolotl, and Bombinator. The present paper

deals with the early development of the egg of Bufo lentiginosus

from the end of cleavage until the closure of the blastopore.

The eggs of this amphibian, although very abundant and easily

obtained, have been but little used either for observation or for

experiment, owing, doubtless, to their small size and to their

relatively deep pigmentation, which makes it very difficult to

follow the fate of living cells.

METHOD.

In preparation for sectioning, eggs were killed in various

fluids: picro-acetic, picro-sulphuric, formalin, and corrosive-

acetic. By far the best results were obtained with corrosive-

acetic (5 per cent acetic). Formalin (3-10 per cent) gives

exceedingly good preparations for a study of surface struc-

tures, but it cannot be relied on to give histological details,

as it usually produces numerous cracks in various parts of

the egg. The sections were stained on the slide with the

mixture of borax carmine and Lyon's blue, recommended in

a previous paper (King, 10). With the use of this stain the

527

528 THE AMERICAN NATURALIST. [Vor. XXXVI.

nuclei appear red, the yolk blue, and the cell outlines are

brought out with great clearness.

THE FORMATION OF THE BLASTOPORE.

The egg of Bufo lentiginosus probably contains a greater

amount of pigment in proportion to its size than that of any

other common amphibian. It is, therefore, very difficult to

study the movements of individual cells before and during the

formation of the blastopore, as has been done to some extent in

the eggs of several other species of amphibians, where the pigmen-

tation is less extensive and cell outlines can be readily determined.

In the egg of Bufo the pigmentation extends in all cases

some distance below the equator, and I have frequently found

eggs in which fully three-fourths of the surface was deeply

pigmented before the appearance of the blastopore. Indi-

vidual eggs, even from the same female, differ greatly in the

amount of pigment they contain. Asa rule, the pigment line

extends farther down on one side of the egg than on the other,

as seen in Fig. 1, agreeing in this respect with the frog’s

egg according to Schultze (24), Morgan and Tsuda (17), and

Wilson (27).

Sections through an egg at the close of the blastula stage

show a large segmentation cavity in the upper hemisphere.

Its dorsal wall is formed, as in the frog, of three or four layers

of small angular cells of uniform size. The cells forming the

outer surface of the egg are almost completely filled with

pigment granules, and a considerable amount of pigment is

scattered throughout all the cells of the upper part of the egg.

The yolk cells below the segmentation cavity are much larger,

more rounded, and stain less intensely than the cells in the

upper hemisphere. There is the same gradual increase in the

size of the cells from the upper to the lower pole that other

investigators have noted in the frog's egg.

The dorsal lip of the blastopore invariably appears some dis-

tance below the equator of the egg, but never in the middle

of the lower hemisphere, as maintained by Houssay (8) for the

axolotl and Jordan (9) for the newt.

No. 427.1] THE EGG OF BUFO LENTIGINOSUS. 529

If the egg has but the usual amount of pigment, the first

evidence of the blastopore, in surface view, is a short, dark,

almost straight line at the extreme edge of the black cells

(Fig. 1). If the pigmentation is unusually extensive, the blas-

topore appears in its same relative position with respect to the

lower pole of the egg but distinctly within the black cells.

Whether there is, as the first step in the formation of the blas-

topore, a “lining up" of the ectoderm cells, as described by

Wilson for the frog, I have not been able to determine.

Sections of an egg at the beginning of gastrulation show

that the dorsal lip of the blastopore is formed primarily by a

sinking in of several of the surface cells (Fig. 9). The shallow

and closure of the blastopore.

depression thus formed is rapidly extended and soon becomes

a pronounced furrow (Fig. ro). The cells involved in this

sinking in are all, without question, large yolk cells which are

decidedly wedge-shaped and contain a considerable amount

of pigment in their smaller ends turned towards the exterior

(Figs. 9, 10, 11).

After the lateral extension of the dorsal lip to form a cres-

cent, a sagittal section through the blastopore shows that the

furrow has deepened considerably and that its inner end is

turned up towards the dark pole (Fig. 11). From this time on

there is a marked difference in appearance between the cells

forming the dorsal wall of the archenteron and those forming

the ventral wall. The cells of the dorsal wall are small, angular,

530 THE AMERICAN NATURALIST. [Vor. XXXVI.

deeply pigmented cells, and they are apparently exactly like

the small cells forming the upper surface of the egg, except

that they do not contain quite as much pigment. The yolk

cells which form the floor of the archenteron are three or

four times larger than the cells of the dorsal wall, and they

are more rounded and

contain little if any pig-

ment. The distinction

between these cells is

made more evident by

the fact that with the

7^5 combination stain used

EJ ) all the yolk cells appear

EENT decidedly blue, while the

SILT Te $ lls of the dorsal wall of

3 XR SI) D cells of the dorsal v

> Se [o] 97)

eS A VIR ^y:

zo E LETT F

S.C.

the archenteron and, in

fact, all the cells of the

upper hemisphere, take

a distinctly reddish tint.

The cells at the anterior

end of the archenteron

are still wedge-shaped at

this stage. They are

intermediate in size be-

tween the cells forming

the dorsal wall and those

forming the ventral wall

of the archenteron, and

they stain like the yolk

gn gus as = a — MA the median plane of cells. a

aT at eed Fe seme In opposition to most

eee ee re; M,cls investigators, Moquin-

Tandon (15), Houssay,

Robinson and Assheton (20), and Marshall (14) maintain that

the archenteron of the amphibian embryo is not formed

by a process of invagination, but by a splitting between

yolk cells which thus form the dorsal as well as the ventral

wall of the archenteron. In a later paper, Assheton (1)

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 531

states that “the formation of the primitive archenteron is

by a process of splitting, and is the direct effect of the pri-

mary center of growth; whilst the continuation of the cavity

produced by an overgrowth is the direct effect of the second-

ary center of growth.”

That the archenteron in the egg of Bufo lentiginosus is not

formed by a splitting between yolk cells is shown, I think, by

a study of a series of well-preserved eggs during the early

stages of gastrulation.

The archenteron

never appears *'slit-

like" at this time.

On the contrary, its

walls are usually some

distance apart and its

inner end is invari-

ably rounded (Figs.

IO, 11). Occasionally

at the stage of Fig.

I2, never earlier, I

have seen an irregu-

lar cleft between the

yolk cells at the ante-

rior end of the arch-

enteron which might, n MOM :

perhaps, be con- Mabdy fier stage than Fig. Kg Mns the character of the

sidered a forward Veil nci spes ncc atrio i dag ?

extension of the :

archenteron, but such a cleft is usually seen only in badly

preserved eggs where the cells are all more or less separated,

and therefore I have always considered that it was artificially

produced by the method used in killing and hardening the egg.

Jordan, Wilson, and Eycleshymer (4) among others, have

watched the disappearance of individual surface cells under the

dorsal lip of the blastopore in the early gastrulation stages of

the living egg. These observations seem to me to afford con-

clusive evidence that invagination of cells plays an important

róle in the formation of the archenteron.

Part of a dian secti

532 THE AMERICAN NATURALIST. [Vor. XXXVI.

The presence of pigment in the inner ends of the cells

forming the dorsal wall of the archenteron has been explained

` by Robinson and Assheton as follows: “The pigmented area

is produced and extended by the deposit of pigment in the

adjacent margins of a double row of yolk cells which eventu-

ally will form the boundary wall of the archenteron, and it

BRE also radiates from this

X^ area along the adja-

. E

ane

Fic. 11. — Part of

pore at the stage of

as in previous figure

di agittal section through the blast

Fig. 2. EC., ectoderm. Other lettering

? cent margins of the

cells of each row. A

slit-like space appears

in the middle of the

posterior portion of

the pigmented area.

This space first limits

the dorsal lip of the

blastopore, and then

extends forward and

ventrally, following

the deposit of pig-

ment, and separating

the two rows of mar-

ginaly pigmented

cells from each

other.” During the

early stages of gastru-

lation there is always

found at the anterior

end of the archen-

teron a marked accu-

mulation of pigment

that is extended

around to include a

few of the yolk cells

in this region (Figs.

IO—12). I have never found more than a few scattered pigment

granules in the lower yolk cells except in this particular part

of the egg, and, if the archenteron is formed by a splitting

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 533

between marginally pigmented yolk cells, then the pigment

must at once almost entirely disappear on one side of the split

and not on the other.

NEAN, PE CO YN

Boge Tu E

x : d =

The pigmentation at CES OLYAN Ete,

INE

the anterior end of the SESE he

archenteron in the egg COREE ERM

of Bufo is not confined AETIA

* S.C. — (ees SECO

to * a double row of yolk € AY ees

cells" anterior to the CI ger SN

à TaT a

actual cleft. Well Ay] SD

marked lines of pig-

ment may extend out in

any direction, even at

right angles to the line

of advance of the arch-

enteron, as seen in Fig.

I3. From thetime that

the blastopore is first

formed, the pigment in

the cells of the dorsal

wall of the archenteron

is always collected

around the cell walls,

INN

Bex

dl G rt of a median sagittal section through the

cleus. Rhumbler (22) blastopore at the stage of Fig. 3. Y., yolk plug. Other

considers this phenom- lettering as in previous figures.

enon to be due to the mechanical effect of pressure. The

suggestion has been offered by Jordan “that the pigment marks

physiological activity, and that the less heavily pigmented cells

of the ventral wall of the archenteron owe their relative lack

of pigment to more sluggish metabolism attendant upon less

rapid cell division.” It is certainly true that the large yolk

cells in the egg of Bufo divide less frequently than the cells

in the upper hemisphere, but there is no evidence that the

deeply pigmented cells of the outer surface of the upper

hemisphere or of the dorsal wall of the archenteron divide

more rapidly than the cells that are found between them. If,

therefore, Jordan’s suggestion is correct, some kind of

534 THE AMERICAN NATURALIST. [Vor. XXXVI.

“ physiological activity " other than that attendant upon rapid

cell division must produce the pigmentation in these cells.

The extension of the lateral lips of the blastopore and the

formation of the yolk plug have been so frequently described

for other forms that a description of these processes is not

necessary here, as they are apparently similar in all respects

to those which take place in the frog. Following the method

used by Pfliiger (20), a number of eggs in the blastula stage

were placed on a mirror in a shallow dish of water and the for-

mation of the blastopore watched in mirror image. Figs. 1-8

Fic. 13. — Part of a frontal section through tl ior end of th

Fic. 2. showine li Ext 1i oci

5 rS 5

+. +h

A R., archenteron.

show the appearance and the location of the blastopore at dif-

ferent times during the day, from 8 a.m., when the blastopore

lip first appeared, to 6 P.M., when the egg began a rotation

around its horizontal axis. There was no evidence to show

that the position of the eggs was changed during the time

they were kept under observation; on the contrary, one egg,

in which a peculiar light spot was seen below the equatorial

region, was watched particularly, and the spot was found to

keep its same relative position until the rotation of the ess

took place. These mirror images show that the dorsal lip of

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 535

the blastopore appears below the equator of the egg at the

edge of the black cells (Fig. 1), and that the blastopore rim

extends at a uniform rate in the form of a crescent around

the lower hemisphere, until finally the yolk plug is formed

(Figs. 2-4). Meanwhile, the dorsal lip of the blastopore has

moved over the yolk, and seven hours after its first appearance

has reached the center of the white hemisphere (Fig. 6). The

closure of the blastopore lips takes place almost uniformly from

all sides as a rule (Fig. 7), although occasionally the blastopore

is somewhat oval when it is in the stage of Fig. 8. At or just

before the stage of Fig. 8 a rotation of the egg around its

horizontal axis takes place, thus bringing the dorsal lip of the

blastopore back to its original position.

In its movement over the lower hemisphere the dorsal lip of

the blastopore seems to pass through about 140° of the surface

of the egg. There is apparently a slight variation in this respect

in different eggs. This measurement is somewhat greater than

that given by Morgan (16) and by Wilson (27) for the move-

ment of the dorsal lip of the blastopore in the frog egg, and

differs considerably from the measurements which have been

given by other investigators. Roux (21) and Pflüger (19) esti-

mate that the dorsal lip moves about 170°, while Kopsch (11)

considers the movement to be but 75°; and this figure is further

reduced by Assheton, who states that much of the so-called

movement of the dorsal lip is only apparent and that the real

movement is only from 60° to 70°.

THE FORMATION OF THE “GERM LAYERS" AND THE

NOTOCHORD.

Observations regarding the origin of the endoderm, meso-

derm, and notochord in different amphibian eggs are so

conflicting that apparently the most plausible interpretation

of them is to assume that there is absolutely no uniformity

in the manner of formation of these structures even in

species that are considered to be very nearly related to

each other. Such an assumption, however, can scarcely be

the correct one.

536 THE AMERICAN NATURALIST. [Vor. XXXVI.

Goette (6), one of the early workers on the development of

the amphibian egg, considered the mesoderm in the egg of

Bombinator to arise from the splitting off of a layer of cells

from the primitive entoblast, the layer thus formed extending

as an unbroken sheet across the dorsal wall of the archenteron.

The rest of the primitive entoblast was said to form the endo-

derm. According to Goette, a central chord of mesoderm in

the mid-dorsal region of the embryo separates from the two

lateral sheets to form the notochord.

A few years later Hertwig (7), after studying the develop-

ment of the eggs of Triton and of Rana temporaria, came to

the conclusion that the mesoderm in the amphibian egg arises

in the vicinity of the blastopore “durch eine paarige Ein-

faltung des Entoblast schon zu einer Zeit wo die Gastrula-

einstülpung noch nicht ganz vollendet ist." The amphibian egg

was thus brought into agreement with Hertwig's observations

on the development of other vertebrates and gave additional

support to his well-known ccelom theory. A few other

writers — Balfour (2), Marshall (14), and Schwink (26)—

agree with the results obtained by Hertwig.

In 1888, Schultze (25), from observations on Rana fusca,

decided that “das mittlere Blatt, sowie die dorsale Urdarm-

wand entstehen aus dem Ektoblast und gehen an der dorsalen

Urmundlippe alle drei Blatter in einander iiber; in den seit-

lichen und ventralen Theilen des Blastoporus setzt sich die

Deckschicht des áusseren Keimblattes mit besonderer Klarheit

in den Entoblast, die Grundschicht des Ektoblast ohne Unter-

brechung in den Mesoblast fort." Perenyi (18), from obser-

vations on the egg of Bombinator igneus, Lwoff (13), from a

study of Axolotl and various Anura, and Brauer (3), from

investigations on two species of Gymnophiona, also came to

the conclusion that the mesoderm is ectodermal in origin.

These investigators, however, differ somewhat regarding the

manner in which this process is supposed to take place.

Perenyi states that, as a result of a turning under, or “ Dupli-

kation," of the three-layered outer wall of the egg at the lips of

the blastopore, “ die aussersten Deckzellen der Blastula in ihrer

ununterbrochenen Fortsetzung nach innen endlich die untersten

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 537

Zellenreihen bilden werden d. h. sie verwandeln sich in Ento-

derm, wahrend die anderen zwei Zellenreihen der Blastula auch

in ihrer Riickwindung beisammen bleiben und eine selbstandige

Schichte, das Mesoderm, bilden."

According to Lwoff, “die Einwanderung der Ektoderm-

zellen und die Verschiebung der Entodermzellen zwei ver-

schiedene und von einander unabhängige Vorgänge sind.

Die Einwanderung von Ektodermzellen beginnt an der Stelle

die das Hinterende des Embryo markirt ; die Verschiebung der

Entodermzellen beginnt im Gegentheil im vorderen Theile.

Wie die weiteren Stadien lehren, bildet das Hineinwachsen

der Ektodermzellen die zusammenhangende ektoblastogene

Anlage der Chorda und des Mesoderms; die Verschiebung

der Entodermzellen führt zur Bildung der Darmhohle, die

infolge des Auseinanderweichens der Entodermzellen ent-

steht."

Brauer's investigations show that there is a turning under

of the outer surface cells, or “animal cells," at the posterior

end of the germ disk and that by the forward growth of these

cells under the upper animal cells a blind sac is formed. This.

blind sac becomes the posterior part of the archenteron, and at

first its dorsal wall is composed of the animal cells invaginated

from the surface. In subsequent development a layer of vege-

tative or yolk cells grows up from either side of the archenteron

and forms a layer of endoderm beneath the invaginated animal

cells which later become the mesoderm.

The origin of the mesoderm in the frog's egg has been

described by Morgan (16) as follows: ** The cells that are to

form the mesodermal layer are present at the time when the

dorsal lip of the blastopore has first appeared, and even just

prior to that time.” The innermost of the cells forming a

ring around the equatorial region of the egg where the black

and the white cells meet are the cells that later become the

mesoderm. “These cells are carried up to the median dorsal

line of the embryo by the closure of the blastopore. They will

then be found forming a layer or sheet of cells that separates

itself on the outer side from the thick layer of small ectoder-

mal cells (that has been simultaneously lifted up) and that is

538 THE AMERICAN NATURALIST. | [Vor. XXXVI.

separated on the inner surface, but not very sharply if at all,

from the dorsal and dorsal-lateral walls of the archenteron."

In the egg of Bufo lentiginosus, when the blastopore first

appears as a slight depression among the yolk cells just below

the equator of the egg (Fig. 9), the cells which form the dorsal

wall of the segmentation cavity are all small, somewhat angular

cells which contain a considerable amount of pigment, particu-

larly the cells which compose the outer surface of the egg.

Near the equator the cells are found to be somewhat larger

and to contain much less pigment, while in the lower hemi-

sphere the cells are-very large, and they have but a few scat-

tered pigment granules. Up to this stage of development

there is no visible separation of the cells into ectoderm, endo-

derm, and mesoderm. The smaller cells resulting from rapid

cell division are found in the upper part of the egg and the

larger cells, which divide less frequently, are grouped around the

lower pole. The mass of cells marked 77 in Fig. 9 undoubt-

edly represent the région of the egg from which the mesoderm

is formed. These cells are in no wise distinctive at this period,

and their later development into mesoderm I consider to be due

solely to their position in the egg during the formation and

closure of the blastopore. The cells which are to become

mesoderm form a layer around the egg at the equatorial region

just inside the cells that are to become the ectoderm, as found

to be the case in the frog's egg, according to Morgan. These

mesoderm cells have many characteristics in common with

the large yolk cells into which they grade, being larger, more

rounded, and containing much less pigment than the small cells

of the upper hemisphere. It seems probable, therefore, that

they were originally produced by divisions of the yolk cells.

When the lips of the blastopore have extended so as to form

a crescent in surface view (Fig. 3) a frontal section through an

egg in the region of the blastopore (Fig. 14) shows that the

dorsal wall of the archenteron is formed of several layers of

small cells which have absolutely no distinction between them

except that the cells of the outer layer, which are more regu-

larly arranged, contain a much greater amount of pigment than

the other cells. At the sides of the archenteron the innermost

No. 427. THE EGG OF BUFO LENTIGINOSUS.

539

of these cells pass into the large yolk cells which form the floor

of the archenteron. A median sagittal section through the blas-

topore at the same or a slightly earlier stage of development

(Fig. 11) shows the beginning of the separation of the ectoderm

from the inner cells of the egg. A tolerably regular cleft extends

some distance around the sides of the egg, on a line usually

with the lower edge of the dorsal wall of the segmentation

cavity, sharply separating the layer of cells forming the

outer wall of the egg from the cells within. The separation

of the ectoderm does not extend as far down as the equator

of the egg at this time, and it is some hours later before

esse mare

coe

Fic. 14. — Part of a frontal section through the region just anterior to the blastopore at the

s of Fi R., archenteron.

the ectoderm at the lips of the blastopore is distinct from

the other cells. It is in the region of the blastopore that the

union of the different layers persists longest, as other inves-

tigators have noted.

At the stage of Fig. 3 the formation of the archenteron is

well advanced, and a median sagittal section through the blasto-

pore (Fig. 12) shows that the endoderm of the dorsal wall of

the archenteron is formed of small, angular, deeply pigmented

cells which, as far as I am able to determine, appear exactly like

the small cells which form the outer surface of the egg. It

seems probable that these cells once formed a part of the outer

surface of the egg in the region just outside of the blastopore,

and that they have been turned under the edge of the blastopore

lip by a process of invagination, thus being changed into endo-

derm. Whether, in later stages of development, the endoderm

540 THE AMERICAN NATURALIST. [VoL. XXXVI.

of the dorsal wall of the archenteron receives additions from the

yolk, as believed to be the case by Assheton and by Wilson, I

have not been able to determine.

After the ventral lip of the blastopore has formed and the

blastopore is beginning to close (Fig. 6), a division of the

ora

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ectoderm into two layers is first noticed. A median sagittal

section at this period (Fig. 15) shows an outer ectodermal wall

composed of a single layer of small, deeply pigmented cells

which are arranged very regularly. The inner ectodermal

sheath is composed of two or three layers of much larger cells

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 541

which are distinctly wedge-shaped and contain a comparatively

small amount of pigment. The inner layer of ectoderm is

sharply separated from the mass of small cells above the

dorsal wall of the archenteron.

When, in surface view, the blastopore has reached the stage

of Fig. 6, or in some cases even as early as Fig. 5, a splitting

is seen in the mass of cells above the archenteron, anda single

layer of endodermal cells is separated from the cells above.

Fic. 16. — Outline of the entire section from which Fig. 15 was drawn.

L., ventral lip of the blastopore.

This process begins some distance in front of the region of the

blastopore and gradually extends forward and backward from

this point, thus completing the separation of the cells in the

mid-dorsal region of the embryo into ectoderm, mesoderm,

and endoderm.

When the blastopore has nearly closed (Fig. 8), a sagittal

section through the middle of the dorsal lip (Fig. 17) shows

that the endoderm cells of the dorsal wall of the archenteron

(Fig. 17, Ex.) have flattened out considerably, and that they

still contain much pigment, which is collected in the end of

542 THE AMERICAN NATURALIST. [Vor. XXXVI,

the cell bordering the cavity of the archenteron. Above the

endoderm is the mesoderm layer (Fig. 17, M.), which varies in

the number of its cells in different parts of the egg. Near

the dorsal lip of the blastopore the mesoderm sheath consists

f Fig. 8. EN.,entoderm.

PIE A n

Other lettering as in previous figures.

of several layers of small cells which are not pigmented. For-

ward from this region the mesoderm gradually thins out until

it is composed of only a single layer of rounded cells lying

between the endoderm and ectoderm; then it gradually

becomes thicker again, and near the ventral lip of the blas-

topore it appears very much as it does near the dorsal lip,

being composed of several layers of cells.

A frontal section through the middle of an egg at the stage

of Fig. 7 shows a single layer of mesoderm over the dorsal

region of the embryo, which gradually becomes several layers

thick at the sides of the archenteron (Fig. 19), and below the

archenteron passes directly into the yolk cells of the lower

part of the egg. Still later stages during the closure of the

blastopore show that the mesoderm is unquestionably extended

at the expense of the yolk cells lying just beneath the ectoderm.

Soon after the medullary folds have appeared the lateral sheets

of mesoderm become fused on the ventral side of the embryo;

thus a continuous sheet of mesoderm is formed around the

embryo, except in the mid-dorsal region where the notochord

has by this time been cut off from it. The extension of the

No. 427.] | THE EGG OF BUFO LENTIGINOSUS. 543

lateral sheets of mesoderm by a process of splitting off from

the outer yolk cells has been noted by Schwink (26) in Bufo

vulgaris, by Scott and Osborn (23) and by Jordan in the newt,

and by Morgan in the frog.

At the close of the blastula stage a very large segmentation

cavity is found in the upper hemisphere of the egg directly

under the black pole (Figs. 9, 11, 12). During the formation

of the archenteron this segmentation cavity decreases con-

siderably in size, and is pushed out of its position, coming to

lie below the archenteron and being separated from it by only

a thin layer of cells (Fig. 16, S.C.). During the closure of

Fic. 18. — Outline of the entire section from which Fig. 17 was drawn.

the blastopore the archenteron increases greatly in size, and

there is then usually but a single layer of large yolk cells

between it and the segmentation cavity (Fig. 19). When the

blastopore is nearly closed, I have frequently found eggs in

544 THE AMERICAN NATURALIST. | [Vor. XXXVI.

which there appeared to be a direct connection between the

two cavities which could not be due to poor preservation of

the material (Fig. 18). It would seem, therefore, that in some

cases the archenteron is extended at the expense of the seg-

mentation cavity, as supposed to be the case by Kupffer (12)

and by Marshall.

A series of frontal sections through the embryo at about

the stage of Fig. 7 will show the various steps in the forma-

"uma.

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4 H CS Ss HN IS 1 A. n Ttc"

n et: the yolk. Lettering as in previous

figures.

tion of the notochord. A section just in front of the dorsal

lip of the blastopore (Fig. 20) shows that the mesoderm in

this region forms a continuous sheath over the dorsal wall

of the archenteron and that it is composed of three to four

layers of small cells (Fig. 20, 77). About eight sections

anterior to Fig. 20 there is a noticeable thickening of the

mesoderm in the mid-dorsal region (Fig. 21), while on either

side of this thickening the mesoderm usually consists of but

two layers of cells. More anterior, the thickened part of the

mesoderm layer is completely separated from the lateral sheets

No. 427.] THE EGG OF BUFO LENTIGINOSUS. 545

as a distinct rounded structure, the notochord (Eig: 22, NJ.

For only a short distance at this stage of development is the

notochord entirely cut off from the mesoderm ; beyond there

DDE

1700

XU A

Fic. 20. — Part

f l section through an egg at the stage of Fig. 7, showing the mid-d

region of the embryo just in front of the blastopore. ÆC., ectoderm; ÆN., endoderm;

JI., mesoderm,

is the same thickening of the mid-dorsal portion of the meso-

derm sheet as seen in Fig. 21. Inthe head region of the embryo

the mesoderm is thinned out to a layer one or two cells deep

Which shows absolutely no thickening in the middle and appears

very much as in Fig. 19. After the blastopore has closed, the

notochord becomes extended forward in the head region and

backward in the region of the tail.

Schultze, Goette, Lwoff, Brauer, Schwink, and Morgan also

believe that the notochord is derived from the mesoderm ; while

«sgg =

SK FQ

Fic. 21. — Part of a section from the same egg as Fig. 1 slightly

EC., ectoderm; ÆN., endoderm; M., mesoderm.

ant to it.

other investigators consider that in the amphibians, as in other

vertebrates, the notochord is endodermal in origin. Although

Field (5) believes that in Bufo vulgaris the notochord arises

540 THE AMERICAN NATURALIST. [Vor. XXXVI.

from the endoderm, in the nearly related species, Bufo lentigi-

nosus, the anterior part of the notochord is certainly mesodermal

in origin. After the mesoderm layer has formed, the endo-

derm in the mid-dorsal region of the embryo is never more

than a single layer of flattened cells. There is a closer con-

nection between the mesoderm and the endoderm above the

center of the archenteron than at the sides, but I have never

found an egg in which these two layers could not readily be

Fic. 22. — Part of a section from the middle region of the same egg. ÆC., ectoderm; ÆW.,

endoderm; M., mesoderm; N., notochord,

distinguished from each other. The endoderm cells are always

much flattened at this period, and they invariably contain a

considerable amount of pigment. The mesoderm cells and

also the cells of the notochord are larger, more rounded, and

contain but little pigment in comparison with the endoderm

cells. »

In conclusion, I wish to express my thanks to Prof.

T. H. Morgan for advice and criticism during the progress

of my work.

BRYN MaAwR COLLEGE, BRYN Mawn, PA.,

December 16, 1901.

-I

—

. 427.] THE EGG OF BUFO LENTIGINOSUS. 547

LITERATURE.

. AssHETON, R. On the Growth in Length of the Frog Embryo.

Quart. Journ. Micr. Sci. Vol. xxxvii 1894.

BaLFour, F. M. Comparative Embryology. II. New York, The

Macmillan Company, 1

80.

. BRAUER, A. Beiträge zur Kenntniss der Entwicklungsgeschichte und

der Anatomie der Gymnophionen. Zool. Jahrb. Bd. x.

EYCLESHYMER, A. C. The Early Development of Amblystoma, with

Observations on Some Other Vertebrates. T€ Morph. Vo

1895.

FIELD, H. H. Bemerkungen über die konii der Wirbelsäule

bei den Amphibien; nebst Schilderung eines abnormen Wirbelseg.

mentes. Morph. Jahrb. Bd. xxii 1895.

3OETTE, A. Die Entwickelungsgeschichte der Unke. Leipzig, 1875.

. HERTWIG, O. Die vie sig des mittleren Keimblattes der Wirbel-

thiere. I, II. Jena, 1881, 3.

Houssay, F. Etudes pereon sur les vertébrés. Arch. de Zool.

Exp. et Génér. Tome viii. 1890.

Jorpan, E. O. The Habits and Development of the Newt. Journ.

Morph. Vol. viii. 18

. Kine, H. D. The Maturation and depen of the Egg of Bufo

lentiginosus. Journ. Morph. Vol. xvii.

. Kopscnu, F. Beitrag? zur Gastrulation beim see und Froschei.

erh. Anat. Gesellschaft. 1895.

KUPFFER, C. Die Entstehung der Allantois und die Gastrula der

Wirbelthiere. Zool. Anzeiger. Bd. ii.

Lworr, B. Die Bildung der primáren Keimblatter und die Entste-

er der Chorda und des Mesoderms bei den Wirbelthieren. Bu:

de la Soc. Imp. des Nat. de Moscou. Ba. viii.

1894.

. MARSHALL, A. M. Vertebrate Embryology. New York, Putnam,

1993.

MoQovuIN-TANDON, G. Recherches sur les premieres phases du dével-

oppement des batraciens anoures. Ann. des Soc. Nat. Tome iii.

1876.

MonGAN, T. H. The Development of the Frog's Egg. New York,

The Macmillan Company, 1897.

. MorGan, T. H. and TsupE, Ume. The Orientation of the Frog’s

Egg. Quart. Journ. Micr. Sci.

ol. xxxv. 1993.

. PERENYI J. Die Entwickelung der Keimblátter und der Chorda in

neuer Beleuchtung. Anat. Anzeiger. Bd. iv. 1889.

PrLUGER, E. Ueber den Einfluss der Schwerkraft auf die Theilung

der Zellen. Pffiigers Archiv. Bd. xxxii. 1883.

548 THE AMERICAN NATURALIST.

20. ROBINSON, A., and AssHETON, R. The Formation and Fate of the

Primitive Streak with Observations on the Archenteron and Germi-

nal Layers of Rana temporaria. Quart. Journ. Micr. Sci. Vol.

XXxii 189r.

. Roux, W. Ueber die Lagerung des Materiales des Medullarrohres im

gefurchten Froschei. Anat. A nzeiger. Bd. iii.

22. RHUMBLER, L. Physikalische Analyse von Lebenserscheinungen

der Zelle. III. Mechanik der Pigmentzusammenhaufungen in den

oe a, der Amphibieneier. Arch. f. Entwickelungs-

mechanik. Bd. ix. 1899.

24. Scorr, W. B. d OsBORN, H. F. On Some Points in the Early

Development of the Common Newt. Quart. Journ. Micr. Sci.

Vol. xix.

24. SCHULTZE; O. Jia Entwickelung des braunen Grasfrosches. Fest-

schrift f. Kölliker. 1887.

SCHULTZE, O. Die Entwicklung der Keimblätter und der Chorda

dorsalis von Rana fusca. Zeitschr. f. wiss. Zool. Bd. xxvii.

iris

26. SCHWINK, F. Ueber die Entwicklung des mittleren Keimblattes und

der Chorda dorsalis der Amphibien. München, 1889.

WiLsoN, H. V. Formation of the Blastopore in the Frog Egg. Anat.

Anzeiger. Bd. xviii. 1900.

NOTES ON THE CCELENTERATE FAUNA OF

WOODS HOLE.

CHARLES W. HARGITT.

THE following notes upon certain faunal features of the

coelenterate life of Woods Hole and adjacent waters pertain

chiefly to the season just past, including records which are

fairly constant at regular intervals throughout the entire year.

For certain of them I am indebted to Mr. Vinal Edwards,

whose painstaking records during many years are matters well

known to many naturalists. I am also under obligations to

Mr. George M. Gray and Dr. H. M. Smith for similar favors.

In addition to notes upon the seasons, habits, etc., brief

accounts are presented of new forms discovered and of the

development of one of the Scyphomedusz.

HyDROMEDUS.

A New Tubularian Hydroid. — In August, 1900, while cruis-

ing and dredging in Muskegat Channel off Marthas Vineyard,

a considerable number of fine specimens of Corymorpha pendula

Ag. were taken by dredge and trawl, many of which were in fine

condition, still imbedded in the sand as in normal life. A

somewhat cursory examination of the specimens showed what

seemed to be young specimens growing among the filamentous :

rootlike holdfasts and apparently budding from the base of

the hydroid, like young polyps, reference to which fact was

made by the present writer in the recently published synopsis

of Hydromedusz. In order to determine more definitely the

apparent anomaly of buds arising from so low among the rhi-

zoids and below the sandy surface of the substratum, and whether

they might show signs of later becoming detached, the speci-

mens were submitted to one of my students, Mr. A. J. May,

with instructions to determine the range of budding, nature of

549

550 THE AMERICAN NATURALIST. [Vor. XXXVI.

development, etc. The results have shown what was only at

first dimly suspected, namely, that the small « buds” were in

fact not buds at all, but a distinct hydroid, apparently parasitic,

growing upon the base of the Corymorpha. This was clearly

demonstrated both from the specific differences which a critical

study of its morphology showed, and by means of sections made

through the points of attachment. The point of attachment

is within a rather limited zone of the base, among the conical

papillz and filamentous rhizoids of the host, where in some

cases as many as half a dozen were growing upon the same

specimen. As will be noted from Figs. 1

and 2, the new hydroid clearly belongs

to the genus Tubularia, having the char-

acteristic form of body, arrangement of

tentacles, etc. Its attachment to the host

was quite close, being inserted within the

filmy perisarc, where its base expanded in

the form of fingerlike absorbent organs.

Briefly its characters may be summa-

rized as follows: Hydranth solitary, from

2 to 5 mm. in height and about 4 mm.

in diameter ; tentacles in two whorls, the

proximal of from eight to sixteen, the

distal of from five to eight. Gonads

were found upon but one specimen, and

in this were immature, but occupying the

characteristic position among the basal

tentacles. So far as known, semi-parasitic upon Corymorpha.

. I propose for it the provisional name Tubularia parasitica, till

further investigations may be had upon additional and more

mature specimens, which may further confirm or modify this

description.

A New Hydromedusa. — On Aug. 10, 1901, a medusa with dis-

tinctively sarsian characters was taken in the tow, similar in many

respects to two species described by Forbes in 1848 (British

Naked-Eyed Medusz), chief among their features being the pro-

liferous development of medusze from the body or manubrium of

the parent medusa. Haeckel likewise described a similar medusa

Fic. 1. — Tubularia parasitica.

No.427.] CCELENTERATE FAUNA OF WOODS HOLE. 551

from the Canary Islands, in which secondary medusz were budded

in great numbers from the very long siphonlike manubrium.

So far as I am aware, no member of the genus Sarsia having

similar characters has been described from American waters.

Forbes's species were Sarsza gemmifera and S. prolifera, as

cited above, pp. 57—59. The former, like Haeckel's S. szphono-

phora, buds its secondary medusz from the walls of the long

manubrium in a spiral-like raceme throughout its entire length.

In the specimen under consideration the proliferous medusz

are budded from the manubrium,

but in a whorl about its base (cf.

Fig. 3. Onlyasingle specimen was

. taken, but its characters were so

definite that there seems little doubt

as to its specific distinctness.

The following characters summa-

rize its more distinctive features :

Bell high, somewhat ovate, or elon-

gate hemispherical; tentacles four,

with prominent basal bulbs, orange-

colored, with marginal fringe of

delicate green, each bulb with a

prominent ocellus densely black and

facing outward. Manubrium long,

pendulous, extending far beyond the

margin of bell, capable of great con- Fte. 2. ~~ ae =e

traction; terminal portion doubly

bulbous, with proximal bulb light sky-blue, terminal green ;

basal portion of manubrium expanding into an orange-colored

gastric pouch which graduates into the body of the bell.

Gonads medusoid, in a whorl about the base of the manubrium ;

ectoderm transparent or faintly tinged with pale blue, entoderm

orange-colored. Height of bell, 1.5 mm.; breadth, 1 mm.

For this medusa the name Coryne producta is proposed. The

description is based upon a single specimen, and while its char-

acters seem sufficiently definite to warrant specific distinction,

it is not impossible that other specimens in larger numbers

May necessitate modification or change at certain points.

552 THE AMERICAN NATURALIST. [Vor. XXXVI.

SEASONAL ASPECTS.

The seasonal range among medusz is noteworthy in many

cases, both by reason of its periodicity in certain species and

by its perennial character in others. The following are among

the distinctively spring medusz.

Hybocodon prolifer Ag. is one of the most restricted in this

respect, ranging from late February to about the middle of

May. It is one of the most beautiful and brilliant of the

early medusa. Its proliferous budding of secondary medusæ

from the large single tentacle is an interesting and well-known

feature. I have also found it producing actinule from the

walls of the manubrium at the same

time. These seem to arise much as

do similar larve, from several other

anthomedusa, and a histological exam-

ination shows essentially similar fea-

tures of oógenesis and spermatogenesis,

though there is no appreciable distinc-

tion of sexes so far as morphological

features are concerned.

Tiaropsis diademata Ag. This is

likewise an early medusa. According

to Agassiz, “ one of the earliest visitors

| of our wharves in spring.” During the

present season it has been taken from

March 20 to May 1, though only in

limited numbers.

Coryne mirabilis Ag. is also one of the earlier medusz, but

with a longer range, from February to May. It has been

reported by Fewkes as occasionally taken at Newport in

early summer.

Lizzia grata A. Ag. Of this medusa I have records during

the present year from March 27 to April 29. Fewkes has

reported it as occurring at Newport during the entire summer,

but this would seem to be somewhat unusual.

Tima formosa Ag. This beautiful medusa, one of the

| largest of the early Hydromedusz, while not specially rare at

Fic. 3. — Coryne producta.

No.427.] C@LENTERATE FAUNA OF WOODS HOLE. 553

this station, is not specially abundant. Its season is compara-

tively brief, ranging from April 12 to May 5. While sometimes

quite abundant at certain localities, its hydroid phase is as yet

unknown, as is also its development.

Trachynema digitale A. Ag. This Trachomedusa is occa-

sionally taken here in considerable numbers in April, but is not

a common medusa. Little appears to be known as to its habitat

or life history, but it is probably more or less pelagic, and

hence subject largely to the influence of ocean currents, etc.

Rhegmatodes tenuis A. Ag. Note is made in this connec-

tion of the occurrence of this medusa in large numbers during

the summer of 1900, the first record during several years,

and its utter absence during the present season. It is usually

recognized as a summer medusa, and the present record only

confirms this. What is more specially worthy of note is the

erratic aspect of its appearance, apparently at long intervals,

and then usually in great numbers, indicating seemingly some

local or environmental condition as governing its occurrence,

rather than purely seasonal mutations. Were Rhegmatodes

a pelagic medusa we might account for mutations of this sort

through the agency of shoreward currents or winds, as often

happens with Physalia, and perhaps also with Trachynema,

as intimated above. But so far as I am aware, this medusa,

in common with most of the Hydromedusz, is more or less

restricted to the littoral fauna, and hence only incidentally

affected by such influences. This is another medusa whose

hydroid yet remains unknown. Possibly when its life cycle

is fully known the apparent capriciousness of its occurrence

may be cleared up.

Staurophora laciniata A. Ag. was likewise taken in consid-

erable numbers in early May by Mr. George M. Gray, and

later in the month by Dr. H. M. Smith. So far as my records

are concerned, this is the first occurrence of this medusa in

several years at this station. One of its peculiar structural

features, the early confluence of the oral lobes with the frills

of the gonads, so that they can only be distinguished with

difficulty, was described by A. Agassiz. No mention, so far

as I am aware, has been made of the origin of actinulz in this

554 THE AMERICAN NATURALIST. [Vor. XXXVI.

medusa by a method. very similar to that of Hybocodon. In

Staurophora, however, they arise within the greatly folded bor-

ders of the gonad pouches beneath the radial canals. As in

the former, no conspicuous differentiation of sex was apparent

in Staurophora, though not having seen them alive, and not

having as yet examined the histology of the gonad region,

this can only be stated tentatively.

Among medusz taken during midsummer, the more common

were Nemopsis bachei Ag., very abundant during August ; Bou-

gainvillia carolinensis Ag., likewise abundant during August ;

Pennaria tiarella McCr., less abundant than during previous

seasons; Ectopleura ochracea A. Ag., unusually abundant dur-

ing August, taken mostly in the evening tow; Luchetlota

duodecimalis A. Ag., Eucheilota ventricularis McCr., not

uncommon during August; Eutima limpida A. Ag. fairly

common, but sexually immature; Lafwa calcarata A. Ag., not

uncommon, and in many cases the several phases in the devel-

opment of the medusa were taken. Dipurena comica A. Ag.

and D. strangulata McCr. were taken, but from a comparison

the specimens so intergraded as to suggest a probability of

their specific identity, with only varietal differences.

Of rarer medusz the following were taken: Willa ornata

McCr., taken in considerable numbers at various times during

the summer, were evidently breeding, as medusz of all sizes

and stages of development were found; Podocoryne carnea

Sars, taken sparingly; Epenthesis foleata McCr., not numer-

ous, colors less brilliant than those figured by Mayer for this

species. Species of Obelia were notably few during the present

summer, in rather sharp contrast with previous records. It

is one of the few medusz with a record extending throughout

the entire year and often occurring in immense numbers, this

latter being common with many species of Obelia.

EO. ScYPHOMEDUS#.

— These medusæ were more than usually abundant during the

summer, a thing not common at Woods Hole except very

. early. Many specimens of Aurelia, Cyanea, and Dactylometra

No. 427.] C@LENTERATE FAUNA OF WOODS HOLE. 555

were taken about the docks of the fish commission during

July, and. even August, which were sexually mature, and

from some of which ova were obtained and the develop-

ment followed as described below. That these were not an

isolated few left over from the earlier numbers of April and

May is evident in that many were taken in the open waters

of Vineyard Sound, and even in the open sea far from land

in the region of the Gulf Stream, mostly of the genus Cyanea.

They may have been caught up by southward currents from a

more northern locality along the coast of Maine or beyond,

and carried to these southern ranges.

Specimens taken into the laboratory and placed in aquaria

discharged developing ova in great numbers, literally covering

and packing the bottom of the aquaria. The following synop-

sis of the development of Cyanea may not be without interest

as having been carried on under the wholly artificial conditions

of the aquarium, from the later cleavage of the egg on to the

freeing of the ephyrae, — a circumstance not hitherto recorded,

so far as I am aware, though Bumpus has recorded the fact

that earlier phases occur readily under such conditions, and

both McMurrich and. Hyde likewise refer to the same fact,

but they do not seem to have succeeded in tracing the entire

development.

The early cleavage phases of Cyanea are passed while the

ova are still within the gastric cavity or while “ nursed " within

the ample folds of the manubrium or frilled oral margins.

A typical blastula results from total and regular cleavage and

appears to be followed by an early gastrulation.and the prompt

obliteration of the blastopore.

Soon after this, and while the larvze are still within the egg

membrane, cilia are developed, and they may be seen slowly

rotating within the membrane, which, however, is early rup-

tured and the embryo escapes as a free-swimming organism.

In shape it is at first almost spherical, but soon takes on the

oval or pyriform shape characteristic of most planule. Details

as to the origin and differentiation of the endoderm are not

yet worked out, but they seem to conform rather closely with

the observations of Hyde and the later conclusions of Smith

559 THE AMERICAN NATURALIST. | [Vor. XXXVI.

on Aurelia, as well as with the recent work of Hein on the

same form.

The larval history seems to vary greatly as to matters of

growth, transformation, etc. A few specimens attached them-

selves to the bottom of the aquaria in from six to ten days and

promptly assumed the typical scyphistoma stage, while others

were swimming planula at the end of as many weeks, showing

no tendency toward transformation.

Encystment. — Both McMurrich and Hyde have recorded an

encysting phase in the life of the planula preceding metamor-

phism. Hyde noted, however, that in one series of experi-

ments this did not take place. My own observations have

probably sufficed to reconcile these discrepancies by showing

that it is incidental rather than essential, — a mere adaptation

to changed conditions.

That encystment may and does occur under certain circum-

stances is not in the least doubtful, if not indeed a more or

less common phenomenon in development under artificial con-

ditions. A very considerable proportion of my own specimens

passed through such a stage, which varies from a few days to

many, and is, indeed, often a permanent and fatal one ; but, on

the other hand, it is not less certain that many passed through

the metamorphoses without the least sign of encystment, while

in others there was the aspect of incipient encystment common

in many hydroid planule, namely, the secretion of a delicate

perisarc-like sheath over the base of the planula in the process

of attachment. There seems little doubt, therefore, that in

these variable aspects we have the clue to the entire matter,

so far as these larva are concerned, and may consider encyst-

ment, as above intimated, to be an adaptation against unfa-

vorable conditions, and therefore comparable with the same

phenomenon among Protozoa and other organisms. May it

not hint the probable phyletic origin of the perisarcal structure

“so largely characteristic of one class of ccelenterates, to which

the facts under consideration may show more or less definite

reversion ?

T) he Scyphistoma. — The young scyphistoma is at first of

vasiform shape, the base small and adherent by the delicate

No.427.] CCLENTERATE FAUNA OF WOODS HOLE. 557

film of horny secretion referred to above. The primary ten-

tacles are usually four in number and arise as small buds from

the broad oral end of the polyp about the same time as the

mouth is formed. In many cases the number varies, only two

arising at opposite sides, two others following at intermediate

positions. More rarely only three tentacles occur in the pri-

mary set, followed by three others likewise in intermediate

positions, giving the polyp at first a trimerous and later a

hexamerous aspect, —a condition usually associated with the

phenomenon of a triangular mouth which continued even dur-

ing the entire history of the polyp. In several cases bifur-

cated tentacles occurred, and continued during the scyphistoma

stage. The average number of tentacles is sixteen, though

this may also vary considerably, many specimens being noted

with twenty or more. The scyphistomz of Cyanea are almost

clear white in color, in rather marked contrast with the dull

flesh color of the polyp of Aurelia, and when viewed in

colonies against a black background present a most beautiful

picture.

Stolonization.— This is a much less common feature in

Cyanea than in Aurelia, though not by any means rare. Sto-

loniferous processes may arise from the bases or sides of the

polyps, and growing laterally may become attached and serve

as points of origin for new buds. They are, however, fewer

in number and give rise to no such complex colonies as are

frequent in polyps of Aurelia. No cases of the direct origin

of secondary polyps from the body of the parent were noted

in the case of Cyanea.

Strobilization. — Owing to the small size of the polyp of

Cyanea, strobilization is comparatively inconspicuous and was |

only at first noted about the time the fully formed ephyra

rendered the fact evident and directed attention to it. The

segments are few in number, in many cases a single one only

occurring at a time, though three or four are not unusual; but

in no case were more than five noted upon any given specimen.

In the cases of polystrobilous specimens the basal portion

seemed to renew its activities, new tentacles arising and the

specimen showing every indication of healthy development,

558 THE AMERICAN NATURALIST. . [Vor. XXXVI.

probably later strobilating as before, though no demonstration

of this was undertaken.

The time involved between the drikoa of the planula

and the assumption of the strobila stage and the discharge of

ephyræ was like that of the planula history, quite variable.

The minimum time noted between the attachment of the

planula and the assumption of the strobila stage was ten days,

making the time from the origin of the planula to that of the

strobila about seventeen days, or about eighteen to twenty

days for the appearance of the ephyra. But as before inti-

mated, there is much variation on this point, probably more

than occurs in natural conditions.

The Ephyra.— Ephyræ liberated in the aquarium seemed

quite normal in morphological aspects. In color they are dull

brownish, the color showing itself as the strobilization proceeds

toward maturity. In habit the young ephyra appears somewhat

sluggish, and, while capable of active movement by the con-

‘tractions of its bell, lies rather quiescent in the aquarium,

seldom moving except as stimulated by agitation of the water

or otherwise. While in rest it lies upon the aboral surface,

with the manubrium extended upward, probably a condition

assumed for the capture of prey.

Several specimens were marked by definite variation in the

number of lobes or other organs. A specimen with nine lobes

had also a similar number of rhopalia, while another with eight

normal lobes had three extra rhopalia irregularly disposed at

intermediate positions. Several specimens taken in the tow

showed also similar variations, as did likewise several adult

specimens taken during the summer and at previous times.

Feeding. — In rearing the polyps several experiments were

made upon the feeding habits. Scrapings of slime, algz, etc.

from the eelgrass, which contained numerous Protozoa, were

found to be among the most successful sorts of diet. Larvæ

of gastropods and starfish were also taken readily by the

polyps, the former especially being apparently quite accept-

able. Diatoms and other micro-organisms taken from the

deep waters of the Sound apparently proved deleterious, the

polyps in aquaria supplied with this food showing evident and

No, 427.] CELENTERATE FAUNA OF WOODS HOLE. 559

rapid decline of vigor and health. In small aquaria numerous

cases of cannibalism were noted, the scyphistome greedily

devouring any planula which came within the grasp of the

vigilant tentacles, the entire process of engulfing the victims

being several times observed.

A Cubomedusa from Woods Hole. — Several specimens of an

interesting medusa of this order were taken in the tow, the

first on August 13, in Great Harbor, a second the following

day at North Falmouth, Buzzards Bay. Two or three others

were taken within the next day in Great Harbor.

In general features the medusa resembles very closely one

figured by Mayer from the Tortugas (Bull. Mus. Comp. Zoöl.,

Vol. XXXVII, No. 2, p. 70), and to a less

degree a medusa described by Fewkes from

the Bermudas (Bull. Mus. Comp. Zool, Vol.

XI, No. 3). In many respects, however, it

differs from both, namely, in its larger size, |

deeper color, apparent absence of gastric fila-

ments, figured by Mayer. Both Fewkes and

Mayer consider the specimens taken by them

as immature, and their identification was

accordingly somewhat doubtful, the former

assigning his to the genus Tamoya, the latter

referring his, together with that of Fewkes, tothe genus Charib-

dea. Certain it seems that unless the specimens are immature

they could hardly be included within either genus, or indeed

within any other of the at present recognized genera of the

family. The absence of velar canals and pedalia, as well as

the very short tentacles, would of themselves be sufficient to

exclude them.

The same is likewise true of the specimens under considera-

tion. They are so nearly identical in morphological features,

size, etc., as to suggest something like maturity, unless, per-

chance, they should be indigenous species, — an exceedingly

doubtful supposition. If borne hither from the Tortugas or

Bermuda by currents, it would seem that the time involved in

transit should have given at least some differentials of size or

other peculiarity.

Fic. 4. — Charibdea

verrucosa.

560 THE AMERICAN NATURALIST.

The following descriptive characters may serve to show in

how far they fail to conform to the type characters of the

genus: bell ovoid in profile, cuboid as seen from either pole;

size from 2 to 3 mm. in short diameter by 4 to 5 mm. in

height. Color a dull amber, somewhat translucent; exum-

brellar surface dotted irregularly with light brownish warty

clusters of nematocysts. Tentacles brown in color, rather

short and spindle-shaped, with deep annulations. Rhopalia

four, radially located, set in rather deep pockets somewhat

above the level of the margin, each with several ocelli near

the tip. Velarium well developed, and strengthened by four

frenulz or thickenings of the subumbrella in the region of the

‘radii. - Absence of any traces of velar canals is noteworthy, as

is likewise absence of mesenterial filaments. Distinct gonads

could not be distinguished, though sections showed a genital

region and ova in process of development. Manubrium well

developed, quadrate in form. In case the specimens may be

referred to the genus Charibdea, it must be upon the assump-

tion of immaturity, as Mayer has suggested, in which case they

compare fairly well with Mayer’s C. aurifera, though in size

somewhat larger and more brownish in color. If, as seems not

unlikely, they must be given specific distinctness, I would pro-

pose for the species the name verrucosa, as significant of the

warty knots of nematocysts which adorn the outer bell.

THE ZOOLOGICAL LABORATORY,

SYRACUSE UNIVERSITY, May 1, 1902.

THE BODY SENSE HAIRS OF LEPIDOPTEROUS

LARV:.!

WILLIAM A. HILTON.

Ir is well known that, as a rule, all arthropods have their

bodies and appendages clothed with hairs, or sete, which are

simply modified parts of the cuticle.

These hairs vary in form and in structure. In the most

common type the hair is hollow, and its lumen communicates

with the body cavity by means of a “pore canal" in the cuti-

cle (Fig. 1). Through this pore various structures may extend

into the lumen of the hair, as a prolongation of a hypodermal

cell, gland ducts, and nerves.

There are published references to solid hairs of insects; but

if we except certain minute elevations of the cuticle, such as

are represented in Fig. 2, and which are more or less hairlike,

I found none in the insects studied. It should be said, how-

ever, that certain scalelike appendages of the cuticle of the

larva of Corydalis (Fig. 3, 2) appear to be solid; but as these

occur, each at the end of a pore canal, they may prove to be

hollow.

The most commonly observed structure that passes through

the pore canal into a hollow hair is a prolongation of a hypo-

dermal cell, which is much larger than the ordinary cells of the

hypodermis (Fig. 4, 7). Such a cell was named a *trichogen,"

by Graber, as it is believed to be the element that produces the

hair. In many cases there are associated with the trichogen

one or more gland cells, which discharge their secretion through

the hair; the stinging hairs of certain larvae are examples of

this type. Such hairs may also have a nerve extending to

them, and perhaps they always do.

A type of hollow hair which occurs much more commonly

than does the glandular hair is one into the lumen of which

! Contribution from the Entomological Laboratory of Cornell University.

561

562 THE AMERICAN NATURALIST. [VoL. XXXVI.

there extends only a trichogen cell and a nerve. The larger

number of the organs of special sense of arthropods are hairs

of this type, more or less modified. Frequently, however, the

modification has been carried so far that the organ of special

sense is not at all hairlike. This is espe-

cially true of many of the supposed organs of

taste and of smell. I have made no attempt

to study these.

The investigation, the results of which are

given in this paper, was confined to a study

Bo hd: ie of those hollow hairs, or setze, which are dis-

the silkworm (Bombyx tributed over the surface of the body and are

an believed to be organs of touch. The primary

object of this investigation was to determine as definitely as

possible just what type of hair on the body surface is a sense

hair; a secondary object was to determine in what ways this

type is modified in various lepidopterous larvae. Some work

was also done on several other orders of insects, with the result

that a close correspondence was found in the structure of these

hairs among even widely separated orders.

METHODS.

In order to definitely demonstrate which hairs were sensory

it was necessary to have recourse to special histological methods,

as the terminations of

the nerves for touch in

insects are exceedingly

delicate, and the usual

microscopical methods

are entirely inadequate

for the successful differ-

entiation of the nervous 244599

tissues. For general

anatomy of the sense Fig. 2.— Sections through hairs from a tiger moth

hairs the usual fixing is aid

and staining fluids were used; but for staining the peripheral

nerve fibers and cells the zuzra vítam methylen blue method

No. 427.] HAIRS OF LEPIDOPTEROUS LARVE. 563

was found to be especially useful; and, as the larger part of

the work was based upon results thus obtained, it may be well

to give the exact method of procedure.

The fluid used for injecting was Griibler’s B.x methylen

blue, į per cent solution in normal salt. The injection by means

of a small syringe was made usu-

ally in the side, back of one of

the last abdominal segments, and

enough liquid was injected to color

the segments near the head, care

being taken to insert the canula

only under the hypodermis and

muscles, and not into the alimen-

42161 g eed dale

IG. 3. — 8

cornuta, showing large and small fla

scales supplied with nerves and cells. tary canal.

i gi

e venas Au a After injection the animal was

nerves demonstrated by methylen blue. left quiet for a period of about

three hours for most insects, but with some, as with Pieris,

two hours was a better time, and four hours seemed to be best

for Datana larve. It is important that the insect remain

preparation, the

alive during this period after injection.

When the proper period has elapsed the

nerves and nerve cells should be well

stained and almost all other tissues free

from stain. After successful staining the

specimen is cut open lengthwise and

pinned out over a hole cut in sheet cork,

the muscles and viscera are removed by

careful dissection, and then, on examining

under a microscope, the nerves and nerve

cells are seen stained upon the surface

of the unstained hypodermal layer. The

bases of the hairs may be seen through

the hypodermis, and it is easy to trace

nerve cells and fibers in connection with them. By keeping

the preparation wet with normal salt solution and using differ-

ent powers of the microscope, much may be learned regarding

the structure and distribution of the peripheral nervous system

without further preparation; for this work the silkworm (Bombyx

Fic. 4. — Section of hair

of Datana.

564 THE AMERICAN NATURALIST. [Vor. XXXVI.

mori) was very good, because the blue nerves on a pure white

background stand out very clearly. Of course, such prepara-

tions last but a short time, fading out in an hour or less, and

the ultimate distribution of the terminal nerve fibers cannot be

seen without sectioning.

The following modification of Bethe’s formula for fixation

was used :

Ammonium molybdate . TRES A

Con HOD. oe OO ee 000 FIO drops

iust Hee eee a one.

This solution was used ice cold and allowed to act on the

tissues from eight to twenty-four hours, after which they were

washed well in cold distilled water and placed in absolute

alcohol for about three hours, then cleared in xylene, and either

mounted whole or imbedded and sectioned.

HISTORICAL.

Setiferous sense organs in arthropods were described by

many early workers; but the first description of a bipolar

nerve cell one termination of which was at the base of the hair

and the other continued with the nerve trunk to the central

nervous system was made by Leydig in 1851, for certain hairs

of Corethra plumicomis. This discovery of Leydig was verified,

and the form described is now generally regarded as the type

of sensitive termination in arthropods. Leydig undoubtedly

mistook the trichogen cells for nerve cells; these two elements

were later distinguished by Hauser (80) and Villanes (81).

Later works, especially of Retzius (90-95) and Vom Rath

(91-96), brought forth the following results:

In arthropods all sensory terminations, the eyes excepted,

are in hairs. To each hair there corresponds at a greater or less

distance from the base one or more bipolar sense cells, of which

the distal prolongation penetrates into the interior of the bair

terminating without ramifying, the other prolongation (proximal)

ina nervous center. These observations, which accord so well

with other classic works on the peripheral nervous system,

were made somewhat doubtful by more recent works.

No. 427.] HAIRS OF LEPIDOPTEROUS LARVAE. 565

In Crustacea, Bethe (95) and Nusbaum (97) recognized the

sensory neuron as a bipolar cell, the distal portion penetrating

the hair without ramifying and the other prolongation con-

tinuing with the nerve trunk; but, at the same time, they also

described a plexus of cells, — not a system of neurons, but a true

plexus. Bohumil Nemec (96) found in certain isopods a system

of peripheral ganglia. These ganglia are centers of a hypodermal

plexus compared by him to the plexus of Bethe. Rina Monti

(93) described a similar plexus in insects, and Holmgren (96)

described and figured such a network of multipolar cells with

their processes in the larva of a sphinx moth.

In '97 and '98 Duboscq summarized very well the results of

many authors and added some of his own observations on

Orthoptera and chilopods. He used both the Golgi method

and methylen blue and discredits much of the former work on

the nerve plexus obtained by methylen blue, and that on the

termination of the nerves by the Golgi method.

Sensory CELLS AND NERVES OF Bopy SENSE HAIRS.

Following the sensory nerve trunks peripherally, they are

seen to run straight out from the central nervous system,

dividing dichotomously, most of the branches coming off when

the region of the back is reached. Aftera number of divisions,

small nerves from bipolar nerve cells join the larger trunks,

and the peripheral parts of the larger nerves may also be traced

to bipolar nerve cells (Fig. 5).

In the silkworm (Bombyx mori) bipolar nerve cells are

spindle-shaped and have a very dark nucleus, which in many

cases nearly fills the cell body (Figs. 5-6). In most forms

studied the cells are more nearly spherical, but yet not mark-

edly so (Figs. 5-6). Asa general rule the sensory cell is

quite a distance from its hair, but again in the myron sphinx

there is a slight variation, the nerve cell being at the edge of

the pore canal, almost in its cavity; this may not indicate

that the nerve is really shorter, but only apparently so, because

of the thickness of the cuticle and consequent length of the

pore canal.

566 THE AMERICAN NATURALIST. [Vor. XXXVI.

The prolongation of the nerve just beyond the sense cell

is of considerable thickness, staining deeply; and abruptly

after this the nerve extending to the hair is very minute.

It was impossible to trace the nerves farther than the bases

of the hairs where whole mounts were made, and Duboscq

(97 and '98), in Orthoptera and chilopods, traced the nerves only

this far. However, earlier workers with methylen blue who

studied simply the surface views represent nerves coming from

the tips of the hairs; but it seems probable that such figures

are in large part diagrammatic. Vom Rath found by the Golgi

oe O Be

£ m=

p e.. ^

ow 7 \

iC -h T gm X . Y

5 \ dl

( (5;

mue ue

OA ;

-~m

woh

Fic. 5. — Surface vi f I lls f the silkworm (Bombyx mori).

: Ba f hairs also shown. Methylen blue.

method cavities of sensory hairs filled with nerves; this result

is regarded as an artifact by Duboscq, who shows clearly how

appearances like nerves may be obtained in the cavity of hairs

due to deposits of chromate of silver; and he shows quite clearly

that when nothing but the nerve cell and fiber is impregnated

the nerve fiber stops at the base of the hair, as was apparently

the case in his methylen blue preparations.

Retzius ('95), in Astacus, by means of the methylen bluemethod,

traced the nerve to the base of the hair. This attracted the

No. 427.] HAIRS OF LEPIDOPTEROUS LARVA. 567

attention of Bethe ('96), whoinvestigated this point and found two

sorts of hairs in Astacus, one as described by Retzius, in which

the cavity of the hair was shut off at its base by chitin, and the

other having its cavity freely open at the base. In the closed

hair a nerve fiber was seen to stop at the base of the hair, but

in the open hair it was continued a short distance into the

shaft, and he suggests that probably this fiber goes nearer

the tip than was observed in the specimens.

It was only by means of sections that it was possible to

trace the nerve in the cavity of the hair, and I found it very

Fic. 6. — The figure in the cen i th t the left views of subhypodermal nerves

= us tror the "silkworm the bie ? th n are also shown. The figures at the

y ) Methylen blue.

difficult to obtain successful preparations. In the silkworm

nerves were traced to the hairs by means of sections, which

showed the nerves passing between the hypodermal cells, past

the gland cell, or trichogen cell, without any branches, and often

as far as the collar of the hair. In a few cases the nerves were

traced a short distance into the shaft; but they appeared to

end not far from the base of the hair on the side of the lumen

(Fig. 7). Either in the hair or just below it a rather large

568 THE AMERICAN NATURALIST. | [Vor. XXXVI.

swelling of the nerve was usually seen ; this is possibly an arti-

fact. In a few cases nerves were traced perhaps one-half the

length of the cavity. In these cases there was a swelling of

the nerve just before its apparent termination (Fig. 7).

I found no evidence to indicate nerves ending in gland cells

or trichogen cells by such branches as have been described and

figured by Blanc (90) but in every case the very fine nerve

termination could be traced up past the hypodermal cell layer

with no branches.

SUBHYPODERMAL NERVE PLEXUS.

Villanes (81), in insects, figures a subhypodermal nerve

plexus made up of multipolar nerve cells. In '93 Rina Monti,

also working on insects, described a similar plexus, and in '95

Bethe described similar structures in Crustacea. Nemec (96),

in isopods, and Holmgren ('96), in a sphinx-moth larva, show

multipolar networks of subhypodermal cells. All the work

thus far was by means of methylen blue or ordinary methods,

and because of the apparent antagonism with the neuron theory

much attention was directed to this subepithelial plexus, and

doubts were raised as to the nervous nature of these cells.

In 1898 Schreiber, who had already in 1897 together with Nus-

baum published a paper on the peripheral nervous system of

Crustacea showing these multipolar cell plexuses, published

another short paper showing these cells demonstrated by means

of a modified Golgi method, and at the close of his article

draws the following conclusions :

I. An identical methylen blue staining of the subepithelial multi- and

bi-polar nerve cells.

2. Like staining of both forms of cells by the Golgi method.

3. True connections of the nerve cells with nerves.

A few months later Holmgren confirms, on the whole,

Schreiber's results, but speaks of many-branched subhypo-

dermal cells, as described by Bethe in Crustacea, and probably

confused by Nusbaum, Schreiber, and other writers on Crus-

tacea with multipolar nerve cells. Holmgren distinguishes

No. 427.] HAIRS OF LEPIDOPTEROUS LARVÆ. 569

these cells from the true multipolar nerve cells of Crustacea,

regarding them as many-branched mesenchymatous cells. In

lepidopterous larvae Holmgren states that there are no mesen-

chymatous cells like those in Crustacea which one would con-

fuse with nerve cells.

In 1897 and 1898 Duboscq criticises the work of many pre-

vious workers upon multipolar subhypodermal nerve cells and

states as his opinion that the bipolar nerve cell is the only true

nerve cell in the hypodermis of arthropods, and that there is

no subepithelial plexus. Although much of the criticism of a

subepithelial plexus by Duboscq is very just, it seems to me

that simply calling all such structures so described connective

Methylen blue.

tissue is hardly justified when we look over the works of

numerous observers. It may in part be true, as Holmgren

points out, that much which has been described, especially in

Crustacea, may come under the head of mesenchyma, but do

we not have multipolar nerve cells as well?

The occurrence of evidently multipolar nerve cells together

with bipolar nerve cells and fibers I found to be very constant

in lepidopterous larvae (Fig. 6). The nucleus of these multi-

polar cells was a very well marked, clear area occupying a large

part of the center of the cell; in the center of this clear

area was a darker staining portion, possibly the nucleolus.

These cells with their many fine branches always take a paler

blue stain than the other nervous structures in the skin. The

very fine branches of these multipolar cells run out long

570 THE AMERICAN NATURALIST. [VoL. XXXVI.

distances, and in the most successful preparations may be seen

to join with similar branches from other multipolar cells and

with other nerves. In some cases, as shown in Figs. 5-6,

branches from these went directly to bipolar nerve cells or to

large nerve trunks. These cells are always either in series

with at least one bipolar nerve cell (Fig. 6) or have some of

their branches joining the nerve trunk of the bipolar cell, or

they are not far from one of these sensory cells. Often it

was seen that these cells occupied a position closer to the

hypodermal cells than other nervous tissues, and sometimes

large nerve trunks would end bluntly, and under the ends of

such trunks would be found a multipolar

cell, perhaps giving one of its branches to

a bipolar nerve cell (Fig. 6).

These cells were best observed in the

silkworm and in the myron sphinx larva,

but they were also seen in Pieris, Datana,

Papilio, and practically in all forms studied.

Whether these multipolar cells are nervous

Fic.$.—Inner face of body tissue or not cannot be fully decided as yet,

saat oe but although they differ somewhat in posi-

tion and staining qualities from other

dn cl etur « undoubted nerve cells, they have many

the hairs. (x22) Methy- characteristics which favor their being

regarded as nervous tissue.

Aside from a network of multipolar cells with their processes

another sort of plexus was observed. In tiger-moth larva the

hairs are on little knoblike elevations. After injection and

staining of nerve fibers there are found to go to each of these

hair tubercles a large nerve, which, after entering the elevation,

may be seen to break up into branches (Fig. 8), and in sec-

tions it may be seen that these branches break up still finer

and each subdivision goes to a bipolar cell connected with a hair.

Now, besides these larger nerve trunks, smaller ones are also

seen entering the elevations, and often in successful prepara-

tions near the edge of the hair tubercles there may be seen

a very complex network of apparent nerve fibers with many

swellings, as shown, much enlarged, in Fig. 9. This net work

No. 427.] HAIRS OF LEPIDOPTEROUS LARVE. 571

is apparently entirely without cellular elements and is at the

same time a part of the nervous system. These networks

in tiger-moth larvae were only observed near the hair tubercles ;

in the silkworm a much less marked network of large and

small fibers was occasionally found.

It is hard to say whether this network is truly a part of the

nervous system or not; but in any case these networks are

Fic. 9. — Nerve plexus from edge of hair tubercle of tiger-moth larva, much enlarged.

Methylen blue.

not at all like the multipolar cells just described, no nuclei or

cells being recognized, but they are well stained with methylen

blue and join undoubted nerves.

CENTRAL ENDING OF SENSORY NERVES.

It was not the purpose of this investigation to make a study

of the central nervous system, but as it became necessary

to trace the nerve fibers to the ganglia, several interesting

conditions were noticed which have a more or less direct bear-

ing upon the previous work; a few of these will be spoken of,

although it is realized that no complete discussion can be given

without considerable further study, and consequently further

57? IHE AMERICAN NATURALIST. [Vor. XXXVI.

investigations will probably bring out more complex relations

than here portrayed.

On each side of the abdominal ganglia of Pieris there are two

large branches or nerve trunks (Fig. 10). The more cephalic of

C these comes from bipolar nerve cells at the

bases of hairs, and the other more caudal

branch goes to muscles; so, as nearly

as could be determined, the cephalic

trunks are sensory and the caudal ones

motor. There may have been a few

motor nerves in the cephalic trunk and

M LLL ganglion Sensory nerves in the caudal one, but if

from Pieris. Methylen blue. there were such, they were inconspicuous

and not noted in the examination of numerous specimens; so

for the present, at least, the more cephalic branch may be

regarded as a sensory branch.

This sensory branch upon entering

the ganglion seems to run as a distinct

tract cephalad without sending branches

to the central cell area of the ganglion.

This tract runs cephalad for some dis-

tance along the outside of the connec-

tive and is finally lost sight of. The iara pone

course of the motor nerve is quite Io caterpillar (Automeris io). a

different ; its fibers seem to come '?""* rok aem

directly from the central cell area of either side of the ganglion.

Aside from the fibers just described, there are nerve fibers

which may be seen to run through

the ganglion longitudinally ; these

take a lighter stain. They run along

the connectives and are just inside

the sensory tracts.

Variations in the number and posi-

Fro. 12.—Small and large hairs from tion of the nerve trunks from the

p DN ganglion occur in different species.

In Io and in a tiger moth the same sensory tract may be noted,

but part of the sensory trunk upon entering the ganglion leaves

the sensory tract and enters the cell area (Fig. r1), but on

n from

No. 427.] HAIRS OF LEPIDOPTEROUS LARVA. 573

tracing the nerve to the periphery it is found that part of the

fibers of the more cephalic nerve come from the muscles,

and so the nerve may be regarded as a mixed nerve, showing

why some of its fibers do not follow the sensory tract.

Fic. 13. — Section of hair from Sphinx chersis.

In lepidopterous larvz the two sides of the ganglia seemed

to be separate, possibly because of imperfect methylen blue

Stain. In Coryalis larva fibers were seen to cross from side to

side.

GENERAL RESULTS.

In the lepidopterous larvz examined two kinds of body sense

hairs were found: first, simple hairs, those having a single

enlarged hypodermal cell at the base (Fig. 12); and, second,

glandular hairs, or those having two rather large modified hypo-

dermal cells (Fig. 12). In the latter case the smaller cell is

probably the trichogen cell, and the other a glandular cell of

some sort. As these figures were drawn from specimens

Prepared in the ordinary way, the nerves are not shown.

Examples of simple hairs are represented by Figs. 13-18.

The insects from which these figures were drawn are indicated

in the explanations of the plates. Glandular hairs are repre-

sented by Figs. 12 and r9.

574 THE AMERICAN NATURALIST. [VoL. XXXVI.

In all species examined both simple hairs and glandular

hairs were found to be supplied with a bipolar sensory nerve

cell and fiber. Sometimes, as in Pieris and Datana, there are

two quite distinct kinds of hairs, large

and small. In this case the large hairs

are supplied by large bipolar nerve

cells, and the small hairs by small

ones (Fig. 20).

In tiger moths and tussock moths

the hairs differ somewhat from those

in other forms; here the single hairs

Fic. 14. — Section of hair from body of other lepidopterous larvae are repre-

e sented by bunches of hairs clustered

together on tubercles. Each tubercle is supplied by one

or more large nerve trunks, which break up on entering the

tubercle. A bipolar nerve cell from each

hair in this case is harder to determine

absolutely ; but as some of the hairs were

found to have such nervation, probably all

do, for all are of the same kind. The

structure of these hairs differs somewhat

from those already described in the form

and shape of the cells, these sometimes F

being hollow or saclike with radiating

nuclei (Fig. 2). The base of the hairs also presents variations,

as shown in Fig. 2; here the chitinous base of the hair seems

to extend down below the level of the cuticle. In a section

through the base of a hair of Notolophus

(Fig. 21) the opening of the hair is seen

to be nearly closed in two places by strong

processes of chitin, and sections taken

just one side of the middle line show

apparently a closed hair, indicating how

small the opening is in the base of the

hair.

All the hairs examined in lepidopterous larvae were open

and undoubtedly sensory, that is, supplied with bipolar

sense cells.

Section of hair of

IG. 15. — Se

Basilarchia archippus.

Fic. 16. — Hair from Samia

cecropia.

No. 427.] HAIRS OF LEPIDOPTEROUS LARVA. 575

Differing from the hairs just described, there are distributed

over the surface of the body of the larva of the Io moth spine-

like clusters, prolongations of the body

wall (Fig. 22); most of these spines are

tipped with short, strong, chitinized caps,

and a few with long, slender, hairlike

processes. These spines when touched

to the tender surface of the hands pro-

duce swellings which are slightly painful.

Numerous specimens stained by methylen :

blue demonstrated fine nerves running ri; simple hair from Chisio-

into these spinous processes and extend- fmit mp:

ing almost to the tips, but no bipolar nerve cells were demon-

strated. Evidently here we have glandular hairs, but not

sensory hairs; however, in the same

specimens there were found hairs of the

usual type, much smaller than these proc-

esses of the body wall or spines; these

true hairs were located both on the bases

of the spines and upon the body wall

between the spine clusters (Fig. 22, A).

No difficulty was experienced in demon-

strating a bipolar nerve cell at the base,

Fic. 18. — Silver-spotted ue of each of these true hairs.

To summarize briefly : all lepidopter-

ous larvas studied have their bodies clothed with hairs, and all

the hairs are sensory, having a bipolar nerve cell at their base

and supplied with a minute nerve fiber, \

with the possible exception of the

poison spines of Io caterpillars.

To carry the generalizations a little

further, the following observations are

useful. In the larvae of May beetles

; : g

the hairs of the body surface are sup- Fic. 19. — Large, probably glandular

plied with bipolar nerve cells (Fig. 23),

and in Orthoptera, as already spoken of by Duboscq (97), the

hairs have a bipolar sense cell at their base. In the larva of

Corydalis, bipolar nerve cells with their fibers supply each of

576 THE AMERICAN NATURALIST. [VoL. XXXVI.

the numerous little, black, scalelike hairs which so thickly cover

the body ; large hairs have larger bipolar nerve cells (Fig. 3).

Günther (01) shows scales of a lepidopterous wing with

nerve cells at the base of both simple and glandular scales,

probably a similar result to that just described in Corydalis.

Q, |

Fic. 20

= S Oe

cae "t -—-—

e Weide

= eee

à :

SENN

ANNANN

X NS

Fig. 21. Fig. 22.

Fic. 20. — Bases of large and small hairs from Datana, showing their bipolar nerve cells.

Methylen blue.

Fic. 21. — Section through center of hair of Motolophus leucostigma.

Fic. 22. — Cluster of spines from Io. (X 17.) Methylen blue. 2, hairlike spine ; 5, usual

type of spine.

Fic. 23. — Large and small hairs from beetle larva with their sense cells and nerves.

Methylen blue.

A SUMMARY.

1. Lepidopterous larvæ are clothed with hollow hairs, each

of which is supplied by a bipolar nerve cell, a process of which

penetrates a short distance into the hair and probably termi-

nates before reaching the tip.

2. In most species all body hairs are sensory; large hairs

. are supplied by large bipolar nerve cells, and small ones by

smaller bipolar cells.

: » Under the hypodermis of caterpillars there is a system

: ar cells more or less intimately connected with nerve

No. 427.] HAIRS OF LEPIDOPTEROUS LARV&E. 577

cells and fibers which (a) stain lighter than the larger nerves

and (7) are closer to the hypodermis than the other cells

and fibers.

4. Nerves from bipolar sensory nerve cells go to the central

nervous system, run to the ganglia, leaving at once to follow

on the outside of the connectives cephalad, forming a well-

marked sensory tract. Motor nerves — those that go to mus-

cles — seem to come directly from the central cell areas of the

ganglia.

5. Almost the only sensory termination of nerves on the

body of insects is by means of hairs.

I wish to make my very sincere acknowledgments to Pro-

fessor Comstock and the Department of Entomology for much

invaluable aid in the preparation of this paper.

BIBLIOGRAPHY.

'96 BETHE ALB. Ein Beitrag zur Kenntniss des peripheren Nerven-

systems von Asticus. Anat. Anz. 1896.

'97 BETHE ALB. Das Nervensystem von Carcinus menas. Arch. f.

Mikr. Anat. 1897.

'90 BLAwc, M. L. La tête du Bombyx mori à l'état larvaire. Extrait du

volume des travaux du laboratoire d'études de la soie. Lyons,

1889-90.

'97 Dunosco, O. Sur la terminaison des nerfs sensitifs des chilopods.

Ann. de l’ Univ. de Grenoble. 1897.

'97 DusoscQ, O. Sur le système nerveux sensitif des tracheates (orthop-

teres, chilopods). Arch. de Zool. Exp. et Génér. 1897.

'98 DuroscQ, O. Recherches sur les chilopods. Arch. de Zool. Exp. et

Génér. Sér.9, tome vi. 1898.

01 GÜNTHER, K. Ueber Nervenendigungen auf dem Schmetterlings-

flügel. Zo/.J/aAró. Bd.xiv. 1 Taf. rg9or.

'80 HAUSER, G. Physiologische und histologische Untersuchungen über

. das Geruchsorgan der Insecten.

'95 HOLMGREN, E. Studier ófver hudens och koertebart hudorg. mor-

folog. hos skand. mikrolepidopterlarven. X. Svenska Vetensk.-

Akad. Handlingar. .

"95 HOLMGREN, E. Zur Kenntniss des Hautnervensystems der Artro-

poden. Anat. Anz. 1895.

578

"93

THE AMERICAN NATURALIST.

HOLMGREN, E. Zum Aufsatze W. Schreibers * Noch ein Wort.”

Anat. Anz. 1898

KUNCKEL et GAJAGNAIRE. Rapport du cylindre-axe et des cellules

nerveuses périphériques avec les organs des sens chez les insects.

C. R. Aud. .Se.. 1881.

LEYDIG, F. Anatomisches über Corethra plumicomis. Zeitschr. f.

wiss. Zool. 1851.

MonTI, RiNA. Ricesche microscopische sul sistema nervoso degli

insetti. Boll. scient. 1893-

NEMEC, BOHUMIL. Zur Kenntniss des peripheren Nervensystems

einiger Crustaceen. Anat. Anz. 6

NUSBAUM and SCHREIBER. Beitrag zur Kenntniss des periphe-

rischen Nervensystems bei den Crustaceen. Biol. Centralbl. 1897-

RATH, O. Vom. Zur Kenntniss der Hautsinnesorgane der Crusta-

ceen. Zool. Anz. 1891

Ratu, O. Vom. Ueber die von C. Claus beschriebene Nervenen-

digungen in den Sinneshaaren der Crustaceen. Zool. Anz. 1892.

Ratu, O. Vom. Ueber die Nervenendigungen der Hautsinnesorgane

der Arthropoden nach Behandlung mit der Methylenblau und Crom-

silbermethode. Berichte der natur. Gesellsch. zu Freiburg. | 1894.

RATH, O. Vom. Zur Kenntniss der Hautsinnesorgane des sensiblen

Nervensystems der Arthropoden. Zeitschr. f. wiss. Zool. 1896.

RETZIUS, G. Zur Kenntniss des Nervensystems der Crustaceen.

Biol. Unt. N.F. 1896

RETZIUS, G. Ueber die Nerven-Prinzipien in der Lehre von der

Einrichtung des sensiblen Nervensystems. Biol. Unt. N.F. 1592.

Rerzius, G. Das sensible Nervensystem der Crustaceen. Biol.

Ont. N.F... 1895.

SCHREIBER, W. Noch ein Wort über das peripherische sensible

Nervensystem bei den Crustaceen. Anat. Anz. 1898.

ViLLANEs. Sur les terminations nerveuses sensitives dans la peau

de quelques insects. cad. .Sc.

VILLANEs. Recherches sur l'histologie des insects et sur les phéno-

ménes histologiques qui accompagnent le développement postem-

bryonnaire de ces animaux. 7h. Pairs. 1883.

ABREVIATIONS USED IN FIGURES.

bm = basement membrane. À = hair. co = connective.

hz hy = hypodermis. m = multipolar cell.

mr = motor root. » = nerve.

s = sensory cell.. sy = sensory r

“st = sensory tract. t = trichogen cell.

. Lines near drawings indicate yọ mm.

HISTOLOGICAL CHANGES IN HYDRA VIRIDIS

DURING REGENERATION.

HANNAH TERESA ROWLEY.

ALTHOUGH a great deal of work has been done on the regen-

eration of hydra, no one has as yet attempted to make out the

histological changes that take place. This point would seem

to be one of special interest, since the old piece appears to

change its form as a whole to produce a new.animal. The

principal question to which I wished to find an answer was

whether, during the period of regeneration, the old cells go

over without change into the tissue of the new animal, or

whether new cells are formed, and if so, in what part or parts.

Green hydras were used almost exclusively, since they regen-

erate more readily and with fewer abnormalities than does the

brown species. Hydras of various lengths were taken, the

foot end and circle of tentacles were cut away, and the remain-

ing middle parts of the body wall were each divided by cross

cuts into four——in some cases two—small rings. These

were allowed to regenerate, and at different periods, ranging

from immediately after the cutting to some days after the

formation of new tentacles and foot, were killed in a solution

of corrosive acetic, hardened, sectioned lengthwise, and stained

in Delafield haematoxylin. In some cases the slides were

dipped for a moment into a weak solution of picric acid in

absolute alcohol, in order to differentiate the endodermal

‘tissue.

Observation of the living piece showed that the ends became

‘rounded, closing in from fifteen to sixty minutes after having

been cut ; that the piece then remained without permanent

‘change: of form for thirty or forty hours, although at intervals

it might change its shape by expanding and contracting. At

about this time small knoblike outgrowths, the tentacles, began

to appear. at one end, the other end fixed itself to the nearest

579

580 THE AMERICAN NATURALIST. [Vor. XXXVI.

support, and in a few days the proportions of a normal hydra

were assumed.

Serious difficulties arose in the microscopic study of the

prepared sections, which it may be well to mention. Although

undoubted karyokinetic divisions were seen in both neuro-

muscular and interstitial cells of the ectoderm, and in th-

endoderm, it was found almost impossible to determine the

exact amount of dividing tissue, for the following reasons.

Throughout the entire period of regeneration there is an

active production of new nettle cells, which are formed from

interstitial cells of the ectoderm. This process begins by a

slight sickle- or moon-shaped thickening of protoplasm along

a part of the cell wall. In certain sections this thickening

appears as a small, darkly stained rod which may easily be

mistaken for a small nucleus in process of division. Again, in

sectioning, this curved, thickened red may be cut at such an

angle as to bring distinctly into view only its extremities, pro-

ducing the appearance of two small groups of darkly stained

material. These could be distinguished from two separating

groups of chromosomes only by bringing into focus the con-

necting line of the rod, which would lie slightly above or below

the ends. In many cases it was found almost impossible to

determine whether such a cell was forming a cnidocyst or was

dividing karyokinetically. These difficulties were increased

by the small size of many of the interstitial cells, by the fact

that the wall of such cells is not sharply defined, and by the

fact that at the time of the formation of nettle cells it is very

often difficult to make out the nuclei of the changing cells.

Moreover, many nuclei of both neuro-muscular and inter-

stitial cells were found to be in a state that distinctly suggested

activity of some sort, — the chromatin mass large and loose, or

even in many interstitial cells broken and scattered throughout

the nucleus. This appearance was very different from that o

nuclei of typical resting cells in the same section, the latter

containing one or two well-defined nucleoli. Yet the large

number of nuclei. with the chromatin scattered as just described,

in sections containing comparatively few karyokinetic spindles,

would seem to indicate that all such cells were not necessarily

No. 427.] CHANGES IN HYDRA VIRIDIS. 581

dividing. Where so many cells were about to divide or had

just completed division, we should expect to find a correspond-

ing number undergoing the actual process. This, however,

was not the case.

A further point should also be kept in mind, vzz., the possi-

bility that multiplication of the interstitial cells may be con-

cerned with the development of new nematocyst cells rather

than with the regeneration of new tissue.

In order to get more light on these points, the tissue of a

normal hydra was examined for comparison with that of the

regenerating piece. Another hydra was irritated with a blunt

needle in order to induce it to discharge the nettle batteries.

It was then left undisturbed for about twenty hours, at which

time it was killed and the tissue prepared as has been described.

It was hoped by these means to obtain further light on the

difficulties of the question, and to be able to distinguish more

clearly between the phenomena of regeneration and those

merely incidental to the formation of new nettle cells.

In the case of the stimulated hydra, nettle cells were found

to be forming as in the regenerating pieces. Moreover, in the

tissue of this hydra and of the undisturbed normal one also I

was surprised to find undoubted evidence of cell division, in

addition to the very general loose and broken appearance of

the chromatin in some of the interstitial cells, which has been

noted as occurring in the regenerating piece. This latter con-

dition, found in a hydra upon which no operation had been

performed, made it still less possible to consider such nuclei

as dividing. On the other hand, a piece that had been regen-

erating seven days, and that had attained to the proportions

of a normal polyp, was found to present the same appearance

under the microscope. It is possible, therefore, that the normal

tissue examined was that of a growing hydra, and that these

conditions are common to the regenerating and growing forms,

while further examination of the tissue of fully grown hydras

might show a somewhat different state.

While these latter experiments, therefore, threw little light

on the amount of division in the regenerating piece, yet the

similarity in appearance between this regenerating tissue, the

582 THE AMERICAN NATURALIST. [Vor. XXXVI.

tissue of the regenerating piece after the normal form had been

attained, and the tissue of normal or growing hydra is of

importance in a consideration of regeneration in this form.

I shall now attempt to make a conservative statement of the

amount of dividing tissue in regenerating hydra, based on

undoubted cell division.

There is very little evidence of the formation of new tissue

during the early stages; an exception was noted in one piece

killed five hours after the cutting, in which a number of divi-

sions were observed in the ectoderm. Some few divisions can

be observed before twenty hours, but from forty hours on, —

the time of tentacle formation, — the division is more active

and the number of dividing cells, especially neuro-muscular

cells, is considerable. Division continues after the tentacles

have begun. Spindles were seen in endodermal cells, and in

neuro-muscular and interstitial cells, These divisions often

occur in groups, especially in the neuro-muscular cells. In

one case as many as five dividing nuclei of neuro-muscular

cells were observed at one spot in a section cut from the side

of the piece and so affording a surface view of ectoderm tissue

alone. Fewer divisions were seen in the endoderm than in the

ectoderm, partly, perhaps, because of the smaller number of

cells in the endoderm.

Unlike the majority of cases of regeneration, however, the

new cells are not formed at the cut surface alone, and the ten-

tacles do not seem to be regenerated solely from new tissue.

Divisions were found to occur as well at the sides as at the

ends of the regenerating pieces, and in almost, if not quite, as

great number. The first appearance of the tentacle, which in

the living animal seems to be brought about by an outpushing

of the tissue of the original piece, appeared under the micro-

Scope to be due very often to contraction in this region,

causing a very slight protuberance of ectoderm, which could

be fully accounted for by an outpushing of old material with-

out the formation of new tissue. Divisions were sometimes

seen at this place, sometimes not. Very frequently, however,

in later stages, when the tentacle had grown to some length,

there was seen to be a group of neuro-muscular cells dividing

No. 427. CHANGES IN HYDRA VIRIDIS. 8

593

at the base of the tentacle, and interstitial cells were seen to

be dividing quite rapidly in the tentacle itself. It would seem,

on the whole, that new tissue is furnished to the tentacle by

division of cells at the proximal end, or base, rather than by

division of cells that have already entered into the new organ.

With the exception of the formation of nettle cells from

interstitial cells, it is not found that cells of one kind give rise

to cells of another, but that neuro-muscular cells of the new

hydra result from neuro-muscular cells of the piece, and inter-

stitial from interstitial.

Considering, therefore, that the first appearance of the ten-

tacle may be unaccompanied by cell division in its immediate

neighborhood ; that cell division, though frequently found in

the region of the growing tentacle, is not confined to this

region but is found elsewhere throughout the length of the

regenerating piece ; that division is found to be present as late

as seven days after the cutting, that is, after the piece has

assumed the proportions of a normal hydra; and that some

division is observed in the tissue of the normal, probably

rapidly growing hydra, — we are justified, I think, in conclud-

ing that the new cells which appear during the regeneration of

hydra are formed by division of the old cells throughout the

entire piece, as in the normally growing animal, and that the

tentacles are formed from old cells and from cells that have

arisen by division of the already differentiated cells of the old

part.

I am very much indebted to Prof. T. H. Morgan, under

whose direction this work was carried on.

Bryn Mawr, January, 1902.

NOTES AND LITERATURE.

ZOOLOGY.

The Fishes of Guam. — In the director’s report of the Bernice

Pauahi Bishop Museum at Honolulu (for 1900), Mr. Alvin Seale,

curator of fishes, gives a record of the birds and fishes collected by

him on the island of Guam. Two hundred and seventy-four species

of fishes were obtained, of which eighteen are described as new,

useful descriptions being given of all the others. The fauna differs

considerably from that of Hawaii, the number of species common

to the two groups of islands being less than one-third of the number

enumerated. The fauna of Guam naturally approaches more nearly

to that of the East Indies. In commenting on this most valuable

and helpful piece of work, the first accurate faunal list of fishes of

Polynesia yet published, two minor criticisms are necessary.

The proof reading should receive more careful attention; Zebra-

soma appears three times as Zabrasoma, and Garres argyeus stands

for Gerres argyreus. Still more important is the form given to new

names. Pseudoscarus platodoni apparently refers to the broad teeth

(platyodon or platyodus). eres cephalopunctatus is unfortunately

named and is, of course, not an agonoid, but a species of Parapercis,

a genus of trachinoid affinities. Gobius deltoides belongs to some

genus other than Gobius. These and other corrections will doubtless

be made in due time by the author, and the work will prove most

useful to future students of the Polynesian fish fauna. Di Se]

Notes on Fishes. — Henry W. Fowler notes in the Proceedings of

the Academy at Philadelphia (Vol. LIII, Pt. IIT) the occurrence of

Myctophum phengodes, collected off the west coast of Greenland by

Dr. Hayes.

In the Zoologischer Anzeiger (Bd. XXV, No. 666) Dr. Poche calls

attention to the identity of the three nominal genera of siluroid

fishes, called Amphilius, Anoplopterus, and Chimarrhoglanis, the .

name Amphilius being the earliest.

In the records of the Australian Museum, Edgar L. Waite records

numerous fishes from western Australia, with valuable notes and

586 THE AMERICAN NATURALIST. | [Vor. XXXVI.

several plates. In another article he gives useful notes on an

Australian shark, Galeus, or Mustelus antarcticus, and its peculiar

placenta-like structure attached to its young.

In the Zoologischer Anzeiger for Nov. 25, 1901, Dr. K. Kishinouye,

head of the Fisheries Bureau of Japan, describes a new shark, Rhino-

don pentalineatus, taken off Cape Inubo in Japan. This huge animal

is over thirty feet in length, and had an oak stick a foot long in its

stomach. The stuffed skin is preserved in Tokyo.

A skin of the whale shark called Rhinodon typicus has just been

received at the United States National Museum from Ormond, Fla.

The skin is eighteen feet long. The species is known thus far from

the type from the Cape of Good Hope and from teeth taken at the

Seychelles Islands. Mr. B. A. Bean notes this discovery in Science.

In the Bullétin de la Société Philomathigue (N.S., Tome III,

Nos. 3, 4) Dr. Pellegrin discusses those fishes which develop with age

an adipose pad on the forehead. This is found in numerous wrasse-

fishes, parrot fishes, snappers, and others. These appendages are

chiefly confined to adult male fishes and are made of adipose tissue.

Pellegrin compares it to the deposit of fat in old age in some human

individuals.

In the Overland Monthly for F ebruary and March Mr. Cloudsley

Rutter of the United States Fish Commission gives the story of the

Sacramento salmon in very unique fashion. A full account is given

of each detail in the life history of the fish, together with photo-

graphs of scenes and places on the salmon’s route, and a very large

number of illustrative drawings. Among the fantastic stories of

animals now in vogue, many of them having no existence in real

nature as distinguished from the forests of Kiplingia, it is refresh-

ing to find a fish story, at once natural and true, the result of years

of patient observation.

In the Proceedings of the United States National Museum (Vol.

XXIV, Nos. 1260, 1261, and 1263) Jordan and Snyder continue their

monographic reviews of Japanese fishes, treating the various forms

combined by Günther under the head of Trachinide. They follow

Boulenger in separating from this group all the species with thoracic

ventrals as being percoid rather than trachinoid in their affinities.

Twenty-six species are enumerated, seven being new. Three new

genera, Pteropsaron, Ariscopus, and Stalix, are described and figured.

Of the Discoboli, four species are enumerated, Lethotremus awe

and Crystallias matsushime being new, the latter the type of a new

No. 427.] NOTES AND LITERATURE. 587

genus near Liparis. The Embiotocidze, or surf fishes, consist of

two species in Japan, Ditrema temminckii and Neoditrema ransonneti.

The Pediculati, or angler fishes, of Japan consist of eleven species,

the following being new: Antennarius nox, A. scriptissimus, A. sangui-

Juus, Malthopsis tiarella. The last two papers are by Dr. Jordan,

assisted by a Japanese student of fishes, Michitaro Sindo.

Mr. William H. Gregg has published a convenient volume on

When, Where, and How to catch Fish on the East Coast of Florida.

The greater part of the book is devoted to a systematic account

of the game fishes of the east coast of Florida, with numerous

figures taken from government publications. The systematic part

is arranged from the writings of Jordan and Evermann, which is

very proper when credit is given, as Mr. Gregg has been careful to

do. To make popular works of this sort possible is one of the duties

of the systematist. After describing the fishes Mr. Gregg tells in an

accurate and readable fashion how and when to catch them.

In the Annals and Magazine of Natural History (Vol. IX, No. 51),

Dr. G. A. Boulenger discusses the classification of the berycoid fishes.

He finds that this group differs from the percoids in no important

respect, and regards the berycoids as merely archaic percoids. The

chief diagnostic character, the increased number of ventral rays, is

found in most berycoids. Pempheris, however, differs from Beryx

almost solely in having the usual number of ventral rays, I, 5, found

in most spiny-rayed fishes. Zeus and Grammicolepis, perhaps not

related to Beryx, have also an increased number. Aphredoderus is

placed among the berycoids by Boulenger. The resemblance of the

fossil genus Asineops to the Berycidz is also noted, while Erisma-

topterus is thought to have no near affinity. To the present writer

Erismatopterus resembles the Percopsidze, having much in common

with Columbia. Boulenger rightly separates Pempheris from the

Kurtidz, and he suggests the close relationship of both Pempheris and

Bathyclupea to Beryx. Stephanoberyx and Malacosarcus he would

place among the Haplomi. Polymixia he regards as an ally of Beryx,

and Morocentris as a more distant relative. Hoplostethus he unites

with Trachichthys, while a new genus, Gephyroberyx, is proposed

for Trachichthys darwini. The genus Paratrachichthys with the vent

advanced is rightly regarded as valid. D. S.J.

Twenty New Pocket Mice. — Pocket mice are mice with pockets,

not mice intended to be kept in the pocket, as the reader may inno-

cently suppose. Dr. C. H. Merriam (Proc. Biol. Soc. Washington,

588 THE AMERICAN NATURALIST. |. [Vor. XXXVI.

March 5, 1902) has given us a most important contribution to the

knowledge of these animals, principally based on the material (some

800 specimens) collected by Messrs. Nelson and Goldman in Mexico.

The genus Heteromys takes the place of our common Perognathus

in the warmer parts of Mexico and in Central America, but comes

north (as Dr. Merriam’s paper shows) into the states of Chihuahua

and Sonora, and to Brownsville, Texas. However, these more

northern animals, along with a number of others, represent a type so

far departing from typical Heteromys that Dr. Merriam segregates

them under a new generic name, Liomys. The type of the new

genus is Ziomys alleni, —the Heteromys alleni of Coues, 1881.

Taking Heteromys and Liomys together, and considering only the

fauna of America north of Panama, no species were known previous

to 1868, when Gray described four. In 1874 Peters described one,

in 1881 Coues one, in 1893 Thomas published four, and two were

made known by Allen and Chapman in 1897. Thus, in all, twelve

were known; and to these Dr. Merriam now adds twenty!

T. DAs C

Osteology of the Flamingoes. — Dr. R. W. Shufeldt ! describes the

skeleton of the flamingo (Phenicopterus ruber) with special reference

to the relations of the flamingoes to the Anseres (ducks, geese, swans)

on the one hand, and to the Herodiones (ibises, herons, storks) on

the other. The author compares minutely the flamingo skeleton,

bone by bone, with the skeletons of representatives of the other

groups. On the whole, the flamingo skeleton presents a mixture

of anserine and ibidine characters, together with certain characters

which are distinctly peculiar to itself. For the most part there is no

marked predominance of either anserine or ibidine features. In

furcula, coracoid, and wing skeleton, the anserine characters are in

excess. The tarso-metatarsus resembles most closely that of an ibis.

e author concludes that, so far as the skeleton is concerned,

the flamingoes should constitute an independent group or suborder

(Odontoglossz, corresponding to Huxley’s Amphimorphz) standing

between the anserine and pelargo-ibidine forms. Hw*Ek

Regeneration in H: ym viridis. — Hydra viridis has been made

the TOT. of a series of regenerating and grafting experiments by

1 Shufeldt, R . W. Osteology of the none Annals of the Carnegie

Museum, vol. i (gos). pp. 295-324, Pls. IX-XIV

No. 427.] NOTES AND LITERATURE. 589

Dr. Helen Dean King.’ It was found that the removal of the oral

end by a cut just below the tentacles was followed by the regen-

eration of fewer tentacles than were possessed originally, while

the diameter of the regenerated hypostome was less than that of

the original hypostome. As this operation reduces the volume of the

body, the result appears to agree with the view advanced by Parke?

(p. 702), “that a certain ratio exists between the size of a Hydra

and its number of tentacles, and that when this ratio is destroyed

by an increase or decrease in size of the Hydra, there will be an

increase or decrease in the number of tentacles of that Hydra.”

When the tentacles were removed by cutting at the base of each

one so as not to diminish the volume of the trunk, in most cases

as many tentacles were regenerated as had been removed.

The severed * heads" remodeled themselves into small polyps,

and, although the hypostomes suffered reduction in diameter, in no

case was a reduction in the number of tentacles observed, in spite

of the smallness of the polyps. This, the author maintains, does

not support Parke’s view. (It should be noted, however, that

Parke’s statement was made with reference only to change of size

resulting from favorable or unfavorable conditions, — not to decrease

in volume by the mechanical removal of part of the body.)

Double-headed forms were produced by splitting the oral end

longitudinally. When the tentacles were first removed, the total

number of tentacles ultimately borne by the two heads together

was an average of 3.4 tentacles per hydra greater than the number

originally borne. When the tentacles were not removed previous to

the splitting of the oral end, the average number of new tentacles

developed by the two heads together was 5.1 per hydra. These

double-headed polyps resolve themselves into two polyps by what

resembles a process of longitudinal division, the final separation

occurring at the extreme aboral end. Some of these double-headed

forms were made to attach themselves oral end downwards. The

Separation of the two parts occurred at the aboral (upper) end as

before, proving that the longitudinal fission is not due simply to the

constant strain exerted by gravity at the point of divergence of the

two branches of the trunk.

1 King, Helen Dean. Observ d Experiments on Regeneration in

Hydra viridis, Arch. pus TAREAS HR der Organismen, Bd. xiii, Hefte

I and 2 (1901), pp. 135-178. 31 text-figs.

? Parke, H. H. Variation and Regulation of Abnormalities in Hydra, Arch.

Sir EEE E der Organismen, Bd. x, Heft 4 (1900), pp. 692-710.

9 text-figs.

590 THE AMERICAN NATURALIST. [VoL. XXXVI,

If the cut edges of a split oral end are permitted to reunite, new

tentacles develop at the regions of union. Polyps with fourteen

tentacles were thus produced, but the repetition of the operation on a

fourteen-tentacled hydra resulted in no further increase of tentacles.

Attempts were made to secure heteromorphosis, or reversal of

* polarity," by grafting. In several cases tentacles were developed

upon an aboral cut surface or a foot upon an oral cut surface, but

in all these cases (which the author interprets as heteromorphosis)

the pieces whose polarity appeared to be reversed were very small

parts of the trunk. In the case of a graft of any considerable length,

the free cut end reproduced parts similar to those which had been

cut away from it. In the cases of apparent heteromorphosis exhib-

ited by very small fragments of the trunk, can it be proved that

there is not a shifting about of the tissues or a migration of cells, so

that the regeneration does not really involve a reversal of polarity ?

H. Wa R

A Revised Classification of the Enteropneusta. — It is nine years

since Spengel's great monograph of this group was published.

A number of important additions to our knowledge have been

made in the meantime, and the author now returns to the subject!

for the purpose of recasting the systematic arrangement of the

species, and rectifying certain violations of nomenclature which the

monograph contained.

The total number of species has been increased by thirteen, and

information about one of the old species, viz., Ptychodera flava

Eschscholtz, has been largely extended since the publication of the

monograph.

A total of twenty-nine species is now recognized by the author.

The arrangement of these into families and genera is as follows:

FAMILY I. HARRIMANIIDE SPENGEL, I9oI.

Genus 1. Harrimania Ritter, 1900.

“2. Dolichoglossus Spengel, 1893.

"o 3. Stereobalanus Spengel, 1901.

FawiLY II. GLANDICIPITIDÆ SPENGEL, IQOI.

‘Genus 1. Glandiceps Spengel, 1901.

"s 2. Spengelia Willey, 1898.

“3. Schizocardium Spengel, 1891:

oe 1 Die Benennung der Enteropneusten-Gattungen, Zoo/. Jahrbuch, Abth. fiir

Systematik, Geographie, und Biologie der Thiere, Bd. xv, Heft 2, 1gol.

No. 427.] NOTES AND LITERATURE. 591

FAMILY III. PTYCHODERID& SPENGEL, 1893.

Genus I. Glossobalanus Spengel, 1901.

* 2. Balanoglossus Delle Chiaje, 1929.

* 3. Ptychodera Eschscholtz, 1825.

The classification adopted by the author in his monograph was

unfortunate in that it conflicted at several points with established

rules of nomenclature. For example, the type species of Ptychodera,

P. flava Eschscholtz, 1825, would have been removed by Spengel to

new genus, Tauroglossus. Delle Chiaje’s well-known species,

Balanoglossus clavigerus, was likewise deprived of its original generic

name and placed in the suggested genus Tauroglossus.

Both these pioneer species are now restored to their rightful

places as types of the genera to which they were originally assigned.

Thislatest scheme of classification of the Enteropneusta undoubtedly

corresponds nearer to the actual relationship of the species than any

other that has been proposed.

As the author remarks, however, the three genera constituting the

Harrimaniidz are certainly considerably less closely related than are

those constituting the other two families.

Dolichoglossus in particular, I would remark, is: much less close of

kin to the other genera of the family than these genera are to each

other. Wm. E. RITTER.

Hymenoptera Parasitica Hawaiiensis. — The part of the Fauna

flawaiiensis (Vol. I, Part III) dealing with the parasitic Hymenoptera

has just come to hand. It is written by Mr. W. H. Ashmead, who

has prefaced to the purely descriptive part some very interesting

remarks on the Hymenoptera of the Hawaiian group. Of the 128

parasitica enumerated, no less than 87 are described as new; these

do not include any which cannot be referred to known families, but

eleven genera appear to be endemic. Several of the previously

known species are easily recognized as introductions from other parts

of the world. Mr. Ashmead suggests that the list as given must

quite inadequately represent the actual fauna, *since many of the

common parasitic families, which must surely occur, are entirely

unrepresented." Since writing these words, Mr. Ashmead has him-

self visited the islands, and no doubt he will shortly publish the

results of his own collecting; but it may be pointed out that the

cond'tion observed is characteristic of oceanic islands in general,

and of the Hawaiian Islands in particular, as may be seen by refer-

ence to Wallace's Zs/and Life, second edition, Chapter XV. Hence,

592 THE AMERICAN NATURALIST. [VoL. XXXVI.

if the missing families turn up, we may subject their representatives

to close scrutiny in respect to their actual origin upon the islands.

The Coccidz of the Hawaiian Islands are quite numerous, and some

have been first described from Hawaii ; but evidence has now accumu-

lated which renders it extremely probable that all the species reported

have been introduced by man, and I am very much inclined to doubt

the existence of a single endemic Hawaiian Coccid. Probably the

same may be said of the Hawaiian ants, which are all of well-known

continental genera. The bees of the islands (excluding the honey-

bee) belong to only three genera, and it is interesting to note that

these are all borers in the trunks or stems of plants. The bees

which burrow in the ground are wholly absent. T5 AC

The Slugs of Borneo. — The naked land-mollusca-of Borneo have

been made the subject of an interesting paper by Mr. W. E. Collinge

(Trans. Royal Soc. Edinburgh, Vol. XL, Part IHE No is)b <The

species known from the island, twenty-seven in all, are enumerated,

two genera (Wiegmannia and Isselentia) and nine species being

described as new. The type species of Wiegmannia (a genus of

four species) is not stated; we may designate as such W. gigas,

Collinge, which is the largest species ; for although W. dubia (Wgm.)

' was the first described, it was not examined by the author of the

genes. E Y

BOTANY.

Meier’s ** Herbarium and Plant Description °! is a portfolio con-

taining twenty-five sheets folded to 11 X812 inches, one inside page

being blank for the attachment of a dried specimen, and the other

ruled and spaced for a description of the various organs of the plant,

drawings, and other notes. On the front cover inside are concise

directions for collecting, pressing, and mounting, and at the back is

a ruled page for indexing the collection. The whole forms a simple

arrangement meeting the usual requirements for pupil's herbariums

and for such meager descriptions as are too often deemed sufficient

in school work. It has the advantage, however, over many similar

Schemes for recording plant analyses, that ample space is afforded

by the outside pages of each folder for additional notes. E. L.S.

. ! Meier, W. H. D., Superintendent of Schools, Griggsville, Ill. Herbarium

_ and Plant Description. Boston, Ginn & Company.

No. 427.] NOTES AND LITERATURE. 593

Leavitt's ** Outlines of Botany ’’! follows substantially the gen-

eral sequence of topics in Gray's Lessons, and retains much of what

must always be highly valuable in that classic text-book. At the

same time so many modifications and additions have been required

to meet the needs of high schools to-day that a really new book is

the result, and one that is fresh and modern in treatment, broad

in scope, yet wisely restricted to parts of the subject appropriate for

beginners.

The most radical departure is in the laboratory studies, which in

important ways improve upon previous attempts to solve the same

pedagogical problems. Instead of being frequent interruptions to

the text they are kept in sections by themselves, each preceding the

section of text to which it relates. They consist of explicit direc-

tions and skillfully worded questions leading the student to intelli-

gent observation of readily obtainable material and to instructive

experimentation. In addition to these aids to laboratory work

many helpful suggestions to teachers are given in an appendix,

while a number of good references for supplementary reading are

included for the benefit of both teacher and pupil.

In the text ecological considerations play an effective though

subordinate part in accounting for peculiarities in the form and

behavior of organs. Primarily, however, the study of the parts of

plants is firmly grounded upon their morphology. A considerable

number of new figures, all of high excellence, supplement or replace

those of the Zessons.

The work is sure to be heartily welcomed by a large number of

teachers, whether among those who are struggling to make the best

of meager equipment and much restricted time, or among those who

are more fortunate in their opportunities. Those who use the book

cannot fail to be impressed by the fine scientific spirit which ani-

mates every page. F. L. SARGENT.

Garden Beans.’ — Few attempts have been made to monograph

the garden beans. The work of von Martens, Die Gartenbohnen,

in 1860 seems to have been the only previous effort in this direction.

1 Leavitt, Robert Greenleaf, A.M., of the Ames Botanical Laboratory. Out-

lines of Botany, for the High School Laboratory and Classroom (based on Gray’s

pm im Botany). Prepared at the request of the Botanical Department of

arvard University. New York, American Book Company. 12mo, 372 pP-

^ figs.

*Irish, H.C. Garden Beans cultivated as Esculents, Report Missouri Botani-

cal Garden, vol. xii, pp. 81-165, Pls. XXX VIII-XLVII.

594 THE AMERICAN NATURALIST. [Vor. XXXVI.

Since that time the varieties of garden beans have greatly increased

in Europe and the United States. All of the leading varieties of

America and Europe were cultivated at the Missouri Botanical

Garden, so that the growth and seed characters were used in the

descriptions. The garden beans are grouped under the following

genera: Phaseolus, Dolichos, Vigna, Glycine, and Vicia. The author

gives a short account of the origin, uses, methods of culture, with

a brief account of Bruchus obsoletus and Colletotrichum lagenarium,

Uromyces phaseoli, and Phytopthora phaseoli. Three species of the

genus Phaseolus are described, P. /unatus, P. vulgaris, and P. multi-

florus. The greatest number of varieties are listed under P. vulgaris.

Dolichos is represented by D. /ab/ab and D. sesquipedalis; Vigna

is represented by V. catjang; Glycine by G. hispida, of which five

garden varieties are listed, but this does not by any means comprise

all of the varieties, as they are numerous in Japan and China, where

the species has long been cultivated. The genus Vicia is repre-

sented by a single species, V. fada.

An excellent feature of the paper is the full citation of the liter-

ature of the genus as well as that of the species, no pains having

been spared to verify references. The half-tone plates accompanying

the paper greatly aid in the botanical study of the garden beans.

The excellent keys for garden varieties also help to facilitate the

determination of the garden forms. This is one of the most impor-

tant contributions to horticultural literature in this country. In point

of thoroughness it is like his paper on Capsicum published a few

years ago in one of the earlier reports of the Garden. It is a model

of excellence in every way, and it may well serve as a guide for much

of the erratic work carried on in this country in listing varieties and

describing the same.

The work carried out by the author is one that has long been

neglected in this country. Mr. Irish is fortunate in having had at

his disposal not only a large amount of material, — and such work can

only be undertaken where this is at hand, — but also a good reference

library, combined with acute judgment in discriminating between the

puzzling garden forms. L. H. PAMMEL.

. Pfeffer's Plant Physiology. — Pfeffer's! revision of his Z/fanzen-

Physiologie has been so thorough and so time-consuming that only the

: Pfeffer, W. Pflanzenphysiologie. Handbuch der Lehre vom iioii -

peers in der Pflanze. 2. Auflage. Leipzig, Engelmann, 1901.

No. 427.] NOTES AND LITERATURE. 595

first half of Vol. II has been made ready for publication. This first

part appeared last summer. The first volume, reviewed in this journal

(Vol. XXXII, pp. 450, 451, 1898), treated the subjects comprehended

under metabolism. The first half of the second volume discusses

growth and the factors that control it, development, variation, and

inheritance — in short, different kinds of work done by the plant,

dependent.upon and made possible by the processes discussed in the

earlier volume. In a book planned as this is, with the first volume

devoted to the transformation of matter and the second to the trans-

formation of energy, more or less repetition is necessary, but it is a

repetition which gives to Pfeffer's treatment of the subjects in plant

physiology the exhaustiveness which the physiologist needs. This

is no book to be put into the hands of undergraduates; it is for the

man who has studied long and is studying hard. However much one

may wish that Pfeffer's literary style were not so difficult, one cannot

help recognizing that it is full of meaning.

The arrangement of matter in Vol. II of the second edition differs

somewhat from the first edition ; consequently comparison of the two

editions as to size is difficult. One sees at once, however, that if

the second half of the volume is to treat the subjects of movements

and the production of heat, light, and electricity in anything like pro-

portional fullness, the book will be considerably larger than in the

first edition. The additions to the first part are many of them the

results of Pfeffer's own work, either investigation or teaching. This

will be equally true of the second part of the volume.

Such a work as this, presenting the status of a science as a whole,

shows where the great gaps in our knowledge are. For instance the

plant physiology of to-day consists of the facts discovered in studying

land and fresh-water plants, and of the interpretations of these facts.

Indeed, the fresh-water alge have taken only a minor place as sub-

jects of physiological inquiry, so that we have to-day a physiology

interpreted by too many in terms applicable to land plants only. T

laboratory guides carry this to the extreme, but they show how

one-sided our knowledge is. Pfeffer's book can contain only a few

references to the marine alge. I am convinced that the careful

physiological study of marine plants, though such study may reveal

no new principles, will modify and correct many of the conceptions

prevailing to-day. The status of the science is satisfactory, but

there is room for much more research. ote po

596 THE AMERICAN NATURALIST. [VoL. XXXVI.

Trees in Winter.— Every observer knows that there are other

ways of recognizing animals and plants than those given by the

books, and we all welcome every addition to the literature of these

occult ways of coming into touch with nature. That trees may, in

general, be recognized as certainly in winter as at other seasons has

long been known, and little handbooks codifying their winter char-

acters have appeared in most European countries, and similar keys,

etc., have appeared several times in our own country, but unfor-

tunately usually in transient pamphlet form.

There is now published a neat and accurate little book! dealing

with the more obviously marked deciduous trees of northeastern

America, the characteristic traits of which are shown by habit half-

tones, and the twig details by three-color plates. Professor Sargent

stands as godfather to the book, which will be an ornament to any

center table, and should do much to give incentive to those walks in

winter that the few enjoy so keenly and the many forego because

they lack a direct object. T.

Notes. — Vol. V of the Annuaire du Conservatoire et du Jardin

Botanique de Genève contains M. Briquet’s administrative report; a

paper by him on the flora of the mountains of Corsica; an enumera-

tion by Hochreutiner of Malvacez collected by Chevalier in central

Africa ; a paper by Wettstein on Gentiana and Euphrasia ; a revision

of Urena by Hochreutiner; a paper by Briquet on Alpine Hieracia ;

notes on Malope and Palaua by Hochreutiner; a description of Poa

balfourii, from the Alps, by Briquet; studies of some American

Nyctaginacez by Heimerl; and an exchange seed list.

The Bulletin of the Torrey Botanical Club for January contains the

following articles: Salmon, “ Supplementary notes on the Erysipha-

cee”; Anderson, “ Dasyscypha resinaria causing Canker Growth on

Abies balsamea in Minnesota” (2 pls.); Anderson, “ Tilletia horrida

on Rice Plant in South Carolina”; and Rennert, “ Seeds and Seed-

lings of Arisema triphyllum and A. dracontium” (1 pl.).

A new edition of Dérfler’s Botaniker-Adressbuch has recently been

issued by the editor, and bears the imprint of Vienna.

The principal articles contained in the Botanical Gazette for

January are the following: Harper, “Binucleate Cells in Certain

Hymenomycetes” (1 pl.); Clark, “On the Toxic Properties of

, Huntington, Annie Oakes. Studies of Trees in Winter. Illustrated with colored

plates and photographs. Boston, Knight Millet, 1902. xviii 4- 198 pp

No. 427. NOTES AND LITERATURE.

597

Some Copper Compounds with Special Reference to Bordeaux Mix-

ture" (7 figs.); Clinton, * Cladochytrium alismatis " (3 pls.) ; Arthur,

“Clues to Relationship among Hetercecious Plant Hosts"; Good-

ding, “ Rocky Mountain Plant Studies, I.”

The January number of Country Life in America is a California

number. Among things of interest to the botanist are excellent

habit photograms of Washingtonia filifera, Sequoia gigantea, Quercus

lobata, Yucca arborescens, and the famous Monterey cypresses, and an

exquisite flower portrait of Romneya coulteri.

In Country Life in America for February Mrs. Comstock writes on

trees, Mr. Davy on the poppyworts, the editor on the nature-study

idea, and Mr. Mowbray on the gardens of the old Hudson River

manors. Many of the half-tone illustrations are not only exquisite

but of botanical value.

The fifth fascicle of Vol. I of Mr. Howell’s Fora of Northwest

America reaches into the genus Plantago, the Benthamian sequence

being followed. Like preceding parts, it contains descriptions of a

few new species.

A phytogeographic paper on Silesia, by Schube, is published as

a complementary Heft to Bd. LXXVIII of the Jahresbericht der

Schlesischen Gesellschaft für vaterländische Cultur.

Vol. XVI of the Acata societatis pro fauna et flora fennica is entirely

devoted to papers discussing the flora of the country.

In Vol. XIII of the Verhandlungen der Naturforschenden Gesellschaft

in Basel Binz gives an interesting comparison of the present flora

of that part of Switzerland in comparison with what Bauhin knew of

it three centuries ago.

Vol. IV of Engler and Drude’s Vegetation der Erde is by Beck von

Mannagetta, and deals with the Illyrian region. It is published by

Engelmann of Leipzig.

A historical bibliography of the flora of Rome is in course of publi-

cation by Pirotta and Chiovenda, in the Annuario of the botanical

institute of that city.

A second part of Ross’ “ Beitráge zur Flora von Sizilien” is pub-

lished in No. 12 of the Buletin de /' Herbier Boissier for 1901.

A paper on Dr. Baldacci’s Albanian collections of 1897 has

recently been separately printed from the Memoire of the Bologna

Academy.

598 THE AMERICAN NATURALIST.

A list of new plants from the Cape Peninsula is published by

Major Wolley Dod in the December number of the Journal of Botany.

An account of the vegetation of the Caroline Islands, by Volkens,

is published in Bd. XXXI, Heft 3, of the Botanische JaArbücher.

Several interesting process plates accompany the text.

Several botanical papers are contained in the recently issued

second part of Vol. XXVI of Proceedings of the Linnean Society of

New South Wales.

Vol XXXIII of the Transactions and Proceedings of the New

Zealand Institute contains the following botanical papers: Cockagne,

“Seedling Forms of New Zealand Phanerogams," “Gunnera and Myo-

sotis in Chatham Islands"; Laing, * Seaweeds of Norfolk Island" ;

Walsh, ** Cordyline terminalis in New Zealand" ; Cheeseman, « Culti-

vated Food Plants of the Polynesians, with Special Reference to

Cordyline terminalis,” “ Recent Additions to the New Zealand Flora” ;

Thomson, “Plant-Acclimatisation in New Zealand”; Rutland,

“Regrowth of the Totara”; Petrie, “Descriptions of New Native

Plants"; Brown, “Notes on the New Zealand Musci.” In the Pro

ceedings of the Affiliated Societies some minor botanical notes are

given, among them a rather curious surmise concerning Selaginella

lepidophylla.

The announcement of the Marine Biological Laboratory at Woods

Hole sets forth an attractive programme of studies and lectures for the

coming summer. Inthe department of botany courses are provided

in cryptogamic botany by Dr. Dacis, Dr. Moore, and Mr. A. c.

Moore; in ecology by Dr. Cowles and Dr. Shaw; in plant physi-

ology by Dr. R. H. True, and in cytology by Dr. Davis and Mr. Wolfe.

In addition to the work of the session, which will extend from July 2

to August 13, an expedition will be organized to Mt. Katahdin and

the coast'of Maine for the purpose of conducting ecological studies

in that region.

CORRESPONDENCE.

Editor of the American Naturalist :

SIR: — In a paper on * The Colors of Northern Polypetalous

Flowers," in the March number of the Watura/ist, p. 295, Mr. Lovell

States that Z/a/ctus nelumbonis confines its visits to the flowers of

Nuphar (Nymphea) advena. In Transactions of the American Entomo-

logical Society, Vol. XVII, p. 316, 1890, it is recorded that this bee was

taken on flowers of Nuphar, Nymphza, and Nelumbo in Illinois, on

Nuphar and Nymphza in Florida, and on N uphar in Wisconsin. It

has been mentioned as a general visitor of Nymphzeacez in Ameri-

can Naturalist, Vol. XXIX, p. 107; Botanical Gazette, Vol. XXVI,

PP: 29, 36; ZIustrirte Zeitschrift für Entomologie, Bd. V, p. 310; Bota-

nisches Centralblatt, Bd. LXXXV, pp. 299, 302; Botanischer Jahres-

bericht, Bd. XXVII, Abth. 2, p. 463. No doubt the female gets

pollen exclusively from flowers of N ymphzacez ; but I have found

her visiting for nectar the flowers of Eriocaulon gnaphalodes, Utri-

cularia inflata, Berlandiera subacaulis, and Verbena urticifolia.

On page 235, in regard to the visitors of Umbelliferz, he says:

"In Germany there have been collected on the caraway 55, on the

wild carrot 61, and on the wild parsnip 118 insects. Probably the

number of visitors to many species exceeds 200.” The plant called

wild parsnip by Mr. Lovell is Heracleum spondylium. In Transactions

of the Academy of Science of St. Louis, Vol. V, p. 459, 1890; Bota-

nisches Centralblatt, Bd. XLVI, p. 110; Botanischer Jahresbericht,

Bd. XVIII, Abth.r, p. 509, visitors are credited to several species as

follows: Zizia aurea, 131; Eryngium yuccifolium, 147; Tiedemannia

rigida, 156; Heracleum lanatum, 174; Sium cicutifolium, 191 ; Cicuta

maculata, 238; Pastinaca sativa, 27 [o

In the literature the status of Müller's view regarding the influence

of dull yellow colors on the visits of beetles is about as follows:

The general proposition was refuted by Bonnier in 1879.! In the

Tyrol, Schulz ? not only found beetles on such flowers, but also dis-

tinctly stated that the general proposition was not true for that region.

» pap m Sci. r Bot., ser. vi, vol. 8, p.

? Beiträge zur Kenntniss e Boitudiuigeciunéhtungen und Geschlechtsver-

theilung bei den {Scape Aa ii à (1800), p- 64.

599

600 THE AMERICAN NATURALIST.

In the same year I discussed the subject in connection with the

Umbelliferz,! enumerating the visits of forty species of beetles to

flowers of Pastinaca, which is nearly twice as many as Müller ever

found on any umbellifer. In the Botanical Gazette, Vol. XXII,

p- 169, 1896, I have cited the statements of Bonnier and Schulz.

Finally, Knuth,? in a volume devoted to the general principles of

anthoecology, abandons Miiller’s view with the statement that Miller

himself and Loew had made observations which did not support it.

He does not mention the observations of Bonnier, Schulz, or mine;

but, of course, when a man feels at liberty to use the literature in

any way he likes, he can record old things as new, and give credit

in any way that suits his fancy. These references ought to be

enough to dispose of a proposition which was never supported by a

reasonable presumption. Chants Rostetsü

CARLINVILLE, ILL., April 4, 1902.

1 Trans. Acad. Sci. St. Louis, vol. v (1890). p. 454-

? Handbuch der Blütenbiologie, Bd. i (1898), p. 224.

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AMERICAN NATURALIST

Vor. XXXVI. August, 1902. . No. 428.

THE ANATOMY OF A DOUBLE CALF.

HENRY LESLIE OSBORN.

THE subject of this article is a Durham calf born near Minne-

apolis, Minn., in March, 1901. It lived only ten minutes; but

excepting that there was a fracture in the hip bone no indica-

tion of injury was seen, and the different organs presented such

a healthy appearance as to justify the belief that the calf could

have been reared by taking special pains to that end. I owe

the opportunity to examine the case to Mr. H. W. Howling, a

taxidermist of Minneapolis, who sent me the carcass after the

removal of the skin. I am indebted to Dr. C. Hart Merriam,

Washington, D.C., for an introduction to the literature of

teratology.

The appearance of the stuffed skin is shown in the two photo-

graphs, Figs. 1 and 2, views from behind and from the dorsal

surface. There are two perfect anterior bodies, each with its

head and thorax and the anterior part of the abdomen, and there

are four perfect anterior limbs. The left body appears in the

mounted condition (see Fig. 2) to be a complete body, into the

side of which the anterior right half has been grafted at an

angle of about ninety ione d 25 hind legs stand nearly at

602 THE AMERICAN NATURALIST. [Vor. XXXVI.

right angles with the axis of the left half, while they are nearly

in the plane of the axis of the right. The internal structure,

however, as will be seen, shows that in reality there are two

nearly complete bodies strangely united to form the animal.

Posteriorly the two bodies are united to form a single region,

standing on a right and left hind leg, each of which belongs

to one of the anterior halves of the body. There is a third hind

leg. It does not touch the ground, but arises from the center

Fic. 1, — View of stuffed skin f behind, from a photograph.

of the rump and projects backward and to the left. This leg

has the usual form and regions. Of course I have no means of

knowing its bony structure, but it shows the same external

subdivisions as the others and ends in two hoofs; the limb,

however, is shorter than the others, as shown in this table of

measurements.

Lzc. Fore Lec. Foor. Hoor-

FKhtid hg ce 1I in. 8 in. 3 in. 1j in.

Mode badeg .. ... . 6 in. 6 in. 3 in. 1j in.

No. 428.] THE ANATOMY OF A DOUBLE CALF. 603

There are two tails, of the same form and size. They are sym-

metrical in position, each located between the functional hind

leg of its own side and the middle leg. The anus is single; it

is located directly under the median hind limb and equally dis-

tant from the tails. No indication of a urogenital opening was

seen, and the structure of the bladder indicates that the penis

had not been formed.

The coloration of the skin exhibits noticeable bilateral sym-

metry ; the general ground color is brown, on which white spots,

Fr- 2. — View of dorsal surface, from a photograph.

or areas, are found closely matching each other as to position

and shape. On the top of each head is a white spot of much

the same position, size, and shape; the two tails end in white,

of the same length in each. The limbs are white below, except

the inner fore limbs, both of which are brown to the carpal

region, and the right to the toe in addition. A girdle of white

encircles the body of each; it is more solid on the left side.

There is a patch on the loin region of each. The following

measurements were made from the mounted animal:

Height of body at hip. . . 27in. | Length of right tail. . . . 13 in.

Base of tail to tip of right 35 in. | “ i =F i

“ s left m a5 in. |

604 THE AMERICAN NATURALIST. [Vor. XXXVI.

The body as it reached my hand had the skin removed, and

the heads and lower limbs. The external points noticed were

these: There was a single anus; a single median umbilical

opening was present; there were two bilaterally placed testes

located on the wall; they had passed to the surface but no

scrotum was formed; they were somewhat widely apart (four

inches?); a penis was not present. There are two complete

spinal columns, as would be inferred from the presence of two

—

Fic. 3. — Dorsal aspect of the bones of the sacral region.

tails. They are, however, both considerably modified in and

behind the lumbar region, as shown in Figs. 3 and 4, which

are dorsal and ventral views of the osteological preparation of

that region. Both columns are evidently present; they come

toward each other and then bend into parallelism, the right

becoming concave on the right side, and the left on the left

side. This is brought about by the outside of each vertebra

being considerably smaller than the inside. The transverse

processes of the lumbar vertebrz are not much more than half

as large on the outside of the vertebra as they are on the opposite

No. 428.] THE ANATOMY OF A DOUBLE CALF. 605

side. The sacral vertebræ are not only bent as described

but there is a twist in them so that the centra come to face

inwards, and instead of looking downward they look toward each

other. Each sacrum articulates on its outer side with an os

innominatum of the usual form, consisting of ilium ischium and

pubis, meeting to participate in the formation of an acetabulum

for the reception of the femur of the paired leg of its own side

of the body. The pubic bones meet ventrally and forma normal

Fic. 4. — Ventral aspect of the bones of the sacral region.

symphysis pubis. The two sacra articulate internally with a

peculiar bone, evidently composed of two incomplete ilia. This

bone presents anteriorly two crests, each with the usual relation

to the corresponding sacrum. In the center of this median

bone there is a ridge which ends bluntly in front, and poste-

riorly meets a broad, thin, and somewhat tapering bone by a

distinct suture. At the junction of these two parts is situated

a shallow but distinct articular surface; it is the acetabulum of

the median hind leg. It is very evident that the anterior bone

606 THE AMERICAN NATURALIST. [VoL. XXXVI.

is composed of parts of two ilia incompletely separated, and that

the hinder bone is in like manner made up of the parts of two

still less developed ischia, while no traces of the pubes have

appeared.

No dissections of the muscular or nervous systems were made,

but we can infer from the osteology that there were two com-

plete spinal cords. The innervation of each of the two functional

hind legs must have been from the spinal cord of its own side.

This separateness of the cord would have seriously affected

coordination. It is related that in the two-headed turtle loco-

motion was very imperfect (Barbour, '96), and in Ritta-Christina

(Hektoen and Riesman, '01) a pinch on the right leg was felt

only by Ritta and one on the left only by Christina. The indica-

tions from a study of the bones are that the musculature of the

outer legs was complete, while the muscles and nerves of the

inner side must have been partly, though very incompletely,

developed.

The viscera of the thorax were not examined. This is to be

regretted, because it would have been important to determine

whether the two hearts and vascular systems were counterparts.

It seems likely from the anatomy of the animal, so far as known,

that they were; still, I am unable to assert the fact. There

were two complete diaphragms, and the abdominal cavity was

Fic. 5. — Junction of md small intestines of the two anterior bodies, and origin of

the single posterior small intestine.

completely subdivided anteriorly. The two livers were entirely

distinct, as were also the two stomachs and the upper parts of

the intestines. The small intestines run toward each other and

meet, forming a continuous tube, as is shown in Fig. 5. There

is no indication of a union of two parts at this junction. The

vascular supplies of each part are totally distinct, the vessels of

No. 428.) THE ANATOMY OF A DOUBLE CALF. 607

each side running out to reach the common territory. The dis-

tance from this junction to the anus is seven feet and two inches,

in all of which the intestine is single. The single small intes-

tine arises obliquely out of the common passage, and not, as

might be expected, squarely from it. After running a distance

of three feet, the small intestine dilates to form the asin intes-

tine. Here two ceca

of unequal size are

located. The two

ceca are exactly op-

posite each other, and

their cavities are con-

tinuous. At their

junction they open

together into the

intestine at the begin-

ning of the large

intestine. The large

intestine runs directly

to the single anus, a

distance of four feet he. T t the junction of the

and two inches. scat airepor aea

There is normally in the calf only a single cæcum. The

presence of two, then, is due to contributions being present

from each of the component bodies of the animal. The

single intestine could be interpreted in either of two ways,

viz., that there was a single intestine into which that of the

other half had been engrafted not far from the stomachs, or

that the intestine is really parts of two fused in the middle

line and thus forming one. The oblique insertion of the single

intestine at its origin seems at first to favor the first view, but

there is considerable evidence against it. Dr. Wyman (’66), in

the case of a double human embryo which he dissected, found

that the intestines meet in the middle line and fuse to form a

single tube, which, however, soon parts, forming two which run

side by side for a time, after which they unite in a single rectum

and terminate in a single anus. The presence of the two cæca

in the case before us indicates that parts of two intestines, each

608 THE AMERICAN NATURALIST. [Vor. XXXVI.

running the entire length of the half body to which it belongs,

meet in the middle line to form one tube. This mode of origin

is in harmony with that seen in the inner iliac and ischial bones

and the median hind leg.

The vascular system shows a similar arrangement. Anteriorly

there are two complete systems, while posteriorly the systems

are complete on the outside, but not entirely so in the central

genital org 1 ls: a.4., anterior bladder ; 4.4/., posterior

bladder; æ., urachus; ur., ureter; s.e., umbilical artery; Z.2., iliac artery; Z.v., iliac

^, Vas ens,

vein ; £., testis ; v. d. defer

area. There are two dorsal aortze (Fig. 7), and the renal and

spermatic arteries are in pairs on both sides, as if the bodies

were independent. Posteriorly, however, the aortz each give

rise to one large iliac artery to go to the paired leg of that side,

and a second which was not traced beyond the root, where it

soon disappeared among the muscles. At the fork of these

' iliacs a single umbilical artery arises from each aorta; it runs

No. 428.] THE ANATOMY OF A DOUBLE CALF. 609

down alongside of the bladder and, passing out with the urachus,

runs to the placenta. A second umbilical artery for each would

be needed to complete two entire bodies; these are wholly

wanting. There are two posterior vena cava, one for each

body half, and similar renal veins were recognized. Posteriorly

—

J -@. bd.

LM wiih

ucts. wise anterior bladder has one os over

; the

Fic. 8. — The two bladders and ee

f the posterior bladder:

forward so as to show the anterior bee

other letters as in Fig. 7.

the venze cavae divide and give rise, after the manner of the

aorta, to an iliac vein for each of the two paired hind legs.

But the inner iliac veins run toward each other and meet and

fuse, forming a single large communicating iliac vein, thus

extending the list of posterior parts common to the two body

ves.

610 THE AMERICAN NATURALIST. [Vor. XXXVI.

There are four complete and independent kidneys; two, by

their position and vascular relations, are clearly indicated as

belonging to each component body. The kidneys are com-

pound, after the form general in ruminants; each has its hilus

looking toward the dorsal aorta, at which three vessels enter

its substance, viz., the renal artery, the renal vein, and the

ureter. There are two bladders; both are median in position,

and one is located in front of the other. They may be desig-

nated as the anterior and posterior bladders. They both spring

from a common neck, where they are attached indirectly, in

common with the end of the rectum, to the body wall. At

this place they partly surround the end of the rectum. They

are in open communication at the neck by a passage running

on the right side of the rectum; but in this region, where

the urethra should appear, none is present, and there is no

communication here with the exterior. No indications of

a penis were found. The posterior bladder had an open

urachus whereby it could discharge, but the anterior lacked

this entirely, and drained through the passage at the neck into

the posterior bladder. The relations of these two bladders is

such that the posterior bladder receives the ureters from the

two outer kidneys, while the anterior bladder receives those of

the inner pair. The ureters of the posterior bladder run down

and open into its anterior or dorsal surface, as shown in Fig. 8,

and in doing this are aberrant, it being the opposite side to

that general in mammals. The anterior pair of ureters are

similarly aberrant. Two views are possible with reference

to these bladders : either they may be considered as belonging

one to each of the two component bodies, in which case we

should be obliged to consider that they had, unlike most of the

organs of the compound, taken a non-homologous and unsym-

metrical position ; or else they may be regarded as a single

formation common to the two bodies, comparable with the

hinder part of the intestine, which has been secondarily sub-

divided, The latter view seems perhaps more in harmony

with the rest of the construction of the body, but decisive

evidence cannot be had from the materials furnished in the

dissection.

No. 428.] THE ANATOMY OF A DOUBLE CALF. 6r1

The genital organs show the same relations as the renal `

system. There are four testes: two are related to the two kid-

neys of one body and the other two to those of the other.

These four testes are, however, paired in the calf not with

reference to their individual bodies, but with reference to the

compound. Two, as already mentioned, have descended: they

are the right of the right body and the left of the left; we

may call them the two outer testes. The two inner testes also

carry out the symmetry, both remaining in their primitive

position beside the kidney. These testes all reach the exterior

by means of ducts which have the same relation as do those

of the corresponding kidney. Those of the outer testes cross

the ureters from the outer kidneys and enter a glandular mass

(seminal vesicles?) in the wall of the posterior bladder. The

two inner vasa deferentia run down and attach to the anterior

bladder, passing as normally to the posterior surface.

The entire urogenital system thus follows the same law of

symmetry of the homologous parts as we have seen in the

other systems of the animal. This is the more interesting in

this case, for in regulating the relations of the ducts from the

organs the law comes into conflict with the symmetries of the

two individuals and replaces it. This is evidence of the pres-

ence of a deeper law of regulation, and indicates the “ fission

theory " (Hektoen and Riesman, '01) of origin for the monster

rather than the “ union theory."

The limits of this article do not permit an extensive notice

of the literature of teratology ; a list of writers consulted is

given at the close. Dr. Fisher (66) gives a figure of a calf

much like this one externally, — a case of *'diplocephalus

tetrabrachius tripus," — but the supplementary hind foot is

ventral and anterior, indicating a difference in the mode of

union of the pelvic bones. No account of the anatomy is

given. He mentions that it was dissected by Dr. Wyman,

but I have not been able to get any information of that dissec-

tion or its results. The case belongs in the line of double

formations, which begins at the minimum with doubling of

digits, runs through every stage of double formation of hands,

arms, and other parts, of whole regions (as the head and trunk),

612 THE AMERICAN NATURALIST. [Vor. XXXVI.

and finally culminates in twins: either incompletely separated,

as the Siamese twins, or completely separated as in ordinary

cases. Authorities in the main assign all such cases to a

“division of the ovum imperfectly effected,” so that “the

resulting bodies instead of becoming complete twins remain

united together, frequently having a greater or less extent of

the body in common”; thus Bateson (94): ‘ Double forma-

tions arise from a fission of the whole or a part of the original

embryonic mass." There are two theories as to the process:

“one holds that they arise by splitting of the original germinal

area of one embryo, and the other that they arise from the fusion

more or less of two distinct embryonic areas lying in one ovum ”

(Hektoen and Riesman, '01, p. 419 e¢ seg.). The present case

indicates a partial fission of the germinal material, that destined

to give rise to the anterior parts being wholly divided, while

some of that destined for some of the hinder organs remains

undivided. This case comes between the Tocci brothers and

the Siamese twins. The former of these I have seen, and

Fig. 9 is from a photograph of them. In these the hinder

part of the body. is apparently strictly single; there is less

division than in the calf specimen. The Siamese twins were

essentially double throughout. In the calf the hind part of

the body is incompletely double. There are cases very nearly

like it, of which a brief mention will be interesting. The

Hungarian sisters, Helen and Judith, had one vertebral column

as far forward as the second sacral vertebra; there was a

single anus and rectum; both felt the same desire to defecate;

there was a single vulva but separate urogenital systems; they

urinated separately by different urethra; menstruation came

at different periods in each (Hektoen and Riesman, p. 430).

In the case of Ritta-Christina the following items are noted

from the account in Gould and Pyle (97, p. 185): umbilical

cord and placenta single; heads, necks, arms, and thorax above

double; abdomen single; lungs imperfectly double, the cen-

tral lungs being underdeveloped ; one pericardium but two

hearts; hearts synchronous; digestive organs separate as far

as lower third of ileum, then single to the anus; stomachs,

spleen, etc., were right and left; livers right and left were

No. 428.] THE ANATOMY OF A DOUBLE CALF. 613

fused centrally ; there were two gall bladders beside each other

in the middle line; the uterus was double; the vertebral

columns were entire and separated by a mass of bone con-

sisting of fused rudimentary ossa innominata ; the sterna were

united at their manubria. Reference has been made in an

earlier part of this

paper to the case

of a human fetus

dissected by Dr.

Jeffries Wyman

(66). It was less

double than the

calf. There were

only three arms,

the median one

made up of two

halves fused; a

single pericardium

enclosing two

hearts; a single

lower vena cava

branching ante-

riorly to enter each

heart ; the umbili-

cal cord had one

vein and two arte-

ries. Theright and

left duodenums

came together

a short distance [SEE

from the two stom- Fic. 9. — The Tocci — ann — in possession

[

achs, whose pyloric

ends were turned toward each other (“symmetrical counter-

parts," Bateson, p. 559), but afterward separated and, though

Side by side, continued distinct nearly to the caecum, which,

as well as the large intestine, was single. There was a single

urinary bladder with a double cavity. There were only two

kidneys, but they were compound, and each — mo TTT

614 THE AMERICAN NATURALIST.

one set opening into the upper bladder and a second, much

larger, opening into the lower bladder. The bladders had

the usual shape, but inwardly the mucous membrane formed

a partial partition, near which the two sets of ureters opened.

There were also two pairs of testes, one of which were resting

on or quite near the kidneys and the other had descended

and were “just at the entrance of the inner abdominal ring." !

This case is much like that of the calf, — the union of the

two intestines, the four ureters, and the two bladders, one

in front of the other; it is not clear that the outer pair

opened into the hinder bladder, but that seems to be intended.

The account of the anatomy of the kidneys leaves somewhat

to be desired ; apparently the fission of the embryonic material

had not gone so far here as in the calf specimen as to the

kidneys, while it had as to the ducts. The relation of the testes

and the ducts seems to have been about the same as in the calf.

BIOLOGICAL LABORATORY, HAMLINE UNIVERSITY,

SAINT PAUL, MINN., April 2, 1902.

LITERATURE REFERRED TO.

'96 BARBOUR, E. H. A Two-Headed Turtle. Science. Vol. IV, p. 159.

'94 BATESON, W. Materials for the Study of Variation.

'00 BRADLEY. Dichotomy as a Factor in Teratogeny. Veterinary

Journal. Vol. li. |

'90 CREIGHTON, CHARLES. Article “Monster.” Zucyc. Brit.

‘66 FISHER, G. H. Diploteratology. Trans. N.Y. Med. Soc.

'97 GouLD and Pyrr. Anomalies and Curiosities of Medicine. W. B.

Saunders.

'01 HEKTOEN AND RIESMAN. American Text-Book of Pathology. W. B.

Saunders. :

'00 LINDSEY, JoHN. A Monstrous Foetus. The Veterinarian. Vol. lxxiii.

'66 WYMAN, JEFFRIES. Description of a Double Fetus. Bost. Med.

and Surg. Journ.

! Partly quoted and partly condensed from Wyman’s paper.

THE METAMORPHOSIS OF SISYRA.

MAUDE H. ANTHONY.

Sisyra, like most of the Hemerobiida, is an insect of whose

life history no thorough study has been made, and very little

is known of either its habits or metamorphosis. It is an

obscure little fly, chiefly interesting on account of some pecu- -

liar structures of its larva. The larva has been known since

Westwood described it in 1839, but it has never been reared

until last summer, at the entomological field station at Saranac

Inn, New York. It passes its larval existence as a parasite on

fresh-water sponges, clinging to their surfaces or descending

into the open osteoles, piercing the sponge tissue with its long

decurved jaws. The food thus obtained is so pure as to be

wholly absorbed, leaving no residuum. At least one-fourth of

the posterior portion of the stomach is atrophied, leaving no

opening from the stomach at that end. Carnivorous habits and

piercing mouth parts are common to other hemerobian larve,

but the decurved position of the mouth parts and the tracheal

gills are peculiar alone to this insect and Climacia, its nearest

ally, which is of similar habits. The singular adaptations of

the mouth parts and alimentary tract to the nature of its food

I shall describe in detail in this paper.

The most remarkable structure of this larva is the silk-

secreting apparatus. In general the silk glands of insects are

metamorphosed salivary glands, but in the Sisyra larva they

are a modification of the Malpighian tubules; and it seems

probable that the nitrogen waste of the body is used, partially

at least, in the manufacture of silk. This seems to be another

remarkable instance of the economy of the by-product occurring

in insect life. No such extensive modification of structure in

adaptation to a peculiar environment has been described in any

other insect ; but two near relatives of Sisyra, Osmylus and Myr-

meleon, have a somewhat similar development of silk glands.

615

616 THE AMERICAN NATURALIST. (VoL. XXXVI.

The species which I have studied for this paper is Szsyra

umbrata Needham, and all my specimens were collected for

me, a few at Lake Forest and the remainder at Saranac Inn.

I have necessarily worked under limitations, not having myself

seen any living specimens.

The adult (Fig. 1) is a smoky brown fly, 6-8 mm. long, very

hairy, and with the characteristic venation of the Hemerobiide,

although the wings are not so delicate and transparent as are

those of the lacewing flies. The maxilla and labium of the

adult, each with peculiar pediform, terminal, palpal joints, are

Fic. 1. — Sisyra umbrata, adult. x 10,

shown in Fig. 2. Other parts of the imago — antenna, leg, and

wing — are figured and discussed later, in comparison with the

same parts in the larva.

The pupa, removed from its double cocoon of finely woven

silk, is shown in Fig. 3. It is 4 mm. in length, has two

clusters of hairs on the dorsal surface of each segment, and

is similar to the larva in color and marking, except that the

light median line on the dorsal surface between two dark ones,

sO apparent in the larva, is not visible. Instead the brown

crescent-shaped markings on either side meet in a dark median

line.

Larva — EXTERNAL ANATOMY.

The larva (Fig. 4) is 6 mm. in length, is yellowish green in

color, and has quite a distinct color pattern in brown on the

dorsal surface. Each of the first seven segments bears, in two

dorsal and two lateral groups, twelve projections, except the

first abdominal, which has only four on the dorsal surface

No. 428.]

M. Anit

x 60; 4, labium.

FIG, 2.

a, maxilla,

THE METAMORPHOSIS OF SISYRA.

617

but six on the lateral. Each

of these is provided with a

long hair. Their purpose

seems to be for protection,

as the floating particles

retained in them make the

insect look very much like

a diminutive piece of aquatic

vegetation. The same thing

is described by Kent for

Hemerobius, where the pro-

tection is furnished by the

dead bodies of its victims,

the aphids which are held

by the long hairs. On the

seventh and eighth abdominal segments there is an additional

pair of hairs on shorter pedicels, near the median line.

eighth segment there are two dor-

sal and two lateral projections, more

elongate than those of the other

segments, each of which serves as

a pedicel for three hairs. The ninth

has only the lateral projections.

The tenth segment is everted at

the end to serve as a spinneret for

ejecting the silk. In sections the

chitin on this part is noticeably

thinner than that covering the rest

of the body. Between the head and

prothorax on the dorsal side is a

small interpolated sclerite, men-

tioned by Grube, and also by Hagen

in his description of Osmylus.

There is no trace of it, however,

on the ventral side. In Chauliodes,

an insect which has the mouth parts

normally directed upward rather

than downward, as is the case with

On the

Fic. 3. — Sisyra umbrata, pupa.

x 2t.

618 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Sisyra, there is an interpolated sclerite on the ventral side.

The head has on the dorsal surface a pair of hairs directed

forward, and on the ventral surface another pair similarly

directed but much more elongate.

The larval respiratory apparatus consists of external gills in

the form of a pair of jointed appendages on the ventral surface of

\ the first seven abdom-

à y inal segments, folded

| under the body so as

to be unnoticeable in

| a dorsal view. They

| / are moved intermit-

| | j tently ina rapid,

| shuttle-like vibration.

Westwood has figured

them with five joints,

but I have found them

uniformly to have only

three. The first pair

seem to have lost the

articulation between

the first and second

joints. These have

thus become two-

jointed appendages.

Near the point of

attachment the first

pair have a curious

hook-like projection

(6, Fig. 5); directed

inward toward the

body. This appears on

the second pair merely

as a knob, and gradu-

; ally decreases in prom-

inence, until it disappears almost entirely on the sixth and

seventh pair (a, Fig. 5). On each pair of appendages except

the first, at its articulation with the second joint the first one

Fic. 4. — Sisyra umbrata, larva. x 15.

No. 428.] THE METAMORPHOSIS OF SISYRA. 619

is forked, the outer branch of the fork being longer than the

inner, and both curved inward toward the body. Between these

two branches the second joint is attached (c and d, Fig. 5).

The purpose of these inward-curving forks is probably to

keep the respiratory filaments from too close contact with the

body, and give them greater access to

the currents passing through the pores

of the sponge. It would seem that the

first pair, having lost the fork at the

terminal part of the first joint, have

developed the hook-like projection

nearer the base for a similar purpose.

Two trachez can be partially traced

in the alcoholic specimens, showing

through the delicate outer membrane

of the respiratory filaments.

The mandibles and maxillae, to fit

them for piercing organs, have attained

a length nearly equal to that of the

antennae (Fig. 4) and are joined to form two sucking tubes. In

à cross section of these tubes (a, Fig. 6) the grooved edges are

shown, by means of which mandible and maxilla fit each other

closely. The grooved edges of both mandible and maxilla are

finely serrated longitudinally, prevent-

ing them from slipping one upon O a e

another when they are used in punc-

turing the sponge tissue. The two

readily pull apart, showing that there

is no adhesion of the chitin, but that

they are merely apposed. As seen in

Fig. 4, these sucking tubes are con-

joined for nearly their entire length,

being separated only at the tip and jyax; 4 tendons; », m, points of

near the base. After they enter the sachment of TEET

head they unite in a Y-shaped joint, thus forming a single tube.

This Y-shaped joint can be plainly seen on the underside of the

head in the chitinous shell obtained by the use of caustic

potash solution. After the junction the single tube passes on

filament of fifth pair.

620 THE AMERICAN NATURALIST. | [Vor. XXXVI.

a nearly vertical line toward the dorsal side, where, after making

a turn at right angles, it joins the cesophagus (Fig. 10).

In 4, Fig. 6, is shown a cross section of the sucking organ

(pharynx) after the point of juncture of the two tubes into one.

On the anterior and posterior side it is

heavily chitinized, these surfaces, together

with the tendons on the anterior side,

being the points of attachment for two

pairs of strong muscle bands. The

remaining portion of the wall of this

sucking pharynx is of thin membrane.

At the contraction of the muscles sur-

rounding it, the cavity within is increased

many times the size it has when the mus-

cles are relaxed. This mechanical con-

trivance forms a very effective pump for

drawing the juices of the sponge tissue

Fic. 7.—Antemz. a,adult, through the sucking tubes into the

x 18; 4, larval, x 36.

cesophagus.

The larval antenna has in most cases sixteen joints, those

beyond the third being usually of equal length. The third is

equal in thickness to the seg-

ments just mentioned, but equals

in length the next three or four

segments. The second is equal

to the fourth in length, but is

twice as thick. The first equals

the second in thickness, but is

only half as long. The eleventh

is frequently somewhat shorter

and the twelfth, somewhat longer

than the fourth, but they are both

equal to it in thickness. The

thirteenth is curiously ovate, and

the three terminal ones are very slender and bear one or two

small spines. At the base of the ovate joint is a much longer

spine. In Fig. 7 the antenna of larva and adult are shown

comparatively, but under different magnification.

Fie. 8.— Legs. 4, larval, x 463

b, adult, x 22; c, enlarged inner

view of foot.

No. 428.] THE METAMORPHOSIS OF SISYRA. 621

Fig. 8 gives a comparative view of the larval and adult leg,

and a view of the lower side of the foot of the adult, showing

the arrangement of claws and pulvilli, and the four large spines

on the inner side of the first four joints of the tarsus. Similar

spines occur on the lower end of the tibia. The larval leg has

but one claw and a one-jointed tarsus. The legs of the larva

are all similar. Those of the adult differ considerably; the

fore legs are thé shortest pair but have the coxa of unusual

length, nearly equal to the tibia; in the middle leg the femur

is the longest joint; in the hind leg the femur and tibia are of

equal length. In all the adult legs the trochanter is much

narrowed at its distal end where it

articulates with the coxa.

Before the pupal stage is reached,

the legs pull entirely out of their

chitinous covering, and the process

of making over begins. In speci-

mens lightly stained with borax car-

mine the adult legs can be distinctly

seen developing under the old chitinous covering. This process

is shown in Fig. 9, the coxa, trochanter, and femur of the adult

leg being fully developed, while the tibia and tarsus with its

five joints are only partially differentiated. The process of

development for the antennz and mouth parts of the adult is

very similar to that of the legs, except that these adult organs,

being so much larger than those of the larva, are early with-

drawn entirely out of their slender chitinous sheaths, and are

developed underneath the larval cuticle.

Fic. 9. — Leg transforming.

INTERNAL ANATOMY.

Fig. 10 is a diagram of a sagittal section of the larva through

the median plane. The position of the brain in the prothorax

has been noted in some other insects. In the very young larva

the brain occupies the normal position in the head, but, as the

pupal stage is approached, it gradually draws backward until it

occupies the position in the thorax shown in Fig. 10. The

prothorax also contains, besides its own, the subcesophageal

622 THE AMERICAN NATURALIST. [Vor. XXXVI.

ganglion. The seventh abdominal segment has two ganglia,

smaller than those of the prothorax but larger than the remain- ,

ing eight, which are distributed — one in each of the inter-

vening segments. The dorsal subsegment, interpolated between

the prothorax and head, and which was previously mentioned,

is shown here. Neither pair of the muscular bands which

expand and contract the pharynx (m, Fig. 10), nor either of

the sucking tubes below their point of juncture (5, Fig. 10),

would be present in a section through the median plane, but I

have figured them here diagrammatically for the sake of clearness.

The most difficult, and, as well, the most interesting part of

this study has been the tracing of the course of the alimentary

Fic, 10. — G l di

g f the larva. a, 2-5, c-c’, three silk glands attached at both ends ;

d, e, two silk glands attached at one end; sr, silk receptacle ; sf, spinneret ; /, fat bodies ;

, brain ; g, subeesophageal ganglion ; », band of regenerative cells of stomach; A, point

of junction of sucking tubes; s, sucking pharynx; », muscle attachment of pharynx;

o, esophagus

canal posteriorly from the stomach. That others have expe

rienced like difficulties in dealing with near relatives is shown

by the contradictory statements made concerning Osmylus by

students of its life history (Suckow, Ramdohr, Loew, Dufour,

and Hagen). I have found the work particularly difficult, as

the alcoholic specimens were too brittle for dissection, and I

was obliged to depend entirely upon the study of sections.

Hagen figures for Osmylus four Malpighian tubules attached

at both ends to the intestine, the anterior attachment near

the base of the stomach, the posterior one near the hinder

end of the intestine. Three other tubules have only the ante

rior attachment and have their posterior end free. All of the

tubules are modified in their middle portion for the secretion

of silk.

No. 428.] THE METAMORPHOSIS OF SISYRA. 623

Meinert found similar conditions in Myrmeleon, except that

there were six tubes with the double attachment and two with

free ends. He also found that there was no posterior exit

from the stomach, but that the small intestine (?) had atrophied

into a compact string of cells with no opening between the

point of attachment of the silk-secreting tubes. The unassim-

ilated portion of residuum is expelled as a mass after the insect

has become an imago.

In Sisyra, as I have before mentioned, there is no such

residuum, since the juices of the sponge are ready for com-

plete absorption. An extensive digestive tract being in this

way rendered superfluous, nature has economized by modifying a

large part of the alimentary apparatus into a silk-secreting organ.

The posterior fourth of the stomach appears merely as a

solid cord of atrophied cells, which ends in the walls of a

dilatation, — “silk receptacle," it may be termed. The walls.

of this receptacle have the structure of those of the Malpighian

tubules. It appears to be the outlet of five tubules, three of

which are attached here at both ends, and two of which extend

posteriorly and end in the body cavity. All are modified in

their middle portions for the secretion of silk. The cells of

the silk-secreting portion are much larger and more irregular

in shape than the ordinary Malpighian tubule cells, and show

the singular branched nuclei characteristic of silk gland cells

in the caterpillar and other insects.

The three tubes having the double attachment proceed

anteriorly to about the middle of the metathorax and are

then modified at or near the point of their backward turn.

Posteriorly they are modified almost immediately after leaving

the silk receptacle at the base of the stomach. The two tubes

having the single attachment have a considerable portion of

unmodified tubule at their distal extremity, intricately coiled

upon itself, as is shown by the number of variously cut sections

which appear at this place in any of the three standard planes

. through the body. The length of the silk-secreting portion is

increased by complicated coils and turns of the tubes, so that

they completely fill the body cavity in the first seven abdominal

Segments, making it next to impossible to accurately trace

624 THE AMERICAN NATURALIST. (Vor. XXXVI.

their course in sections. It seems probable, however, that

there is no constancy in the nature of these coils, as the num-

ber of sections of the tubes varies greatly in different speci-

mens. Iam reasonably certain, however, of the three attached

and two free tubes, as the cross sections show quite uniformly

in the anterior portion of the abdomen multiples of three, and

in the posterior, multiples of five. The variations from this

number can be accounted for by turns in the tubes.

Connecting the silk receptable with the spinneret, which

occupies the last three abdominal segments, is a tube which

seems to have a straight course except for a bend ventrally in

the sixth segment. This is the outlet for the silk secretion

which collects first in the silk receptacle. The walls of this

outlet tube have somewhat the structure of those of the Mal-

pighian tubules, yet the cells are more regular in shape and

have greater uniformity in the size and number of their

nuclei. They also show, in a slight degree, the columnar

nature of intestinal cells. This fact, together with the ventral

bend in the course of the tube and its terminations, indicates

that it is the small intestine, entirely diverted from its original

function and modified so as to serve merely as an outlet for

the silk secretion.

After passing through this tube, the silk collects in the

spinneret, the walls of which are surrounded by bands of strong

circular muscles, which aid in ejecting the secretion when the

cocoon is spun.

In Fig. 10 the nature and position of the spinning glands

are shown diagrammatically. The complete course of one of

the three which are attached at both ends, and of one of the

two which are attached at one end, is figured.

Fig. 11 shows longitudinal and cross sections of the silk

glands with their fantastically formed nuclei. The typical

structure of the Malpighian tubule is also shown and its grad-

ual modification into silk-secreting cells. In a, Fig. 11, 4

section of a functional silk gland is shown, the walls being much

thinner and the nuclei of simpler forms than in å of the same

figure, where no secretion is visible. Some of the sections

figured show traces of vacuolization, a common sign of the

No. 428.] THE METAMORPHOSIS OF SISYRA. 625

degeneration which is characteristic of the first stage of meta-

morphosis. Sections of older larvae show the silk glands in a

much more advanced stage of degeneration, and numerous

leucocytes in their vicinity or

clinging to their walls. Inthe

pupz there are only a few frag-

ments of the larval silk glands,

but adult Malpighian tubules

are seen forming, for which the

material from the degenerated

silk glands is probably again

used.

Digestive Epithelium.—

Sections through the digestive

epithelium of a normal, feeding

larva show vacuolization, even

n showing gradual m

fi n into silk-secreting cells ; e, longitudinal

section of silk gland, showing vacuolization. at this early stage At the

anterior end of the stomach is a circular band of small cells

which remain active and functional during the period of degen-

eration of the other epithelium. In the process of pace

all traces of cell boundaries disappear, in some

places the cell substance being a liquid mass

in which float the nuclei. The disintegrated

epithelium is finally cast off from the walls

into the center of the cavity and the new

rapidly grows out from the regenerative band

to take its place, new nests of regenerative

cells appearing at frequent intervals. It would

seem that the solid cord of cells at the poste-

rior end of the stomach serves as another

regenerative center, since at this stage there

is an opening from this end of the stomach pepe

— Larval wing.

A,hypodermis; 2,

through what was the solid cord in the larva. peripodal membrane ;

The adult intestine seems to be developing ^A% * whs

from the cells at this point as a regenerative center. The adult

Malpighian tubules are probably also developed in part from these

cells. I have seen clear evidences of a new set of these organs

forming in the normal position at the base of the stomach.

626 THE AMERICAN NATURALIST. ` (VoL. XXXVI.

DEVELOPMENT OF THE WING.

Some time before the larva spins its cocoon the wing

buds begin to appear, showing through the thin skin. These

form a very convenient index of the age of the larve. Fig. 12

represents a longitudinal section through a wing bud at an

(e v?

| j TAM

Fic. 13. — Section of older wing. 4, basement membrane; c, cuticle; 7, trachea; 7, leucocytes.

early stage, before it is clearly visible from the outside. The

two layers of closely packed cells have not yet united. The first

formation of the trachez is also shown.

The next stage figured is much later, when the two layers

have united, leaving large openings through which pass the

trachez, now fully developed (Fig. 1 3). Leucocytes are found

frequently in these openings. The great stretching of the

wing tissue by its growth both longitudinally and laterally has

drawn the upper part of the cells into slender processes, all of

Fic. 14. — Section of older larval wing. 5», basement membrane ; c, cuticle;

“ £, trachea; 7, leucocytes.

which join the basement membrane, separating the two cell

layers. Many of the nuclei have been drawn up by the strain

from their normal position along the layer of chitin, so that in

some cases they nearly touch the basement membrane. The

subsequent growth of the wing is nearly all in a longitudinal

direction.

No. 428.] THE METAMORPHOSIS OF SISYRA. 627

Fig. 14 shows the effect of a greater stretching in a

longitudinal direction. Most of the nuclei have returned to

their position next the cuticle. A very few have progressed

too far for return, and are left clinging to the tracheal open-

ings or standing alone

and apart from their

fellows. The cell pro-

cesses are much more

ragged and broken

than in the last stage,

and the basement

membrane is pulled

out of its straight line,

being very uneven and

irregular. Traces of

FiG. 15. — Pupal wing. c, rudimentary chitin; 4, mother the old connection

cell of hair; 7, leucocytes ; z, trachea.

may be seen, however,

in the slight thickenings in the cell processes. Along the

tracheal openings where the adult veins are to be, we see

preparation going on for the task of depositing extra layers of

chitin. At these points the cells become aggregated in two

or more apparent layers and have larger nuclei.

The next section figured is from the wing of a pupa

(Fig. 15). Here the wing has increased greatly in length and

decreased in thickness. The slender cell processes have

almost entirely disappeared, all the cells are on one level and

have secreted a thick

layer of material,

which, after the final

molt, will harden

into chitin. Owing P

to the close quarters :

within the pupal rre. epe E cs a, soon after molting ; 2, older.

skin, the wing has trachea; A, cee cel of hair; 7, leucocytes.

become extensively fluted and crinkled to permit the now rapid

growth longitudinally. In Fig. 15 a hair cell is shown develop-

ing in the cuticle, —a large vacuolated mother cell, with nucleus

at one side and small cells clustered about it.

628 THE AMERICAN NATURALIST. [Vou XXXVI.

The next stage shown is from a recently transformed adult

(a, Fig. 16). The crinkling has disappeared, leaving the wing

nearly flat. The processes are contracted within the cell body,

drawing the chitinous layers close together. Near the trachez

are shown the fully developed hair bulbs. The tracheal open-

ings and spaces between the cells show numerous leucocytes,

and all the hypodermis is engaged in the work of thickening

the chitinous covering.

In 4, Fig. 16, a section of an older wing is represented.

The upper chitinous layer has become much thickened and the

only traces of cells are the occasional nuclei and the thin line

Fic. 17. — Wings of adult, x 15. a, fore wing; 4, hind wing.

of protoplasm between the chitinous layers. In the wings of

some insects nothing but the chitinous plates persist, but

Sisyra seems to retain throughout this slight vestige of cellular

tissue.

Fig. 17 shows the venation of the fore and hind wing of Sisyra.

It is rather simple for this family. There are few cross veins.

The course of the principal veins may be easily followed. There

is, however, quite an extensive variation in the tips of the main

branches. A study of this variation should throw some light

on the method of evolution of the peculiar hemerobian type.

I have tabulated the extent and nature of the variation in fifty

wings. Fig. 18 shows the type of branching of the principal

No. 428.] THE METAMORPHOSIS OF SISYRA. 629

veins, as far out as the point where they cease to be uniform,

and the ten different forms of tips appended to these branches,

found in the specimens studied. The branches of R2, R, etc.,

are lettered (a, 4, c, etc., Fig. 18), and the different types of

tips numbered (1-10, Fig. 18) to correspond with the letters

and numbers of the table. . The variants and their frequencies

are shown for each lettered tip in the table; e.g., tip No. 7—

the most common type— occurs, in the fore wing, four times

on branch a, sixteen times on branch å, etc.

FORE WING. | HIND WING.

LEFT. SYMMETRICAL. | RIGHT. LEFT. SYMMETRICAL. | RIGHT.

I * 5.45 o 1 B 4 10 1 4534 5 B T 5 9 Io

a 3 4-4 1 3 2 2 15 3

b 31 216 4 5 20

€ ta if or 4

d 5 19 I 31 3

e 6. | 2 17 [32

J I 22 2 4 2i

s I 24 I LII I3

A 7 18 14 II

; eee d uz 4

H H I 19 6

k 2 12 9 2 n br4 3 2

É 9 2:5 4 7 Hj 3 4

- 9 7.5 4 5 29

^n II 7 7 2 22 I

o 4 16 I 4 2314 42

Ż 4 2I 25

From a glance at the table and Fig. 18 one sees the

tendency is toward extensive multiplication of symmetrical

branches on R2, and that back of R; the branching becomes

increasingly asymmetrical toward the left, this asymmetry cul-

minating in Cz:, especially in the hind wing. No variation in

the cross veins of the hind wing was observed in the twenty-

five specimens examined, and only a slight variation in those of

the fore wing. In every case the variations noticed were in

the disk of the wing, except that in two cases there was a

Cross vein between the first pair of secondary branches of R2

630 THE AMERICAN NATURALIST. [VoL. XXXVI.

(a, 6, Fig. 18). Such a case is shown in Fig. 17. The cross

vein in the basal part of the subcostal space, said by some

to be absent in Sisyra, was present in every specimen. The

other variations seen were in the number of cross veins between

KA: and R2, which is normally three. In two cases four were

present and in one other instance there were five. But these

were all of a nature easily explainable, as there were two or

three faint veins standing close together, showing that the

material usually forming one vein had formed several. Between

Rz and R3, the only other space showing any variation, the

Fic. 18. — Diagram of variation in wing veins, illustrating the table.

normal number is one. In seven cases there were two, in one

three, and in one four. The same tendency to have several

weak veins close together was shown here. All the added

cross veins were on the proximal side of the normal one. The

variation in cross veins seems to have no connection with that

in the tips. The specimen having the most erratic form of tip

(9, Fig. 18), which occurred only once, had the cross veins

entirely normal.

SUMMARY.

The points of chief interest developing from this study may

be summarized as follows :

1. In Sisyra the mouth parts are not in the usual opposed

position, but are approximated in pairs to form two sucking

tubes, excessively elongated, and decurved.

2. The brain is in the prothorax in old larvae.

3. The stomach has no posterior opening in the larva.

No. 428.] THE METAMORPHOSIS OF SISYRA. 631

4. Three Malpighian tubules are attached at both ends,

and two are attached at one end, —all metamorphosed into

silk glands in their middle portion.

5. The small intestine is modified into an outlet for the

silk secretion.

6. The spinneret is formed from the terminal part of the

alimentary canal.

7. A completer account than has hitherto appeared is here

given of the late stages of wing development, and the variations

in venation of fifty wings are tabulated.

BIOLOGICAL LABORATORY, LAKE FOREST COLLEGE.

BIBLIOGRAPHY

1771. DE GEER. Mém. pour servir à l'histoire nat. ins. Tome ii, pt. ii.

1839. Westwoop. An Anomalous Insect found in the Vene fluvi-

atili. Mag. Nat. Hist. (2d series). Vol. iii, pp. 200. Amn. Sci. Nat.

(2d series). Vol. xi, pp. 380.

1844. GRUBE. Beschreibung einer auffallenden, an siisswasserschwammen

lebenden Larve. Wiegm. Arch. f. Natu Bd. xi, pp. 331-337. 1 pl.

2. HAGEN. Die Entwickelung und fe innere Bau von Osmylus.

Linnea Entomol.

Vol. v, pp. 454-457.

5. VINE. Anatomy and Habits of Aphidivorous Neuroptera. Zt. Jour.

Micros. Vol. v, PP. 254—267, 395-408.

1901. NEEDHAM. Aquatic Insects in the Adirondacks. Bulletin N.Y.

State Museum. No. 47: PP. 555-557.

THE REGENERATION OF THE PHARYNX IN

PLANARIA MACULATA.

HENRIETTA F. THACHER.

In a paper on Planaria maculata, published in 1898, Morgan

called attention to an important feature of the regeneration of

the pharynx, namely, that in regenerating pieces which were

cut anterior to the old pharynx, the new pharynx forms “at

the edge of the old tissue, but lying for the most part within

the zew material," while if the cut were made posterior to the

pharynx the new organ forms in the anterior end of the o/d

tissue. During the next two years there appeared a number of

papers describing experiments on the power of regeneration in

planarians, but giving no further knowledge of the develop-

ment of the pharynx except that to be derived from general

statements of the various positions it may assume in the regen-

erating pieces. More recently a paper by Bardeen on the

physiology of Planaria maculata has taken up the formation

of the pharynx in this species and given a somewhat detailed

account of it.

In studying the origin and growth of the new pharynx, I

wished to find out if there were any marked difference between

its formation in the zew tissue of the anterior piece and in the

old tissue of the posterior piece, since the conditions for regen-

eration seemed so unlike in the two cases. At the same time

I was interested in observing in the posterior pieces what might

be the influence of the reproductive organs when present in the

region where we should expect the pharynx to appear, and if,

under these conditions, the already existing cavities and external

opening can be made use of. In this connection I examined

s^ries of regenerating posterior pieces in which the cut had been

made (1) just posterior to the pharynx, but at a time of year when

the reproductive organs have disappeared ; (2) anterior to these

organs when they are fully developed ; and (3) posterior to them.

634 THE AMERICAN NATURALIST. [Vor. XXXVI.

The greater part of the material for my experiments was

collected from ponds near Woods Hole during the months of

June, July, and August, and was preserved for study in the

following winter. Later in the year more planarians were

found near Bryn Mawr, and were kept in aquaria until needed.

Each worm was cut into three pieces, of which the middle, con-

taining the pharynx, was rejected, and the anterior and posterior

pieces were allowed to regenerate. The latter were killed at

intervals of twelve hours for about eight days, at the end of

which time those remaining had begun to feed. Where it

seemed advisable, series were killed at closer intervals, but

as a rule this was not necessary. The pieces remained in cor-

rosive acetic for about ten minutes; after cutting they were

stained on the slide in Delafield’s haematoxylin, and then

dipped for a few seconds into a strong solution of aniline

orange. The latter differentiates the endoderm, muscle bands,

yolk cells, and the lining of the reproductive system.

Soon after the worms are cut, the edges of the cut-end

draw together and matter from the digestive tract and loose

parenchyma tissue collect about the injured region. Within

a few hours the surface opens out again, until only a slight

depression marked by a dark line of pigment is noticeable.

In the mean time parenchyma cells collect at the cut surface

and form a narrow sheet across it, while cells from the margin

of the uninjured epithelium push out and cover the new growth.

This overgrowth of epithelium is comparatively rapid, as the

cells in this region flatten out and cover individually a much

larger area. By the end of the’ first day the tip has become

rounded and the anterior end is covered with an epithelial

layer; later, with the increase in their number, the flattened

cells resume their columnar form.

The parenchyma of the normal planarian appears on exami-

nation as an ill-defined mass filling in the spaces between the

organs. Its protoplasm stains very slightly, but the tissue is

conspicuous owing to the many deeply colored nuclei that are

present in it. These nuclei are occasionally seen in small

groups, but they are usually scattered and separated from each

other by the loose spongy tissue. Among the parenchyma

No. 428.] PHARYNX IN PLANARIA MACULATA. 635

cells are found other cells which are specialized in function, —

large rhabdite cells, lying just below the ectoderm and show-

ing the rhabdites buried in the protoplasm, and the mucous

cells, staining an intense blue; also, along the digestive tract,

a few large granular cells for which Bardeen has suggested a

digestive function. In addition, I have noticed some large

cells staining with aniline orange, which seem to contain some

such material as yolk. Whether this is a fourth kind or merely

the digestive cells I have not been able to decide. There are

also present elongated connective-tissue cells.

Although the majority of the parenchyma cells show little

protoplasm, scattered through the tissue of the normal unin-

jured animal there are a good many cells whose nuclei resemble

those of the ordinary parenchyma, but which have gathered

about them an irregular mass of granular protoplasm. As

many of these are in various stages of division, while I have

found no division among the cells with less protoplasm, it is

probable that the accretion of protoplasm is the usual fore-

runner of division in the parenchyma cells, and that the number

of the latter is constantly increasing even in normal planarians.

While the number of these dividing cells differs individually,

they are often very numerous and are confined to no one region

of the body, except that they are more rare in the extreme

tips. The dividing cells stand out very distinctly as they are

larger and seldom have other cells near them.

After the animals are cut, the amount of division increases

somewhat throughout the whole body, but very markedly near

the injured surface. Cells also migrate down into this region

and form there a thickening which rapidly enlarges. That

migration takes place is indicated by the long trails of proto-

plasm that give to the cells a bipolar appearance with the axes

directed towards the growing mass (Fig. 1), while in the normal

planarian there is no such effect and the cells are more irreg-

ular and rounded in outline. After the thickening has reached

a certain degree of concentration there is rarely any cell divi-

- Sion in the proliferated mass, but mitotic figures can frequently

be found in the region near by, where the cell mass is less

dense. Cells also continue to migrate into the new part and

636 THE AMERICAN NATURALIST. | [Vor. XXXVI.

add to the new growth. The number of cells in the parenchym-

atous tissue of the body itself is also usually much increased

as the nuclei throughout continue dividing, and by the time

the pharynx thickening appears, which is in the course of the

third day, the tissue is filled with cells conspicuous on account

of their greater amount of protoplasm.

I shall consider first the growth of the pharynx in the ante-

rior piece and then that in the posterior piece. By the time

the proliferation of cells in the regenerating posterior tip

is quite marked, a slight shift in the direction taken by

some of the cells denotes the beginning of the new pharynx.

The cells now collect just on the line between the. old and new

tissue at the posterior end of the digestive tract, which has

rounded off, and usually has begun to bud out side branches.

As is true of the growing end, so here, when the thickening

becomes marked, no division of cells can be found in it, but

in the less dense tissue around it there are many dividing cells

and apparently considerable cell migration towards the new

growth. The collection of cells stains very deeply in compari-

son with the old tissue near by, due probably to the greater

amount of protoplasm in this region.

The thickening for the pharynx grows until it is quite large,

and then the split for the pharynx chamber begins. This

opens up very rapidly as a narrow cavity running across the

base of the thickening and up on the sides, being usually

more advanced on the ventral than on the dorsal side. It !5

at first irregular in outline, but the cells along the edge soon

flatten out into a thin lining epithelium. As development

proceeds, cells are added at the anterior end of the new

pharynx, while there is no division in the compact mass of cells

already collected (Fig. 2). About the fourth day the lumen

of the pharynx itself appears in the center of the proliferation,

due to an elongation and pulling apart of cells which become

larger and thin walled and form the lining of the pharynx.

This cavity gradually enlarges posteriorly to open into the

pharynx chamber, and anteriorly to unite with the wall of

the digestive tract, which then breaks through to join with ie

The new region has in the mean time steadily increased !n

No. 428.] PHARYNX IN PLANARIA MACULATA. 637

length, and the branches of the intestine have pushed back

into it, while the central digestive cavity has also somewhat

enlarged. Shortly after the lining of the lumen has formed,

the muscle bands of the pharynx differentiate. They arise

from cells just between the two epithelial layers ; they elongate

and undergo some change in their constitution, since they

stain with aniline orange very shortly after the lengthening

process has begun. The change is first visible in the region

where the pharynx is continuous with the body of the animal

Fic. 2. Fic. 1.

Fic. 1. — Large parenchyma cells migrating towards regenerating region.

Fic. Que dies in anterior piece at close of fourth day. The lumen has just appeared.

lumen ; Z, lateral digestive branch; C, pharynx chamber.

Fic. = S ltd in n posterior piece at cione of third day. The lateral digestive branches have

y ) There are many dividing cells present.

and extends gradually posteriorly. So far as I could deter-

mine, the circular muscles develop before the longitudinal,

but if so, the difference in time is very slight. In the course

of the fifth or sixth day the pharyngeal chamber opens to the

exterior by an ectodermal invagination from the ventral sur-

face, and shortly afterwards the animal begins to feed.

The shape and size of the pharynx vary comparatively little

after the inner lumen appears. At first, before the formation

of this cavity, the shape is slightly pointed, but it soon becomes

more regular and assumes the form of the normal pharynx,

except for its length, which is only about once and a half or

twice the breadth. The addition of cells at the anterior end

638 THE AMERICAN NATURALIST. | [Vor. XXXVI.

does not continue after the structure is well organized, and the

latter part of the time needed for the regeneration of this

organ is occupied in changing the parenchyma cells into the

different tissues. Up to the time when the muscles appear,

the cells still remain crowded together and show by their deep

color the presence of unusually dense protoplasm. After the

muscles are well formed the tissue assumes a less dense appear-

ance, and this change arises apparently at the same time in

the cells throughout the new tissue. Deeply staining masses,

as of mucous, can also be seen, so that the tissue soon resem-

bles that of the normal worm.

The process of formation of a new pharynx in the old tissue

of the posterior piece seems to be very similar, and equally

simple during the nine months of the year when the repro-

ductive organs are absent. The overgrowth of the cut sur-

face and the increase in the number of dividing cells is just

as was found in the anterior piece. The pharynx thickening,

however, appears about twelve hours later than in the former

case, z.e., about the end of the third day, and is usually notice-

able shortly after the two branches of the digestive tract have

joined. But that the pharynx is formed with relation to the

“axial gut," as Bardeen states, I am not prepared to say, as

occasionally a case is found in which the branches have not

met, although a thickening is present, and often in a slide

showing a conspicuous gathering of cells the union between

the intestines has barely taken place. The pharynx forms 4

short distance posterior to the cut, in the old tissue. It is

first made visible by the shifting of the axes of some of the

cells, which now direct themselves toward the new point of

activity, and the thickening increases rapidly (Fig. 3). The

cavities open up just as in the anterior piece, but the chamber

is apt to be much longer and more irregular in outline than

in the other case. Later the enlarged cavity rounds off

so that the chamber resumes its normal size. The central

cavity of the digestive tract enlarges in a posterior direction,

and at the same time the pharynx thickening is added to by

cells at the anterior end; thus, by the time the pharynx !5

ready to open up, the original space between it and the

No. 428.] PHARYNX IN PLANARIA MACULATA. 639

digestive tract is bridged over (Figs. 4, 5). The appearance and

development of the lumen, muscle bands, and normal spongy

tissue is identical with that in the anterior pieces, and the

posterior piece is also ready to feed in seven days.

Summing up these results, it appears that the regeneration

of the pharynx in the new and in the old tissue is much the

same, (1) as to the origin and migration of the cells, (2) as to

the formation of the two cavities (the chamber and the lumen

of the pharynx), and (3) as to the differentiation of the cells

into the normal tissues. They differ (1) in the length of time

that elapses before the pharynx thickening shows itself (about

twelve hours more being required for its appearance in the old

tissue than in the new), (2) in the position of the developing

— ko

Fic. 4. Fic. 5.

Fics. 4, 5. — Stages in development of pharynx in posterior piece.

pharynx with regard to the central digestive cavity, and (3)

in the size of the pharyngeal chamber as shown by its early

irregularities in the posterior piece. The last two differences

are probably due to the disposition of the digestive branches

in the two cases.

In the early part of the summer the reproductive organs of

this species are fully developed, and regeneration under these

conditions is especially interesting. The genital apparatus lies

for the most part in the region just between the pharyngeal

and genital pores, so that when an animal is cut just posterior

to the pharynx these organs occupy the region where the

new pharynx should develop. Under these conditions there

is no sign of the growth for the new pharynx until towards

the end of the fourth day, when a small collection of cells can

640 THE AMERICAN NATURALIST. | [Vor. XXXVI.

be found in a ventral but not necessarily median position.

Also this mass is usually at a normal distance from the fused

intestine. Morgan (1900) has noted this delay in the forma-

tion of the pharynx when the worm is cut anterior to the

genital pore at this time of year (July 2) and finds about

twelve days instead of seven required for complete regenera-

tion. His later experiments, begun July 31, gave different

results, but this was probably due to the degeneration of the

reproductive organs between these two dates, as he found that

the time required for this set of pieces to become complete

animals was only seven days.

This delay may be due to the scarcity of parenchyma cells

in this region, which is almost completely filled by the tissue

of the genital organs. When these organs are fully present

the parenchyma cells collect as shown in Fig. 6, migrating

principally from the anterior end into the region of paren-

chymatous tissue that lies just ventral to the penis. In this

drawing are shown in the tissue along the edge of the geni-

tal organs certain irregular breaks that seem to appear at

the beginning of the degeneration of these structures. The

pharynx forms ventral to such cavities, and the split for the

pharynx chamber is at first entirely independent of them

(Figs. 6, 7). Later the wall between may break down, and

thus greatly enlarge the chamber, but if so it is a secon-

dary process; there is also a tendency towards degeneration

in the thin line of tissue between the pharynx and reproduc-

tive chambers, as shown by the dotted line in Fig. 7. In no

case, however, does the old genital pore become the new

pharyngeal pore.

As the genital organs degenerate, which I have found takes

place during the last two weeks in July and the beginning

of August, the region in which the pharynx appears becomes

quite irregular. The only feature that seems at all constant is

the relative distance between the cross-branch of the intestine

and the pharynx thickening. That this is true may be seen

by a study of its position with relation to that of the genital

pore. The main genital duct opening to the exterior persists

longer than any other part of these structures, but its presence

No. 428.] PHARYNX. IN PLANARIA MACULATA. 641

seems to have little or no influence on the position of the new

pharynx. The latter may appear far anterior to it, posterior,

or, as in Fig. 8, it may assume an eccentric position and lie on

the same level; in this case the duct and pore are shown by

dotted lines, as the pharynx lies about eight sections to the side

of the median line, z.e., of the genital pore. Thus the new

pharynx can be found in all possible positions, and the varia-

Fic. 6.

Fic. 7.

Fics. 6, Lm Development of pharynx during presence of reproductive organs. A, penis; B,

genital chamber ; C, digestive tract (in 6 the wall has broken down — it and the

degen erating cavity); D, eieo due to degeneration of reproductive organs ; Æ, pharynx ;

F, pon on oi hn rel

Fic. 8.

* 4h

duct and

rs to one side of the pore buton the same level. The dotted lines itin di direction

of it ts following sections

tions, such as lateral, ventral, and oblique, should be noted in

connection with any hypothesis of the relation of the pharynx

to the axial gut.

As regards the process of regeneration in the posterior pieces

of worms cut behind the genital pore when the reproductive

organs were believed to be fully developed, the process of

formation seems identical with that which takes place during

the absence of these organs. Consequently there is no need

to describe what takes place.

I wish to express my indebtedness to Prof. T. H. Morgan,

under whose direction this work has been done.

BIOLOGICAL LABORATORY, BRYN MawR COLLEGE.

A STRUCTURAL FEATURE CONNECTED WITH

THE MATING OF DIEMYCTYLUS

VIRIDESCENS.

WILLIAM A. HILTON.

THERE seems to be very little doubt at present that the

fertilization of Urodela is internal. Since Spallanzi in 1785

proved internal fertilization must take place with Triton, many

investigators have shown conclusively that such is the case in

other species.

The mating habits of a number of salamanders have been

observed, but very much is yet to be learned about many forms

concerning which we now know little more than the mere fact

that fertilization is internal; however, it seems very probable

that in all Urodela fertilization is accomplished by a similar

method. In most cases the courtship is more or less compli-

cated and terminates with the female following the male, who

emits a spermatophore containing zoósperms, which is received

by the cloaca of the female as she passes over it.

There are many variations in the mating habits of different

forms described, but unfortunately few species are easy to

observe. Of American forms practically nothing is known

of the more terrestrial species. Diemyctylus viridescens

and D. torosus of the aquatic species are about the only

ones in which a satisfactory idea of the mating has been

obtained.

A number of observations have been made upon European

salamanders by Robin (74), Gasco (80), and Zeller (90), who

have given good accounts of their habits. With Triton, for

instance, Zeller has carefully described the antics of the male

before the female ; these may extend over a period of hours;

he rubs against her and rushes before her. At length she is

attracted by him, and as he moves off she follows and receives

643

644 THE AMERICAN NATURALIST. [Vor. XXXVI.

in her cloaca the spermatophore which he emits. This method

of mating is very commonly described among salamanders,

but variations occur; for instance, in Salamandra maculosa,

and in Salamandra atra, Siebold (58), Pfitzner (80), and others

describe the male perched upon the female's back; he clasps

her for a period before spermatophores are voided. In these

cases the male is usually perched upon the center of the

female's back. In JMegapterma montana and several others

Bedriaga (82) says the male grasps the female and emits

spermatophores while still clasping her.

Ritter (99) in D. torosus described, so far as he was able,

a method of fertilization similar to that given by Bedriaga in

several species; that is, the male emits spermatozoa while

clasping the female. The mating of D. viridescens has been

carefully described by Zeller (90), Gage (91), and Jordan (91),

and, so far as the final result is concerned, the process may be

said to be similar to that described by Zeller and others for

many forms of Salamandridz, but in several particulars the

mating habits are peculiar and deserve further attention.

As breeding season approaches with Diemyctylus, a crest

appears upon the already broad tail of the male, and the cloacal

regions of both male and female enlarge. It is stated by some

that during this time the colors of the male become brighter;

but this is probably exaggerated, as there is but little difference

in the color of the male and female at this time, except possibly

one or the other may have recently passed the red land stage

and not yet attained the deeper color of the more mature

individuals, or it may have just shed its skin. With Diemyc

tylus there is much less difference in the sexes than is found

in many European Urodela, yet there are several striking dif-

ferences, most of which have been noted. The female differs

from the male at all seasons of the year by having more slender

hind legs, and during breeding season there are on the under-

sides of the male's hind legs black, wartlike, horny elevations,

probably developed to aid the male in clasping the female.

These have often been described, but as this method of clasp-

ing the female is rare with European species, the occurrence of

these organs is seldom mentioned with them.

No. 428.] DIEM YCTYLUS VIRIDESCENS 645

Cope ('89) in his work on Batrachia describes a series of

three to four pits along the sides of the head just back of the

eye; he regards these as not constant for the species (Fig. 1).

After examining a considerable number of animals I have

come to the conclusion that these pits are constant structures.

In adult males there are usually three to four quite large, deep

pits, which can be seen without difficulty extending from just

back of the eye along the side of the head. In adult females, as

a general thing, one would say there are no pits, but on a more

careful examination one to four small depressions are often seen ;

these usually appear as pin pricks, or minute depressions in

the skin. They are similar in structure to

thosefound in the adult males, but are much

smaller and less developed in the female.

In early larval forms there are no signs

of depressions or pits in either males or Ak its

females, but in medium-sized to large red ge i E d Diediyc:

males there are along the sides of the head T ii O ER ud

minute depressions which look very much

like those found in the adult females, and more or less advanced,

according to the size of the animal.

In early red females, as a usual thing, no pits are found,

but in later red forms small beginning pits are sometimes

present and sometimes absent ; but judging from all the indi-

viduals examined it may be stated that as a usual thing these

small pits of the female make their appearance at about the

time the red land form changes to the viridescent water form,

and that the corresponding structures in the male make their

appearance in the small red forms about the time that their

sex can be determined by dissection.

Before speaking of the significance of these structures which

are so well developed in the adult male and so rudimentary in

the female and young, it will be necessary to speak somewhat

further of the mating habits. Usually in captivity during the

fall, winter, and spring the male at once settles upon the

female’s back and clasps her just in front of her fore legs with

his powerful back legs in such a manner that it is impossible

for her to escape and, at the same time, possible for him to

646 THE AMERICAN NATURALIST. | [Vor. XXXVI.

bring his head down in contact with hers. Although she is

unable to escape, in many cases she may struggle, carrying the

male with her for a short distance ; but usually it is the other way,

and the male jerks the

female about after hav-

ing been perched quietly

on her back for a short

time. This jerking about

is not without system,

for almost invariably the

2. — Section of a pit and glands of an adult male rd male jerks the female

Fic.

myctylus. g, gland tubules; e, epithelium of pit. x3

somewhat and places one

side of his head just below the top and a little back of the eye,

in contact with that of the female, leaving his head there for a

few seconds and waving his tail with a gentle fanning motion.

After a short period he jerks the female about in such a way

that the other side of his head occupies a

similar position in regard to the female's

nose, pulling her violently in order to do

this. This touching one side of his head

and then the other side to the female's

snout, accompanied by the more or less

necessary jerking about in the water and

fanning movements of the tail, continue pe. 3. — Section of a tubule of

for some time, and after a considerable pears Cee Den

number of these jerkings the male slowly — '

leaves the female, his cloaca expanded, and is usually followed

closely by the female, whose nose is near his tail. The male `

moves slowly forward and throws his body into serpentine

undulations, and a spermatophore is emitted which may come

to the cloaca of the female as she follows him farther along.

The question naturally arises as to what relation these can

stant habits have towards the accomplishment of fertilization;

of course it is evident that it is necessary for the male to be

followed by the female in order that she may receive the

spermatozoa which he emits in masses or spermatophores, and

her following him may be due to an attraction which he exerts

over her, but what is the nature of this attraction ?

No. 428.]

DIEM YCTYLUS VIRIDESCENS.

647

As indicated by Jordan (91), and as simple experiments in

feeding captive Diemyctyli show, probably the strongest sense

of the newt is either the sense of smell or

the tactile sense ; the sense of sight is not

particularly well developed. Now, to return

to the pits on either side of the head ; upon

sectioning these pits it was found that a

number of simple gland tubes were col-

lected about the bases of the pits, and these

tubules (Figs. 2, 3, 4), although not open-

ing widely in the pits when not active,

may be seen to have places where the cell

nuclei were drawn out and arranged paral-

GR, .-

Fic. 4.— Section of a tubule

and i

inactive gland of an adult

mal iemyctylus. 7, tu

bule; e, epithelium. x 210.

lel with the epithelial nuclei; but when the glands are filled

with secretion a few well-marked openings into the pits may

be found (Fig. 5).

peor

These pits are formed

2 from simple depressions or ingrowths of

from an adult male

tylus, showing the opening

int bi. g; helium;

g, gland tubule. x 1s.

this broadening and

epithelium, and the epithelium which lines

the pits does not differ in any way from

that which covers the surface of the body

(Fig. 6). The pits were found to begin

in about half-grown red males or very late

red females. As adult life is reached

these pits or depressions become broader,

and usually deeper, and together with

deepening there is an increased develop-

ment of the glands, which become very numerous in adult males.

The glands are first formed when

the pits are hardly more than narrow

insinkings of the skin, and, as Ancel :

(1901) states for other skin glands */ ^,

of Amphibia, they are derived from . -dp A

the ectoderm. The cells which form * era t

the rudiments of these glands are E

derived from those of the insinking,

or directly from the adjoining sur- *

£ A

ates eB

BED iE a

a bt

»- a

Fic. 6. — Section of a pit in a young red

male Diemyctylus. e, epithelium; g,

udim y gl 3 dabunt x 105.

faces of the skin (Fig. 7). The little masses of cells which

develop into glands become nearly separated from the epithelium

648 THE AMERICAN NATURALIST. (VoL. XXXVI.

from which they were derived, and the communications with

the exterior by means of minute ducts are not formed until

much later. The cells of the early rudiments of glands soon

begin to be arranged in a more and more columnar manner,

and after they are arranged in this way the duct to the

exterior is formed partly from cells of the gland itself and

partly from the epithelium of the

E TE r pit. The fully developed gland

é Pd is simple, saccular, or slightly

T $ de tubular, with a very small duct.

" Pd uf s$ : Sometimes glands are pei

B ^C RC E "um kN b which seem to be parts y

M s DATY or : kte divided into several portions ;

eo Pado that is, have one, two, or more

»- Sete os picts pong whale omens branching off from the

"Diem wt sey sees main one (Figs: 8, 9). New

fee Par coe rui Bue glands are formed from the old

g gland; c, still later stage of developing

gland. x 20. ones in this way, and although

these dividing tubules are found most abundantly in half-grown

specimens, a few of them are often found in the adult. These

glands are more perfectly formed in those individuals where

the pits are best developed; but those in a very active con-

dition, with their lumens widely distended, were only found

in adult males during the breeding

season. During the mating season,

either in fall or spring, these glands

were very much enlarged in the adult |

males, their epithelial cells were much

like low cubes, and the tubules pos-

sessed large lumens (Figs. 3, 10).

When these glands were sectioned red male Diemyctylus of neatly

out of breeding season a very marked sl size, 20,

contrast was noticed; their lumens were much diminished,

the cells reached and nearly touched in the center and no

colloid secretion was present (Fig. 4). These little pits of

the immature animals or of the adult females have at their

bases a few masses of cells, or a few small gland tubules,

which are undoubtedly the beginnings or remnants of the

Fic. 8. — Section of a tubule from a

No. 428.] DIEMYCTYLUS VIRIDESCENS. 649

glands so characteristic of the adult males (Fig. 11); sometimes

one or two tubules appear to have secretion in the lumens, and

some have openings into

the pits.

To recapitulate: The pits

in the adult males are almost

invariably well marked, while

those in the females, when

they occur, are rather diffi-

cult to detect ; and then, too,

the gland tubules in the male Fe. o. — Section of a tubule from a red male Diemyc-

are very ninebrous (Fig. 2). = of nearly adult size. 7, lumen. x 260.

Adult males taken at other times than the breeding season,

as a usual thing, show the gland tubules in a resting state;

the lumens of the glands commonly have the cells almost touch-

ing each other at the center, but males

taken when breeding show these gland

lumens filled with secretion and the cells

appear as a low epithelium.

Se It may be that these glands which are

bame Siopa emi developed so extensively in the male, and

Siei ooi T5! placed in a position which favors them,

epe Tre with secre- are for the purpose of attracting the

female by means of some secretion.

Although, as already spoken of, similar mating habits have

in general been described, they are not exactly the same as in

D. viridescens, nor have similar glands or pits been described

for other forms. Ritter, in an

interesting paper on Diemyctylus

forosus, describes no such struc-

tures, nor was I able to detect

any trace of such pits or glands

in several specimens examined ;

and, in fact, as already stated,

the mating habits are entirely Fc 11-— Section of a pit from an adult female

dissimilar in the two species. earns Me rnn E cio dunt

650 THE AMERICAN NATURALIST. [Vor. XXXVI.

GENERAL SUMMARY.

1. The glands on the side of the head of D. viridescens reach

their complete development only in adult males, and according

to my observations are most active during the time of mating.

2. The pits are formed from invaginations of the epidermis,

and the gland rudiments are derived from these invaginations

or from the adjoining epidermal surfaces.

3. Adult males always possess three to four large, well-

marked pits. Adult females sometimes have one to four pits

on a side, and also a few small gland tubules, some of which

may open into the pits.

4. Early red larval males have small pits about as soon as

the sex can be determined by gross dissection, and at the bases

of the pits there are usually a few small gland tubules.

5. Pits when present in red females make their appearance

late, usually just before the red terrestrial changes to the viri-

descent aquatic form; at this time the pits are seen to be

shallow depressions of the skin, and no gland tubules are

present.

6. No pits or glands were found in the early larval form of

D. viridescens.

7. The glands described occur only when pits are present,

and the tubules have openings into the pits in the adult males,

and sometimes in the females.

8. In mating, the female follows the male only after the

usual courtship, in which first one side and then the other side

of the male's head touches the female's snout.

In the preparation of this paper I have become greatly

indebted to the Department of Histology of Cornell University

and the professors in charge, and also to Mrs. Gage for lending

me her slides of Diemyctylus heads to complete my series

of stages.

No. 428.] DIEM YCTYLUS VIRIDESCENS 651

MORE IMPORTANT REFERENCES.

ANCEL, P. Developp. des glands de la peau des batraciens.

Arch. de Biol. Tome lxviii.

BEDRIAGA, J. VON. Ueber die Begattung bei einigen geschwanzten

Amphibien. Zool. Anzeiger. Bd. v, pp. 265-268, 357-359.

COPE, E. D. Batrachia of North America. Bull. No. 34, U.S.

‘ational Museum, Washington, D.C.

GAGE, S. H. Life History of the Vermilion Spotted Newt (Diemyc-

tylus viridescens Raf.). Amer. Nat. December, 1891.

Gasco, Tr. Les amours des axolotols. Zool. ditiis Bd. iv;

JORDAN, E. O. The Spermatophores of Diemyctylus. Journ. of

Morph.. Vol. v, p. 263

The Habits and Development of the Newt (Diemyctylus

viridescens). Journ. of Morph. Vol. viii.

KINGSBURY, B. F. Spermatheca and Methods of Fertilization in

Some oo Newts and Salamanders. Proc. Amer. Micr. Soc.

Vol. x

RITTER, W. E. Diemyctylus torosus. Proc. Cal. Acad. Sci. Ser.3,

o 2

Rosin, CH. Observations sur la fécondation des urodèles. Journ.

de lanat. et de la phys. norm. et path. de homme et des animaux,

tome x, pp. 376-390; Annals and: Mag. Nat. Hist, vol. xiv

(1874), p. 96.

SPALLANZANI, L. Expériences pour servir à l'histoire de la géné-

ration des animaux et des plants. Pp. 53, 97. Genève, 1785.

SOME HITHERTO UNPUBLISHED OBSERVATIONS

OF ORESTES ST. JOHN ON PALEOZOIC

FISHES. |

C. R. EASTMAN.

Mr. Orestes Sr. Joun, the only pupil of Professor Louis

Agassiz who took up the study of fossil fishes, and whose

researches in this class of vertebrates, published in Vols. VI

and VII of the Minois Paleontology, are among the leading

contributions to the literature of American Paleozoic forms,

passed his early life in the vicinity of Waterloo, Iowa, and

during a period of fifteen years, dating from 1863, brought

together one of the finest collections of Meso-Devonian fish

remains ever obtained from that state. This collection has

recently been deposited in the Museum of Comparative Zoólogy

at Cambridge, where Mr. St. John was formerly assistant in

paleontology, and it is understood that further collections from

the Carboniferous of the Mississippi valley are to follow. In

the same institution are also preserved some of the type speci-

mens which are described in Vol. VII of the ///inozs Reports,

With casts of others that are now in the State Museum at

Springfield, Illinois.

Accompanying the St. John collection of Devonian fishes is

à manuscript description and drawing of a nearly perfect man-

dibular (gnathal) plate of Dinichthys pustulosus, an interesting

Species whose occurrence in Iowa, Illinois, and Wisconsin has

only recently been made known. As this species was originally

described from portions of the dermal armor, examples of the

dentition being at that time unknown, and as doubt has been

expressed ! whether it really pertains to Dinichthys rather than

to some other coccostean, it is of interest to find its dinichthyid

nature confirmed by this early discovery of St. John’s, the credit

! Dean, D. Palaontological Notes, Mem. JV. Y. Acad. Sci., vol. ii (1901), p. 122.

: 653

654 THE AMERICAN NATURALIST. [Vou. XXXVI.

for which is after a long period of oblivion finally secured to

him. The actual specimen, whose description follows, appears

to have dropped out of sight altogether ; but if still in existence,

the present account may perhaps lead to its resurrection.

Other interesting observations of St. John with reference to

Edestus and Cochliodus are contained in the form of a personal

letter to Professor Agassiz, written in 1871, extracts from

which are quoted in the following, and his sketch of Edestus

minor is here reproduced. Detached segments of Æ. heiurichi

have been figured by Newberry! and Trautschold,* and a cor-

responding example of E. minor from the vicinity of Moscow

is described by Karpinsky,® but St. John's specimens are the

only ones, so far as the writer is aware, that have been found

of the latter species in this country.

Professor Agassiz was one of the first to pronounce upon the

nature of these problematical fossils, and compared them with

the rostral prolongation of Pristis. Leidy saw in their seg-

mented condition a resemblance to the compound maxillary of

Lepidosteus, but later regarded them as elasmobranch spines,

an opinion concurred in by Sir Richard Owen, Newberry,

Woodward, and most modern writers. They are now definitely

determined, however, to be the symphysial dentition of cestra-

ciont sharks.*

I. ON THE OcCURRENCE OF DINICHTHYS IN THE HAMILTON

LIMESTONE OF ILLINOIS.

(August, 1882.)

* Professor A. H. Worthen has obtained from the Devonian

limestone of Andalusia, Rock Island County, Illinois, a nearly

perfect example of the right mandibular dental plate of a small

species of Dinichthys, which is quite distinct from either of the

two forms already described by Dr. Newberry from the Ohio

l Ann. N. Y. Acad. Sci, vol. iv (1888), PI. v Vu 2a, 26; Monogr. U. 5.

Geol. Surv., vol. xvi (1889), Pl. XXXIX, Figs. 2

Ha Soc. Imp. Nat. Moscou, vol. lviii (383, fs v Figs.

3 Verh. b. russ. pee Ges. St. Pétersbourg [2], vol. xxxvi dre pP- 381,

450, Text-figs. 1c,

* Geol. Mag. an ix, p. 148; Bull. Mus. Comp. Zoöl., vol. xxxix (1902), No- 3-

No. 428.] PALEOZOIC FISHES. 655

Shale [D. herzeri and D. terrelli]. The age of the beds from

which this specimen was derived is unquestionably not later

than the Hamilton, as is shown by associated Brachiopoda,

etc., characteristic of the Hamilton of Iowa and Illinois. The

present specimen, therefore, affords the earliest indication of

this genus yet discovered.

“The characters displayed by the Illinois specimen, which

serve to distinguish it from the Ohio forms, may be summarized

briefly as follows:

“1, The dental plate under discussion, while resembling

D. herzeri in general form and proportions, differs from the

latter in the character of the cutting edge, which is destitute

of the series of conical denticles occurring in that species.

The dentary portion of the mandible is similarly well demar-

cated from the spatulate posterior shaft, and is similarly rounded

along the front margin. The anterior beak is much produced,

irregularly lozenge-shaped in transverse section, with a sharply

rounded anterior angle and a sharp posterior cutting edge. A

deep notch separates the cusplike beak from a low, round-

crested prominence in advance of the cutting edge of the

dentary, which is deeply beveled along its outer face by attri-

tion against the upper dental plates. To the same cause is prob-

ably to be ascribed the gently concave curve of the trenchant

border-itself. The latter terminates abruptly behind, and along

the posterior slope of the dentary are to be counted the bases

of five small downwardly directed denticles. The lower portion

of the posterior slope is smooth and meets the shaft in an

obtuse angle. In this denticulation of the posterior slope of

the dentary, and perhaps also in the presence of an elevated

prominence or denticle behind the anterior beak, are to be

found the principal differences between the present specimen

and D. terrelli.

“2. The form of the shaft is not unlike that of D. herzeri,

except that it is more contracted immediately behind the den-

tary portion, and more regularly arched along its lower border

before passing into the acutely rounded posterior extremity.

The general outline of the shaft is spatulate, its outer face

moderately convex, and it is separated from the dentary portion

656 THE AMERICAN NATURALIST. |. [Vor. XXXVI.

by a shallow, curved depression. D. terrelli has a relatively

shorter and more massive posterior shaft. As contrasted

with the compact structure of the dentary portion, which

has almost the density of enamel, the spatulate shaft is

finely striato-punctate, and presents more the condition of

osseous tissue.

“3. In D. herzeri, the secondary: prominence or denticle

behind the anterior beak is buttressed on the inner side by a

prominent angulation or ridge [this is really part of the den-

ticle, and occurs in other species as well], which sweeps from

the apex of the denticle downward and backward until it merges

with the thickened portion of the dentary bone. But in the

Fic. 1.— Dinichthys pustulosus Eastm, Hamilton limestone; Andalusia, Ill. Right gnathal

plate. X 4} (nearly).

present species the entire inner surface is deeply excavated,

and there is no evidence of a buttress having occurred.

*4. The present specimen evidently belonged to a species

of much smaller size than either D. herzeri or D. terrelli, since

the total length of the mandible is only 20 cm., while both o

the Ohio forms exceed 60 cm. That it does not pertain to an

immature individual seems plainly indicated by the marks of

wear, and general appearance of the bone.

* The salient features which have been pointed out in the

foregoing are sufficiently characteristic to warrant the estab-

lishment of a new species, which we have pleasure in naming

in honor of Dr. J. S. Newberry, to whom we owe the admirable

notice of the two first discovered species. In the accompany ing

figures are shown (4), view of right mandibular ramus from

No. 428.] PALEOZOIC FISHES. 657

external aspect; (B), inner aspect of anterior portion of the

dentary ; and (C), inferior view showing external outline and

channel separating the dentary portion from the shaft.

“ Horizon and Locality: — Hamilton Limestone; Andalusia,

Rock Island County, Illinois."

(Note. — The specific title which St. John had intended to propose has

already been applied by Dr. J. M. Clarke! to a gnathal plate from the

Hamilton of western New York. Moreover, although St. John clearly has

the priority of discovery, the species which he here recognizes as distinct

has been described under the name of D. pustulosus? Only one of the

Fic. 2. — Dinichthys P

Eastm. Hamilton limestone; Milwaukee, Wis.

Left gnathal plate. 32.

illustrations to which he refers is now extant. It is reproduced in the

accompanying Fig. r, and in Fig. 2 is shown a corresponding dental plate

belonging to an individual of about equal size from the Hamilton of Mil-

waukee, the original being preserved in the Milwaukee Public Museum. A

fragmentary gnathal of D. pustulosus, from the Hamilton of New Buffalo,

Iowa, is also preserved in the Museum of Comparative Zoólogy at

Cambridge.]

1 Bull. U. S. Geol. Surv., No. 16 (1885), p. 17, Pl. I, Fig. 1.

? Bull. Mus. Comp. Zoél., vol. xxxi (1897), p. 38, Pl. III, Fig. 4; Amer. Nat.,

vol. xxxii (1898), p. 748, Figs. 1,2; Journ. Geol, vol. viii (1900), p. 32, Fig. 1.

658 THE AMERICAN NATURALIST. (VoL. XXXVI.

II. OBSERVATIONS ON EDESTUS AND COCHLIODUS.

(From a letter addressed to Professor Louis Agassiz, dated Springfield,

Illinois, June 29, 1871.)

* MY DEAR PROFESSOR :

* Excuse this postscript, but I met with rare success yester-

day morning, in the finding of what appears to be the terminal

denticle of an immature Edestus vorax [sic] Leidy. This is

Fic. 3. — Detached segment of Edestus minor N. and W. Coal measures; Illinois.

the second specimen known to me in which only a single den-

ticle occurs — the other one, from a similar horizon in the

Coal Measures, being referable to Edestus heinrichi N. and W.,

and readily distinguished from the former species by the

stronger [development] and more erect position of the denti-

cles. The slight depression at ‘a’ [Fig. 3] indicates the area

occupied by the overlapping of the posterior extremity of the

succeeding [z.c., preceding] denticle, which, however, is not

a a developed in the present individual.

* «I also received last evening à

remarkably beautiful specimen of

Cochliodus sp. from the St. Louis

limestone. The specimen is very

small, and presents two, or a pair, of

the large posterior teeth in their

relative position ; but what is particu-

larly interesting, it shows the coarsely

osseous [z.e cartilaginous] posterior

prolongation of the rami, which appear to be terminated in

articular processes much in the same manner as occurs in the

modern Cestracion. The two nodes a, a [Fig. 4] are appar-

ently the anterior prolongation of the rami for the support of the

Fic. 4. — Cochliodus sancti-Indovici

(St. J. and W.). St. Louis lime-

stone; Alton, Ill. Enlarged.

No. 428.] PALEOZOIC FISHES. 659

dental plates in that portion of the mouth. And of these anterior

teeth I believe we have specimens —a very small, narrow, enrolled

form — much resembling the ‘second’ tooth of C. contortus.

“The genus Trigonodus of Newberry will have to be aban-

doned, it being identical with his Sandalodus. The described

forms of the latter probably represent the superior dentition.

| * Very respectfully,

HO BEF

[NorE. — It is evident from St. John's pen-and-ink sketch, reproduced

in Fig. 3, that his specimens of Edestus belong to Æ. minor N. a

instead of to E. vorax Leidy. The confusion probably arose from the

fact that Leidy's name is inadvertently applied to the type of E. minor by

Newberry and Worthen in their explanation of Plate I of the fourth volume

of the Z7nozs Paleontology, an error which was subsequently corrected.

The type specimen of E. vorax is now preserved in the Museum of the

Academy of Natural Sciences of Philadelphia; that of Æ. minor in the

Cabinet of Amherst College; and that of Æ. giganteus, which is scarcely

distinct from Æ. vorax, in the Columbia College Museum. The location

of the type specimen of Æ. heinrichi has been ascertained to be in the State

University at Urbana, Illinois.

The depressions on either side of the median projection at a, in Fig. 3,

are not marks of contact with an adjoining denticl2, as supposed by St. John,

but are plainly channelings corresponding to the buttressed condition of the

crown in Campodus variabilis (N. and W.). In fact, a comparison of the

symphysial series of the latter with the type of Edestus minor proves that

the coronal apices of the two forms are surprisingly alike. The detached

segments of Edestus are often pyritiferous, and their decomposition is best

arrested by treating them with a film of collodion.

The spccimen of Cochliodus referred to in the second paragraph above

is figured by St. John and Worthen in Vol. VII of the ///inois Paleon-

tology, Pl. viii, Fig. 8 a, under the name of Pecilodus sancti-ludovici.

There seems to be no sufficient reason, however, for its removal from

Cochliodus. The complete dentition of this genus is known in at least one

jaws were found in natural iation with the anterior series, the latter

having the form of “ Helodus” teeth. The symphysial series of the same

Species has been described by Newberry (Ann. N. Y. Acad. Nat. Sci.,

Vol. xvi, p. 301, Pl. xxiv, Fig. 24) under the name of He/lodus coranus.

In C. contortus the anterior and symphysial series have not yet been defi-

nitely recognized as such. ]

CAMBRIDGE, Mass.

THE MARSH OR RICE-FIELD MICE OF THE

EASTERN UNITED STATES.

S N: RHOADS.

THE status of the rice-field mice (Oryzomys) found in the

marshes of the Atlantic coast from Delaware Bay to the Gulf

of Mexico has been of uncertain fixity since our first knowledge

of their existence in 1837.

In that year a certain Rev. Dr. Harlan of the Academy of

Natural Sciences of Philadelphia is stated by the Rev. John

Bachman to have used the skull of one of these mice, sent by

him for identification to the Academy, as the basis of the first

description of the species, published in .Sz//fimam's Journal.

The name there given was Mus palustris, and the professed

type a specimen in the Academy’s collection labeled as comi-

ing from “Fast Land," near Salem, N.J. Until recently no

other specimens had been secured in New Jersey, and natu-

ralists had so far concluded that the animal was never found

there that Mr. Outram Bangs, in a recent paper on the mam-

mals of Florida and Georgia (Proc. Bost. Soc. Nat. Hist., p. 188,

1898), decided to ignore Harlan's statement as to the type

locality and refer it to South Carolina, whence the Bachman

Specimens were procured. But recent investigations, based on

the chance rediscovery of this interesting mouse in the coast

marshes of Cumberland County, N. J., as announced two years

ago by Mr. Witmer Stone, have enabled the writer to secure a

sufficient series of specimens of Harlan's typical O. palustris

to settle some points in this controversy, as indicated in the

nr brief synopsis.

. Harlan's marsh mouse, Oryzomys palustris ( Har lan).

136 Mus poss Harlan. .Sz/ipkam's Amer. Journ. Sci. Vol. xxxi,

Pp. 386.

D. hoa lity: ^ Fast Land” (a term having no meaning to present

inhabitants except as applied to upland, or land not needing reclamation

from tides), near Salem, N. J.

661

662 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Geographic distribution (of palustris, typicus): Southern New Jersey

to northeastern South Carolina.

Habitat: Tide marshes of the coast, inland to the limits of brackish

water. Wintering in muskrat houses in New Jersey.

Description of species : Size, smallest of the Atlantic coast group. Skull,

relatively large and massive. Colors, above grayish black with pale tawny

or ashy suffusion. Underparts wholly grayish white. Fur dense and soft.

Average measurements of ten old adults from New Jersey and North

Carolina: total length, 237 mm.; tail vertebre, 108 mm.; hind foot,

29 mm.; ear from crown, 12 mm. Skulls of two old males, total length

(basilar, of Hensel), 37 mm. ; zygomatic width, 18 mm.

2. Bachman's marsh or rice mouse, Oryzomys palustris oryzivorus (Bach-

man).

1853 Arvicola oryzivora Bachman. Quad. N. Amer. Vol. iii. p. 214

1893 Oryzomys palustris natator Chapman. Bull. Amer. Mus. Nat.

ist. Vol. v, pp. 43-48.

1894 Oryzomys palustris texensis Allen. Bull. Amer. Mus. Nat. Hist.

Vol. vi, pp. 177-179.

Type locality: Southeastern South Carolina, between Charleston and

Savannah, Ga.

Geographic distribution : Northeastern South Carolina to Orange and

Citrus Counties, Florida, Gulf coast to southern Texas.

Habitat : Same as palustris, but penetrating farther inland to fresh water.

Description of species: Compared with palustris of New Jersey and

northern North Carolina (Bertie County) the differences of Georgia and

north Florida specimens, alike typical of oryzivorus and natator, are too

slight to allow of the recognition of both the latter-named races. In fact,

no palpable departure from palustris is manifest until north Florida speci-

mens are examined. As the zafator form is thus synonymous from an

anatomic and zoógeographic standpoint with Bachman’s type, I am forced

to so consider it in this review.

Oryzivorus, as thus restricted, may be distinguished from palustris by

somewhat larger size, a relatively much lengthened tail, and a narrower

skull. It is a more slender, elongate animal. The colors of upper parts

are browner (less gray) than palustris, the subapical third of the hairs of

back and sides being pale ochraceous brown instead of pale tawny. T9

buffy shade invades the underparts and gives a slightly soiled wash thereto

even on breast and belly, as contrasted with the uniform clear ashy white

of palustris. Average measurements of three adults from St. Mary's, Ga-:

total length, 255 mm.; tail vertebra, 118 mm.; hind foot, 30 mm. ; ear

from crown, 15. mm. Of five males from Gainesville, Fla. (from type

locality of zatator, fide Chapman): total length, 286 mm.; tail vertebra,

136 mm. ; hind foot, 33 mm. ;

. Tę will be seen by above figures that the difference in length consists

chiefly in the great relative lengthening of the zail, the body being not

No. 428.] MARSH OR RICE-FIELD MICE. 663

was taken is 20 mm. longer than average palustris. This illustrates the

tendency of the southern animal to elongate in body without corresponding

3. Bangs' marsh mouse, Oryzomys palustris coloratus Bangs.

1898 Oryzomys palustris coloratus Bangs. Proc. Bost. Soc. Nat. Hist.

Vol. xxviii, p. 189.

1901 Oryzomys natator Jforidanus Merriam. Proc. Wash. Acad. Sci.

Vol. iii, p. 277. (Monroe County, Fla.)

Type locality : Cape Sable, Florida.

Geographic distribution : Tropical life zone of southern F lorida, south

of Caloosahatchee River and Lake Worth.

Habitat : Not given; probably similar to that of its allies,

Description of Species: From Bangs: “ Upper parts rich, reddish

brown, about between hazel and ferruginous. . . . Under parts white,

often suffused with cinnamon. . . . kull similar to that of O. palustris

natator, but slightly larger."

r. Bangs gives much stress to cranial differences between coloratus and

“natator” = oryzivorus and palustris, some of which are rather exaggerated,

and that of the greater size of the supraorbital bead in oryzivorus is not held

out by my New Jersey series. He admits the greater slenderness of the

skull in southern examples. Consonant with this I find the supraorbital

bead to be no more pronounced in these, if anything less heavy than in

Georgia examples. Measurements (fide Bangs) : total length of three adult

males, 301 mm.; tail vertebra, 142 mm. ; hind foot, 34.93 mm.

No cranial measurements are given by Mr. Bangs. I include

Dr. Merriam’s O. natator Jlortdanus as a synonym. He seems

to have omitted coloratus entirely in his recent synopsis. The

small size of foridanus given by Merriam is puzzling.

In conclusion I may acknowledge my indebtedness to

Mr. Outram Bangs for loan of specimens of Oryzomys from

South Carolina and Georgia.

AUDUBON, CAMDEN County, N. J.,

pril r4, 1902.

NOTES AND LITERATURE

GENERAL BIOLOGY

Iron and Living Matter.' — The title together with the first para-

graph of the preface gives a fairly correct idea of the contents of this

book. A somewhat free translation of the paragraph referred to is

as follows: In the course of my bacteriological and morphological

Studies I, like so many other observers, felt myself simply over-

powered with the infinite variety of the vital phenomena and felt an

almost irresistible necessity to gather all these phenomena together

under some one fundamental principle. In the course of my

attempts to find such a principle I came finally to the conclusion

that all vital phenomena are due in the last instance to the oxidation

of the iron of living matter.

One scarcely feels like taking the author’s discussions of the vital

phenomena and their causes seriously because they seem so much

more like the dreamy wanderings of an imaginative but untrained

mind than like a scientific presentation of a coherent set of facts.

It is not clear just why or how the author came to decide upon the

oxidation of iron as being the one fundamental process back of the

life phenomena. He states in the preface that what helped more

than anything else to fix this view in his mind was the fact that he

had succeeded in explaining by means of it all the enzyme reactions.

Indeed the only experimental work recorded in the book is that

which is intended to prove that the characteristic enzyme reactions

are nothing more or less than the effects of oxidations and reductions

of the iron which they are supposed to contain.

The experimental work which is supposed to prove this surprising

result consists simply of qualitative tests for iron and phosphoric

acid in Merck’s commercial enzyme preparations ! The author does

not seem to understand that the occurrence of traces of iron and

Phosphorus in commercial enzyme preparations may be only impuri-

ties and not a part of the active enzyme, nor does he seem to know

! Sacharoff, U. Das Eisen als das thatige es der Enzyme und der leben-

digen Substanz. Translated into German by Dr. M. Rechtsamer in Odessa.

ena, Fischer, 1902. 8vo, 83 pp., 2 pls.

665

666 THE AMERICAN NATURALIST. | [Vor. XXXVI.

that enzymes have been the subject of considerable careful investi-

gation during the past fifteen years and that their nucleoproteid

nature which he has just * discovered " is now in fact being given

up by the men who are most prominent in the chemistry of the

enzymes. Only a few months ago Pekelharing prepared a pepsin

which contained no phosphorus ; this pepsin still contained a minute

trace of iron, as shown by qualitative tests, but it was too small to be

determined quantitatively and Pekelharing paid no further attention

to it. There is no reason for supposing that Pekelharing could not

purify this pepsin still further and thus get rid of the last traces of

iron, as he has already removed the last traces of phosphorus, should

there be any sufficient object in his doing so.

Since the only experimental evidence cited by the author is thus

meaningless there is scarcely any point in following him in a review

of this kind through other subjects, such as synthesis of living matter,

growth, cell division, reproduction, muscle contraction, the activity

of the sense organs, and the chemical processes of the central ner-

vous system. The discussions seem all like the products of a wan-

dering mind whose scientific bearings have been completely lost.

It seems surprising that a firm like that of Gustav Fischer at

Jena could have been induced to publish a work of this kind.

A New Laboratory Manual of Biology. — The teacher of biology

is confronted not only with the problems of his own science, but with

those of how best to teach it, and a laboratory manual is a provi

sional answer to many questions of the second kind. From this

standpoint Hargitt's! Outines will be of much service to teachers.

It abandons the older method of beginning with the simplest repre-

sentatives of animals and plants which are at the same time least

familiar and most difficult of study, and adopts the more recent

practice of introducing the subject by well-known types, in this

instance the frog and the fern. "Then follow exercises on the animal

and the vegetable cell, and finally a series of type animals and plants

ranging from hydra and the molds to the grasshopper and flowering

plants. The mingling of plants and animals in the latter part of the

work, though a time-honored practice, destroys the unity which the

plant and the animal kingdom ought to show and substitutes nothing

of special value for it. The text, which is for the most part clear, 1S

1 Hargitt, C. W. Outlines of General Biology. C. W. Bardeen, Syracuse

N.Y. 164 pp.

No. 428. |J NOTES AND LITERATURE. 66

unaccompanied by figures, thus throwing the student more completely

on his own resources. Here and there it is perhaps too descriptive,

as, for instance, on page 74, where the form of the jellyfish might

have been left for the student to make out for himself. Occasionally

terms could be improved. Thus, on page r9, in the account of the

external apertures of the frog, anus is used for cloacal opening ; and on

page 98 the plates of the starfish are described as 2ozy instead of

calcareous. Except for the wrong font of s's on page 38, the proof

reader's work seems to have been done with much care. These

defects, however, are insignificant compared with the good qualities

of the book, which will undoubtedly find its way to many laboratories

where plants and animals are dealt with in a single course.

ZOOLOGY.

Gegenbaur’s Comparative Anatomy of Vertebrates.'— The

second and concluding volume of this masterly work deals with

the digestive and respiratory organs, the organs of circulation, and

the urogenital system. The present volume is only a little over

two-thirds the size of the first one, and its real subject-matter is

still further restricted, in that about one-fifth of its 7oo pages is

given up to an index of some 20,000 entries, covering both volumes.

There are 354 text illustrations. One impression made by the

perusal of this volume, as compared with the first, is some lack of

completeness. Thus, in the section on the pancreas, though the

ducts of Wirsung and of Santorini are described, no exact state-

ment is made as to their relations to the anlages of the gland, and

the interesting and important phases presented by them in different

mammals is passed over without comment. The lungless condition

of many salamanders is only briefly noticed (p. 302). The descrip-

tion of the arterial system is very fragmentary. Almost no mention

is made of the coronary arteries, whose conditions in the fishes

and in the higher vertebrates present many important modifications.

The exact comparison of the aortic arches of the amniota with

those of fishes is nowhere very clearly brought forward. Although

the relations of the azygos and hemiazygos veins of mammals to

lGegenbaur, C. Vergleichende Anatomie der Wirbelthiere. Bd. ii. Leipzig,

W. Engelmann, rgor. viii + 696 pp.

668 THE AMERICAN NATURALIST. [VoL. XXXVI.

the post-cardinal veins of the lower vertebrates is sketched, the

instructive variations which these veins show is not even alluded

to. lus, in many places the second volume shows deficiencies

where, from the completeness of the first, one would expect to find

an ample and well-balanced account. As in the first volume, so in

the second there is a strong disinclination to accept the results of

embryology. Though this is in some respects an advantage, for in

the last twenty years embryological evidence has certainly been

given much more weight than it should have had, still its almost

complete exclusion is by no means a wise course. The account of

the origin of the complex teeth of mammals and the question of

the number of generations of teeth in this group, lose much by the

omission of embryological facts. So, too, the discussion of the rela-

tions of the pronephres to the mesonephros is left in a very unsatis-

factory form because of the absence of embryological data. While

embryology may have been too exclusively followed by many, its

results are too important to be lightly cast aside. Notwithstanding

the defects that have just been pointed out, the volume before us

and its companion certainly represent the high-water mark among

modern comparative anatomies of the vertebrates, and the author

is to be congratulated on his good fortune in having completed a

book that will bear comparison not only with the best of contem-

porary work but with the best that has gone before. P.

Animal Life in the Deep-Sea. — Professor Seeliger's? pamphlet

of fifty pages presents a very readable popular account of the animal

life of the deep sea. After a brief historical résumé of the growth

of deep-sea investigations, the effect of the environment on the

animals is considered. The icy temperature of the deeper waters

is contrasted with the variable temperature of the surface. The

influence of the enormous water pressure is well illustrated by com

parison with that of atmospheric pressure. The presence of free

oxygen and of carbon dioxide and the influence of the latter on

skeleton building in deep waters are discussed. Finally, the absence

of sunlight and its relation to the sense organs, luminous organs an

food supply of the deep-sea animals is dealt with at some length.

The account, though brief, is carefully compiled, and is followed by

an excellent series of note references, in which some of the questions

brought forward in the text are more fully considered.

1 Seeliger, O. — Zierlebém der Tiefsee. Leipzig, W- aumente r9ol. 49 PP»

1 Tafel.

No. 428.] NOTES AND LITERATURE. 669

Direction of Hair. — Students of general biology will be inter-

ested in Kidd’s* studies on the direction of hair as evidence of the

inheritance of an acquired character. The author points out that

the hair slope of an individual may be modified during life. The

hair streams of mammals are disposed in lines of least resistance,

and yet are of so little importance to the animal that they cannot be

regarded as produced by selection, — natural, sexual, or germinal.

As they have arisen through the mechanical influence of the envi-

ronment and have been handed on from generation to generation,

they afford what the author believes to be a good example of the

inheritance of acquired characters.

Alcyonaria of the Azores.? — The explorations of the Hironde/le

about the Azores, the Gulf of Gascogne, and Terre Neuve brought to

light thirty-eight species, of which nineteen were new. The Alcyo-

naria from the Azores are increased from six to twenty-nine, of which

twelve were new, and the total for the Atlantic between Madeira and

the arctic circle reaches one hundred and sixty-six. Six species

found at the Azores have been taken in the West Indies or off the

coasts of North America, which is about the same proportion of

American species found among the mollusks of the Azores in the

collections of the Hirondelle. The American species are all from

depths below 500 meters, while those peculiar to the eastern parts of

the Atlantic come mainly from levels above 400 meters. The abyssal

forms are thus the only ones common to America and the Azores.

The occurrence near the Azores of several species from the Mediter-

ranean serves to emphasize still further the dependency of its fauna

upon that of the Atlantic. The genus Calypterinus is rejected and

its species transferred to St hyodes, the supposed generic distincti

being due to a pathological condition caused by the presence of an

annelid, one of the Eunicidze, in a furrow of the trunk. CAKE

Corals of the Pacific.* — The author finds that the corals of Hawaii

and Laysan are principally Porites, Pollicipora, and Montipora. The

! Kidd, W. Use-Inheritance illustrated by the Direction of Hair on the Bodies

of Animals. London, A. and C. Black, 1901. 47 pp.

? Studer, Th. Alcyonaires provenant des campagnes de l'Zrende//e, 1886—

1888, Result. des campagn. scient. Albert de Monaco, fasc. 20, 1901. 4to, 64 pp.,

II pls.

? Studer, Th. Ergebnisse einer Reise nach dem Pacific (Schauinsland,

1896-1897 ), Madreporarier von Samoa, den Sandwich-Inseln und Laysan, Zoo/.

Jahrb., Abth. f. Syst., Bd. xiv (1901), pp. 388-428, Taf. XXV-XXXI.

670 THE AMERICAN NATURALIST. [VoL. XXXVI.

Madrepores and Astræidæ common in more southerly waters and

carried northward in the eastern Pacific by the Japan current do

not characterize these northerly reefs of Hawaii and Laysan. The

reef-building corals of this region show more affinity to those of

lower California and the Mexican coast, to the Panamic fauna, due

in the author’s opinion to the weak westerly current in which the

islands in question lie. This not only tends to establish Panamic

influences here but prevents the access of the southern fauna,

eg., the Madrepores. ECAR

The Lepidopterous Genus Depressaria. — The tineid moths of

North America, owing to their great number, small size, and the

difficulty of preserving them in good condition, have not received

the attention they deserve. Yet there are few groups which offer

such good opportunities for original research, our country being

full of undescribed forms, many of interesting habits, remarkable

form, or exquisite colors; while not a few are destructive to culti-

vated plants. Under these circumstances we rejoice to find that

Mr. August Busck, of the Department of Agriculture, has taken up the

study of these insects with remarkable energy and enthusiasm. Our

pleasure in Mr. Busck’s studies is the greater because the material

he works upon is in our National Museum, where it can always be

found and examined by the student. It is to be hoped that collect-

ors in all parts of the country will contribute material and biological

notes, all of which will be carefully acknowledged by Mr. Busck in

publication, if we may judge of his methods by the paper before us.

This paper (Proc. U. S. National Museum, Vol. XXIV, pp. 731-7 49) is

arevision of the North American species of the genus Depressaria, of

which thirty-nine are recognized as valid. It is not necessary in

this place to enter upon any detailed discussion of the paper, but we

may be permitted to point out that two of the specific names are

wrongly spelled, owing to the erroneous spelling (so common among

entomologists !) of the food plants from which the names are derived.

Depressaria psoraliella Wlsm. (p. 740) should be 2. psoraleella, the

plant being Psoralea. D. senicionella Busck (p. 742) should be

D. senecionella, the plant being Senecio. The food plant of D. arnt-

cella, from Mt. Shasta, is said to be A. angustifolia (i.e. A. alpina);

but it seems that this species does not occur on Mt. Shasta, and the

plant was more likely 4. /ongifolia Eaton. T. D. A. €

No. 428.] NOTES AND LITERATURE. 671

BOTANY.

A University Text-Book of Botany.

differs from the same subject as taught when many of the older

teachers of to-day were students, more in its many-sidedness than

in any other respect, — a result largely due to the publication and

use of Sachs' text-book in the seventies, and of the translations,

adaptations, and abridgments of that book that quickly followed its

appearance. Since then numerous and in some cases excellent

laboratory and reference books, prepared for elementary class or

laboratory use or for students of special branches of the science,

have been brought out, but no distinctly original comprehensive book

of large size has appeared from an American author before this

work by Professor Campbell.

In his preface the author clearly states his purpose to have been

the preparation of a reference book, and not a laboratory manual,

presenting in as compact a form as possible an outline of the

essentials of modern botany and drawing its illustrations as far as

possible from American material. A large part of the figures are

original; and while, of necessity, anything like full references to the

enormous literature of even the last few years have been impossible,

each subject is provided with a short bibliography, opening the way

to other references.

The book consists of fifteen chapters, headed: Introduction, The

Plant Body, The Plant Cell, Classification, The Algz, Fungi, The

Archegoniate, Pteridophyta (two chapters), Spermatophyta (three

chapters), Physiology, Relation to Environment, and Geological and

ographical Distribution. The treatment is interesting and direct,

yet conservative in debatable matters, and half-tone plates scattered

through the text, representing types of vegetation and plant societies,

add much to the effectiveness of the ample illustration. W. T.

The Cyclopedia of American Horticulture.? — In the summer of

1900 the first volume of this large and comprehensive work was put

1 Campbell, D. H. 4 visent Text-Book of Pa New York, The Mac-

millan Company, 1902. xv + 579 pp» 15 pls., 493

Bailey, L. H., and Miller, W. Cyclopedia of prca i —Ó Com-

Prising suggestions for Za on of horticultural plants, descriptions of the

species of fruits, vegetables, flowers, and ornamental plants sold in i United

States and Canada, together TA geographical and eam pebes sketches. Illus-

trated with over 2000 original engravings. In four volum New York,

Macmillan Company, 1900-1902. $20.

672 THE AMERICAN NATURALIST. [Vor. XXXVI.

in the hands of subscribers, and the early spring of 1902 sees the con-

cluding volume in print. What has been said earlier in the Maturalist

about the quality of the first three volumes applies equally well to

the one now issued. . Whether the work be consulted by the gardener

for cultural methods, by the amateur for the names of cultivated

plants, or by the schoolmaster for information as to the horticultural

resources and possibilities of a given state or territory, it will be

found to offer a ready answer to most questions and to indicate how

the more obscure ones may be answered by one having the patience

to follow them up. Throughout the Cyclopedia the personality of

its editor is manifest, although very many of the articles have been

written by others; and perhaps the most interesting reading in it is

the introduction to the fourth volume, in which, from his own pen,

we learn how the work was conceived, planned, and executed. The

Cyclopedia is a notable piece of book making, and it is gratifying to

know that the editor hopes, by means of annual supplements, to

round it out with analytical keys for the determination of genera, —

which in the body of the work are alphabetically arranged, — an

extended bibliography, and the current chronicles of horticultural

change. W. T.

Notes. — In the Ottawa Naturalist for April Professor Greene

describes five new species of Ranunculus, from various parts of the

United States and Canada.

In the Botanical Gazette for February Professor Sargent publishes

a fourth paper on * New or Little-Known North American Trees,"

among which are several notable species of Crategus and one of

Prunus.

Prunus virginiana and P. serotina, as cultivated ‘in France, are

contrasted by Guinier in Nos. 1-2 of the current volume of the

Bulletin de la Société Botanique de France.

The Lespedezas of Missouri are reviewed by Mackenzie and

Bush in No. 2 of the current volume of Zransactions of the Academy

of Science of St. Louis.

Lieferungen 4, 5 of Schumann’s * Blühende Kakteen” have

appeared.

_ A voluminous study of Cirsium arvense, by Lund and Rostrup, with

French abstract, has been published as Vol. X, No. 3, of the

Mémoires del’ Académie Royale des Sciences et des Lettres de Danemark.

No. 428.] NOTES AND LITERATURE. 673

Professor Branner, who paid particular attention to palms while

connected with the geological survey of Brazil, twenty or more

years ago, and who then published an exhaustive account of the

Structure of the palm stem, has once more taken up his notes,

and in the Popular Science Monthly for March gives a well-written

illustrated account of the ecological and economic characteristics of

the Brazilian palms.

No. 22 of the current series of “ Contributions from the Gray Her-

barium of Harvard University," constituting Vol. XX XVII, No. 17, of

the Proceedings of the American Academy of Arts and Sciences, is by

Mr. Fernald and deals with species of Carex.

A memoir on Ustilago reiliana, by Mottareale, has been separately

issued from Vol. IV, fascicle 2, of the Annali della R. Scuola Superiore

d’ Agricolture in Portici.

Hefte 7, 8 of Engler's Das Pfanzenreich are devoted respectively

to Naiadacez (by Rendle) and Aceracez (by Pax).

The fourth part, concluding Vol. I, and the second part of Vol. II,

of the British Museum Catalogue of the African Plants collected by

Dr. Friederich Welwitsch in 1853-61 have been distributed by the

trustees of the Museum.

Vol. III, Part III, of J. Medley Wood’s Wata? Plants, comprising

Pls. CCLI to CCLXXV, inclusive, has recently been issued.

Vol. VII of the * Flore de France," by Rouy and Foucaud (con-

tinued by Rouy and Camus), constitutes the 1900 volume of the

Annales de 1’ Académie de La Rochelle, recently issued. It extends

from Rosacez to Cornacez.

Another of Dr. von Schrenk's important contributions to the

economic study of mycology constitutes Bulletin No. rg of the

Bureau of Plant Industry of the U. S. Department of Agriculture,

and deals with the decay of timber and methods of preventing it.

In addition to the results of his own study, the author includes a

Summary of the preventive results reached in Europe. As in his

. earlier papers, illustrations are both full and good. _

A detailed account of the raising and manufacture of vanilla, by

Lecomte and Chalot, is published from the press of Naud of Paris.

A paper on the Caoutchouc-yielding Landolphiacez, by Hua and

Chevalier, is issued by Challamel of Paris.

674 THE AMERICAN NATURALIST. [Vor. XXXVI.

If Mr. Wells’s artist could have illustrated his hypothetical lunar

landscapes from the vegetation of the central African * mountains

of the moon,” as figured by J. E. S. Moore in some cuts reproduced

in Nature of January 23, he would have gained rather than lost in

the uniqueness of the effect.

In Forest Leaves for February Professor Rothrock figures habit and

bark of Pyrus coronaria.

An illustrated paper on Ochnacez, by Barteletti, is published in

fascicle 4-6 of Malpighia.

Professor Greene publishes three new species of Senecio from

British Columbia in the Ottawa Naturalist for February.

Liatris pycnostachya, as a garden plant, is illustrated in Die Gar-

tenwelt of January 11.

The orchids of eastern Asia, as represented in the herbarium of

the Muséum d’Histoire Naturelle, form the subject of an illustrated

paper by Finet, in the Revue générale de botanique of December r5.

The conclusion of Spegazzini’s “ Stipeæ Platenses ” constitutes

. No. 22, Vol. IV, of the Anales dei Museo Nacional de Montevideo.

Fusicladium dendriticum is the subject of Bulletin No. 67 of the

Illinois Experiment Station, by Mr. Clinton. It is illustrated by a

number of reproductions of photographs, and eleven pages are given

to a full bibliography, —- a feature as useful as it is unusual.

In the Botanical Magazine of Tokyo, No. 178, Vyeda has an illus-

trated paper on the “ Benikoji fungus " of Formosa, — used in the

production of a red fermented rice beverage.

Dr. Peglion publishes an article on the cereal Peronospora (Sclero-

spora graminicola) in L’ Italia Agricola of January 15.

“The Algz of Jamaica” is the title of a paper by F. S. Collins,

published as Vol. XXXVII, No. 9, of the Proceedings of the American

Academy of Arts and Sciences.

The life history of Oscillaria prolifica is sketched by Isabel F.

Hyams and Ellen H. Richards in the Technology Quarterly of

ember

An ecological study of the heath formations of northern Germany,

by Graebner, constitutes Vol. V of Engler and Drude's Die Vegeta-

tion der Erde, and is the first of a series of volumes that are to

deal with the plant formations of middle Europe. Though Germans

No. 428.] NOTES AND LITERATURE. 675

commonly use the word Zeide in the sense of a wood, Graebner

agrees with classic English usage in employing it for the open bushy

formations characterized by Erica, Calluna, Empetrum, Juniper, etc.

Professor Selby, in the Buletin of fhe Torrey Botanical Club for

December, records another series of experiments with seeds sub-

jected to the low temperature of liquid air, with the customary result

that their viability appears not to be affected by even a forty-eight

hours' sojourn at a temperature of — 190* C.

Dr. Goodale has a short note in the American Journal of Science

for February on the memorial greenhouses at the Harvard Botanic

Gardens and some of the physiological work being done in them.

The effects of water and of certain aqueous solutions on foliage

are discussed by J. B. Dandeno in a lengthy and well-illustrated

paper, reprinted from Vol. VII of the Zransactions of the Canadian

Institute.

The mapping of botanical data is discussed by Blanc in the Bule-

tin de l Herbier Boissier of December ši:

A practical little handbook of greenhouse methods, that should be

in possession of every “ nature-study " teacher, is Green and Mackin-

tosh’s Outline of greenhouse laboratory work, issued as Class Bulletin

Vo. 12 of the Experiment Station of the University of Minnesota.

A paper by Mr. Chesnut on plants used by the Indians of Men-

docino County, Cal., constitutes No. 3 of Vol. VII of Contributions

Jrom the U. S. National Herbarium, and contains numerous illus-

trations.

Lieferung VI of the new edition of Wiesner’s Rohstoffe des

LYlanzenreiches, issued in December, begins a consideration of

€conomic woods and contains numerous illustrations of structural

detail.

Vegetable powders and the means of knowing their composition

by aid of the microscope are being treated by Greenish and Collin

in current numbers of the Pharmaceutical Journal.

M. de Wildeman has recently distributed an account of late-

Producing Apocynacez, collected in the Congo country by Gentil.

The cause of white-topped meadow grasses in Finland is discussed

by Reuter in Vol. XIX of Acta societatis pro fauna et flora fennica, and

a bibliography of the subject is given.

676 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Dr. White publishes a note on the use of Solanum heterodoxum in

Mexico for the curdling of milk, in Science of December 13.

A long list of decorative plants hardy in South Dakota is given

by Mr. Hansen in Bulletin No. 72 of the experiment station located

at Brookings.

An interesting illustrated article on rattan and its preparation for

the market, by Preyer, is published in Der Zropenpflanzer for January.

In Bulletin No. 6, dealing with Capsicums, by Mr. Tracy, the

Bureau of Plant Industry of the U. S. Department of Agriculture

begins the publication of a series of catalogues of the trade names

of American vegetables.

In the concluding fascicle of Vol. XIV of the eighth series of

the Annales des sciences naturelles, botanique, M. Van Tieghem sums

up the results of his studies on the ovule as a basis of classification

in the flowering plants, giving at the end a résumé of the vegetable

kingdom classified on this basis.

Dalla Torre and Harms’ Genera Siphonojamarum in the fourth

fascicle reaches the genus Cochleanthera of the Guttiferz.

Engler and Prantl’s Die natürlichen Pflanzenfamilien in Lieferung 211

reaches Isoetacez among the pteridophytes and in Lieferung 212,

Pottiaceze among the bryophytes.

An account of the dates at which the parts of Elliott’s Botany of

South Carolina and Georgia were issued is contributed to the Decem-

ber Bulletin of the Torrey Botanical Club by Barnhart.

The double fascicles 14-15 and 16-17 of Ascherson and Graeb-

ner’s Synopsis der Mitteleuropüischen Flora deal respectively with

Rosacez (in part) and Graminez (in part).

In Coste's Flore descriptive et illustrée de la France, the second vol-

ume of which is now in course of publication by Klincksieck of

Paris, the description of each species is accompanied by an excellent

small cut, showing habit and often essential detail.

. An anatomical study of Hippocrateaces, connected with the

occurrence of caoutchouc in that family, by Fritsch, occupies Heft 5

of Vol. XI of the Beihefte zum botanischen Centralblatt.

The aid afforded by calcium oxalate crystals in the identification of

vegetable drugs is the subject of a paper by Kraemer in the Journal

of Pharmacology for December.

No. 428.] NOTES AND LITERATURE. 677

The development, structure, and properties of the epidermis of

certain dicotyledonous plants which persist for several years, forms

the subject of a paper by Damm in Vol. XI, Heft 4, of the Beihefte

zum botanischen Centralblatt.

Professor Heckel publishes, through the house of A. Challamel

of Paris, a valuable paper on “ Les graines grasses . . . des colonies

françaises.”

Part VII of Wiesner's Rohstoffe des Pflanzenreiches, appearing from

the press of Wilhelm Engelmann of Leipzig, is devoted to fibers.

The pollination of Solanum rostratum and Cassia chamecrista, both

of which produce right- and left-handed flowers on the same plant,

has been reinvestigated by Harris and Kuchs, whose paper is pub-

lished in the February number of the Kansas University Science Bul-

etin, ‘The earlier conclusions of Todd are not confirmed.

A paper by Haberlandt, entitled “ Sinnesorgane im Pflanzenreich,”

issued from the Engelmann press of Leipzig, deals with irritable

stamens, pistils, foliage leaves, insect traps, and tendrils.

A biographic sketch of Schimper, by Schenck, with portrait, has

been separately printed from the Berichte der deutschen botanischen

Gesellschaft for 1901.

A Eulogy of Unger, delivered in connection with the unveiling of

the Unger bust at the Vienna University last July, has recently been

distributed by Professor Wiesner, from the Verhandlungen der k.k.

2001.-bot. Gesellschaft in Wien.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS, AND DEATHS.

EDUCATIONAL GIFTS.

Albion College, $21,000, from Gov. A. T. Bliss.

Adelphi College, Brooklyn, $125,000, from John D. Rockefeller ; $125,000

from other donors.

Amherst College, $25,000, for the library, from Col. Mason W. Tyler;

$65,000, from alumni and friends, for an observatory.

Bryn Mawr College, gifts amounting to $256,000, thus securing the con-

ditional gift of $250,000 from John D. Rockefeller.

Columbia University, $50,000, from Mrs. Lena Currier.

Cornell University, $250,000, from John D. Rockefeller ; $250,000, from

ther sources.

Dartmouth College, $32,500, by the will of F. W. Daniels.

Hamilton College, $5000, by the will of Prof. Anson Judd Upson.

Lick Observatory, $2500, from Mrs. Phoebe A. Hearst.

Massachusetts Institute of Technology, $5000 a year, for three years, for

work in sanitary science.

Northwestern University, about $200,000, by the will of James F. Robinson.

Pennsylvania State College, $100,000, from Andrew Carnegie ; $60,000,

from Mr. and Mrs. C. M. Schwab ; $20,000, from James G. White.

Radcliffe College, $5000, by the will of Mrs. Sarah A. Rand.

Rutgers College, $20,000, from F. M. and Ralph Vorhees.

Swarthmore College, a conditional gift of $100,000, from Isaac H. Clothier ;

$300,000, from other sources.

Smith College, $100,000, from various sources, securing the Rockefeller

gift of the same amount.

University of California, $20,000, from Mrs. Phoebe Hearst ; $5000, from

Henry Weinstock; $8000, from Dr. M. Herzstein; $50,000, from

D. O. Mills; $30,000, from other sources.

University of Pennsylvania, $300,000, from Joseph Wharton, for the school

of finance ; $100,000, for new buildings, from an anonymous donor.

University of Vermont, $6000, from Dr. Seward Webb, for the purchase

of the Pringle herbarium. :

Union College, $40,000, from Andrew Carnegie ; $125,000, by compromise, .

from the estate of Thomas Armstrong.

Vassar College, a library building, from an anonymous donor.

Wells College, $50,000, from Henry A. Morgan; $25,000, from N. L.

. Zabriskie.

678

GIFTS, APPOINTMENTS, RETIREMENTS. 679

Wesleyan University, $75,000, from Charles Scott, Sr. and Jr., for a physi-

cal laboratory ; $75,000, from an anonymous donor.

Western Reserve University, $100,000, from A. A. Pope.

Yale University, the residue of the estate of Edward W. Southworth,

estimated at $200,000 to $300,000; a chemical building, costing, with

the land, $96,000, from Mrs. Thomas G. Bennet ; a building for the

geological department, from an anonymous donor.

APPOINTMENTS.

Dr. Florentino Ameghino, director of the National Museum of Buenos

Aires. — A. P. Anderson, curator of the herbarium of Columbia University.

— J. H. Bair, assistant in anthropology in Barnard College. — Dr. F. A.

Bather, assistant keeper in the department of geology in the British

Museum. — Dr. A. N. Berlesi, assistant professor of plant pathology in

the university at Milan. — Dr. Robert S. Breed, professor of biology and

geology in Alleghany College. — Jean A. Brodhurst, assistant in botany in

Barnard College. — Dr. R. E. Buffington, assistant in histology in Columbia

University. — Dr. F. Carara, assistant professor of botany in the university

at Catania. — Dr. D. Carazzi, assistant professor of zoólogy in the uni-

versity at Sassari — Dr. Adolf Cluss, professor of bacteriology in the

university at Halle. — Dr. W. C. Coker, associate professor of botany in

the University of North Carolina. — Dr. Carl Erich Correns, professor

extraordinary of botany in the university at Leipzig.— Arthur L. Dean,

assistant in plant physiology in Yale University. — Dr. R. B. Dixon,

instructor in anthropology in Harvard University. — Dr. J. E. Duerden,

interim professor of zoólogy in the University of North Carolina.— Dr.

B. M. Duggar, professor of botany in the University of Missouri. — Dr. A.

Fischer, professor of botany in the university at Basel. — Dr. P. A. Fish,

professor of physiology and pharmacology in Cornell University. — Dr.

Fournier, professor of geology and mineralogy in the university at

Besancon. — Professor F. Graff, head of the mineralogical institute at

Freiburg i. B. — Dr. Caswell Grau, associate in zoólogy in Johns Hopkins

University. — Dr. Arthur W. Greeley, assistant professor of zoólogy in

Washington University. — Dr. J. Henscher, professor extraordinary of

parasitology in the university at Zürich. — Dr. William Hunter of London,

government bacteriologist at Hongkong.— Dr. Giuseppe Jatta, docent

for zoólogy in the university at Naples. — Dr. E. C. Jeffrey, assistant pro-

fessor of vegetable morphology in Harvard University. — Herbert Stanley

Jevons, lecturer in mineralogy in the University of Sydney, N.S.W. — Dr.

B. F. Kingsbury, professor of human physiology in Cornell University. —

Dr. Ludwig Koch, honorary professor of botany at Heidelberg. — Dr.

Henry B. Kiimmel, state geologist of New Jersey.—Ivey F. Lewis,

assistant in biology in the University of North Carolina. — Haven Metcalf,

680 THE AMERICAN NATURALIST. | [Vor. XXXVI.

professor of botany in Clemson College. — Dr.. P. Mingazzini, professor of

zoólogy in the university at Messina. — Dr. F. W. Neger, professor of

botany in the forestry school at Eisenach. — Dr. Russell B. Opitz, demon-

strator in physiology in Columbia University. — Dr. Charles Palache,

assistant professor of mineralogy at Harvard University. — Dr. Florence

Peebles, assistant professor of biology in the Woman's College of Balti-

more. — C. G. Pringle, keeper of the herbarium of the University of Ver-

mont. — Dr. F. Raffaele, professor of zoólogy in the university at Palermo.

— Hugh D. Reed, instructor in vertebrate zoólogy in Cornell University.

— L. F. Rettiger, instructor in bacteriology in Yale University. — Ralph

W. Richards, instructor in geology in Tufts College. — Dr. H. Ries,

assistant professor of geology in Cornell University. — Dr. D. Rosa, pro-

fessor of zoólogy in the university at Modena. — Dr. Rudolf Rosemann,

professor of physiology in the university at Greifswald. — Dr. A. Russo,

professor of zoólogy in the university at Catania. — Dr. Florence R. Sabin,

assistant in anatomy in Johns Hopkins University. — Dr. D. Saccardo,

assistant in the royal institute of plant pathology at Rome. — Camille Sau-

vageau, professor of botany at Bordeaux. — Dr. Victor Schiffner, professor

extraordinary of botany in the university at Munich. — Dr. R. J. Schubert,

assistant on the Austrian Geological Survey. — Dr. George B. Shattuck,

associate professor of physiographic geology in Johns Hopkins University.

— Dr. Yngve Sjóstedt, custodian of the entomological collections at Stock-

holm. — Dr. Hans Soledever, professor of botany in the university at

Erlangen. — Dr. R. M. Strong, interim professor of zoólogy in Haverford

College. — Dr. Albert R. Sweetser, professor of biology in the University

of Oregon. — Dr. G. B. de Toni, professor of botany in the university at

Sassari, Sardinia. — Dr. Karl von Tubeuf, professor of forestry in the

university at Munich. — Dr. Frank R. Van Horn, professor of geology at

the Case School of Applied Science at Cleveland. — Dr. L. Waager,

assistant on the Austrian Geological Survey. — Dr. Adolf Wagner, docent

for botany in the university at Innsbruck. — William A. Willard, instructor

in zoólogy in the University of Nebraska. — Dr. I. Wendell Williston, head |

professor of paleontology in the University of Chicago. — Dr. Alfred W. G

Wilson, demonstrator in geology in McGill University. — Dr. E. Zacharias,

director of the Hamburg City Institute of Botany. — J. V. Zelizko, assistant

in the museum of the Austrian Geological Survey. — Dr. Carl Zimmer,

custodian of the zoólogical institute and museum in Breslau. — Professor

. A. Zimmermann, botanist to the biological station at Tanga, German

East Africa.

RETIRED.

Dr. Max von Vintschgau from the chair of physiology in the university

at Innsbruck.

No.428.] GIFTS, APPOINTMENTS, RETIREMENTS. 681

DEATHS.

Sergius Alpheraky, Russian entomologist. — Prof. Ernst von Ballion,

entomologist, at Novorossiisk, Russia, Sept. 9, 1901, aged 84.— Dr.

C. Berg, director of the National Museum in Buenos Aires, and a well-

known zoólogist, January 19. — Dr. Alexander Bittner, head geologist of

the Austrian Geological Survey in Vienna, April t. — Dr. Hans Buchner,

professor of bacteriology in the university at Munich, April 5, aged 51.—

Stanislaus Clément, zoólogist and former head of the museum at Nímes,

February 7, aged 73. — J. Ottoman Dietz, student of Coleoptera, in New

York, December 25. — Henri Filhol, vertebrate paleontologist of the Paris

Natural History Museum, April 28, aged 6o. — Dr. Friederich Leopold

Goltz, professor of physiology in the university at Strassburg, May 4, aged

— C. J. Emil Haglund, student of Hemiptera, at Lommaryd, Sweden,

Dec. 9, 1901, aged 64. — Emil Holub, African traveler and ornithologist, in

Vienna, February 21, aged 55.— Dr. Joseph Kriechbaumer, student of

Hymenoptera, in Munich, May 2.— T. J. Léger, professor of botany in

College, Washington, Pa., Sept. 28, 1901. — Dr. Emil Adolf Lóvendal,

conservator of the soblogical museum at UM entomologist and skill-

ful engraver on copper, July 6, 1901, aged 6 . C. Mausel-Pleydell,

zoólogist, in Whatcombe, England, May 3, disi pe — Carl Nehring, col-

lector, in Piracicaba, Brazil, Januar ary 3. — Joseph Nolan, assistant on the

geological survey of Ireland, in Dublin, April 19. — . Penning of the

geological survey of the United Kingdom, in London, April 20. — Andreas

Reischet, naturalist and head of the Linz Museum, aged 55. — Dr. Carlo

Riva, instructor in petrography in the university at Pavia, killed by an

avalanche on Monte Grigna, June 3. — Baron Friedrich Rosen, professor

of mineralogy in the university at Kazan, aged 68. — Dr. Richard Burton

Rowe of the U.S. Geological Survey, at Los Angeles, Cal, May 26, aged

39. — Dr. Ferdinand Sommer, formerly professor of anatomy at Greifswald,

aged 74. — Baron E. von Tróltsch, archeologist of Würtemburg, June 29,

I9or, aged 73.

(No. 427 was mailed July 15.)

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THE

AMERICAN NATURALIST

Vor. XXXVI. September, 1902. No. 429.

THE DEVELOPMENT AND HOMOLOGIES OF

TRE MOUTH PARTS OF INSECTS.

VERNON L. KELLOGG.

THE problem of the homologies of the mouth parts of insects

is one long worked at by zoólogists. Since Savigny's first

Statement in 1816 of his conclusions regarding the homologies

of the arthropod appendages, this problem has been a favorite

one with insect morphologists, and in this century of work

much has been accomplished. There is a practically complete

agreement as to the homologies of the parts of the biting

mouth as this mouth is variously composed in the Orthoptera,

. Coleoptera, Neuroptera, eż aZ, and a fair agreement obtains

~ With regard to the interpretation of the homologies of some

of the more modified kinds of mouth parts possessed by the

Piercing and sucking insects. This is true especially of those .

.. insects, like the Hymenoptera and the Lepidoptera, among which

there are generalized forms showing the essential biting type

(as with the sawflies among the Hymenoptera and Eriocephala

_ and Micropteryx among the Lepidoptera), together with a series

_ 9f gradatory forms leading plainly up to the highly specialized

conditions exhibited by the higher members of these orders.

684 THE AMERICAN NATURALIST. [Vor. XXXVI.

These homology-determinations were first made by a study

of the comparative anatomy of the fully developed mouth parts

(those of adult insects), and indeed have a fairly safe ground-

ing on this comparative anatomical study alone. But with the

development of embryological studies of insects came the con-

firmation of these determinations, or some of them, by the study

of the development of the mouth parts. From their origin as

budding appendages, arising on the successive segments of the

embryonic head, their development has been readily and cer-

tainly traced to the definitive mouth-part condition ; and man-

dibles, maxillae, and labium are as certainly serially homologous

with each other and with the legs and antennz as are the more

obviously homologous segmental appendages of the crustaceans.

But this ontogenic development of the insectean mouth

parts, simple and continuous as it is in the case of insects with

an incomplete metamorphosis, is a very complex and difficult

subject of study in all of those insects which undergo what is

termed a complete metamorphosis, and this for the reason,

now familiar to entomologists, that in the late larval and early

pupal life of such insects a more or less radical histolysis, or

breaking down of the larval organs and tissues, occurs, with a

building up of the imaginal organs from small, primitive cen-

ters called histoblasts (imaginal disks), which are not derived

from the corresponding larval organs but (for the appendages

as legs and mouth parts) from the larval derm or cellular skin

layer. Thus we have in the development of the mouth parts

of insects with complete metamorphosis a discontinuity which

sadly interferes with the determination of homologies by

ontogenetic study. Indeed, so serious has this obstacle proved

that we have as yet practically no complete tracing through

both embryonic and post-embryonic development of the

growth and development of the mouth parts of any insect of

complete metamorphosis. And they are, for the most part,

precisely those insects of most radical post-embryonic meta-

morphosis which possess in adult condition the most highly

modified and specialized mouth parts, and which present to us the

most serious task in the interpretation of the mouth-part homol-

ogies. The Diptera, of course, best exemplify these conditions.

No. 429.] THE MOUTH PARTS OF INSECTS. 685

There is no special difficulty, outside of the general difficul-

ties which the study of insect embryology commonly presents,

in tracing from beginning up to completed larval condition the

development of the mouth parts of insects with complete meta-

morphosis; and the homologies of these larval mouth parts

with the mouth parts of adult insects with incomplete metamor-

phosis can accordingly be determined on a basis of ontogenic

study (also, of course, on a basis of comparative anatomy).

The biting mouth parts of the more generalized flies, of the

lepidopterous caterpillars and coleopterous grubs, can be

homologized with the mandibles, maxilla, and labium of the

adult cockroaches and locusts, constituting the generalized

biting or so-called orthopterous mouth. But when the attempt

is made to carry the homologies on to the adult piercing and

sucking mouths of the flies and butterflies we lose in the pre-

pupal stage our grip on the continuity of embryonic and adult

mouth conditions and find ourselves forced to rest our interpre-

tation of the homologies of the adult dipterous, lepidopterous,

and hymenopterous mouth on the basis of comparative anatom-

ical studies. And fortunately for us the persistence of certain

generalized forms already referred to enables us to make a

pretty secure determination of these homologies for all of the

orders except the Diptera. To my mind, indeed, the study of

the comparative anatomy of the mouth parts of the generalized

flies (families of the Nematocera) enables us to be pretty cer-

tain even in that order, but such an attempt! of mine in 1899

has certainly failed to be convincing to several entomologists.

There is necessary, then, the completion of the tracing of

the development of the mouth parts; nothing less, under the

circumstances that the most generalized of dipterous mouths

are not at all generalized (if one may be so paradoxical), but

are so specialized that no safe determination of the homologies

can be made on the basis of comparative anatomy, — nothing

less will be convincing or satisfactory for the solid grounding

of an interpretation of the homologies of the mouth parts of

1 The Mouth Parts of the Nematocerous Diptera, Psyche, vol. viii (1899) : I,

PP- 303-306, January; II, pp. 327-330, March; III, pp. 346-348, April; IV,

PP- 355-359, May ; V, pp. 363-365, June; with 11 figs.

686 THE AMERICAN NATURALIST. [Vor. XXXVI.

the Diptera, and if this tracing can be effected for the other

orders of holometabolous insects, it will put the homology-

determinations on a much better foundation than they now

have. It is the beginnings of such an attempt that is outlined

in this paper.

NEUROPTERA.

The Neuroptera belong to the holometabolous insects, 7.e.,

insects with complete metamorphosis, but this metamorphosis

in many forms is of a very simple and straightforward kind as

compared with the radical metamorphosis of a fly or butterfly,

for example. The mouth parts of the

adult insect are, too, of the orthop-

terous or biting type, and there is

no question regarding the interpreta-

tion of the mouth-part homologies.

Mandibles, maxilla, and labium of

the neuropterous mouth are obviously

homologous with the similarly named

parts of the orthopterous mouth.

Furthermore, the differences between

the larval and adult mouth parts are

comparatively slight, and no question

i Ge e e ii made regarding. the homologies

removed. /5., labrum; med. dures between the two sets. Yet it 1s

per ats së ron pip rA, worth while to trace the development

labium; Z4. labial palpus; a»^, of the imaginal parts in its more con-

tntenha.

spicuous features, and get a first sight

at the relation between larval and imaginal mouth parts in a

holometabolous insect. This relation is readily made out in

the large and familiar neuropteron called the ‘dobson fly,”

or “ hellgrammite,” Corydalis cornuta.

Corydalis cornuta (Figs. 1-5).—The mouth parts of the

larval Corydalis are shown in Fig. 1i, and their orthopterous

character, together with the details of the various parts, are SO

. readily apparent that little description is needed. The man-

. dibles (md.) are very heavy and long ; the maxillae (szx.) have

a short proximal segment, cardo (not visible in the drawing),

No. 429.] THE MOUTH PARTS OF INSECTS. 687

and a usually elongate parallel-sided stipes (s¢.) bearing at its

terminal extremity the much-reduced three-segmented palpus

(mx.p.) and a stil smaller two-segmented terminal lobe, or

galea (ga.), the lacinia being wholly

wanting; the labium (/Z) has the

glossze (g/.) (inner terminal lobes)

fused but emarginate, the para-

glossze (outer terminal lobes) want-

ing, and the palpi (Zf.) three-

segmented and well developed. The

mouth parts are similar in both

sexes.

When the larval dobson is ready

to pupate (at the probable age of

three years) it leaves the stream it '

has lived in, crawls under some

stone near the water's edge, and

changes into a quiet, non-feeding

pupa, which, however, is not enclosed

in a hard, opaque cuticle, but retains

the power of violent wriggling, and

bears the wing pads and legs only

loosely appressed to the body. The

mouth parts of the pupa (Fig. 2)

show slight yet obvious differences

from those of the larva (and also

from those of the imago) The

mandibles (md.) show a difference

from the larval mandibles in the

Character of the dentation and in

outline of the whole sclerite ; the

maxilla have short, five-segmented

; 5 Fic. 3. — Dorsal aspect of head of old

palpi and two short terminal lobes, larva of Corydalis cornuta, with body

r Li . di d

1.€., both galea Cee) tnd aome (E) fere o nce De o e

and the labium (/.) has its free Z4., larval head wall; Zmd., larval

margin more emarginate and less larval Utam: lant., larval antenna

truncate or blunt, the palpi (/i.p.) 4. wall of pupal head ; f.#d., pupal

iaining three-segmented. pupal bbinn: p-ant., pupal antenna

688 THE AMERICAN NATURALIST. | [Vor. XXXVI.

If one dissects away the cuticle of the head of an old larva

about to pupate, the pupal mouth parts will be found formed

fairly within the old larval ones, and thus in perfect corre-

spondence with them. Rather it would be truer to say that

they are apparently the transformed larval parts minus the

to-be-shed larval cuticle. This is shown in Fig. 3, in which

the larval cuticle of the right-hand half of the head (including

the whole of the labium) has been dissected away, exposing the

Fic. 5.

Fic. 4.— V l aspect of head of adult male Cor-ydali: ta. lb., labrum; md., mandible;

mx., maxilla; m2z.p., maxillary palpus; ga., galea; Za., lacinia; li., labium; Z., labial

pus.

Fic. 5. — Ventral aspect of head of pu -ydali: ta, the pupal body wall being dis-

sected away on right side (in figure), showing formi

i al

ng imaginal head and appendages.

maxilla; /./i., pupal labium; #.4., pupal body wall

, imaginal mandible; i»x., imaginal maxilla; 7./é., imaginal labium;

ih., body wall of imaginal head.

still soft, unchitinized pupal cuticle, while the left side of the

head is still wholly larval. From the right pupal mandible has

been slipped the larval mandibular sheath, from the right pupal

maxilla has been slipped the larval maxillar sheath, and from

the whole pupal labium has been removed the larval covering.

But the slight changes in outline and character of the pupal

mouth parts are plainly apparent, while the identity of larval and

pupal mandibles, maxillæ, and labium is unmistakable. There

_is yet no apparent difference in the mouth parts of the sexes.

No. 429.] THE MOUTH PARTS OF INSECTS. 689

In the adult (Fig. 4) we find mouth parts still of simple

orthopterous type, with parts plainly homologous with the

various orthopterous parts, and also as plainly with the parts

of its own larva

and pupa; but in

the male the

familiar but ex-

traordinary modi-

fication of the

mandibles, con- A

verting them

from biting and ric. Mouth parts of larva of Anatis 15-punctata. md., mandi-

mast i catin g "s oe ae E veh ee lobe; »x.P., XS Mg icon:

organs into a pair

of long, non-dentate, pointed, clasping organs (md.) for holding

the female, attracts our special attention. But these organs

are certainly mandibles; the maxilla (zr. and labium (/.),

changed slightly to be still more thoroughly orthopterous in

type, are in their own places, and no other mandibulate organs

except the claspers are present. In the female the imaginal

mandibles are of usual biting dentate type. To make sure of

the mandibulate character of the long claspers we have but

to dissect the head of an old pupa, as shown in Fig. 5. In

this figure the pupal cuti-

cle has been removed from

the right-hand half of the

head, while left intact on

the left side. Removing

the pupal labial cuticle, the

imaginal labium, practi-

cally identical with: the

pupal one, is exposed, with

Fic. 7.— Mouth parts of adult A za£is T,

5-Punctata.

md, mandible; mx., maxilla; mrj., maxillary be palpi shortened by

cri n » galea; Za., lacinia; Zi., labium; Z^, — « telescoping” but ready

to expand to full length;

within the pupal maxillar sheath the imaginal maxilla in its now

thoroughly orthopterous character is found, and within the

comparatively short, strongly dentate, pupal mandible is found,

690 THE AMERICAN NATURALIST. (VoL. XXXVI.

strongly “telescoped,” the strange adult mandible, with its

lack of dentation, its pointed tip, and its great length (easily

attained by extension of the longitudinally compressed organ

as discovered within the pupal sheath). Thus the transforma-

tion of larval parts into pupal, and of pupal into imaginal, is

obvious, and the homologies between larval and imaginal parts

are firmly founded on ontogenic basis.

COLEOPTERA.

The Coleoptera, like the Neuroptera, have biting mouth

parts in both larval and imaginal stages, but the differences

are usually greater, and the general metamorphosis is on the

whole more radical.

Anatis 15-punctata (Figs. 6-8). — The accompanying figures

made from a study of the mouth parts of Anatis 15-punctaia

illustrate the relations between larval and imaginal mouth

parts of a member of the order. The larva (Fig. 6) have

strongly chitinized, sharp-toothed mandibles (md.), maxilla

(mx.) with single terminal lobe (mx./.), rather large four-

segmented palpus (mzx.f.), and fleshy liplike labium (Z.), with

fused terminal lobes and short one-segmented palpus (Z-)

inserted on a segment-like projection. In the adult (Fig. 7)

the mandibles (md.) are shorter and heavier, the maxillae (zx.)

have both terminal lobes, galea (ga.) and lacinia (/.), distinct,

and four-segmented palpi (mr.p.), the distal segment being

much broader than the others. The labium (//.) is rather elon-

gate, with distinct basal sclerites (submentum and mentum),

fused terminal lobes, and short three-segmented palpi (Z.).

The small size of the larval head precludes such dissections

as were easily made in the case of Corydalis, and the thick-

ness and opacity of the chitinized cuticle of the head makes

it impossible to clear specimens and study the forming imagi-

nal head within, a method very successfully used in the cases

of the honeybee and digger wasp (see postea). The develop-

ment of the imaginal head and mouth parts had to be studied

by means of sections, and here again the firmness of the head

wall offered a serious obstacle to satisfactory work. I have

No. 429] | THE MOUTH PARTS OF INSECTS. 691

been able, however, to get series showing plainly the later

steps of the development of the imaginal parts within the head

of old larva. The developing imaginal parts, their definitive

outlines already so strongly indicated as to make them recogniz-

able (apart from their position), lie within the corresponding

parts of the larval head (Fig. 8), imaginal mandibles with their

tips within the larval mandibles,

imaginal maxilla with their two

terminal lobes lying partly within

and corresponding to the single

terminal lobe of the larva, and

imaginal palpi lying almost wholly

within the larval palpi, and finally

imaginal labium lying in the base

of the larval labium. All of the

forming imaginal parts are plainly

seen to be folds or evaginations of

the forming imaginal derm layer,

which shows in sections as a con-

tinuous broad cellular line lying

just underneath the larval integ- ilary palpus; ZZ, larval labium; 24.

ument. imaginal labium

Thus in Anatis we have practically the same conditions of

development of the imaginal mouth parts within, and corre-

sponding to, the larval mouth parts as we found in Corydalis.

LEPIDOPTERA.

Among the Lepidoptera we find a great range in degree of

specialization of the mouth parts. In Eriocephala and Microp-

teryx, as described by Walter! and myself,? the mouth parts

are really of the biting type, the mandibles being short, heavy,

and dentate, true jaws, the maxillae showing a cardo, stipes,

short galea, and lacinia, and long six-segmented palpus, and

the labium being liplike, with plainly distinguishable submentum

! Walter, A. Beiträge zur Morphologie der Schmetterlinge, Jenaische Zeitschr.

Í- Naturwiss., vol. ioe (1885), pp. 751-807

2 Kellogg, V. L. The Berrea Parts of the Lepidoptera, Amer: Nat., vol. v

(1895), pp. 546-556, PI. X

692 THE AMERICAN NATURALIST. | [Vor. XXXVI.

and mentum and prominent three-segmented palpi. But in

all the Lepidoptera above the Eriocephalide, Micropterygide,

and Tineidze, from a considerable to a very pronounced speciali-

zation is present, manifested

: 1 | x Wh E bya complete reduction of

AS LAWN WE DN the mandibles, by the reduc-

2 Sad i) IN tion of the labium to a small

Gi V Sup LY rigid plate on the ventral side

d ii NS ZAA of the mouth bearing the per-

nx € sisting three-segmented palpi,

Fic. 9. — Frontal aspect of head of imago of and by s remarkable modifica-

oaa Iconian wa asilia, tion Of the maxillæe whereby

Zi.~., labial palpus. a

the galeæ (or laciniæ) are pro-

longed, grooved on their inner surfaces, and apposed to form

the familiar sucking proboscis, while the other parts of the max-

illæ are reduced and fused to form a rigid supporting base for

the proboscis. In numerous moths no food is taken in the

adult condition, and here the proboscis itself is reduced slightly

or much, even to complete atrophy, and in extreme cases there

is no mouth opening at all.

Notolophus leucostigma (Figs. 9-12).

— In the white-marked tussock moth,

Notolophus leucostigma, the mouth parts

(Fig. 9) of the adult, although function-

less, or at least apparently incapable of

taking food, show all the usual parts

peculiar to the typical specialized lepi-

dopterous mouth. The labium isa small

fixed plate, forming part of the ventral Fic. ro. — Frontal aspect of head

wall of the head and bearing the conspic- a wah T^ —— jud

uous hairy three-segmented palpi (/f.) ; mandible; mæ., maxilla; a^

2 i maxillary palpus; /., labium.

the maxillze (wzx.) are simply two slender

tapering processes, the halves of the usual proboscis, but in

this case not applied to each other and hence not forming à

sucking tube ; the maxillary palpi are wholly reduced, and the

mandibles entirely wanting.

In the caterpillar (Fig. 10) the biting mouth parts common

to lepidopterous larvæ are present, with full complement of

No. 429.] THE MOUTH PARTS OF INSECTS. 693

distinct and readily recognizable mandibles (zzd.), maxillze (zzx.)

with short but distinct three-segmented palpi (mx.f.), and

labium (/.) with very small ex-articulate palpi.

If an old larva, nearly ready to pupate, be taken, and its

head dissected, as illustrated in Fig. 11, it will be found that

underneath, or within, the larval labium,

or labial cuticle, will be found the form-

ing imaginal labial palpi ; within the lar-

val maxilla will be found the forming

imaginal maxilla, while within the larval

mandible will be found nothing at all.

In Fig. 11 the larval cuticle of the left

side of the head has been dissected away,

showing this correspondence between

larval and imaginal parts; the larval max-

illary sheath has been slipped off of the

forming imaginal maxillar process, while

on that part of the forming imaginal head

from which the larval mandible was taken

there is not a trace even of a forming

organ. Fig. 12

shows the entirely

dissected-out

pupal (equals subimaginal head), with the

already unmistakably recognizable imagi-

nal mouth parts.

Thus in this representative of the

Lepidoptera we find the imaginal mouth

parts developing in perfect correspond-

ence with the larval parts, imaginal

maxillae within larval maxilla, imaginal

labium in larval labium, and within the

well-developed larval mandibles nothing, with a corresponding

- total absence of mandibles in the fully developed moth. By

sectioning the heads of old larvz, it is readily perceivable that

these developing imaginal mouth parts lying within and corre-

sponding to the various larval parts are evaginations of the new

or imaginal derm which forms a continuous layer underneath the

i.7:x., imaginal maxilla

maxilla

labial palpus; ant., Koi

694 THE AMERICAN NATURALIST. [Vor. XXXVI.

larval integument. Similarly, it is apparent that the imaginal

-antennze and compound eyes are in the one case evaginations and

in the other simply modified portions of this imaginal derm ;

and although I have not made cuttings of a complete series

of heads from young to oldest larvae, enough of the younger

stages have been studied to show the simple dermal origin of

al these parts by a continuous process of evagination and .

modification. We are sufficiently acquainted with the origin

and mode of development of the legs and wings of insects

from histoblasts to recognize in these histoblasts, or develop-

mental centers, simple invaginations of the derm, which later

become evaginations. Whether an organ, as wing, leg, antenna,

or mouth part, shall begin as an invagination or an evagination

of the derm is chiefly a matter of mechanical necessity or ease,

and of degree of radicalness in the metamorphosis. In either

case the ultimate origin, that of being simply a particular

portion or area of derm, is the same; the invagination must

become an evagination; the difference lies in the mechanical

factors of the developmental process.

HYMENOPTERA.

In the order Hymenoptera there is to be found, as in the

Lepidoptera, a wide range of degree of specialization of the

mouth parts, varying from the biting, orthopterous mouth of

the sawflies to the highly modified sucking mouth of the honey-

bee ; but throughout the order the mandibles persist in plainly

jawlike character, and are always recognizable landmarks in

mouth-part dissections. The only questions in the homology-

interpretation occur in those cases where the labium and

maxillae are much modified and more or less completely fused

or bound together. But these questions are not very serious;

entomologists are fairly agreed, on a basis of comparative

anatomical study, on the interpretation of the homologies of

the hymenopterous mouth parts. But the results of a study of

the post-embryonic development of the mouth parts, 7.¢., the

development of the imaginal mouth parts, undertaken by one

of my students, Mr. M. H. Spaulding, illuminate too beautifully

No. 429] | THE MOUTH PARTS OF INSECTS. 695

and effectively the whole study of the development of imagi-

nal mouth parts in holometabolous insects to be overlooked

because of the lack of any crying need for an ontogenic con-

firmation of the hymenopterous homologies. Mr. Spaulding

has been admirably successful in so clearing and staining the

heads of variously aged larve of the honeybee and of a digger

wasp, Ammophila s., that the developing imaginal head

within the larval integument may be as easily studied as the

exterior of the larval head itself. The bee and wasp larve, it

will be recalled, are both “inside feeders,” z.e., lie during their

life enclosed in a protecting cell, in one case of wax, in the

other of hardened mud, and thus may and do dispense with the

heavily chitinized opaque head cuticle common to exposed

insect larvae. And both larva have full complements of mouth

parts, namely, mandibles, maxilla, and labium, — a condition

not common to all larvae in those two orders, Hymenoptera

and Diptera, in which the post-embryonic

metamorphosis is most radical. This con-

dition is a necessary one for the determi-

nation of the relations of the imaginal to

the larval parts.

Ammophila sp. (Figs. 13-15). — The

larval mouth parts (Fig. 13) consist of

well chitinized crushing mandibles (sd.),

short fleshy maxillze (mr.) with very small

one-segmented palpus (mr.p.) and smaller ^ afar o digger vam

terminal lobe (mzx./.), and short liplike 4 — la sp. md.,

. ae MeL M s dame A,

labium (Z) with pair of very small one- di: all bike’ med,

segmented palpi (/L?.). The adult wasp ppap cer —

also has a complete complement of mouth

parts (Fig. r4), all very elongate and slender, the mandibles

(md.) heavily chitinized and toothed, the maxille (mr.) long

and slender with distinct cardo and stipes, five-segmented palpus

(mx.p.), and simple terminal lobe composed of the fused galea

and lacinia, and the labium (/7.) also long and narrow with fused

submentum and mentum, four-segmented palpi (/7.f.), slender

ligula formed of the fused glossz ( gl), and distinct slender para-

glossae (p.¢.) less than half as long as the fused glossae.

696 THE AMERICAN NATURALIST. | [Vor. XXXVI.

It is obvious that these long slender imaginal mouth parts

cannot be contained within the very much shorter and alto-

gether smaller larval parts. As a matter of fact, the whole

imaginal head is for simple mechanical reasons forced to lie

during its development chiefly in the anterior larval thoracic

segment, the anterior portions, including the antenna and

mouth parts, projecting forward into the larval head capsule.

But still there is indicated perfectly the correspondence

between particular imaginal parts and particular larval parts

Fic. 14.

Fic. 15.

Fic. 14. — Mouth parts of adult digger wasp, A mmophila sp. aan mer and labial palpus

. left side, in figure, not drawn). szd., mandible; zx., m .p., maxillary palpus;

sies maxillary lobe; 77. labium; /2.Z., labial ET. pe pvc pg, paraglossa ;

Fie. 15. cam a old larva of digger wasp, Ammophila sp., cleared to show forming imaginal

à within. /.c., larval head wall; Z4, forming imaginal head; z.e., imaginal eye;

z.ant., imaginal antenna; Zmed., larval mandible; 7.»:d., imaginal mandible; Z.»x., larval

maxilla ; erage 2 tmx. Ż., imaginal er palpus; ZJj., larval labium;

i.li., imaginal labium; zJZ5., ginal labial palpus

by the fact that the projecting tips of the elongate imaginal

parts penetrate or lie within the short larval parts. This is

shown clearly in the cleared and stained heads prepared by Mr.

Spaulding, as well as in series of sections. Fig. 15 is drawn

with camera lucida from one of the whole head preparations,

and, as indicated by the lettering, those parts of the imago

which we have, on the basis of comparative anatomy, assumed

to compose the labium, do project into and correspond with

the larval labium; the case is similar with maxilla and man-

dibles. But, in origin, these imaginal mouth parts arise as

No. 429.] THE MOUTH PARTS OF INSECTS. 697

dermal modifications and outgrowths which for simple demands

of space become far removed from the larval mouth parts, the

bases of the developing imaginal parts lying, indeed, in late

larval life in the first thoracic larval seg-

ment. But in earlier larval life the begin-

ning imaginal parts lie almost wholly

within the larval parts, and no one study-

ing the series of whole head preparations

and of sections can fail to be convinced

of the certain correspondence and homol-

ogy between larval and imaginal parts,

although there may be said to be no perfect

transformation or development of the one

into the other, the evidence being that of a

correspondence in position on the head and of part for part.

Apis mellifica (Figs. 16—18). — The beautiful series of cleared

and stained heads of honeybee larve of different ages, and the

series of sections of similar heads pre-

pared by Mr. Spaulding, show a condition

in the development of the imaginal mouth

parts of the bee wholly identical with

that just shown for the digger wasp.

The larval mouth parts (Fig. 16) are very

weakly chitinized, but are complete and

readily distinguishable. They resemble

in general the mouth parts of the digger

wasp larva, but are smaller, weaker, and

the short fleshy maxilla bears only the

minute one-segmented palpus, having no

tiny lobe as in the wasp maxilla. The p. ee oe sak

imaginal mouth parts (Fig. 17) of the bee, honeybee, Apis mellifica. md.,

familiar to all entomologists, are com- maxillary palpus: dpi

posed of horny, trowel-like mandibles 1277 lobe; st., stipes; cd., cardo;

(md.), long maxillae (mx.) with cardo (cd), m., mentum; Ag. paraglossa ;

stipes (s/.), small one-segmented palpus *^9**: 4, bi palpus.

(mx.p.), and with galea and lacinia fused to form a single

flattened, pointed, bladelike terminal lobe (mzx./.), and of

labium (/.) with long, tapering subcylindrical ligula formed

bium; Z., labial palpus.

698 THE AMERICAN NATURALIST. [Vor. XXXVI.

of the fused glossa (g/.), short but distinct flaplike paraglossae

(pg.), three-segmented palpi (/.f.) borne on a long palpiger,

and at the base a distinct mentum (s.) and submentum (sm.).

| As with the digger wasp,

the developing head of the

imago, with its long

antennz and mouth parts,

demands more space than

is afforded within the lar-

val head segment, so that

it is crowded backward and

occupies part of the first

and second larval thoracic

segments. But the form-

ing imaginal mouth parts

are to be found with their

tips projecting into the

corresponding larval parts,

imd., imaginal mandible; Zx., larval maxilla;

i.mx., imaginal maxilla; ZZi., larval labium; ZZ, as shown in Fig. I8. The

imaginal labium avt

conditions of the develop-

ment of the imaginal parts, and of their perfect correspondence

with the larval parts, are wholly like those already explained

for the digger wasp.

DIPTERA. `

In the case of the Diptera, — and it is here that the neces-

sity of ontogenetic study is most important, indispensable

indeed, for the determination of the homologies, — we have,

as in the Hymenoptera and Lepidoptera, a great variety of

mouth-part conditions culminating in the extreme specialization

characteristic of the muscid forms. In most Diptera it is

obvious that a total reduction of at least one pair of the buccal

appendages has occurred, with a large reduction and complete

modification of the remaining parts. From a considerable

study of the anatomy of the fully developed mouth parts in a

long series of dipterous forms, including representatives of all

except one (the Ornephilidze) of the nematocerous families, —

No. 429.] THE MOUTH PARTS OF INSECTS. 699

those families by common agreement held to constitute the

more generalized portion of the order, —I came to the conclu-

sion that the old and most widely, if perhaps uncritically,

accepted interpretation of the homologies of the dipterous one

is the true one. This interpretation homologizes the labella-

bearing proboscis common to all the more specialized flies with

the labium of other insects, finds the maxillae represented in

these specialized forms chiefly or only by a pair of palpi, and

finds the mandibles wholly wanting in all but the females of a

few families. In the case of most of the nematocerous families

the labium retains a truly labiumlike character and has not

developed the pseudotrachez-bearing labella, while the max-

illa are represented by a well-developed bladelike terminal

lobe as well as by the palpi. The mandibles when present are

of the character of elongate blades or stylets, never of the

Character of true crushing or biting jaws. The structural

character of the mouth in each of the nematocerous families

is described and illustrated in my series of papers (1899) in

Psyche, previously referred to.

But several interpretations of the homologies of the mouth

` parts widely at variance with the above have been offered. In

these various interpretations the possession of mandibles by

any flies at all is denied; the so-called labium is considered to

be composed of modified parts of the maxilla, and the so-called

maxillae are believed to be parts of the labium ; in fact, most of

the possible changes which an active speculation could invent

have been rung on the theme. Nor are these interpretations

based on mere speculation ; they are the results, in several cases,

of prolonged and disinterested examination of considerable

series of specimens.

In the face of such differences of opinion, and with the

apparent limits of the method of the comparative study of the

fully developed mouth parts of various members of the order

reached, it becomes imperative to seek the clue to these lost

homologies in the facts of development. And this is really the

first object of this present study. Can the homológies of the

dipterous mouth parts be discovered by the study of the devel.

opment of the parts?

700 THE AMERICAN NATURALIST. [VoL. XXXVI.

For a complete developmental study of the mouth parts of

any dipteron it would be necessary to begin with the budding

appendages of the head segments in early embryonic life, to

trace the development of these appendages to their definitive

form in the hatched larva, and finally to follow the transforma-

tion, if it occurs, of these larval parts into the ultimate imaginal

ones. Asa matter of fact, such actual transformation does not

occur, so that the study of the postembryonic development of

the mouth parts con-

sists of noting the ec-

dysis of the larval parts

y and determining the

ontogenic relations of

the new imaginal parts

to the old larval ones.

As for the embry-

onic development of the

mouth parts, —7.e., the

development from bud-

ding appendages to

definitive larval parts, -

— that has been done

for several Diptera,

and in particular by

Metschnikov for Simu-

+h d. 1 rem d

Fic. 1g. — T bryonic stages i p

imulium sp. (after Metschnikov): younger stage at

left, older at right; in upper row whole embryos from lium ; One of the two

lateral aspect, in f l f heads of

flies whose postembry-

onic development I

shall describe. These embryonic studies make certain the

homologies of the larval parts; in those flies like Simulium,

whose larvæ are provided with a biting mouth with full com-

plement of parts, it is easy to note plainly the development of

mandibles, maxillæ, and labium from the successive pairs

of budding head appendages (Fig. 19), and thus to homologize

these parts certainly with the mandibles, maxillæ, and labium

of adult insects of incomplete metamorphosis. There remains

to determine the relations of the larval mouth parts of Simu-

lium with its very different imaginal mouth parts.

stages. £4., ephalic lobes » anf. , antenna ; snd.,

mandible; mr., illa; Zi, labium.

No. 429.] THE MOUTH PARTS OF INSECTS. 7901

In selecting flies for the study of the postembryonic devel-

opment of the mouth parts I have chosen two which in the

me condition possess all the parts possessed by any fly,

and these parts in as generalized

condition as is to be found in the

order, and which also possess in

the larval stage a similarly full

complement of mouth parts. Such

larvae as those of the Muscidz,

with their problematical hooks and

lack of other parts, and such imag-

pow 20. — Mouth aed * adult — i à à

, female. Zep. ines as the muscid flies, with no

th bona Km. nae ar parts left except proboscis and

mk palpus; Z., labium; Az, para- maxillary palpi, are impossible for

3 the determination of the relation

between larval and imaginal parts. From the mouth parts of

the imaginal Simulium and of other nematocerous forms it is

not difficult to trace the evolution to the specialized muscid

conditions, and if the mouth

parts of Simulium and similarly

equipped flies can be inter-

preted, the various members of

the dipterous series culminating

in the muscids can. So in

Simulium and Blepharocera I

have found suitable forms for

study; both with females pos-

sessing the so-called mandibles,

both with maxille and labium

well developed in both sexes,

and both with larvae equipped

with biting mouths with unmis-

takable mandibles, maxillae, and s- . md, mandible; sa, mas mel

: s : maxillary lobe; #x.f., maxillary palpus ;

labia, and in one case, that of z., labrum; eA, epipharynx; Zi, labium;

Simulium, with the embryonic %4» bypopharax.

development of the larval mouth parts fully traced and the

homologies certainly! determined.

1 Metschnikov, E Nut logische Studien an Insekten, Zeifschr. J wiss. Zool,

P eo

702 THE AMERICAN NATURALIST. [Vow. XXXVI.

Simulium sp. (Figs. 20-23). — In the female imago! the

mouth parts (Fig. 20) consist of a short liplike labium (Z.) com-

posed of a short basal sclerite and three terminal lobes, being

the two large paraglossze (fg.) and a median short membranous

lobe, the fused glossz ; of a pair of maxillae (»x.), each consist-

ing of a basal sclerite, a long five-segmented palpus (mx.p.), and

a single pointed, bladelike terminal lobe (;zx./.) reaching nearly

to the end of the third palpar segment, serrate on its inner

margin at the tip and better developed than in most Nema-

tocera; and of a pair of short mandibles (md.), broad, thin, and

weakly chitinized. As in other nematocerous flies, there is a

well-developed labrum-epipharynx (/ef.) and an elongate flat-

tened hypopharynx (/yp.). In the males the mandibles are

wanting. |

In the larva (Fig. 21) the mouth is of the biting type, with

short-toothed and heavy mandibles (md.), short, jawlike maxilla

(mx.) with distinct one-segmented palpus (sx. .), and a small,

strongly chitinized labium (Z.) or labial plate. In addition,

labrum (/é.), epipharynx (z7.), and hypopharynx (/yf.) are all well

developed.

The head of the larva having a thoroughly opaque, strongly

chitinized cuticle, it was impossible to clear whole heads suffi-

ciently to make visible the developing imaginal head and its

parts, so that the method of sections had to be relied on to

reveal the internal conditions. These sections of heads of

larvae of various ages show plainly that the general method of

development of the imaginal parts within the larval head, and

the correspondence between forming imaginal parts and the

corresponding larval parts already noted in the other orders of

holometabolous insects, hold good in the Diptera. Fig. 2?

shows in sagittal longitudinal section the forming imaginal

head parts within the larval head. This section shows partic-

ularly well the relation of the forming imaginal antenna to the

vol. xvi, 1866 ; embryonic development of mouth parts of Simulium described on

PP. 392-421.

In the adult mouth I shall assign to the various parts those names

which, from my earlier study of the comparative anatomy, seem correctly used,

the use of which is confirmed by the results of this ontogenetic study.

No. 429.] THE MOUTH PARTS OF INSECTS. 793

larval antenna. In the larva the antenna are very small com-

pared with their size in the imago, and the imaginal antenna is

thus forced, in its development, to occupy a region in the larval

head not included in the larval antenna. But the tip of the

imaginal organ lies fairly within the larval organ, thus indi-

cating by correspondence in position, what is plainly obvious

from anatomical consideration, the homology between the larval

and imaginal organs. Similarly the forming imaginal mouth

parts are to be found in unmistakable correspondence or

homologous relation with the larval parts: By tracing the

development of the parts, marked in Fig. 22 as the forming

—

SINUS RTT >

5 : "app.

Fic. 22. — S 1 1 of old 1 (y f 1

head parts within. Zc., larval head wall; z.d., iiL derm ; Za. , larval anten enna; t.a.,

imaginal antenna ; z.e., imaginal eye; Zmd., terval mandible; Z2 €. imaginal mandible;

lmz., larval maxilla; z.1x., imaginal maxilla ; /./., larval labium; 2./7., imaginal labiu:

imaginal mouth parts, through larvae of successively older ages

to pupation and the achievement of the definitive imaginal con-

dition of these parts, it is certain that the parts marked respec-

tively imaginal mandible, imaginal maxillz, and imaginal labium,

lying respectively in the larval mandibles, maxilla, and labium

(with homologies firmly based on ontogenic basis), do develop

into those definitive imaginal parts named mandibles, maxille,

and labium in Fig. 20, illustrating a dissection of the mouth

Parts in a female adult Simulium. F ig. 23, a horizontal, frontal

section through the head of a Simulium larva, shows also the

forming imaginal maxilla and mandibles within corresponding

parts,

704 THE AMERICAN NATURALIST. [VoL. XXXVI.

Bibiocephala doanei! Kellogg (Figs. 24-26). — The Blepha-

roceride, or net-winged midges, agree with the Simulidz, or

black flies, in having the females equipped with mandibles,

which in the Blepharoceridze are well developed as long, slender,

bladelike saws (see Fig. 24, md.) used to lacerate the bodies (as

I have observed) of the tiny midges caught as prey by the

bloodthirsty females. In addition the adult females have max-

illae (Fig. 24, mx.) with well-developed lobe (mx./.) and long

five-segmented palpus (mzx..), and a labium (/Z) consisting of

strong elongate basal sclerite which presents indications of a

line of fusion of submentum and mentum, and a pair of free

fleshy terminal lobes, the paraglosse (pfg.). The males are

equipped like the females except for the mandibles.

The larva of Bibiocephala doanez has a biting mouth (Fig. 25)

composed of short, stout, crushing mandibles (md.), weaker

jawlike maxillae (#x.) without palpi, and a soft liplike labium

(lij. In addition there are well-developed labrum-epipharynx

(Z.ep.) and hypopharynx (Zy5.).

The development of the imaginal head shows the same phe-

nomena as in Simulium. In Fig. 26, from a vertical transverse

section through the head of an old larva, the derm of the form-

ing imaginal head is plainly seen in continuous layer, modified

at z.c. to produce the developing compound eyes and at mx. and

md. the forming imaginal mandibles and maxilla. In this

section the imaginal parts of the maxille visible are the

forming palpi, and their definitive, long, segmented condition

is plainly to be seen in these telescoped organs tucked tightly

inside the larval maxilla. The forming mandibles do not yet

show their definitive character, but in tracing these organs

through a series of older larva: the gradual taking-on of the

slender sawlike character is manifest. The series of Blepha-

rocera preparations which I have show even more plainly than

the Simulium preparations the perfect correspondence and

* box-in-box " sort of relation which exists between the larval

1 This blepharocerid fly was described by me in Psyche, vol. ix (April, 1900)»

PP. 39-41, 2 figs., under the name Lifoneura doanei. In a recent revision of the

North American Blepharoceridz, now in press, I refer this species to the genus

Bibiocephala.

No. 429.] THE MOUTH PARTS OF INSECTS. 705

FIG. 25.

Fic. 24.

Fic. 23. — Frontal "iie through the head of old larva of Simulium sp., showing forming

imaginal parts. imaginal derm; /.d., larval mandible; imd.,

imaginal man ean dai larval maxi íi.mzx., imaginal maxilla; Z.»«&x.5., evil wack

lary palpus; Tad id ypopharyn

Fic. 24. — Mouth parts of adult Bibiocephala doanei, female; md., mandible;

axilla;

mx.d., maxillary lobe; mx a ., maxi palpus; /2., Bian; A. a. a ep., la-

brum-epipharynx; 4y., hypopharynx.

Fic. 25. — Mouth pa of larva of PEAS doanei. md., mandible; mæ., maxilla; ZZ,

PA, qiiia pipharynx; yypopharynx.

Fic, 26. — Frontal section, through head of old larva of Bibiocephala doanei, showing forming

imaginal head ithin. Zc., larval head wall; z.d., imaginal derm; z.e., imaginal eye;

Zmd., larval — t.md., imaginal mandible; Z.»x., larval maxilla; z.»x.., imagi-

nal maxillary palpus

706 THE AMERICAN NATURALIST.

mouth parts, of whose homologies no doubt can exist, and the

forming imaginal parts, of whose homologies, in definitive con-

dition, I thought myself long ago able to speak confidently on

a basis of comparative anatomical study, but of which now on

a basis of ontogenetic study I am simply without doubt.

STANFORD UNIVERSITY, CALIFORNIA.

January, 1902

AN ANALYTICAL KEY TO THE GENERA OF THE

FAMILY FORMICIDZE, FOR THE IDENTI-

FICATION OF THE WORKERS.

CARLO EMERY.

(Translated by WILLIAM MORTON WHEELER.!)

SINCE the publication by Mayr, in 1865, of the volume on

the Formicidz of the voyage of the JVovaza, no comprehensive

work has appeared which could be of service in the identifica-

tion of the genera of this family. The number of these genera

has increased considerably in the mean time, and the defini-

tions formerly given by Mayr for a number of them are no

longer exact, as their characters have been modified by the

discovery of new species. Moreover, certain genera have been

subdivided, others fused together; and all of this is to be

found scattered about in a host of detached publications, so

that it is almost impossible for any one who is not a consum-

mate specialist to find his way about in the labyrinth.

I originally began, for my own personal use, to construct

analytical tables for the workers of the subfamilies Myrmicinz

and Ponerinz ; but I now believe that I would render a serv-

ice to entomologists by publishing these tables, after having

revised and completed them. I have added tables of the

genera of the other subfamilies, together with a table of the

characters of the subfamilies themselves, likewise in analytical

form

l For the present authorized translation Professor Emery has carefully revised

the tables of the Myrmicinz, Dolichoderine, and Camponotine, of his “Clef

Analytique des Genres de la Famille des Formicides, pour la Determination des

neutres ” (Ann. Soc. Entomol. Belgique, tome xl (1896), pp. 172-189), and has, more-

Over, permitted me to translate the as yet unpublished German table for the

Dorylinz and Ponerine which he has been preparing for Das Thierreich. The

WOrk has thus been brought up to date and cannot fail to be of great service to

myrmecologists the world over. — W. M. WHEELER.

707

708 THE AMERICAN NATURALIST. | [Vor. XXXVI.

I trust that these tables will facilitate the labor of identifica-

tion and enable those who are beginning to study the exotic

ants to find their way more easily. I am well aware of the

fact that one may find one’s self in doubt at certain bifurcations

of the path. How is one to know, e.g., whether the worker is

dimorphic, when one has only a single specimen of the species?

I have made no use of such characters except when I had noth-

ing better to present, and then I have tried to reinforce them

as much as possible with accessory characters. Sometimes I

have cited characters peculiar to the males and females, as these

are often the most important in distinguishing certain genera,

the workers of which present only feeble or insignificant differ-

ences. Nevertheless, I decline to attempt for the present an

analysis of the sexual forms, which are still too imperfectly

known.

In the enumeration of the segments of the abdomen I

include the one or two constituting the pedicel, so that the

segment following this, and usually designated by other authors

as the first abdominal, is for me the third in the Myrmicine,

the second in the Camponotinz, etc.; the last visible segment

is, therefore, always the sixth in the females and workers, the

seventh in the males. In the male I designate as “ subgenital

lamina " what is usually, but improperly, called the hypopygium

and is in reality the ventral lamina of the eighth segment.

For the anatomy of the gizzard and the poison apparatus I

would refer the reader to the works of Forel, Dewitz, and

myself.!

Having adopted the dichotomic form for the identification of

the genera, it follows that the order cannot express their

natural affinities; but this can lead to no inconvenience in à

1 Forel, A. Etudes paps age en 1878. Anatomie du Gésier des

Fourmis, Zu Soc. Vaudoise Sc. Nat., vol. xv (1878), pp- 339-362) Fl. XXIII ;

Der Giftapparat und die Analdruesen yes Ameisen, Zeitschr. f. wiss. Zool., Bd. xxx,

Suppl. (1878), pp. 28-68, Taf. ITI-IV.

| witz, H. Ueber Bau und Entwickelung des Stachels der Ameisen, Zeitschr.

Jf. wiss. Zool., Bd. xxviii (1877), pp- 527-556, Taf. I.

Emery, C. Ueber den sogenannten ers einiger Ameisen, Zeitschr. f

wiss. Zool., Bd. xlvi (1888), pp. 378-412, Taf. XXVII-X

In the Loss will be found figures of all the forms of w gizzard in the

No. 429.] THE FAMILY OF FORMICID.E. 709

work the aim of which is essentially practical. I have marked

with an asterisk the names of the genera represented in the

Palearctic fauna.!

CHARACTERS OF THE SUBFAMILIES.

I. Cloacal orifice in the shape of a slit; sting well developed or rudimental.

$ Sting developed, though sometimes very small, but capable neverthe-

less of being exserted from the abdomen. The first two segments of

the abdomen usually modified, either forming together a two-jointed

pedicel, or the first alone (petiole) forming the pedicel, the second

(postpetiole) being merely constricted posteriorly and articulating

ith a spheroidal surface of the third segment, which is usually

transversely striated (stridulatory organ) ; rarely the second segment

is not appreciably modified.

T Nymphs usually enveloped in a cocoon; pedicel consisting of a

single segment, more rarely of two, but in this case the frontal

Carine are very close to each other and do not cover the inser-

tions of the antenne (Doryline) or the mandibles are linear

and denticulate (Myrmecia).

a. Frontal carinz very close to each other, almost vertical, not at

all covering the antennal insertions (except Acanthostichus) ;

abdominal pedicel of one or two segments. In the male the

genitalia are completely retractile and the subgenital lamina

is usually (perhaps always) furcate; cerci absent

Ist subfamily, DoRYLINE

b. Frontal carinze separated or close together; in the latter case

they are anteriorly to form an oblique or horizontal

lamina, covering in part the insertion of the antenne;

abdominal sediel of a single segment (except Myrmecia) ;

copulatory organs of the male incompletely retractile; sub-

enital lamina never furcate (except in Paraponera); cerci

nearly always present — . . 2d subfamily, PONERIN.E

tt bene naked; pedicel of two segments; rarely the postpetiole

attached to the following segment over its whole extent.

F rontal carinz usually separated from each other. In the male

the copulatory organs are almost always exserted (being entirely

retractile only in certain genera of the group Solenopsidii);

cerci nearly always present (except Anergates)

3d subfamily, MyRMICINE

! In addition, I have indicated by heavy type, in the translation, the names of

all genera known to be represented in the ant fauna of the United States. —

W. M. WHEELE

710 THE AMERICAN NATURALIST. [Vor. XXXVI.

§§ Sting rudimentary (except Aneuretus) ; abdominal pedicel consisting

of a single segment; no constriction between the second and third

segments of the abdomen; the poison glands are often rudimental

and there are anal este which secrete an aromatic product of

characteristic odor (Tapinoma-odor). Nymphs without a cocoon

4th subfamily, DoLICHODERIN

II. Cloacal orifice round, terminal, surrounded by a fringe of hairs; sting

transformed into a sustentacular apparatus for the orifice of the

poison vesicle, whic as a peculiar structure — called by Forel

“pulviniferous vesicle” (vessie à coussinet). Abdominal pedicel

consisting of a single segment; no constriction between the second

and third segments. Nymphs rarely naked, most frequently enclosed

in a cocoon. Male genitalia not retractile

5th subfamily, CAMPONOTINE

Ist and 2d Subfamilies: DORYLINE and PONERINE.

1. Eyes large, three ocelli, mandibles slender, denticulate; postpetiole

campanulate, narrower than the succeeding segment (Australian)

T Fab.

Of a different conformation .

Antenne 6-jointed ; first joint of hind tarsi dilated (Aficun)

Melissotarsus Emery

Antenne with more than six joints; tarsi simple .

Postpetiole hardly wider than the petiole, and much narrower RE the

anterior border of the following segment, and on this account

appearing as the second segment of the abdomen so g

Postpetiole of a different conformation . . . . . . . . . . 6

Antenne 12-jointed Seren: VUE TUI ENTIS ae mas

Antennz 9-10-jointe

Hind legs without spurs, [onis not A E (Asia “Abics. Aus-

tralia) nictus Shuckard

Hind legs with spurs ; fast arapal at sonicated e as a club (Asia,

Australia) . DOM ; . Cerapachys F. Smith

(Ciba Oöceræa and Cysias)

6. Postpetiole shorter than the following segment and somewhat narrower,

but not distinctly separated from the latter; mesoëpinotal suture

obsolete, promesonotal suture distinct ; pygidium 3- ipM antenna

7-12-jointed ; no eyes (Asia, Africa) . P Dorylus Fabr.

Postpetiole as in the preceding; incscépinonl kare distinct, pro-

mesonotal suture obsolete ; antennæ 12-jointed (neotropical)

Cheliomyrmex gu

Of a different conformation .

7. The frontal carinz, which are fused with eb athek d with ihe pdt

form a plate projecting out over the mandibles; the antennz are

inserted close to the anterior margin of this structure . 8

No. 429.] THE FAMILY OF FORMICIDE. Tis

Frontal carinz of a different conformation EON ou

. Antenne 12-jointed ; abdomen stretched out straight (Africa)

Probolomyrmex Mayr

Antenna 9-jointed, tip of the abdomen deflected down and forward

(America, Africa, Australia) . . . . . Discothyrea Roger

Frontal caring very close to each other; antenna inserted very near

the oral margin. Tip of abdomen Meus deflected downward to

Frontal carine of a different conformation, or the tip of the abdomen

not deflected. . "— n

. Clypeus in front projecting in "he middle; petiole odin (America,

Europe, Australia) . . . *Sysphincta Roger

Clypeus not projecting in Goki petiole sce like (America, Europe)

ratium Roger

11. Abdomen straight and constricted behind each segment; pygidium

impressed or furcate (South America, Australia)

oo ie

—

Abdominal segments not constricted

12. Body cylindrical with an elongated bud; usually with one ocala on

the vertex IT : ee ge GR)

Of a different chukemádion pou pss 14

13. Antenne 12-jointed (eciabtenicaly. oe TUN UR Cjlnitosiyivex May

Antenne I11-jointed (Africa) . . Simopone Forel

14. Petiole inserted behind on the EA Herai its whole breadth ;

antennz 12-jointed . . ea KC

Petiole constricted off from the jjoutpétiolé, PAR separate, eo qa

15. Middle and hind tibize without spurs, claws very large, eye well devel-

oped (Australia) . . . Onichomyrmex Lr

Middle and hind tibiz Maid: eye very dal: 16

16. Pygidium with a row of prickles on its lateral badei niie square ;

antennz robust (South America). . . . . Acanthostichus pe

Pygidium without prickles on its border .

17. Very small (at most 214 mm.); funiculus with a Guinn als: -

one spur well developed on the middle and hind —À Vorne America,

Australia) . Re gar

Larger, middle al hind tibiae tach ici two spurs 18

18. Integument shining . . E tea To :

Integument at least in SE opaque, ies nip tured

I9. Antennz very thick, the whole funiculus club-like iis. dao din)

Myopopone Roger

Antennz with filiform funiculus, but slightly thickened towards its tip.

(Australia) Amblyopone Erichson

Mandible bine: at its aoc or Ld a dui dilatation (Madagascar,

ia) . . Mystrium Roger

Mandible poisid a at its itip (Amirin, kanpi hiin Australia)

* Stigmatomma

oger

712 THE AMERICAN NATURALIST. [Vor. XXXVI.

Mandibles inserted on the anterior corners of the head . a

Mandibles inserted in the middle of the anterior border of the edi 55

. Frontal carine approximated and almost perpendicular, not covering

the insertions of the antenna. — deme constricted off

from the following segment . 23

Frontal carinz more or less dilated, pie covering E insertions of the

antennz at least in part; antennz always 12-jointed. . . . 26

Petiole sharply margined laterally; last joint of antenna not particu-

larly large (Asia, Australia, Africa). . . .. Phyracaces Emery

Petiole not margined laterally

24. Last antennal joint much longer ba de: ibam joint; otsdng a

—

one-jointed club (Asia, Australia, Africa) . Cerapachys F. Smith

The last two or three joints of the antenna form an indistinctly sepa-

rated club (India) . . . Lioponera Mayr

25. Frontal carine farther from tah dier du in the sides of the head.

Mandibles with very long thorn-like teeth (South America)

Thaumatomyrmex Mayr

Frontal carine nearer to each other than to the sides of the head;

mandibles of a different conformation . . 26

26. Mandibles slender, when closed first strongly converging, pen directed

straight forward, beak-like, below with a powerful tooth, eye very

large, placed anteriorly (India) . . . . aint Jerdon

Mandibles of a different conformation . 2

27. Middle legs with two simple spurs; hind "- with: a seit nd a pets

tinate spur (America, Africa, Asi ia) . . . Centromyrmex Mayr

Spurs of the middle and hind = Of like structure . i. > 28

28. Claws pectiniform . . . . Ve vici ei wu NT aM. N

Claws not pectiniform 3°

29. Mandibles slender with a fen lange ecth eng the medist bords

(Australia) Prionogenys Emery

Mandibles AGE d soto. or brosder with dentate edges (warm

regions of the whole world) . . . . Leptogenys Roger

. Antennal fovea continued back into a groove sowie bends around the

eye and is capable of enclosing the antennal scape and a portion of

the funiculus (South America) . . . . . Paraponera F. kgs

Of a different conformation .

31. Margin of clypeus denticulate ; MOSS on Leiter side vide. a idi

like projection (India) . ; . . Odontoponera Mayr

Clypeus not denticulate . 32

Frontal carinz not boosdened into ions anieribify; bn sabe ‘slightly

dilated, widely separated from each other ; middle and hind legs with

well-developed median spur ; lateral spur, when present, very small

F rontal carine converging posteriorly and there usually closely approx-

imated. Anteriorly they are dilated to form a horizontal lobe . 39

33- Antennæ with 3-jointed club ; claws simple . M

ion. |

Es]

- 429.] THE FAMILY OF FORMICIDE. 713

Antennz without a differentiated club; claws usually toothed or

split

. Petiole distinctly pedunculate; thorax above with distinct sutures

South America . . Typhlomyrmex Mayr

Petiole not pelina; ; Pass above without sutures (New Guinea)

Rhopalopone Emery

. Third abdominal segment strongly fornicate dorsally, so that its pos-

terior margin is directed downward, or even somewhat forward;.

thoracic dorsum without sutures. . . . . . Se nee, |

Of a different conformation 37

Eye small ; third abdominal — sstonighs deflected (South Ainii

Alfaria Emery

Eyes larger; third abdominal segment but aksi deflected (southern

Asi Stictoponera Mayr

i Paoa suture ohda d or distinct as a deep depression, which

oes not, however, interrupt the sculpture of the integument (South

America) . Ectatomma F. Smith

Promesonotal suture ‘distinct wed sharply c ibn — the sculpture

of the integument . 8

. Antennal fovea elongated DackWarda as a groove ; yiii with Reeth

or spines; hind coxa unarmed (South America, Australia

canthoponera Mayr

Antennal fovea not elongated € epinotum unarmed ; hind

coxa unarmed (Australia) . yti —€— Mayr

Antennal fovea not elongated ; eot eden hind c with a

spine (South America) . nen Mayr

Episternum of mesothorax balloiete out ; : petiole behind with two teeth

or spine (India and Australia) . (ue: eo Diacamma z

Episternum of mesothorax not hefuwed out .

. Medial spur of the middle and hind legs alone developed, the el

spur is lacking, or very small (Trapeziopelta) :

Both spurs of the middle and hind legs well developed . . . . 47

. Integument smooth or sculptured, without pubestence . . 42

Integument delicately sculptured ; at least the abdomen puliseciot 45

. Clypeus in the middle with a slender projecting lobe ; lateral spur small

but distinct (Malasia and daten: . . . . Trapeziopelta Mayr

C igpeos without a aF : ; 43

ae flat and point 4(Africa) b paslidourpieiek Andee

Mandible slender, with. a few teeth on the PER border ^ 44.

Mandible obtuse at the end (Africa) Plectroctena F. Smith

andible pointed, its medial border with two eeth (Ceylon)

Myopias Roger

. e) in front with a needle-shaped process (South America)

Belonopelta Mayr

Clypeus without a point in front. . . peo Gree WE

714 THE AMERICAN NATURALIST. (VoL. XXXVI.

46. Antenna with a 4-jointed club (Asia and Papuasia) Cryptopone Em.

Antenna without a club, or with an indistinctly marked off 5-jointed

club (warm and temperate regions of the whole world) * Ponera Latr.

47. Clypeus flat, separated from the frontal carina by a scarcely percep-

tible suture or not at all; body opaque, with fine gray pubescence

(warm regions of the whole world) . . . . . Platythyrea Mayr

Clypeus separated off by a distinct suture Wee DULL V S

48. Anterior border of clypeus with two teeth . . . . . . . . 49

Clypeus not bidentate . . eo ow Pe

Me]

. Epinotum with two teeth ; settle - fuis (Africa)

Streblognathus Mayr

Epinotum unarmed ; petiole not pointed (South America)

Dinoponera Roger

. Clypeus in the middle with a raised piece E on either side

frica FC — pa

Clypeus arched or carinate .

51. Cheek in front of the eye with a longitudinal taHd 2 0205454 2 ae

Cheek without a carina. . . KM CE M, 53

pe Crewe Genta (ARa s o... n ‘get Mayr

Claws simple (South America) . Neoponera Emery

53. Eye in the middle or behind ihe middle et: thé side of the head

(Africa) , d Forel

Eye in front of the middle of the side of the head . =.

54. Mesoépinotal suture obsolete or not impressed; mesonotum not arth

in profile (warm regions of the whole world) Pachycondyla F. Smith

Mesoépinotal suture impressed ; mesonotum distinctly arched in pro-

file . Euponera Forel

55. Antennal kar EE with Pat iiber behind ‘the frontal carine ;

petiole prolonged above into a thorn-like point (warm regions of the

globe) . : . . . Odontomachus Fabricius

Antennal iind not condideni bebini j6-

Antennal fovea bordered by a carina jail ebd the ejes head

not emarginate posteriorly ; petiole pointed (Ma €

psomyrmex Emery

Antennal fovea without a lateral keel behind - eye; head always

emarginate posteriorly; petiole usually not pointed

* Anochetus Mayr

3d Subfamily : MyRMICINJE.

1. Clypeus not prolonged back between the frontal carina! which are

closely approximated to each other; antenna roce = Pseu-

domyrmii)

rin some species of Pseudomyrma the clypeus seems to be continued back

y between the frontal carinæ, but this AISEA is the equivalent of

the : frontal area; it is often distinct from the clypeus.

. 429.] THE FAMILY OF FORMICIDE. 715

Clypeus almost always prolonged between the frontal carina, which

are more or less separated; in the — case, the antenna are

II-

-jointed

. Clypeus suddenly discendingd in Troet, or as if inflected c or pubtrüicatéd,

usually armed with teeth at the level of this inflection ; rarely it is

uniformly sloping and deeply emarginate at the anterior border

(Africa, Asia, Oceanica) . . Sima Roger

Clypeus neither inflected nor deut. not or duly ieebly emarginate

(America) . . . . Pseudomyrma Guerin

. Antenne 7-jointed, withe&t 3 a distinct club (13-jointed in the male) ;

frontal carinz, as usual, distant from the lateral borders of the head ;

thorax spinose (tribe. Myrmicarii) . Myrmicaria Saunders

Antennz of a different conformation Grien 7- ioiai the last joint is

enlarged or forms part of a differentiated club, or the scape may be

enclosed in a deep groove, or the thorax is without spines)

. Antennal fovea or groove placed at the side of the head; the carina

formed by its dorsal margin (and which does not poreapoíud to the

frontal carina of other ants) passes outside of the eye; posterior

angles of the head pointed or prolonged or denticulate; antennz

I1-jointed in all the sexes yn oup . Cataulacus F. Smith

Of a different conformation

. The antennal fosse terminate behind on thé: dia bi the ek pass

above the eye and are sufficiently deep to conceal the whole antennal

scape ; antennz I I-jointed, without differentiated club ; gizzard a

form, of peculiar structure (tribe Cryptocerii) (America) . 6

nissan] fosse differently placed, or the antennz of a diferent « con-

formation ; gizzard of the usual form . 7

Antennal foveæ approximated in front, P ES male behlüi, not

reaching the sides of the head except at their extremities

Procryptocerus Emery

Antennal fovez covered throughout their length by the lateral border

of the head Cryptocerus Fabricius

Postpetiole aricadated to Abe ind Putas ot y a segment

(tribe Crematogastrii) . Tet atogaster Lund

Postpetiole inserted at the anterior pem e the Ab asi segment . 8

- Head cordiform, emarginate behind, with the posterior angles strongly

rounded and devoid of spines; last joint of antenna very much

smaller than the preceding joint (tribe Dacetii) . . . . . . 9

Head of a different conformation Zo secs d. e x RS

Antennal fovez short; antenne irjeined au S I

Antennal foveæ as long as the sc d a

- Only the last joint of the antennz gum saa the receding joint

Daceton Perty

Last two joints of the antennz longer than the graine g

Acanthognathus Mayr

716 THE AMERICAN NATURALIST. [Vor. XXXVI.

. Antennal fovez shallow, at the medial side of the eye; antenne

5-jointed, the third much elongated . . Mp dito. F. Smith

Of a different conformation . 12

12. Antennal fovez placed at the oval. or medial side of the eyes. . .13

Antennal fovez placed at the lateral side of the eyes . 14

in Antenne Gjointed 44 orn ee Sienai F. Smith

BM MM E ns iby oak out Eprtbus Emery

14. Antenne 6-jointed. . . . . . . . . . + Epopostruma Forel

Antenne 7-G-jointed . . . . « «s s Rhopalothrix Mayr

Antenne I2-jointed . . Ceratobasis F. Smith

15. Antenne II-jointed, nihoni distinct clots or a club consisting of a

single joint (tribe Attii, America) . .

Club of several joints, or the antennz not I pAn 20

Frontal carinæ very close to each other and dilated at ies anterior

extremity ; clypeus not distinctly prolonged between them . 17

Frontal carina Donee embracing the posterior — of the

clypeus 18

. Integument opaque "m even, bristling with tng, bs parts

Apterostigma Mayr

sprig are bristling with tubercles and spines, with hooked and scale-

like hairs . . . Myrmicocrypta F. Smith

18. No erect in on the bodri $ eion idt usually prolonged to the

posterior corners of the head . . . . . Cyphomyrmex Mayr

Body bearing erect hairs . i 19

Integument even, bearing adiy délicate eique haies

Sericomyrmex Mayr

Integument rough, bearing stiff or hooked hairs . . Atta Fabricius

20. The shallow antennal foveæ bordered laterally by an abrupt carina ;

antennæ 11-jointed, with a club of three joints, the last of which is

decidedly predominant . . . . = +++ - * - . 2I

Of a different conformation x

Lad

“I

kam

Cie arched both longitudinally and transversely Wasmannia Forel

22. Club of the antennæ 2-jointed, the last joint much larger than the other

(tribe Solenopsidii and the genus Phacota) euo 23

Antennal club of a different conformation or indistinct è 28

. Antennæ 9-jointed (1o-jointed in the female and 1:3-iointed. in in the

male)!; no dimorphism among the workers . Carebara Westw

Antennz 12-jointed . : . . .-. Adelomyrmex Emery

Antenne 10- or fy jetted”. du:

1 Professor Forel refers to the genus Oligomyrmex Mayr, the worker of which

is unknown, an undescribed Australian species with strongly — workers.

The antennze are 9-jointed, as in Carebara worker and in Oligom yrmex fe male.

—

- 429.) THE FAMILY OF FORMICIDA. 717

v ADtent T630IBfed . «convi cpu coe wrote RE

Antenne II-jointed . (58

. Dimorphism of the al ae ordi iik slightly Widi or, in the

opposite case, the head of the worker major is subquadrate or broader

than long (antennæ 1o- to t 1-jointed in the female)

* Solenopsis Westwood

Dimorphism of the workers very marked; head of the worker major

elongated (antennz 11-jointed in the female) . Aéromyrma Forel

Thoracic sutures indistinct. . imo od ooo M POM tip

Mesoépinotal suture strongly stn ind

. Ninth antennal joint conspicuously indir tiam m eighth, ed

much shorter and especially narrower than the tenth

Diplomorium Mayr

Ninth joint of the antennae not distinctly longer than the eighth;

workers polymorphic ; soldiers with enormous heads

Pheidologeton F. Smith

. Antennal fovee deep, capable of containing the whole scape, and

placed along the sides of the head . . - 29

Antennal fovez less deep or differently ud FOE 30

Antenne 9-jointed (ro-jointed in the male as in E 1 à meso-

notum with a blade-like posterior edge and usually armed with spines

Meranoplus F. Smith

Antenne 1 I-jointed, the last joint very E Calyptomyrmex Eme

. Erect hairs on the body trifid . . . * Tri R Forel

Hairs not trifid .

. Posterior border of the dope teed t in the Kis of a eit ridge

which borders the antennal foveæ in front. . . . . . . . 32

Posterior border of the clypeus not forming a ridge

- Mandibles pointed, without apical border. * Strongylognathus M ayr

Mandibles with dentate apical border .

33. Portion of the clypeus in front of the goed insertion narrow, but

not reduced to a mere ridge (antennz of the male ro-jointed) (form-

ing with the four preceding genera the tribe Tetramorii) . 4

Portion of the clypeus in front of the antennal insertion reduced. toa

trenchant ridge “ea of male b pos ue i rw

34. Antenne 12-jointed . cu | 4 y v uc. 2 X

Antenne 11-jointed . m m ences 788

35. Epinotum armed with spines or + Gah . . . . *Tetramorium Mayr

Epinotum rounded, unarmed . Rhophomyrmex Mayr

36. Thoracic dorsum deeply coi a at the inesoéptootal suture

Dacryon Forel

Thoracic dorsum scarcely or not at all ipu at the mesoépinotal

suture . Rhe uis EE . Xiphomyrmex Forel

37. Antenne era ees nw 2. . Pristomyrmex Mayr

Antena jomed .. . . : Pe ce . 38

718 THE AMERICAN NATURALIST. [VoL. XXXVI.

[I2

Petiole pedunculate in front ; dimorphism of the workers very marked

Acanthomyrmex Emery

Petiole not EPEE in front; no — ine in the

workers . Myrmecina Curtis

. Antenne 7-10- jointed, jh ask dide: very eem tiaras without spines

or teeth; eyes present. . . Allomerus Mayr

Antenna 10-12-jointed; when dee are SPD ahi eyes are wanting or

the epinotum is armed with two spines. . . . . = à v 4o

. Without eyes i doque un raa EN

With eyes, sometimes aoa SER quiis distinct ; iw

. Antennz 12-jointed, without distinct club ; insects very sm

all

* Leptanilla Emery

Antenne 1o-11-jointed, with 3-jointed club 42

. Postpetiole armed with a spine on its lower kitice oe Mayr

Postpetiole unarmed ; size very small Monomorium decamerum Emery

. Antenne 11-jointed

Antennz 12-jointed rsen in Pheidole Wirsuiilla dade:

Thorax and petiole without any trace of teeth or spines; pronotum

never angular 45

Metanotum ndy sleays LS d with edi or spines; aiia they are

absent, the pronotum has angular humeri . 48

. Clypeus bidentate in front 0 cia oen 46

Clypeus unarmed . E E M E

Petiole distinctly pedunculate i in irott . . . * Monomorium Mayr

Petiole not pedunculate . . . Xenomyrmex data

. Thorax unarmed, impressed in de, region rdàt the mesoépinotal su

Vollenhovia Mayr

Thorax sima with spines and without an impression at the meso

epinotal su ; Stereomyrmex Emery

. Eyes Mig bin Beaart wai Forwards

Se André

Eyes round or oval 49

. Thoracic dorsum jetioundiy lide at the inso suture 59

Thoracic dorsum little or not at all impressed .

. Homen of pronotum rounded. . =. . . . . . - Huberia Forel

Humeri of pronotum angular .

. Antennal club 3-jointed, at bat | as Jar as ‘Be iiaii of the

funiculus yrmex Emery

Club indistinct, ie last (Ei ‘Sines wack ral than the remainder

of the funiculus . Podomyrma F. Smith

. Frontal carinz as TAE as ste idein scape . * Tomognathus Mayr

Frontal carinze much shorter than the sca

Workers strongly dimorphic; integument shining, petk pedunculate

achomyrm orel

1 This group was established by Professor Forel as a subgenus of Liomyrmex:

it has seemed to me to deserve elevation to the rank of a genus.

No. 429.] THE FAMILY OF FORMICIDA. 719

Workers not dimorphic . 54

. Petiole scarcely pedine in irt biipdüde acad below with a

spine,’ integument in great part smooth and shining

* Formicoxenus Mayr

Petiole with a short peduncle, postpetiole unarmed below; body in

great part opaque . . . . *Leptothorax Mayr

. Workers dimorphic, -— wibo PENE intermediate between the

large-headed soldiers and the workers; antennal club 3-jointed,

longer than the remainder of the funicle Gpjdiatedi in PA. granulata

Pergande) ; sting very feeble . . Pheidole Westwood

Workers monomorphic or T 3 in the latter case the extreme

forms are connected by intermediates, and the antennal club is

usually indistinct or shorter than the remainder of the funiculus 56

36. Petiole armed with spines above? o uone S nos eoe e lu. 57

Pellole without-a spine a 5... no e accepere o ee EB

57. Petiole with one spine- . . . . . . . . . Lordomyrma Emery

Petiole with two spines . . . Atopomyrmex André

58. The last three joints of the antenne are are shorter than the

remainder of the funiculus and do not form a very distinct club — 59

The last three joints of the antenne form nosse a club about as

long as the rest of the funiculus . 6

Thoracic dorsum impressed at the e mesoëpinotal suture ; promesonoral

suture usually distinct a o

Thoracic dorsum without any trace of lohan: orimpression. . . 62

60. Posterior spurs pectinated . . . . . S . o "NETNA

Posterior spurs simple or absent . 61

61. Middle of clypeus projecting in an kies epibohun unnið, with a

median impression which can receive the petiole; integument in

great part shining (America) . : . Megalomyrmex Forel

Middle of clypeus angular in droit; pinoia armed with spines ;

antennæ thick (Madagascar). . . . Eutetramorium Emery

ypeus of a different A T bios usually armed with

teeth or spines (always without spines in —

namma Westwood

* Holcomyrmex Mayr ?

62. Posterior spurs simple ; petiole very long ene Ocymyrmex Emery

Posterior spurs pectinated (America) . Pogonomyrmex Mayr

1In Z. corsicus Emery, the Perte of which is unknown, the postpetiole of the

female has only an obtuse too

? In Atopomyrmex cranes d nodifer the node of the petiole is merely

angular in front.

* All the characters of the genus Holcomyrmex are found singly in one or the

other forms of the genus Stenamma as I have defined this genus in my work on

the ants of North America (Zool. Jahrb. Syst., Bd. viii, p. 297). The teeth of the

clypeus are indistinct in Æ. muticus Emery.

720 THE AMERICAN NATURALIST. [ VoL. XXXVI.

Clypeus armed with two ridges, which project forward in the form of

teeth, rarely without teeth, but then the epinotum is quite unarmed ;

mesoépinotal suture marked * Monomorium Mayr

Clypeus of a different Caoin. rarely 2- Paun but then the

mesoëpinotal suture is indistinct 6

64. Inferior angles of the pronotum pointed . . . . - Rogeria fad

Inferior angles of the pronotum rounded .

65. Postpetiole campanulate, attached Cision io. means of its shed

posterior surface to the einig segment; thoracic dorsum with

neither suture nor impression . Macromischa Roger

Postpetiole distinctly constricted postery : Ce hea ae

66. The abdomen, seen from the side, is licia its anterior angle

attached to the postpetiole; epinotal spines recurved forwards

Trigonogaster Forel

Of a different conformation

. Without erect hairs, petiole dc with a a sudes DONE

petiole usually very large . * Cardiocondyla Emery

With erect hairs; which are sini ees and microscopically den-

E ne es OQ Saas s ns MAE: Mayr

€

The following genera are not included in this table : * Anergates Forel

and Epæcus Emery, parasitic ants, which have no workers; Trichomyrmex

Mayr, Tranopelta Mayr, * Oligomyrmex Mayr, Rhopalomastix Forel, Crato-

myrmex Emery, of which the workers are unknown; and finally Pheidola-

canthinus F. Smith, which was insufficiently characterized by its author

and is unknown to me in nature.

4 Subfamily: DOLICHODERIN#.!

1. Sting well developed; petiole with a long peduncle in front

Aneuretus any

Sting rudimental ; AREE squamiform or nodiform, not pum d in

tont. . p

2. Chitinous integument stiff d britde, often KERET Puer ‘horas

and scale often spinose or angular; gizzard without a calyx and with

delicate cuticle, not furnished with cilia at the eaman

Dolichoderus Lund

Integument thin and flexible, finely sculptured; thorax and pee

never spinose ; gizzard of a different conformation . 3

3. Body very slender ; legs and antennæ much elongated, qune or gua

sized species; gizzard without a Ce furnished with cilia at the

entrance (Australia) . a . Leptomyrmex Mayr

1 The genera of the Dolichoderinz are in large part established on anatomical

a characters (structure of the gizzard) and the wing neuration; this renders the

identification of isolated workers very difficult.

unl

Fal

—

- 429.] THE FAMILY OF FORMICIDA. 721

Body less slender ; species Foie always small; gizzard with a reflected

cix. Ck Wee

. Eyes very lige; occupying enicditid the ide óf the bent

Turneria Forel

Eyes much smaller. . WES

. Cloacal orifice apical; penike very p withaat a distinct este

heepepn mie Mayr

Cloacal orifice inferior

. Scale of petiole very small add — piela; or even shegethior

absent . . i ee Ee |

Scale more or ias D ri: but a seve t 9

Maxillary palpi 2—4-jointed, labial palpi 2-3- T in ‘the anterior

wing of the female and male the transverse nervure joins the external

branch of the cubital nervure (Europe, India, Australia)

* Bothriomyrmex za

Maxillary palpi 6-jointed, labial 4-jointed

Scale of petiole small but distinct in the dorka (well devetopett in ihe

female); in the anterior wing the transverse vein joins the external

branch of the cubital vein; there is no closed cubital cell in the

male, one only in the female; no discoidal cell; gizzard with a

convex, 4-lobed calyx (a monotypic American genus) Forelius Emery

Scale rudimental or none; the transverse vein joins the cubital at the

point of bifurcation; a single closed cubital cell, eue a single

discoidal; gizzard with a depressed calyx wit e

apinoma Foerster

Metanotum bearing a conical projection more or less distinctly devel-

oped ; wings as in Forelius (American species) Dorymyrmex Mayr

Metanotum of a different conformation ; wings with a discoidal cell 10

Gizzard very short, with a great reflected calyx; no ocelli, stature but

slightly variable . ridomyrmex Mayr

Gizzard at least as long as eon stature y highly veka ocelli usually

present in the large workers II

. Thorax not impressed at the tubéblplüotil vtae i no ésiiskable

dimorphism in the workers ; wings with two closed cubital cells

* Liometopum Ma

Thorax impressed at the mesoépinotal suture; workers often remark-

ably dimorphic; with a single closed cubital cell; transverse vein

meeting the bifurcation of the cubital (American species)

Azteca Forel

The genus Linepithema Mayr, of which only the male is known, is not

included in the above table.

722 THE AMERICAN NATURALIST. [VoL. XXXVI.

5th Subfamily: CAMPONOTINE.

Mandibles long, linear, denticulate ; eyes very mid Myrmoterus Forel

Mandibles of a different conformation . . . es. vp iL

Antenne II-jointed, or less . . ee ee ern ee le

Antenne 12-jointed . i.

Clypeus projecting Feeder ove he vsadiblnk: eyes inci enormous,

anana BIO S. a RO. o own esomyrmex Mayr

Of a different conformation

Last joints of the antennz bem Ns a differentiated lada: antennz

g-Io-jointed . . . . . -Myrmelachista Roger

Antenne without a differentiated b

Frontal carine separated from each other by a paci daat dá

from the lateral borders of the head

Frontal carinz closer to each other chad to sie TRE tordi of the

Eyes reniform, very large . . . . . Dimorphomyrmex Er. André

Eyes oval, of medium size . . . Aphomomyrmex Emery

Maxillary palpi 2-jointed, labial dn pag VS ‘inact Roger

Maxillary palpi 6-jointed, labial palpi 4-jointed 8

Metanotum and scale more or less 2-toothed or actu

* Acantholepis Mayr

Metanotum and scale unarmed 9

Antenna I11-jointed (Old World gi : ioc * Plagiolepis Mayr

Antenne 9-jointed, exceptionally 11-jointed ; tioni short and thick-

set (American species). . Brachymyrmex Mayr

hym;

Eyes very large, occupying iy the whole 7 the sides of the head

Gigantiops Pe

Eyes occupying less than one-half of the sides of the head .

. Prothorax with an angular crest on either side; mesonotum ME

nent,in the form of a boss ; gizzard as in dos Notoncus Eu

Thorax of a different conformatio

Antennz inserted some distance behind tne ae gizzard w ik bod

straight sepals 13

tennz inserted very near ihe poterint edge of the epe eus . 20

Maxillary palpi 5-jointed; petiole elongate, narrow ; stature saddle

but not dimorphic in the form of the head . (Ecophylla F. Smith

Maxillary palpi 6-jointed ; petiole short, squamiform or nodiform, me

spinose or dentate . .

Dimorphism clearly sedi in be size, dors; nd ien in i die scale

of the head; stature usually very variable . . * Camponotus wee

No marked hace phen 1 in the workers ie de di

Eyes placed towards the posterior PONR of the ith d

Opisthopsis Emery

Eyes on the sides of the head Qc me 9

No. 429.] THE FAMILY OF FORMICIDE. 723

—

. Thorax and petiole without spines or teeth . . oles |

sin and petiole, or the latter alone more or TA spinose or den-

: 18

: bed thickset ; vu BENO behind (hirik Pápiiasfi)

Calomyrmex Emery

Body slender; head narrowed behind (America)

Dendromyrmex Emery

Body thickset; thorax without spines, its anterior angles rounded;

petiole prolonged into a point on either side . Echin nopla F. Smith

Body less thickset; thorax usually dentate or spinose (when this is

not the case the body is T in black, and the scale of the sena

is quadridentate) . . 19

In profile, the mesonotum occupies dà TN of a TNR cleft over-

arched by the metanotum ; eyes supported laterally by a lobe of the

head in the form of a blinder (cillére) . . . Hemioptica Roger

cit pdm not depressed; eyes usually free, rarely with a distinct

blinder . ; . . * Polyrhachis F. Smith?

« The sind 4 to 5 Multi joints distinctly differentiated to form a club

ei esa Er. André

No differentiated club 2

. Calyx of gizzard reflected id a E a Md ring as in

Plagiolepis (Australia, New Zealand, Chile). Melophorus Lubbock

Sepals of calyx distinct and little or not at all reflected (except * Pren-

olepis, none of these genera are found in Australia or Hi) oeza

- Clypeal fovea distinctly separated from the antennal fovea

* Prenolepis Mayr

ys: fovea confluent with the antennal fovea, or feebly separated in

Pseudolasius . 23

. Joints 2-5 of the hiii diir or not fiare ds the succeeding

joints ; ocelli usually a| 24

Joints 2-5 of the i longer us: the Maple ux ioni: cilii

distinc 25

Mandibles a. with stique PEEN hladas ; doki very mike :

head large in the worker major; clypeal fovea slightly separated

from the antennal fovea (Sunda Islands and Moluccas)

Pseudolasius Emery

Mandibles shorter, with less oblique blades; dimorphism scarcely or

not at all perceptible (holarctic region) . . . *Lasius Fabricius

- 4th joint of maxillary palpi nearly twice as long as the sth

* Myrmecocystus Wesmael

ins = of the maxillary palpi a ks lta ge es mein; s. . 6

late * Formica Linne

Mandibles narrow and pointed vow ee Co F Pelyepus Latreille

at ag! Simplex Mayr has been found in Palestine. Mr. Edward Saunders sent me

e provenience of which appears to be authentic.

724 THE AMERICAN NATURALIST. [VoL. XXXVI.

ADDENDA.

Since the manuscript of the preceding paper was received,

Professor Emery has made some changes in the subdivision of

the genus Cerapachys. He now divides this genus into five sub-

genera as follows : (1) Cerapachys (sensu stricto), with 12-jointed

antennz ; (2) Parasyscia, with 11-jointed antenne; (3) Odcerza,

with 10-jointed antennz; (4) Syscia, with 9-jointed antenna

and the basal segment of the gaster but little longer than the

postpetiole; (5) Cysias, with 9-jointed antennz and the basal

gastric segment very large. The more homogeneous genus

Phyracaces is not cut up into subgenera.

In another recent paper? Emery describes a new genus,

JEnictogiton, based on a male specimen of a peculiar doryline

ant from the Congo (A. forsiceps Emery). As the name indi-

cates, this insect is allied to /Enictus.

As a further addition, I may mention that Forel? has very

recently described a remarkable new genus of Ponerinz from

Haiti under the name Emeryella. It resembles the extraor-

dinary genus Mystrium in the structure of its mandibles.. In

other respects it is allied to Ectatomma. The following is a

translation of Forel's diagnosis :

* Emeryella gen. nov.

* Mandibles, at first sight, very similar to those of the genus Mystrium,

but without the two rows of teeth along their inner borders. They are

linear, slightly depressed, longer than the head, feebly curved inwards,

especially at their distal half, the basal half being nearly straight. Their

bases are a little longer than their tips, which are obliquely truncated.

There are only three teeth on the mesial border; the first is very broad,

short, and obtuse, and not far from the base; the second, situated in the

middle, is short and obtuse ; the third is still smaller, and near the tip. The

mandibles are nowhere canaliculated. They are inserted, like those of

Mystrium, on the widely separated anterior angles of the head, so that they

enclose a large empty space.

1 Note mirmecologiche, Rendiconto delle Sess. della R. Accad. delle Scienze dell’

Istituio di Bologna (Nov. 17, 1901), pp. 3-15.

? Note sulle Doriline, Bull. della Soc. Ent. Ital, anno xxxii, trim. 1 (1901)

PP 43-03,

et da # , 1

Ann. Soc. Ent. Belg. , tome xlv (1901), pp- 334-382-

No. 429.] "IHE FAMILY OF FORMICIDÆ. 725

** Apart from the above, all the characters are very similar to those of Ecta-

tomma, especially of the subgenus Gnamptogenys. Antenne 12-jointed.

Eyes large, lateral. Frontal carine widely separated, short. Clypeus

rounded behind. Promesonotal suture only slightly visible. Mesometa-

notal suture very deep, constricted. Pedicel of the abdomen like that of

Gnamptogenys. Middle and hind legs with but a single spur, which is

pectinated. Tarsal claws bidentate.

* This genus is undoubtedly very closely related to Ectatomma ; but the

structure of the mandibles is so peculiar and recalls so forcibly the group

of Mystrium and Myrmecia that I feel fully justified in establishing the

genus."

The type of the genus Emeryella is E. Schmitti Forel.

W. M. WHEELER.

PECULIAR STAGES OF FOLIAGE IN THE

GENUS ACACIA.

CARLETON E. PRESTON.

Tute development of foliage leaves in general appears, from

the results of investigations made thus far, to be a regular.

increase in complexity, almost the same for the ontogeny of a

single plant as for the race in historic development, as disclosed

by fossil remains. As an especially good example may be men-

tioned a series given by Jackson ('99) illustrative of the history,

both phylogenetic and ontogenetic, of Liriodendron. For the

genus Acacia, Reinke ('96) gives a very comprehensive review

of the development based upon the seedling forms and com-

parative study of the adults of the several subgenera, noting

especially the phyllode-forming tendency. :

As the first point which this paper aims to bring out deals

with the bipinnate leaf and its development regardless of the

phyllode, it may be well to call to mind the regular course of

seedling leaf forms. The first leaf, after the cotyledons, is

normally singly pinnate, with about four pairs of leaflets;

the next is generally bipinnate, with but a single pair of pinnze ;

further change is a mere addition of more such pairs along the

main axis. A peculiar transition stage between the singly

pinnate and the bipinnate is sometimes found in seedlings

of A. leprosa Sieber when growing under cultivation. The

shadow prints (Figs. 1 and 2) annexed show the nature of this

peculiarity. The lower pair of leaflets only is replaced by a

pair of strongly developed pinnz, while the rest of the axis

runs on singly pinnate and rather weak in structure. As a

rule, no such continuation of the main axis is to be found.

In the case of Liriodendron, Jackson found some stages which

could be regarded as progressive beyond the normal condition

of the present species. Here the complexity was increased

simply by augmentation of the number of lobes to the leaf.

747,

728 THE AMERICAN NATURALIST. [VoL. XXXVI.

This augmentation, as has often been shown, takes place in

Acacias in the passage from the customary seedling leaf to the

majority of bipinnate adults, a large number of pinnz being

added before the full-sized leaf is reached. But this mere addi-

tion of pinnz scarcely seems to be the right interpretation of

the increase in complexity if one compares the seedlings of

Liriodendron and Acacia. What corresponds to the increase

of lobes in the former is not so much the addition of pinnz in

the latter as the increase in degree

of pinnation. Starting with a singly

pinnate leaf, one arrives at a bipin-

nate form of remarkably fine struc-

ture in such species as A. decurrens

Willd. But may there not exist a

tendency toward triple pinnation

in some of the forms? This ques-

tion, suggesting itself, led to a

rather careful search for evidence

in its favor, with the result that in

A. decurrens not a few leaves were

discovered in which such a tend-

ency was manifest. Of one of these

leaves a shadow print is given

(Fig. 3), showing clearly the third

degree of pinnation on some of the

Fic. 1. — Seedling of A. Zerosa, showing basal leaflets of the pinne. How

abnormal third leaf. (Natural size.)

much of a prophecy this is of a fully

tripinnate leaf in certain Acacias of some future epoch no one

can really tell; the tendency, however, cannot be without

significance.

In his paper, which deals with animals as well as with plants,

Jackson carries the same principle so far involved one step in

advance in point of application. As a leaf develops base and

tip first of all, he reasons that any change in shape which may

add to its complexity will not appear at those points, but rather

upon the proximal portions of the lamina. This he shows to

be the case in Liriodendron, citing also several other examples

in substantiation of his view.

No. 429.] FOLIAGE IN THE GENUS ACACIA. 729

In the case of the A. /eprosa seedlings with abnormal devel-

opment the lateral pinnae may be regarded as further modifica-

tions of the proximal pair of leaflets of the ordinary first leaf of

seedlings.! If this is, as seems probable, the right interpreta-

tion of such peculiar forms, it almost necessarily follows that

the change in all the species at this stage is of the same nature,

and that as a rule the higher, or more distal, leaflets upon the

main axis lose the power to develop, on account of the greater

strength of the newly developed

lateral pinnz. It may be worth

while to state that these stages

were found only after their predic-

tion as the result of a course of

reasoning, and were the direct

objects of the search which led to

their discovery.

The tendency toward triple pin-

nation is likewise, as may be easily

seen by reference to the figure,

only present upon the proximal

leaflets of the pinnz, and not at

all to be found on the pinnz near

the tip of the leaf.

The final point to be discussed

is the gradual metamorphosis to

the phyllode. In seedlings where ; leanai im)

the change is gradual, as is the

case in the great majority of species, but one pair of pinnæ

is present at the time of its occurrence. In the abnormal

seedlings shown, the prolongation of the axis is suppressed

as the transition stages appear. In such cases it is possible to

say with a high degree of certainty that the phyllode represents

merely the flattened petiole. Next to be considered are the

conditions of such forms as experience these changes later

in life. 4. rubida A. Cunn. and A. heterophylla Willd. have

already been described by Reinke, and in his article one stage

in the transition as it occurs in A. heterophylla is figured.

Fic. 2. — Abnormal seedling of A . Zeprosa

! Cf. seedling of Gleditsia, where such transition stages are frequent.

730 THE AMERICAN NATURALIST. [Vor. XXXVI.

According to that author the change is merely a gradual flat-

tening of the petiole, accompanied by the reduction of parts

more distal. The stage represented in his paper is explained

in the following words: ** Wo es zu breiten Phyllodien mit nur

einem Fiederpaar an der Spitze kommt, entsprach in den von

mir gesehenen Fallen das Phyllodium einem nur oberseits

geflügelten Blattstiel, das Fiederpaar dem untersten Fieder-

paar eines Fiederblattes, Spindel und höhere Fiederpaare

würden somit als abortirt anzusehen sein, Steht auf der

Spitze eines Phyllodiums einmal ein zweipaariges Fiederrudi-

ment, so pflegt die Spindel zwischen den beiden Fiederpaaren

ungeflügelt zu bleiben ” (p. 568).

Reinke is naturally giving only the results of his own obser-

vations, yet one cannot but draw the inference that a certain

law is at least suspected, czz., that only the petiole flattens in

each case, and that at the same time the distal parts gradually

cease to develop, until at last the stage is reached which this

writer figures, after which the last pair of pinnz fails to develop,

and the phyllode is complete.

In the botanic garden of Harvard University there are two

species of Acacia which show these transition stages in great

abundance. They agree well in leaf characters with A. rubida

and A. heterophylla, and will be discussed under these names,

though full identification was not possible, owing to absence of

flowers and fruit. Whether or not the determinations are

right, the principle which they illustrate remains the same.

Of both of these species the writer made a special examination

during the spring of 1900, with a view to observing the different

transition stages, their sequence and character. Upon re-read-

ing Reinke's article in the summer of igor, and finding there

certain statements which in a way did not correspond with his

results, he repeated the examination and ran across more pecu-

liar forms which tend rather to disprove than to confirm the

existence of any law governing such changes.

. The first thing which appeared from these two examinations

was that the method of reduction of the more distal parts

described by Reinke, though indeed frequent, is perhaps not

the most common of all. Rather more often an equal decrease

No. 429.] FOLIAGE LN THE GENUS ACACIA. 7251

in size and final simultaneous disappearance of all the pairs of

pinna was noted, as opposed to an unequal reduction starting

Fic. 3. — Leaf of A. decurrens showing tendency toward a triple pinnation.

(Slightly reduced.)

at the distal end. In this A. rubida seemed to be very regular ;

In A. heterophylla various methods of change were shown. As

732 THE AMERICAN NATURALIST. [VoL. XXXVI.

far as sequence is concerned, there appeared to be no definite

order, series of transition stages running in both directions.

There was also found a fairly large number of stages which

lead one to doubt greatly whether in all cases it is the petiole

only which is transformed to the phyllode, and whether, if one

pair of pinnze persists at a certain stage, that pair is always the

proximal one. The prints which follow may, to be sure, repre-

sent mere anomalies, but from their number, at least, they can-

not but raise in one's mind a certain amount of hesitation to

Fic. 4. Fic. 5.

Fics. 4 and 5. — A. heterophylla, variation in transition to phyllode. (l4 natural size-)

consider the existence of a law as to method in any way estab-

lished. Here the flattening appears in some cases entirely on

the distal portions without affecting the petiole, in others both

petiole and rachis are involved to varied extents. How these

are to be interpreted under one definite law seems incompre-

hensible.

It is hardly to be expected, however, that the process of

réduction of parts is going to follow the same strictly regular

line as that of advancement. The two are brought about by

entirely different causes and are in no way comparable. There-

fore the fact that such changes as those last described occur

does not greatly invalidate the testimony of the earlier presented

No. 429.] FOLIAGE IN THE GENUS ACACIA. 733

data in support of a principle dealing with a progression. On

the whole, the conditions of the seedlings of A. /eprosa and of

the leaf of A. decurrens stand strongly in favor of the existence

Fic. 6. Fic. 7.

Fic. 8, FIG. 9.

Fics. 6-10. — A . heterophylla, variations in transition to phyllode. (l!4natural size.

of a regularly increasing complexity in leaf form, governed

by fixed laws and repeated in an abbreviated form in plant

ontogeny. To a certain extent also they add weight to Professor

734 THE AMERICAN NATURALIST.

Jackson’s theory regarding stages in the development of a

single leaf. The peculiar phyllode form rather discourages the

idea of formulating any law covering phyllode formation, and

lends an extremely potent meaning to Reinke’s statement,

«Der Wechsel von Phyllodien und Fiederblattern ist bei

dieser Art [probably, as regards phylogeny, in others also] ein

sehr mannigfaltiger.”’

REFERENCES.

'99. JACKSON, R. T. Localized Stages in the Development of Plants and

"Animals. Mem. Bost. Soc. Nat. Hist. Vol. v, pt.

96. REINKE, J. Untersuchungen iiber die Aiaia do der

"Leguminosen. Pringsheim’s Jahrb. f. wiss. Bot. Bd. xxx. 70 pP-

THE RELATION OF WIND TO BIRD MIGRATION,

C. C. TROWBRIDGE.

THE influence of meteorological conditions on the migra-

tion of birds has been studied by a number of observers.

Among the important monographs on the subject is that of

Prof. W. W. Cooke, “Bird Migration in the Mississippi Val-

ley," in which it is shown clearly that migratory waves of

birds accompany changes of temperature. The wind direc-

tions and velocities were recorded in the paper mentioned but

the author did not regard the migratory movements to be due

to the direct effect of the wind, which was considered merely

as the chief cause of the temperature changes.

The same view is held by Dr. W. Stone in his “Birds of

Eastern Pennsylvania and New Jersey," in which by a series

of curves the fact is illustrated that migratory waves of birds

follow closely changes of temperature.

The author of the present paper believes that the bird waves

described by the above observers were largely due to the direct

influence of favorable winds, and that the wind is a more impor-

tant factor in bird migration than it is generally regarded.

That the subject has not received much attention is shown

-by the recent valuable review on bird migration by Dr. F. H.

Knowlton in Popular Science Monthly» In the paper referred

to no mention of the relation of the wind to bird migration is -

made. In fact, the word “wind” is not used in the thirteen

pages of the article.

In considering the relation of meteorology to bird migra-

tion, general as well as local atmospheric conditions must be

studied, and it is important that the atmospheric conditions

Where the birds start on their migratory journeys be deter-

mined, if possible. Furthermore, the number of new arrivals

! Knowlton, Dr. F. H. The Journeyings of Birds, Pop. Sci. Mon., vol. 1x,

No. 4, p. 323.

735

736 THE AMERICAN NATURALIST. [VoL. XXXVI.

recorded in any locality is not always a safe criterion of the

migratory movement of birds, because adverse meteorological

conditions often tend to retard or stop the migrations and

bring certain species under notice which, with favorable con-

ditions, would have passed by unobserved.

A previous paper by the author, *Hawk Flights in Con-

necticut,” ! contained an account of flights of hawks which had

occurred along the southern border of Connecticut during

the autumn migrations, in which the results were presented of

observations covering a period of nine years, from 1885

to 1894.

The present paper is an account of further observations on

the migrations of hawks and on the effect of the wind on the

migrations of various other species of birds. It is necessary

to refer to the contents of the previous paper on the subject,

and the principal facts given in it are as follows :

It was shown that flights of hawks occurred in September

of almost every year in southern Connecticut, consisting of

different species, some of which passed along the coast singly,

others in flocks of from twenty to two hundred. The hawks

were most numerous at from one half to one mile inland

from Long Island Sound, along the first low hills back from

the coast.

Twenty-two hawk flights were recorded in nine years, and

in each case notes were made on the relative abundance of

the different species of hawks seen migrating. A record was

kept of the meteorological conditions on the days when the

flights in question took place. It was found that on the

days when the flights occurred the wind blew from the north

or northwest, and usually with considerable velocity. On

seven days the wind blew at the rate of over twenty miles

an hour, and on eight other days not less than fifteen miles.

The greatest flights occurred on three days, when the velocity

of the wind was about forty miles per hour during part of the

day.

An explanation of the cause of these autumn flights of hawks

was given, and the theory advanced has been fully verified by

1 The Auk, July, 1895.

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 737

subsequent observations. The cause of the flights was stated

to be as follows:

Hawks drift with the wind when soaring,

continually, their movements during migrations are largely

dependent on the direction of the wind. In the present case

hawks migrating in autumn in New England drift south or

and as they soar

WIND N.W. b

^w,

Mar I.— Coast LINE oF THE NEW ENGLAND STATES AND NEW JERSEY.

The long arrows indicate the general direction of migration of hawks and many other land

d ng why these birds congregate

vente | in the autumn, when de wind is northwest, illustratin

The peces circles drawn about a point off the comet of Maine are low-p isobars indi

causing the inds over ipe: north-

pea hawks f ollow a narrow path and

g higher as the id becomes

enstern part of the United States.

low; at B the hawks gradually separate, es

more favorable.

738 . THE AMERICAN NATURALIST. [Vou. XXXVI.

southeast until they reach the Atlantic coast line. They then

turn westward and follow the shore until they have reached

New York and New Jersey, where they gradually separate and

pass on southward. Thus the hawks migrating from a large area

of country — eastern Canada, New England, and perhaps north-

eastern New York — are forced to travel along a narrow coast-

line path at right angles to the main direction of their migration.

A condensed table of flights which occurred from 1885 to

1895 is given below. It has already been published in part,

but it is necessary in the present paper for the purpose of

showing certain new facts that will be presented.

The meteorological data in the table as formerly given was

that for New Haven, Conn., where the observations on hawk

flights were made. In the table as given at present the obser-

vations of the U. S. Weather Bureau taken at Boston are sub-

stituted for those taken at New Haven. This change is made

because the former station is nearer the locality where the

hawks start on their migratory journey than the latter place.

Another column has been added to the table for the purpose

of showing the effect of temperature changes on the migratory

movements of the hawks. The meteorological data in the

tables have been kindly furnished by the Boston office of the

U. S. Weather Bureau.

It is difficult to determine the number of hawks that pass

a given point on the Connecticut shore, during a moderately

large flight in September. An estimate, which appears to the

author to be conservative, is that over 15,000 of them pass

New Haven in one day. The sharp-shinned hawks outnumber

the other species several times over. The broad-winged hawks

are next in abundance, then the ospreys and marsh hawks which

are about equally numerous.

Observations show that many ospreys and marsh hawks cross

to Long Island, but it is evident that the main flight of hawks

is along the Connecticut shore.

Since 1894 observations have been continued on flights of

hawks in Connecticut in autumn, and in New Jersey in spring,

Which have resulted in bringing to light additional facts relat-

ing to the effect of wind on the migration of birds.

No. 429.] RELATION OF WIND TO BIRD MIGRATION.

TABLE I!

739

LIST OF MIGRATORY FLIGHTS OF HAWKS WHICH HAVE OCCURRED IN

SOUTHERN CONNECTICUT DURING

METEOROLOGICAL RECORDS,

THE YEARS 1885-95.

BOSTON, MASS.

TEMPER-

WIND ELO- ATORE

DATE. WEATHER. | DIREC- AS CHANGE REMARKS.

10 IN Past

P ed 24 Hours

Sept. 23, 85 | Rain N.W. are —7 | Moderate Highs: falco spar-

verius commo

Sept. 18, '86 |' Clear N.W.| 21 —8 ght of ll hawks and

us am land birds.

Sept. 22, '86 | Partly N.W.| 1o —2 sas aii abundant.

cloudy

Sept. 16, '87 | Clear N. 12 — 5 |A great flight all day.

Sept. 17, '87 | Clear N.W. 8 —7 | Buteo saan abundant early

in the morning.

Sept. 24, 87 | Clear N. 14 re Baas latissimus abundant, also

Accipiter velox.

Oct. 19,'87 | Cloudy N. IO — 11 |No large flight, but omen all

the migrant hawks observed.

Sept. 10, '88 | Cloudy W. 9 —2 | Accipiter velox abundant.

Sept. 22, '88 Cloudy N. 24 —3 | Moderate flight.

Sept. 22, '89 | Clear W. 14 —8

Sept. 28, '89 | Clear W. 14 — 5 | Very large flight.

Oct. 15,'89 | Clear N. 19 +5 | Moderate flight ; Accipiter velox

lentiful.

Sept. 18, ’90 Cloudy W. 13 —2 |40 hawks killed; Buteo latissi-

s abundant.

Sept. 21, 'go | Clear N.W.| ta — 16 | Moderate flight.

Sept. 25, ’90 | Clear W. 14 +2 |Pandion halietus carolinensis

abundant.

Sept. 24,’90 | Clear |N.W.| 20 —8 | Small ga

Sept. 8,'91 | Clear W. ÉI- —2 | Hawks increasing in numbers

Sept. 9,'91 | Clear W. IO —5 Ade flight ; killed over 20

Sept. 14, ’91 ear W. II 2 |Small flight.

Oct. 21, ’92 Cloudy W. 9 —3 |Small flight of Accipiter velox.

Sept. 20, ’93 ear W. 8 di ha od "Ww Buteo latissimus

Sept. 21,'93 | Clear N.W.| 12 py sn flights of adult Buteo latis-

simus

Sept. 13, %95 | Cloudy | N. 14 | —10 | Large flight.

Sept. 14, '95 | Clear N.W.I-n — 1I |Great flight of Buteo latissimus.

1 The flights of hawks recorded in the above table were observed at New

Haven, Conn., and the meteorological data given is that for Boston, from 1885 to

1887 at 7 A.M., from 1888 to 1

ogical conditions for Boston has been already explained.

1895 at 8 A.M. The reason for giving the meteor-

740 THE AMERICAN NATURALIST. (VoL. XXXVI.

During September, 1895, daily observations were made on

the migration of hawks at New Haven, Conn., when as far

as possible the variety, as well as the number of hawks which

were seen on each day, was noted. A record was also kept of

the meteorological conditions for the same period, particularly

the state of the weather and the velocity and direction of

the wind.

The observations extended over a period of twenty-three

consecutive days, from September 6 to September 29, this

being the month during which five or six species of hawks

migrate southward. The object of these observations was to

ascertain if wind from a southerly quarter not only retarded

the migration, but practically stopped it, and also to determine

if the migratory movement ceased when there was little or no

wind. The observations were as follows :

Sept. 6. Fair; wind N.E., r1 miles. 1 bald eagle; 1 fishhawk ; 2 sharp-

shinned hawks.

Sept. 7. Fair; wind N.E., 16 miles. 1 broad-winged hawk.

Sept. 8. Cloudy; wind S.W., light. 1 sharp-shinned hawk.

Sept. 9. Cloudy; wind S., light. 1 large hawk; 1 sparrow hawk.

Sept. 10. Warm and hazy; wind S., very light. No hawks seen.

Sept. 11. Cloudy; wind S.W., 10 miles. 1 fishhawk; 1 sparrow hawk;

3 sharp-shinned hawks.

Sept. 12. Fair; wind S.W., light. No hawks seen.

Sept 13. Clear, cool; wind N.N.W., 22 miles. Several flocks of broad-

. winged hawks of from ten to fifteen observed; 4 adult bald

eagles; hundreds of sharp-shinned hawks, and fish, sparrow,

and marsh hawks common.

Sept. 14. Clear, cool ; wind N. by W., fresh. All the hawks which were

observed yesterday, common to-day. A number of eagles

also seen. Broad-winged hawks exceedingly abundant ; flock

after flock passed westward over New Haven, Conn. Eighty-

five were counted in one flock, while another equally large

body of them was observed at the same time. These hawks

passed over the city continually from 7 A.M. until 3 P.M.

Sept 15. Clear, cool; wind N., light. A few hawks observed migrating

at a high altitude.

Sept 16. Cloudy; wind N. to S.W., light. No hawks seen.

Sept. 17. Clear; wind S., 5 miles. 2 fishhawks; 1 sharp-shinned hawk.

Sept. 18. Light rain; wind N.E. No hawks seen.

Sept t9. Light rain; wind S., light. 1 fishhawk.

Sept 20. Cloudy; wind S.W., light. No hawks seen.

RELATION OF WIND TO BIRD MIGRATION. 741

No. 429.]

Sept. 21. Clear, warm; wind W. No hawks seen

Sept. 22. Clear; wind N.W., light. A few sharp-shinned hawks.

Sept. 23. Wind W. by N, light. No hawks seen.

Sept. 24. Cloudy; wind N., 12 miles. A small number of broad-winged

and sharp-shinned hawks and one each of the following

species: fish, marsh, and sparrow hawks.

Sept. 25. Stormy; wind S.E., light. No hawks seen.

WIND NE.

= *

ums E ma

p—_— ees >

2 a : ae

J ails

Map IL —Coasr LINE oF THE NEw ENGLAND STATES AND NEW JERSEY.

long arrows indicate the direction of migration of hawks i

when the wind is northeast. A storm center is rire by aeta iso!

i rtheast in New England.

small flight of 1 hawks along the Connecticut shore, as

is northeast there is usually a

indicated.

742 THE AMERICAN NATURALIST. [Vor. XXXVI.

Sept. 26. Fair; wind S.W., 10 miles. 2 fishhawks; 1 sparrow hawk.

Sept. 27. Cloudy; variable light winds. No hawks seen.

Sept. 28. Clear; variable light winds S.E. 1 fishhawk.

Sept. 29. Stormy; wind E. 1 fishhawk.

In the table given below are placed the days, thirteen in

number, taken from the above list, when not more than one

hawk was seen. The meteorological conditions at New Haven

on these days is also recorded in the table. An inspection

of the observations shows that no strong northwest winds

occurred on any of these days.

TABLE II.

IT NuMBER | DIRECTION

DATE, 1895. Was tik, |Winp Drrection,| VELOCITY, |. Hawks| or UPPER

Lower STRATA.| MILES PER :

dion: OBSERVED. | STRATA.

Sept. 7 Fair NE. 15 I —

Mo Cloudy S.W. 18 I

ieee Fair Variable si o Mu

y I2 Fair y Oo SN

€. 3B Cloudy S.W. 7 o uv

Soar 89 Rain N.E. 6 I N.E.

"c 30 Cloudy S.W. 5 o W.

I SE Clear W. 2 o nem

M AS W. 1 o w.

* o 6 Cloudy E. Light o N.

x RT Cloudy N. 4 o eS

To Clear N. 9 I Ti

= 8g Cloudy N.E. 7 I S. by W.

It has been observed that the direction of the movement of

the upper strata does not influence the migratory movement,

and that the surface currents are those in which the hawks

usually migrate. In the last column of Table II the direc-

tion in which the upper strata were moving is given in a few

cases. The table, of course, is not meant to show the number

of birds migrating, but it gives an idea of the number of hawks

observed by one constantly watching for them.

In the table given below, the U.S. Weather Bureau obser-

vations are those for Boston on the days given in Table H.

It is evident from a comparison of these two tables (II and

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 743

III) that the meteorological conditions on nearly all days were

very unsettled in New England, and that although in a few

cases the wind direction, for a time at least, was favorable

for a migration of hawks near Boston, the adverse conditions

in Connecticut stopped the migratory movement.

TABLE III.

UNITED STATES WEATHER BUREAU OBSERVATIONS AT

BOSTON, MASS., 8 A.M., SEPTEMBER, 1895.

Winn Winp VELOCITY, TEMPERATURE

DATE, 1895. WEATHER. Dore dro. MILES PER "HANGE IN Past

Hours,

Sept. 7 Cloudy. N. 13 +3

ee Cloudy. N.W 14 —I

ainat Cloudy. N.W 3 +1

eee Cloudy. N.W 5 —2

“. 46 Cloudy. S.W. 12 t3

“o I Cloudy. N. 8 +4

EE ae Partly cloudy. W. 12 +6

uir i Clear. N, 2 —I

mo Clear W.: 7 +2

eee Clear N. 3 —2

To Cloudy W. 12 3

" Clear N. j —1O

cm) Clear. E. 18 +2

In Table II, as well as in Table III, the directions and veloci-

ties of the winds were taken from the local U. S. Weather

Bureau records.

There is a slight discrepancy between some of these obser-

vations and those taken by the author and given previous to

Table II. These differences are due to variations in the direc-

tion of the wind on certain days. For example, part of the

morning of September 28, Table II, the wind direction was

S.E. and light, while at the time of the U.S. Bureau observa-

tion it was N., 9 miles. The table on the following page

shows the days of September, 1895, when flights of hawks

occurred.

744 THE AMERICAN NATURALIST. [VoL. XXXVI.

TABLE IV.

WIND IND

DATE, 1895. chess’ qn DIRECTION VELOCITY Noen —

(New Haven). (N.H.) OBSERVED.

Sept. 13 | Clear. N.N.W. 22 Large flight.

"o ner] Clear. N.W. to N. 9 Very large flight.

IIS Clear. N. I3 Small flight.

S SE Cloudy. N. 12 " "

The observations made of flights of hawks that have occurred

during a period of fifteen years show that they have taken place

on nearly every date during September, after the sth of that

month. This is readily seen from Table I, which gives the

flights of hawks which occurred during the years 1885 to 1895.

It is evident that there is no special date on which the hawks

migrate during the month of September. Their appearance in

large numbers must be due therefore to some special influence.

This has already been shown to be the wind from the north-

west quarter, in which the birds drift southeastward to the

coast line, making necessary their flight along the shore of

Connecticut.

The observations made during September, 1895, show an

additional fact; if hawks migrated regularly in New .England

throughout September without regard to the direction of the

wind, then there would be a small continual flight of hawks

in southern Connecticut during the entire month, due to the

peculiar shape of the coast line. Such would be the case,

because hawks migrating in a southward direction through

the New England States would eventually reach the south-

ern border of Connecticut, where it would be necessary for

them to fly westward along the coast, producing the small

daily flights referred to.

It has been proved that no daily flight takes place, only

a few stray hawks being observed. Moreover, at any time

1In Table IV the velocity of the wind at 8 A.M., September 14, is given as

9 miles at New Haven. At Boston, however, at the same time the wind was

N.W. 14 miles, and at New York 12 miles

On September 24 the wind at Boston was N. 18 miles, and at New York

N.W. 14 miles,

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 745

during the month a flight of many thousands of hawks will

occur if the wind turns to the northwest quarter, the hawks

appearing within several hours after the change of the wind

takes place. This is shown by the following example :

The morning of September 18, 1890, was warm and calm.

In the most favorable station for observations near New Haven,

between sunrise and nearly eight o’clock, only two hawks were

observed. About eight o’clock a breeze started up from the

northwest, and a great number of hawks soon appeared, over

forty of which were shot. Again, on September 12 and 23,

1895, no hawks were seen, but on the dates immediately

following, September 13 and 24 respectively, hawks appeared

in large numbers.

The significance of these facts is that the wind not only

changes the line of flight of the migrating hawks, but that it

is also the immediate cause of their migratory movement.

In further proof of this statement it has been observed that

when a northwest wind blows for three days in September, on

the third day hawks are not abundant ; for all of those hawks

which are ready to fly southward start at once when the wind

begins to blow in a favorable direction.

A second wind from the northwest quarter, after a week of

southerly winds or calm, will produce a second flight of hawks,

usually of shorter duration than the first.

A third favorable wind will seldom cause another large

September hawk flight, although every wind from the north

or northwest throughout the autumn produces a greater or less

abundance of hawks along the Connecticut coast. .

The expression *the immediate cause of migratory move-

ment," used in the present paper, perhaps requires an

explanation: If a favorable wind, acting as a physical agent, is

used by birds as a means of migration, it is also an immediate

cause of their migratory movement because it determines the

time of their migration.

By the expression “favorable wind ” is meant a wind which

when resolved into components with respect to the migratory

746 THE AMERICAN NATURALIST. [Vor. XXXVI.

EFFECT OF TEMPERATURE CHANGES ON THE MIGRATORY

MOVEMENTS OF Hawks.

To what extent these flights of hawks and other land birds

are due to changes of temperature is difficult to determine.

An examination of the data given in Table I, however, shows

the following facts :

1. The direction of the wind at Boston at the 7 and

8 A.M. observations on days when twenty-four flights of

hawks occurred was as follows: north, six days; north-

west, eight days; west, ten days. Total, twenty-four days.

2. The mean velocity of the wind for the twenty-four days

(morning observations) was thirteen miles an hour, maximum

twenty-four miles per hour, minimum eight. (In many cases

the wind increased considerably during the day.)

3. Mean change of temperature in past twenty-four hours

of twenty-four days on which flights occurred was — 5.7 degrees.

4. The number of days with higher temperature than the

preceding day was four (+ 5 -- 2 4- 2 + 1).

5. The number of days with lower temperature than pre-

ceding day, but less than three degrees change, was six

(—2—2—3—2—2-— 93)

From the above it is seen that on ten days of the twenty-

four in the table the temperature was either higher than the

preceding day (in four cases) or lower by not more than three

degrees. It seems evident, therefore, that the chief causes of

the great migratory movements that occurred on the days given

in the table were the favorable winds and not diminution of

temperature.

Time or Day DURING WHICH HAWKS MIGRATE.

When hawks occur in flights during the autumn migrations

they usually make their appearance some time after sunrise and

continue flying all day, the maximum of the flight occurring

in the forenoon in southeastern Connecticut and in the after-

noon near the New York state line. In the former locality

the sharp-shinned hawks begin to appear soon after sunrise and

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 747

the broad-winged hawks usually not until about eight o’clock,

unless there has been a favorable wind the previous day, in

which case the flight begins earlier. Many other species of

birds, — woodpeckers, thrushes, blackbirds, etc., — apparently

acting under the same influence that produces the migration

of hawks, namely, the favorable winds, make their appearance

before sunrise near the Connecticut shore and are abundant

for a few hours only, because they soon fly back into the

country to feed. The birds mentioned are known to migrate

at night. Most species of hawks, on the contrary, appear to

migrate during daylight only. This point is in agreement

with observations made by others.

THE MIGRATION OF THE BRoaAp-WiNGED Hawk.

The behavior of the broad-winged hawks when they are

migrating in flocks is peculiar. If, for example, these birds

are soaring together in a flock of from twenty-five to fifty,

one bird will be seen to separate from the flock, set its wings,

and sail away in the migration direction (west in southern

Connecticut). Presently another will follow at an interval of

a few hundred feet ; then another, until finally the entire flock

is observed to be sailing with set wings in single file. When

the hawks have sailed from a half mile to two miles this way,

they collect together and begin to soar again in an area of

about one or two acres and continue circling until they have

attained a considerable height. The peculiar manceuver is

then repeated.

The broad-winged hawk is the only one of the entire family

that shows a decided tendency to collect in large flocks during

the migration. Red-tailed hawks are sometimes observed in

small flocks.

HAWKS BLOWN OUT TO SEA.

Strong northwest winds are undoubtedly the cause of

many birds being blown out over the Atlantic Ocean and lost

during the southward migration. Hawks form no exceptions,

for instances like the following are not unfrequently reported :

748 THE AMERICAN NATURALIST. [Vor. XXXVI.

When the Munson line steamship Curitiba, bound for New

York, reached lat. 30° N- long. 70° 30’ W., on Oct. 11, 1900,

*five ospreys came on board; two of them were shot, one

was captured, one alighted on the after-deck boat and then fell

into the sea, and the remaining one fell down the funnel. At

dusk on the following day two more hawks were captured.”

These facts were given to the author of this paper by Captain

Hoppe of the Curitiba.

When the birds were first seen the Curitiba was 500 miles

from Florida and 400 miles from Cape Hatteras. According

to the U.S. Weather Bureau observations taken at 8 A.M. on

October ro, at Cape Hatteras, the wind was north, 28 miles per

hour, and on October 11, north, 14 miles per hour. At Boston,

on October 11, the wind was northwest, 24 miles per hour.

These strong offshore winds were unquestionably the cause

of the appearance of the hawks far out at sea. The Curitiba

also encountered a large number of bats which had evidently

blown out to sea by the northwest wind.

MIGRATING PERIODS OF VARIOUS SPECIES OF HAWKS IN

AUTUMN.

In the table on the opposite page the periods when a num-

ber of species of hawks migrate southward in the latitude of

Connecticut are given.

WIDESPREAD EFFECT oF STORMS.

It is well known that high north and northwest winds are

directly due to centers of low barometric pressure lying to the

eastward.

In the northern hemisphere the motions of the lower strata

of the atmosphere about centers of low barometric pressure

are counter-clockwise. These low areas are known as storm

centers, and in the eastern part of the United States they

moderate intensity is shown by Map III, which is an approxi-

mate representation of the U.S. Weather Bureau map of

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 749

TABLE V.

SPECIES. DATE. ABUNDANCE,

esu s OTA Me SOR Í HI dde

Fishhawk Sept. 1-Oct. 15 | Abundant.

(Pandion halietus caroli-

nensis

Marsh hawk Sept. I-Oct. ro. | Very common.

(Circus hudsonius)

Pigeon hawk Sept. 10-Oct. ro | Regular, but not common.

(Falco columbarius)

Sparrow hawk Sept. 1o-Oct. ro | Very common.

(Falco sparverius)

Sharp-shinned hawk Sept. 5-Oct. 15 | Very abundant.

(Accipiter velox)

_Cooper’s hawk Sept. 5-Oct.15 | Very common.

(Accipiter cooperi)

oshawk Oct. 25-Nov. 25 | Occasional, sometimes com-

(Accipiter atricapillus) mon in November.

Broad-winged hawk Sept. ro-Oct. 1 | Very abundant.

(Buteo latissimus)

Red-shouldered hawk Oct., Nov. (?) Common.

(Buteo lineatus)

Red-tailed hawk Oct. 20-Dec. 1 Very common.

(Buteo borealis)

September 18, 1900. The map shows the extent of coun-

try throughout which north or northwest winds prevailed at

8 a.m. on that date. These winds were reported through-

out an area of over 800,000 square miles, and their recorded

velocity varied from ten to forty miles per hour.

Owing to these winds a large southward flight of birds must

have occurred on the given date in eastern Canada and in the

eastern part of the United States.

This map is employed to show also the general migration

directions of various water birds in the eastern part of the

United States. The arrows 4A show the probable line of

flight of many ducks, swans, and shore birds from the interior

of Canada to the North Carolina coast. Those marked C show

the sea flight of many water birds from Labrador, as noted

in the Gulf of St. Lawrence region by the author, and those

marked B indicate the direction of the coast line migration.

750 THE AMERICAN NATURALIST. [Vor. XXXVI.

The converging arrows AA and C explain the abundance of

water fowl along the coast of North Carolina.

FLIGHTS oF HAWKS IN SPRING.

From the middle of March until the first part of May flights

of hawks occur along the Atlantic coast. These flights appear

to be greatest over the hills near the New Jersey coast, but occur

also at some distance in the interior. Near Paterson there is

a hill about 500 feet high, part of the Watchung range, over

which large numbers of hawks pass in the spring. On the west

slope of this hill many pits and brushwood blinds are made every

year, both in the woods and in the open ground, and are occu-

pied during March and April by men and boys who make a prac-

tice of killing hawks for New York and New Jersey taxidermists.

The author has witnessed several flights of hawks on this hill,

which is the first high land back of the coast.

The most favorable wind for a flight is west, or a little south

of west. The red-tailed hawks are the first to appear in the

spring, and the sharp-shinned and the broad-winged hawks are

thelast. The periods when some of the hawks may be expected

are given below :

. Fishhawk, latter part March to May r.

Marsh hawk, April.

Sharp-shinned hawk, April 10-May Io.

Cooper's hawk, April.

Broad-winged hawk, April 10-May 5.

Red-tailed hawk, March 1-April ro.

—

Observations described above seem to warrant the following

conclusions :

I. That the migratory movements of hawks are largely deter-

mined by the direction of the wind, hawks regularly depending

. on favorable winds as a help in migration.

2. That an adverse wind not only retards the migratory

movement, but that it almost completely arrests it.

3. That the migratory period of the various species of hawks

lasts for from about fifteen days to one month ; during this time

the migratory movements take place on days when favorable

winds occur.

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 751

4. When the wind is favorable and approximately parallel to

the direction of migration, hawks fly and sail at a "eu altitude

and occasionally soar in circles.

5. When the wind is favorable but sna perpendicular to

the migratory direction (the favorable component being small),

riot

Map III. — EASTERN CANADA AND EASTERN UNITED STATES.

The dotted lines are the isobars of September 18, 1900. A low barometric pressure is

centered off the Maine coast. The small arrows indicate the direction ot the wind, and

its velocity in miles «si hour is given in a number of cases. The mean velocity of the

wind records on the map as it is given here is over seventeen miles per hour.

hawks fly low and soar continually, often alternating soaring

with the wind and flying or sailing nearly against it.

6. That hawks migrate during the daylight, and, other con-

ditions being the same, they are most abundant in migratory

flights when the atmosphere is clear.

752 THE AMERICAN NATURALIST. [Vor. XXXVI.

. When a migratory flight of hawks takes place, continued

favorable winds exhaust the number of hawks ready to make the

migratory journey, but a second favorable wind about one week

later may cause a second flight equal in magnitude to the first.

Map IV.—Coast LINE OF SOUTHERN NEW ENGLAND, NEW JERSEY, AND

DELAWARE

The long arrows indicate «ms direction of migration of hawks and various other birds in the

spring, when the wind is west or southwest. Owing to the fact that the coast line is

nowhere at right ang o the direction of migration, the flight is not in a narrow

emet as along the Cometic shore in autumn, but ten or fifteen miles wide in the

m part of New Jersey.

The arrows marked A = a "e up the Hudson enya -- marked Z a flight up the

Housatonic valley i and those marked eneral northeast flight in New

England. The haies range, along which many leoi s migrate, is indicated by D.

8. That a favorable wind, when the favorable component is

small, may cause decided deviations of the course of the migrat-

ing birds from the main migratory direction.

No. 429.] RELATION OF WIND TO BIRD MIGRATION. 753

These conclusions apply to both the northward and south-

ward migrations. Most of the effects of the winds stated are

intensified when the wind is strong.

It has been observed that after a period of adverse or light

winds, during which no migratory movement was noticed, a

moderate wind in a direction favorable to the migration causes

a marked migratory movement, especially if such a wind is gen-

eral and not local. It thus seems evident that birds wait for a

favorable wind to carry them on their migrations.

Changes of temperature, without doubt, affect the migratory

movements of hawks, but, as has already been shown, a number

of large flights have occurred in autumn on days when rises of

temperature occurred near where the hawks started on their

journeys. Moreover, flights of hawks often occur on cool days

in spring, lower temperature usually accompanying west winds

at that season of the year.

Water birds, particularly the Limicolz, migrate in large num-

bers from the north in July, before the hottest period of the

summer has passed. It is not surprising, therefore, to find

causes affecting the time of migration of birds other than

changes of temperature.

It is probable that many varieties of birds regularly make

use of the wind as a physical agent in carrying them on their

migratory journeys, the habit being well formed in the case

of the Falconide.

PHYSICAL LABORATORY, COLUMBIA UNIVERSITY,

NEW York.

NOTES AND LITERATURE,

ZOOLOGY.

North American Deer. — The Deer Family! by Theodore Roose-

velt and others, is a popular account of the deer found in North

America, written by men eminently fitted by experience and personal

knowledge to deal with the subject from the double viewpoint of

the naturalist and sportsman. The first half of the book is by

Mr. Roosevelt, who furnishes the introductory chapter (pp. 1-27),

and treats of the mule deer, or Rocky Mountain blacktail (pp. 28-64),

the whitetail deer (pp. 65-97), the pronghorn antelope (pp. 98-1 30),

and the wapiti, or round-horned elk (pp. 131—164) ; Mr. Van Dyke

writes of the deer and elk of the Pacific coast (pp. 167—191),

Mr. Elliot of the caribou (pp. 257—287), and Mr. Stone of the moose

(pp. 288—325). Mr. Roosevelt, through his many years of ranch

life on the Little Missouri and his numerous protracted hunting trips

in the great game regions of the West, writes not only from abundant

opportunity for observing the habits of the animals, but with the

sympathy and intelligence of a keen naturalist as well as an ardent

Sportsman.

In the introductory chapter reference is made to the misapplication

of names * by all peoples of European descent who have gone into

strange lands," not only in reference to some of our deer and prong-

horn, but to other game animals and birds. The deer of North

America are grossly classified as forming six kinds: ‘The moose,

caribou, wapiti, whitetail, and the two blacktails," minor differences

being very properly disregarded. Their geographical ranges and

leading characteristics are briefly stated, with some comment on the

€quipment required by the hunter of large game.

There is reference also to “the most striking and melancholy

feature in connection with American big game, — the rapidity with

! Roosevelt, Theodore; Van Dyke, T. S.; Elliot, D. G.; and Stone, A. J.

The Deer Family. New York, The Macmillan Company, 1902. 8vo, ix + 334 PP-

10 illustrations by Rungius, 7 maps by C. Hart Merriam, and numerous half-tone

cuts of antlers of moose and caribou. American Sportsman’s Library Series,

edited by Caspar Whitney.

| 755

756 THE AMERICAN NATURALIST. [VoL. XXXVI.

which it has vanished ”; and, says Mr. Roosevelt, “ if we are a sensible

people, we will make it our business to see that the process of extinc-

tion is arrested. At the present moment the great herds of caribou

are being butchered as in the past the great herds of bison and

wapiti have been butchered. Every believer in manliness, and

therefore manly sport, and every lover of nature, every man who

appreciates the majesty and beauty of the wilderness and of wild

life, should strike hands with the far-sighted men who wish to pre-

serve our material resources, in the effort to keep our forests and our

game beasts, game birds, and game fish — indeed, all the living

creatures of prairie, and woodland, and seashore — from wanton

destruction. ... But this end can only be achieved by wise laws

and by resolute enforcement of the laws."

The work is thoroughly non-technical in character, even to the

omission of the scientific names of the species throughout most of

the chapters, yet it loses nothing in scientific interest or value

thereby. As a contribution to the natural history of the big game of

North America its importance cannot easily be overestimated, there

being here brought together more first-hand knowledge of the life

histories of the species treated than has ever before been placed on

record. Mr. Rungius's illustrations are effective and appropriate,

while the numerous half-tone cuts of heads and antlers of moose and

caribou are important and interesting features. The seven maps, by

Dr. Merriam, showing the ranges of the deer, elk, and pronghorn in

the year 1900, have a melancholy interest and great permanent

value. That showing the present range of the elk is a sad com-

mentary on the wasteful destruction of big game in North America.

A hundred years ago this animal ranged from near the Atlantic sea-

board westward to the Pacific coast, and from southern Canada

into Mexico, forming, in some sections of the country, bands of

thousands. Now its range is restricted to a narrow strip of country

in the main Rocky Mountains, from Alberta to the northern

border of New Mexico, with a few isolated areas of very small

extent in California, Oregon, Arizona, Montana, North Dakota, and

Minnesota.

The Deer Family is thus not only a book of special interest io the

sportsman and the general reader, but an invaluable contribution to

the natural history of the deer tribe in North America. It should

also stimulate interest in the protection by all rational means of the

waning remnants of the former great herds of these noble and

attractive creatures. hob

No. 429. NOTES AND LITERATURE.

9 757

Upland Game Birds.'— This volume, by Edwyn Sandys and T. S.

Van Dyke, is the second in the American Sportsman's Library

Series, edited by Caspar Whitney, and, like the first, Zhe Deer

Family, by Roosevelt and others, shows the editor's good judgment

in the selection of his authors for this noteworthy series of books.

The scope of the work includes not only the turkey, grouse, partridges,

and mourning dove, as would be expected, but also the cranes,

woodcock, the upland and golden plovers, and introduced “foreign

game," but not the snipe. Although written ostensibly for sportsmen

by sportsmen, it has a much broader interest. Its attractive style,

varied incident, and personal reminiscences must render it attract-

ive to the general reader, and especially to those interested in the

wild things of nature. The successful sportsman is necessarily a

keen observer, and long experience in his craft renders him familiar

with the ranges and life histories of his favorite objects of pursuit ;

but few have the literary gift to impart attractively to others the fund

of information gained through such varied and extensive opportuni-

ties of observation. The authors of the present volume are well

equipped for their task, as regards both experience and literary

ability.

Mr. Van Dyke's portion of the work (pp. 377—41 7) relates exclu-

sively to * The Quail and Grouse of the Pacific Coast," with which

he shows himself especially familiar. The rest of the field (pp. 1-376)

is covered by Mr. Sandys, whose several chapters contain much that

is detailed and explicit regarding the habits and peculiarities of the

game birds he treats, and in many ways admirably supplement the

much that has been written by ornithologists. The spirit of the natu-

ralist pervades the pages of Upland Game Birds, although they are

amply enlivened by personal reminiscences that should prove espe-

cially attractive to the sportsmen ; and the authors also neglect no

opportunity to castigate the “ game hog," and to urge more effective

protection for game birds. Five of the nine very effective and

pleasing illustrations are by Mr. L. A. Fuertes. TAA

Nestlings of Forest and Marsh,? by Irene Grosvenor Wheelock,

contains accounts of the home life of about twenty birds, most of

! Sandys, Edwyn, and Van Dyke, T. S. Upland Game Birds. New York,

The Macmillan Company, 1902. 8vo, ix + 429 pp., frontispiece and 8 full-page

plates. American Sportsman’s Library Series, edited by Caspar Whitney.

2 Wheelock, Irene Grosvenor. Nestlings of Forest and Marsh. Chicago,

CA C. McClurg & Co., 1902. 8vo, xvi + 257 pp., 22 pls., text-figs.

758 THE AMERICAN NATURALIST. [Vor. XXXV I.

them common and familiar, e.g., the oriole, robin, crow, brown

thrasher, a few less well known, such as the long-billed marsh wren

and the yellow-headed blackbird. By commendable energy and

patience the author has obtained notes describing the selection of

the nesting site, the search for materials for building, the care of the

young, and their efforts, successful or tragic, to make a start in life.

In her “ Foreword " the author claims that her book is “as accurate

as careful observation in the field can make it"; nor does anything

in the book seem inconsistent with this claim. It is not, however,

safe to accept all her conclusions from the facts recorded. She says

further : * So far as reading human characteristics into animal life is

concerned, can any one tell where the brute ends and the human

begins? Many of the emotions of man's heart find their counter-

part in the life of birds. "That we do not perceive this proves only

how dull is our sight." The most hazardous attempt to read bird

minds is the account, on pp. 119 and 120, of a supposed successful

attempt of chickadees to poison a young one which had been made

a captive. The story is told in the best faith, but it is an excel-

lent example of what Prof. Lloyd Morgan has felicitously termed

*the inability to distinguish the observed fact from the observer's

inference." It is curious that the author seems wholly ignorant of

Professor Herrick's book, which has laid the foundations of that

part of the study of birds which she has chosen for her field. An

acquaintance with Professor Herrick's work would have saved her

from assuming that a parent bird “seemed to know instinctively

which one [of the nestlings] had been fed." The study of the marsh

birds is the most interesting part of the book, but several of the other

chapters contain bits of valuable information, e.g., the account of the

destruction by red-headed woodpeckers of the nests and young of

cliff swallows. In an attempt to write brightly and entertainingly,

the author too frequently sacrifices dignity, nor can her humor be

said to be of a high order. The illustrations are, with a few excep-

tions, poor, though it should be said that this fact is partly due to the

evident reluctance of both the author and the photographer to

interfere with the home life of the birds under observation.

_A New Genus of Nemerteans.' — An important paper by Miss

Thompson describes the anatomy, histology, and relationships of

a new heteronemertean, of especial interest because it apparently

1 Thompson, Caroline B. Zygeupolia litoralis, a New Heteronemertean, Proc.

-~ Acad. Nat. Sci. Philadelphia (December, 1901), pp. 657-739, Pls. XL-XLIV, 1902.

No. 429. NOTES AND LITERATURE.

9 759

forms a connecting link between two formerly well-differentiated

families. The species Zygeupolia litoralis is not uncommon at

Woods Hole, Mass., and because of its transparency would make

an admirable form for class study wherever the living worms can be

obtained. Most of the principal features of its anatomy can be

demonstrated, on the living worm under the low power of the

microscope.

The general color of the body appears to be independent of sex

and of sexual maturity, while that of the intestinal caca depends

upon the amount and character of the food therein. In external

appearance the species so closely resembles Micrure ceca, with

which it is sometimes associated, in size, shape, and color, “that the

two can scarcely be distinguished withoüt a hand lens which reveals

the absence of lateral slits in the former and their presence in the

latter.”

In the excellent and detailed description of the different organ .

systems the following points are of especial importance: The pro-

boscis has no retractor muscle, its posterior end lying entirely free

in the rhynchoccel, exactly as has been described for Cerebratulus

lacteus? The musculature of the proboscis consists of two principal

layers arranged as in many of the Lineidz, and there are also more

or less distinct muscular crosses so characteristic of this family.

A strong layer of circular muscles surrounds the posterior end of

the stomach, but ends abruptly at the beginning of the intestinal

region. This forms a definite inner layer of circular muscles which

the author looks upon as homologous with the inner circular layer of

Carinella, and which is so highly developed in exactly the same

region in Carinoma. Her views are well supported by diagrams

and figures, and seem to be conclusive. A similar, but less highly

developed, muscular layer has been described for Micrura.

The cesophagus exhibits two well-marked regions differing greatly

in histological features, the posterior cesophagal cavity, or stomach,

being lined with cells which resemble far more closely those found

in the intestine than those of the cesophagus proper. This suggests

that the anterior cesophagal cavity only may have arisen from the

ectoderm, while both stomach and intestine are derivatives of the

entoderm.

The caudal cirrus, or caudicle, has been studied in detail, and the

statements of various observers who have described this organ erro-

neously are corrected. The posterior opening of the intestine lies

1 Coe. Trans. Connecticut Acad., vol. ix (1895), p. 488.

760 THE AMERICAN NATURALIST. [Vor. XXXVI.

above the base of the cirrus and not, as has sometimes been

described, beneath it. In section the caudal cirrus shows the outer

epithelium, a thin circular and comparatively strong longitudinal

muscular layer, and a zone of mesenchyme cells surrounding a large

central blood space. The two lateral nerves are here situated in the

outer epithelium, the outer longitudinal muscular layer and cutis

being absent. Neither do the gonads, intestine, nor proboscis sheath

continue back into this organ. This description agrees closely with

the characters given by Punnett for the caudal cirrus of Micrella

rufra, except that he finds only rudimentary blood vessels.

The distribution of cutis glands has been very fully studied. A pair

of neurochord cells was found in the ventral ganglia. A longitu-

dinal groove on each lateral margin of the body just anterior to

the intestinal region is looked upon as a sense organ which may

be homologous with the lateral sense organs recently described by

Punnett for Micrella,? where they are situated just back of the

excretory pore, as are the well-known sense organs in Carinella.

Blood lacunz were not found anterior to the brain; lateral blood

vessels are united at intervals below the cesophagus. A large lacuna

without definite walls passes into the caudal cirrus. Parasitic greg-

arines were found in the intestinal epithelium and in the ova.

Infected eggs grew to several times their normal size.

The evidence that the genus Zygeupolia should be placed among

the Lineidz seems conclusive, the only feature distinguishing it

from other members of the family being the absence of cephalic

furrows. It is rightly regarded as the most primitive or aberrant mem-

ber of that family which has yet been described. With Punnett’s

new genus Micrella there is certainly a close relationship, and both

forms serve to bridge over the gap between the Eupolida and the

Lineide. In many points of anatomy the two genera are very

similar, and while in Zygeupolia the cephalic furrows are wanting,

yet an approach to this condition is found in Micrella, where the

furrows reach only half way to the brain. Punnett's new genus

Oxypolia likewise forms a connecting link in the series, although

this clearly belongs to the Eupolidz, being more closely related to

Valencinia. W. R. C.

! Punnett. Quart. Journ. Micr. Sci., vol. xliv (1901), p. 553-

? Punnett. Loc. cit., p. 551.

No. 429.] NOTES AND LITERATURE. 761

BOTANY.

Elementary Plant Physiology.

revised edition of the same author's earlier book, Lxferimental

Plant Physiology, will without doubt be welcomed by the teacher.

The experiments chosen are almost all of the simpler kind,

being qualitative rather than quantitative, for which reason they will

especially recommend themselves for use in elementary courses.

In the introductory chapter is given a series of selected courses

which will serve as useful suggestions to the inexperienced or hurried

teacher. At the same time the number of experiments described is

sufficiently large to allow of independent choice, so that any course

selected from the book may have a certain elasticity — a point of no

little importance.

In starting out with the consideration of growth the author has

undoubtedly chosen that phase of plant physiology which appeals

most strongly to the average student, a fact which warrants the

choice, when it is remembered that the object is to introduce the

beginner to the subject. The chapters on the exchange of liquids

and gases and on the general subject of nutrition are full and seem

entirely adequate for the scope of the book. The wisdom of leaving

the consideration of the effects of stimulation and correlation (the

various trophic phenomena) to the last, in a place removed from the

related topic of growth, might be questioned if the evident intention

of the author were not taken into consideration. It is clearly his

object to impress most strongly upon the student the phenomena

connected with general and, on the whole, less complicated problems

of plant physiology and to touch lightly upon the more intricate ones

connected with correlations of growth. There are many reasons jus-

tifying such a treatment in an elementary course, particularly in a

course which presumably would not be accompanied by many

explanatory lectures or by much reading.

The text is terse and clear, the typography and form of the book

excellent, both the author and publisher are to be congratulated on

having produced a very acceptable and practically useful laboratory

aide. HMR

Notes.— A new botanical publication, the Recueil de /’ Institut

Botanique ( Université de Bruxelles), has been launched by Professor

1 MacDougal, D. T. Elementary donee Physiology. New York, Longmans,

Green & Co., 1902. 8vo, xi + 38 pp. I gs.

762 THE AMERICAN NATURALIST. | [Vor. XXXVI.

Errera, after the plan of the “Contributions ” from various other

scientific establishments and laboratories. It is to contain original

contributions and reprints of studies from the laboratory that have

been published elsewhere. Rather curiously, it begins with Vol. V,

but the first four volumes of the series are definitely planned and

are promised for the early future. In form and style the Recueil

is excellent. As would be expected from Professor Errera's own

studies, the contents of this volume, as well as of the other four that

are promised, are largely physiological or cytological.

Under the title 72e Mew Phytologist, a new British journal, edited

by Professor Tansley, has made its appearance in London. Among

other things, the first number contains *A Revision of the Classifi-

cation of the Green Algae," by Blackman and Tansley.

A new botanical journal, entitled Magyar Botanikai Lapok ( Un-

garische Botanische Bilitter), is published at Budapest, beginning

with January of this year.

The index to new species and varieties published in Engler's

Botanische Jahrbücher, Vols. XXVI- XXX, occupies 45 double-

column pages of the concluding number of the last-named volume.

The April Bulletin of the Torrey Botanical Club contains the fol-

lowing papers: Salmon, “Supplementary Notes on the Erysipha-

cem"; Cook, “Development of the Embryo Sac and Embryo of

Castalia odorata and Nymphea advena" ; Piper, “New and Note-

worthy Northwestern Plants, VI"; Arthur, * New Species of Uredi-

nex, II"; and Rydberg, * Studies on the Rocky Mountain Flora,

Vink.”

The Ottawa Naturalist for March contains papers on the botany

of the West and Northwest by Macoun and Greene.

No. 2 of the Biltmore Botanical Studies, dated April 30, 1902;

contains the following papers: Beadle, *New Species of Thorns

from the Southeastern States, IL"; Boynton, “Studies in the Genus

Amorpha" ; Boynton, *Two New Southern Species of Coreopsis” ;

Boynton, “Notes from a Collector's Field Book"; Harbison, “A

Sketch of the Sand Mountain Flora" ; Harbison, *New or Little-

Known Species of Trillium, II" ; Beadle, * Studies in Philadelphus ie

Beadle, «Two Drupaceous Ties pU australis and P. mitis)

from Alabama."

The January number of the Buletin of the Wisconsin Natural

ies Society contains the following botanical articles: Brunchen,

* Studies in Plant Distribution” ; ; Graenicher, * Flowers adapted

No. 429.] NOTES AND LITERATURE. 763

to Flesh-Flies”; Bennett, * Additions to the Flora of Milwaukee

County"; and Brown and Fernekes, * Contribution toward a List

of Milwaukee County Fungi."

A number of illustrations of botanical interest are contained in

the recently published seventh Report of the Forest, Fish and Game

Commission of New York

The Engelmann Botanical Club of St. Louis has issued an excel-

lent bulletin on tree planting, giving practical instruction as to what

not to do, as well as what to do, in attempting to beautify a city by

the growth of trees,

Dr. Urbina, of the Museo- Nacional of Mexico, has distributed

from the Anales of that institution a botanical study of the various

Zapotes of Mexico.

In a separate recently issued from the Z»*azsactions of the Royal

Society of Canada, Dr. Fletcher writes on the value of nature study

in education.

The principles of generic nomenclature are discussed by Mr. Shear

in the Botanical Gazette for March.

The leaf forms of Liriodendron are discussed and figured by

Berry in Zorreya for March.

An account of the acaulescent violets of central New York, by

H. D. House, is published in Zorreya for May.

A new species of Viola, V. angelle, from New Jersey, is described

by Polland in Torreya for February.

A yellow-fruited form of Zex myrtifolia is noted by R. M. Harper

in Zorreya for March.

Ailanthus grandis, a new Indian species, is described and figured

by Prain in Zhe Indian Forester for April ; he also gives comparative

notes on the other species of the genus and their distribution.

The morphology and anatomy of Azorella Selago are treated

by Charlotte Ternetz in Heft 1 of Abtheilung I of the Botanische

Zeitung for this year.

Continuing his studies of Ochnacez, Van Tieghem, in the April

Journal de Botanique, separates from the old genus Ochna the new

genera Ochnella, Porochna, and Discladium, giving differential keys

for the genera of the tribe Ochnez as so constituted.

A new Zauschneria, Z. arizonica, is described by Dr. Davidson in

the Bulletin of the Southern California Academy of Sciences for Janu-

ary, — the initial number of this publication.

764 THE AMERICAN NATURALIST. [VoL. XXXVI,

Dr. Weber, who in former years studied Opuntias with Engel-

mann, has issued separates, from the Buletin de la Société Nationale

d’ Acclimatation de France, of an article on the species (O. /eucotricha)

which furnishes the prized “ Duraznillo,” or peach prickly pear

of the high table-lands of Mexico, and its immediate relatives.

An interesting account of the hallucinations caused by taking an

infusion of Axhalonium lewinii is published by Havelock Ellis in the

Popular Science Monthly for May.

A photogram of Zchinocactus texensis is published in the Monats-

schrift fur Kakteenkunde for April 15.

In No. 7 of the current volume of botanical Proceedings of the

California Academy of Sciences, Miss Eastwood gives a key to fifty-

seven species and varieties of Ribes recognized as occurring on the

Pacific coast, and describes nine species as new.

Matsumura publishes a list of the wild and cultivated Leguminosz

of Japan, Loochoo, and Formosa, in the Tokyo Botanical Magazine

of March 20,

M. Theuriet, an amateur of roses, published an illustrated classi-

fied list of the 6781 species and varieties cultivated at l'Hay, France.

Blanchard has a note on the Vermont chokeberries in Rhodora

for March.

In Pharmaceutical Archives for April, Mr. Denniston publishes a

paper on the general and bark characters of Viburnum ellipticum.

A large and fully illustrated paper on the comparative embryology

of the Rubiacez, by F. E. Lloyd, constitutes Vol. VIII, No. 1, of

the Memoirs of the Torrey Botanical Club.

An account of Primula parryi, with an excellent half-tone illustra-

tion, is published by Knowlton in Zhe Plant World for February.

Scrophularia glabrata, from Arizona, is described and figured by

Davidson in the March Bulletin of the Southern California Academy

of Sciences.

The seeding of Plantago fastigiata is the subject of a paper by

Griffiths in the Buletin of the Torrey Club for March.

Peperomia davisii, from St. Kitts, is described by Britton in

Torreya for March. -

A cockscomb fasciation of the pineapple is noted by Harshber-

. ger in the concluding part for 1901 of the isum d of the Academy

jx of Natural Sciences of Philadelphia.

No. 429.] NOTES AND LITERATURE. 765

In the Botanical Gazette for April, DoNNELLIA C. B. Clarke is

proposed for the reception of Calisia grandiflora Donnell Smith.

The new genus constitutes one of the many segregates of Trade-

scantia.

An interesting account of the harvesting, curing, and cleaning of

the seed of oa pratensis, the standard lawn grass of the United `

States, forms Buletin No. 19 of the Bureau of Plant Industry of

the Department of Agriculture, by Pieters and Brown.

E. D. Merrill publishes some notes on Sporobolus in Rodora for

March

A study of the Zamias of Florida, by Wieland, is published in ZZe

American Journal of Science for May.

The North American genera of Aspidiez are discussed by Under-

wood in the Budletin of the Torrey Botanical Club for March. A key

is given to the genera.

The evergreen ferns of New England are considered by Daven-

port in Ržodora for March.

In The Fern Bulletin for April Mr. Maxon proposes the restora-

tion of Liebman’s name Asplenium resiliens for what is commonly

known as A. parvulum.

In an article on * The Logfern” (Dryopteris Cristata Clintoniana),

Mr. William Palmer gives a key to the cristata and Goldieana groups

of that genus, in Zhe Fern Bulletin for April.

The mechanical expulsion of the brood bodies of several species

of Lycopodium is noted by Lloyd in Zorreya for February.

Several species of Lycopodium, and the propulsion of the gemmz

of some species, are considered by Mr. Leavitt in Rhodora for

March.

The tenth of Mr. Alvah Eaton’s papers on the genus Equisetum

in North America, in Zhe Fern Bulletin for April, deals with the

varieties of Æ. Zitorale.

A monograph of the Lejeunez of the United States and Canada,

by Evans, constitutes Vol. VIII, No. 2, of the Memoirs of the Torrey

Botanical Club.

Two new western mosses are described and figured by R. S.

Williams in the February Bulletin of the Torrey Club.

766 THE AMERICAN NATURALIST.

PETROGRAPHY.

Weinschenk’s *«Die Gesteinsbildenden Mineralien ” ' affords the

most satisfactory introduction to petrography that has yet appeared.

Its first 47 pages are devoted to the physical and chemical methods

employed in the separation of rock constituents and in the determi-

nation of the character of isolated mineral particles. The remainder

of the volume describes briefly the principal rock-forming minerals

as they appear under the microscope. In spite of the brevity of the

treatment necessitated by the small size of the book nearly all the

essential facts relating to the characteristics of the individual min-

erals are to be found in its pages. The reader is supposed to be

already acquainted with the general principles of optics, so no space

is devoted to this subject.

The special feature of the book is the collection of r8 plates,

bound in two brochures and enclosed in a pocket from which they

may be removed for ready reference. They contain summaries of

the physical, chemical, optical and crystallographic characters of

the minerals discussed in the text ; a list of minerals classified accord-

ing to their color, in thin sections, and their crystallographic habits ;

lists of those exhibiting optical anomalies, twinning structures, dis-

tinct cleavages, and of those whose powder reacts alkaline ; and lists

in which the minerals are arranged according to their magnetic

strength, their solubilities, their fusibility, their density, their refract-

ing indices, the strength of their double refraction, and their optical

character. ‘These tables are so well arranged and they embody

such a mass of information in small compass that they will prove

not only useful to younger students but also extremely valuable to

those of maturer experience.

So far as the writer knows, the present volume is the only modern

elementary treatise on the microscopical characters of rock-forming

minerals that can safely be recommended to beginners in the study,

Ww. S. B.

E nk, Ernst. Die Gesteinsbildenden Mineralien. | Freiburg, Herdersche

Verlagshandlung, 1901. 146 pp., 100 figs., 18 tables.

(Vo. 428 was mailed August 19.)

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ait

The American Naturalist.

ASSOCIATE EDITORS:

J. A. ALLEN, PH. D., American Museum of Natural History, New York.

J. H. COMSTOCK, S.B., Cornell ieri 1i

"WILLI I

ALES HRDLICKA, M.D., A:

-. . S. JORDAN, LL.D, Stanford U. eraty

- CHARLES A. KOFOID, Px.D., Lion ‘of Cali ifornia, Berkeley.

ake For.

LHE

AMERICAN NATURALIST

VoL. XXXVI. October, 1902. No. 430.

HISTORICAL EVIDENCE AS TO THE ORIGIN OF

THE PAIRED LIMBS OF VERTEBRATES.

BASHFORD DEAN.

WHATEVER be the results of embryologists, the real test of

the problem of the mode of origin of the paired limbs of verte-

brates belongs obviously to paleontology. For if the paired

limbs, as most morphologists believe, arose as highly specialized

lappets of a continuous lateral skin fold, it is quite evident

that sooner or later these conditions will be demonstrated

convincingly among the earlier fossil fishes. In point of fact,

the paleontological evidence bearing upon this matter is of

such great value, even at the present time, that we can justly

regret that it has received scanty attention at the hands of

embryologists. It is indeed so imperfectly noticed in the

recent voluminous papers which bear upon the general problem

of limb origin that I have been led to the following comments.

I. THE WEIGHT oF EVIDENCE.

If we accept the general rule that earlier forms are of greater

value for purposes of determining primitive characters than are

later ones, and if we admit that a study of many genera and

767

768 THE AMERICAN NATURALIST. | [Vor. XXXVI.

many species is of greater value than an examination of few

specimens of few species, we must, I believe, conclude that the

doctrine of the origin of the paired limbs from lateral dermal

folds rests upon strong foundations. Reducing to simplest

terms our knowledge of the earliest sharks or shark-like fishes,

among which the ancestral jaw-bearing vertebrates are reason-

ably sought, we find that of forty odd species, representing

fifteen genera (Paleozoic), whose fin characters are known, all

but those of a single genus, and possibly a second, are dis-

tinctly in favor of the lateral-fold theory. And the exceptions

to the rule are the most recent forms and in important struc-

tural regards the least primitive.

More concretely, such genera as the Upper Silurian and

Lower Devonian Parexus (two species), Climatius (eight spe-

cies), Diplacanthus (five species) (=family Diplacanthide),

Cheiracanthus (three species) (Acanthodida), and the long-

enduring Acanthodes (twelve species) (Lower Devonian to

Permian); as the Upper Carboniferous Acanthodopsis (but we

may put the last aside, since its fin structures are not as well

known as in the neighboring forms); as the Permian Prota-

canthodes and Traquairia, — all of these ancient sharks have

their paired limbs in the form of distinct longitudinal skin folds,

and these paired fins, furthermore, are essentially similar to the

unpaired ones. This conclusion is based upon numerous and

well-preserved specimens. The next ancient sharks whose fin

structures are described are the (now accepted as) Upper

Devonian cladoselachids (about two genera and six species).

These have the paired fins extended along the body, their

hinder web narrowing gradually and merging with the body

wall, showing no trace of a lobate fin base projecting from the

body. Here, too, the paired fins closely resemble the unpaired

fins. Structural details in these forms are known from a score

or more of well-preserved specimens. A kindred Upper Devo-

. nian shark is Ctenacanthus, whose fin structures have hitherto

been unknown; but an interesting specimen of this form, show-

ing pectoral ám and spine zz 5274, has now been discovered, and

_ it shows convincingly that this shark had a fin-fold type of fin.

he radials pass to the margin of the fin, as in Cladoselache,

No. 430.] PAIRED LIMBS OF VERTEBRATES. 769

each with a corresponding basal; unlike cladoselachids, there

are no intercalated rays. Next ancient is Traquair’s specimen

of Cladodus (Lower Carboniferous) (cf Fig. 1), whose pectoral

gives evidence of a line of basal plates extending backward like

T T

Fic. 1. — The Paleozoic shark Cladodus neilsoni Traquair. The fossil is shown surrounded

mtem e teme f Cladoselache of of similar size, — indicating that the fins of

a rayless axis of an “archipterygial’’ fin. This specimen has

been taken in evidence of the non-primitive nature of the fin-

fold type of limb (vide infra). Last of the ancient forms are

the Permian symmoriids and xenacanthids. Of these Symmo-

| Tium, judging from the single fragmentary pectoral described

T70 THE AMERICAN NATURALIST. [Vor. XXXVI.

by Cope, is similar to cladoselachids, but remarkable in that its

hindmost rays apparently coalesce at their bases so as to form

a stout supporting plate. There is no evidence, however, that

this projected from the body wall, nor have we any knowledge

of pelvic or unpaired fins. In xenacanthids, on the other hand,

many and well-preserved specimens (of two or three species)

show a biserial archipterygium in the pectoral fin, and this

condition has been the main paleontological stay of those who

oppose the fin-fold hypothesis. They lay no emphasis, how-

ever, on the late appearance of this form (for there is no

evidence forthcoming that the Coal Measure “ Xenacanthus ”

had fins like the Permian form), or on the facts that its ven-

tral fin is zo£ a biserial archipterygium but resembles closely

the unpaired fins immediately behind it, and that this form has

already developed a mixipterygium, and has specialized an

amphistylic attachment of jaw to skull, and that it has largely

lost its integumental defenses.

Balancing the matter fairly, I think that we must conclude

that the weight of the evidence of Paleozoic sharks falls

emphatically on the side of the longitudinal fin-fold hypoth-

esis. Indeed the only way, as far as I can see, in which this

evidence can be denied is by a frank rejection of the well-

accepted view as to the primitive nature of the earliest elasmo-

branchs (Proselachii).

Il. Tue Fin-Fotp CHARACTER OF THE EARLIEST FORMS,

ACANTHODIANS AND CLADOSELACHIDS.

The dermal-fold nature of the paired fins of the acantho-

dians requires little comment. Excepting in a brief note of

Dr. Fritsch,! no attempt has ever been made to explain them

in accordance with the theory of Gegenbaur ; that is, that their

origin is due to outgrowth of elements belonging to serially

homologous gill bars. Continuous fin folds they certainly are;

1 Fauna d. Gaskohle Formations Bóhmens (1893, p. 71), in which the spine le

regarded as the archipterygium. Dr. Jaekel homologizes shoulder girdle to gill

arch, but does not consider the relations of the fins themselves. Verhandl.

Deutschen Zool. Gesell. (1899), pp. 256-258.

RUN

No. 430.] PAIRED LIMBS OF VERTEBRATES. 771

whether or not we accept the origin of their supporting elements

as equivalent to the coalesced radial supports of a more typi-

cal shark-like fin. On the other hand, the paired fins of the

cladoselachids are distinctly shark-like, although in pattern

unlike those of later selachians. They are remarkable in that

they are built on the plan of balancing organs; ze, the fin

was attached keel-fashion, along the side of the body, and prob-

ably functioned by means of ventrad and dorsad movements

(undulatory at fin margin ?). And the blade of the fin could

not be rotated (twisted) sideways. This is attested structur-

ally by (1) its graded series of radials, — larger, most numer-

ous, and most crowded in the anterior region ; (2) by the absence

of a metapterygial lobe, and in the continuation of the fin web

hindward along the margin of the body ; physiologically (3) by

its close analogy with the dorsal and anal of such a form as

Mola, in which we know that this undulatory movement has

been specialized. That the paired fins of Cladoselache, more-

over, cannot be interpreted as **due to bottom living " on the

plan, e.g., of Rhina, becomes clear when we consider that the

radial cartilages are unjointed throughout their length, that

the ventral and pectoral fins are not drawn together in a ray-

like way, and that the fins, in spite of their remarkable charac-

ter, are as small in proportion to the size of the body as those

of modern selachians (Pristiurus, Scyllium), which are known

to be free swimming. (Cf. Natural Science, Vol. VIII (1896),

pp. 250-252, and Anat. Anzeiger, Bd. XI, pp. 673-679. Vide

also infra.)

III. Tue Evipence or PALEONTOLOGY AS TO THE DEVEL-

OPMENT OF THE UNPAIRED FINS.

In recent work tending to support the hypothesis of Gegen-

baur no reference is made to the mode of origin of the

unpaired fins. This, we must grant, would have an important

bearing upon the problem of the extremities if evidence existed

to show that in a late stage in the evolution of the unpaired

fins an archipterygium arose, me that of the paired fins. As

a matter of fact, paleontol trates that such a condition

772 THE AMERICAN NATURALIST. [Vor. XXXVI.

is represented in the case of the ancient ganoids (crossop-

terygians); for in certain Devonian forms (cf also Mesozoic

genera) the dorsal fins show clearly concrescence of radial and

basal elements in a way which parallels strikingly the biserial

archipterygium (of the paired fins)! This condition, although

well known to the paleontologist, appears to have escaped the

notice of the adherents to the Gegenbaurian school. It is per-

fectly evident that if the biserial archipterygium arises in the

unpaired fin from the concrescence of lateral fold-like support-

ing elements, the similar structure in the paired fins must have

arisen in a similar way, or, to say the least, the burden of dis-

proof lies with those who believe that such closely similar

structures arose by diametrically opposite processes.

IV. CONCLUSION.

The evidence of Paleozoic sharks, then, is most distinctly in

favor of the Thacher-Balfour teachings. The oldest forms,

acanthodians and cladoselachids, represented by the best-

preserved fossils present lateral-fold fins. Moreover, there is

evidence that as the series advances from the Lower Devo-

nian, the structures of the biserial archipterygium are gradu-

ally acquired. The Carboniferous Cladodus meilsomi, Fig. 1,

of Traquair shows for the first time a definite segmentation

of the supporting elements of the base of the pectoral fin?

Then in the Permian Symmorium Cope, the fin bases not only

are formed, but show apparent fusion in the metapterygial

region, and even perhaps an extension from the body wall of the

metapterygial terminal, a condition which would be best corre-

lated with a change of function in the fin. And finally, in the

xenacanthids, whose structures are known only in this later

_ horizon, the pectoral fin acquires a biserial archipterygium. The

conditions, however, of Cladodus neilsoni and of Symmorium

1 Cf. also the anals of Xenacanthus.

? Judging from the size of the shark and the character of the radials, the

pectoral fin was probably larger, and there is thus ground to believe that the

posterior radials of the fin have been lost. This is better understood by reference

to the accompanying Fig. t, where the Scottish specimen is shown overlaid with

an outline of Cladoselache of similar size.

No. 430.] PAIRED LIMBS OF VERTEBRATES. 773

are still doubtful, the present knowledge of them having been

based upon single, imperfectly preserved, and fragmentary

specimens. The conditions in Cladoselache, on the other hand,

are known from upward of thirty specimens, some of which are

admirably preserved.! In conclusion, if the Gegenbaurian

adherents are to contest the sum and substance of the paleon-

tological evidence, they have, it seems to me, the difficult task

of demonstrating (1) that acanthodians, the most extensive

group of Paleozoic sharks, do not possess pleuropterygia ; (2) that

Cladoselache, in spite of its earlier occurrence and peculiar

structures, is not as archaic a type of shark as Xenacanthus ;

and (3) that the *archipterygium " formed in the unpaired fins

as the terminal member of a series whose original structures

were of fin-fold character, is formed in exactly the opposite way

from the archipterygium in the paired fin. Until these contra-

dictions have been disposed of, no evidence based upon recent

forms can, I believe, attack the main trenches óf the lateral-

fold theory.

Postscriptum. — Objections which have been urged to the

foregoing conclusions.

As far as I am aware, no objections have been raised to the

conclusion that the paired fins of the acanthodians represent

definite lateral dermal folds, whatever may be regarded as the

homologues of the supporting elements of these fins. (Thus

Fritsch endeavors to find in the pulp canals of the anterior fin

spines rudiments of an entire biserial archipterygium (Fauna d.

Gaskohle Formations Bóhmens, Bd. III, p. 71). (As a corollary

to this suggestion, I take it, the fin spines of 2// acanthodian

fins are equivalent to archipterygia, since the spines are sim-

ilar in structure in all fins!) Cladoselache, however, has been

subjected to adverse criticism by several advocates of Gegen-

baur’s hypothesis. Professor Jaekel, the first of these critics, is

the only one who appears to have had the opportunity of exam-

ining actual specimens of this form, for during his visit to

1 Sections of the muscle plates have been found to show transverse striz in

the fossilized muscle cells. Their description will shortly be published.

774 THE AMERICAN NATURALIST. [Vor. XXXVI.

New York he inspected some of the material obtained by

Professor Newberry for the museum of Columbia University.

He regarded * Cladodus " as a shark which had become highly

specialized and adapted to bottom living. Against this view

it is merely necessary to note that Cladoselache possessed

dorsal fins, one of which was situated well forward on the

trunk region, its anterior rim lying above the pectoral fin, a

condition entirely unknown in any depressed type; that the

caudal fin was of a peculiar form, extreme in its heterocercy,

and provided on either side with broad, stout lateral flanges

closely comparable to those of such swift-swimming forms as

Isurus, or even scombroid fishes. It is evident that none of

these characters can be associated with a bottom-living form

such, for example, as Rhina. Moreover, if Cladoselache were a

bottom-living form, its fins would obviously have been larger in

proportion to the size of the animal than those of recent sharks.

In this connection, however, it is only just to Dr. Jaekel to

State that at the time of his visit to Columbia University none

of the best material of this form had yet been obtained, — that

the caudal and dorsal fins, for example, were not known.

A second writer, Professor Semon, in one of his studies on

Ceratodus (Forschungsreisen, Bd. I (1898), pp. 105—106), sub-

scribes to a page of general denial as to the primitive nature of

Cladoselache. As he does not, however, refer to the detailed

literature upon this form, quoting only from a single and semi-

popular article, and in view of other omissions, I am inclined to

believe that his criticism is inadequate. Thus I note that he is

unaware that the horizon of Cladoselache is now conceded

(cf. Orton and others) to be Upper Devonian instead of Lower

Carboniferous, and that he puts this form in the Coal Measures

side by side with xenacanthids, whose structural characters are

known only from the Permian ; also that he states that dipnoans

(he does not mean by “dipnoans,” I take it, the very problematical

arthrodires, since these are naturally included in the term“ placo-

derm," which he specifically mentions) and crossopterygians were

already present in Silurian times,— a statement I can only account

for on the basis either that he has collected data which, when

published, will throw light upon this empty page of the geological

No. 430.] PAIRED LIMBS OF VERTEBRATES. 775

record, or that by Silurian he means Devonian, an age in

which, by the way, we must admit that Cladoselache occurs.

He surely cannot refer in the above terms to the minute frag-

ments of Ostracophores of Oesel, which were once, about a

decade ago, referred to as ganoidean and dipnoan.

And most recently Dr. Braus! has considered in a special

paper the later discoveries in fin structures in fossil selachians,

and has even concluded that “through paleontology the theory

of Gegenbaur has been given new life." Braus, however, makes

no reference in this connection to the archipterygial characters

of unpaired fins, nor yet to acanthodians, and furthermore, he

treats all extinct forms without reference to their different

geological horizons. The pectoral fin of Cladoselache he regards

as essentially similar to that of Chlamydoselachus, believing that

its metapterygial region has been obscured by the overlying

body wall during the process of fossilization, and that the fin in

reality possessed a lobate base which projected from the body

wall, as in the recent shark. He accordingly must believe that

the continuous posterior dermal margin of the fin in the fossils,

as several writers have figured it,? is not a just one. I think,

however, that I am justified in stating that this reactionary

view finds no support in any known? specimen of this form.

And I may further add that the line which continues the web

of the lappet-shaped fin along the side of the body, which

Braus is skeptical about, and which is hostile to the Gegen-

baurian theory, is preserved in faultless condition in as many as

à dozen specimens which are now under my hands. Braus

further believes that the ventral fin of Cladoselache shows a

trace of a distinct metapterygium (as of Chlamydoselachus),

inasmuch as radials occur in the hinder portion of the fin web,

1 Verhandlungen d. Phys.-Med. Gesellsch. zu Würzburg (N.F.), Bd. xxxiv (1901),

PP- 177-192 ; cf. also Zool. Forschungsreisen, Bd. i (1901), Lieferung III, PP- 197,

? The fin margin has thus been figured — rounding in along the body wall —

undissentingly by at least four authors. Braus does not consider this important

feature of the fin, which I am sure he would have felt bound to do had he

examined actual specimens.

*I speak confidently, since I have good reason to believe that all cladose-

lachids have hitherto been obtained through three collectors in Ohio, whose out-

Put I have had the opportunity of examining.

776 THE AMERICAN NATURALIST.

a conclusion to which I do not understand the seguztur. The

absence of mixipterygium in cladoselachids (as well as in acan-

thodians, in fact the whole matter of the acanthodians) he does

not consider in this connection.

Braus, in summary, teaches that paleontological discoveries

lead to the “overwhelming conclusion" that the recent shark

fin has acquired its typical form of skeleton through a process

of reduction wherein the original terminal and post-terminal

axial portions have become lost, while the pre-axial region has

developed and with it its basal plates. The reviewer, how-

ever, must confess that he finds it rather difficult to determine

exactly how Braus has obtained these paleontological results,

unless by deriving the Devonian forms from the Permian ones,

— which, in view of the many remarkable structural characters

of Cladoselache (which Braus also, by the way, does not con-

sider), appears as unconvincing as a process which derives the

Miocene Protohippus from the Pliocene horse.

RECENT INVESTIGATIONS UPON THE EMBRYO

SAC OF ANGIOSPERMS.

DOUGLAS HOUGHTON CAMPBELL. t]

THE development of the embryo sac of the angiosperms has

engaged the attention of many botanists from Hofmeister on,

and the homologies of its parts have been widely discussed.

All of the earlier researches seemed to indicate an extraordinary

uniformity in the structures of the embryo sac, very few devia-

tions from the type being noted, and the importance of these

variations being frequently ignored.

With the improved histological methods developed in recent

years there has been a renewed interest in the subject. The

important paper by Treub on Casuarina! called attention to

several striking deviations from the ordinary angiospermous

type. This paper was followed by a long series of investiga-

tions by many botanists both at home and abroad, which have

extended materially our knowledge both of the morphology and

physiology of the embryo sac.

The uniform results of the earlier investigations may be

explained, in part at least, by the selection of the more special-

ized forms for study. These would naturally conform, for the

most part, to the structure characteristic of the embryo sac of

the typical angiosperms. The more generalized and presum-

ably more primitive forms were neglected, and the significance

of such important variations as the largely increased number of

antipodal cells in many grasses was usually overlooked. So

great, indeed, was the uniformity assumed to be, that all efforts

to explain the origin of the angiosperms from lower forms was

considered well-nigh hopeless.

The studies of the last dozen years have shown that there is

much more deviation from the type than was supposed to be the

1 Sur les Casuarinées et leur place dans le systéme naturel, Annales du Jardin

Botanique de Buitenzorg, tome x, pp. 143-231. Leyden, 1891.

: 777

778 THE AMERICAN NATURALIST. | [Vor. XXXVI.

case, and these have encouraged further researches upon the

simpler types of angiosperms. Some of these have yielded

interesting results, and already give some hints of the possible

origin of the structures of the angiospermous embryo sac.

While it cannot be said that this very important point is

likely to be explained satisfactorily in the near future, never- <

theless it can be said that not unim-

portant advances have been made, and

the purpose of this sketch is to bring

together the most noteworthy results

of these recent investigations, and to

indicate their bearing upon the ques-

tion of the origin of the angiosperms.

In the great majority of investigated

angiosperms the embryo sac, as is well

known, exhibits extraordinary uniform-

ity in its structure. In most instances

the embryo sac arises from a subepi-

dermal cell (Fig. 2 A), which may either

develop at once into the embryo sac,

or, as is more commonly the case, it

divides into several cells, one of which

becomes the embryo sac.

With the first division of the nucleus

of the young embryo sac, the polarity

which is so marked a feature becomes

established. Of the two nuclei result-

mui ing from the division of the primary

stabs “hs de saco er nDüclns one: moves to the apical

is the egg apparatus, consisting of ^ (micropylar) end of the sac, the other

he two synergidz,.

s; at the chalazal end, the three to the basal (chalazal) end. Two

Sec X ill shows du peo nuclear divisions follow, resulting in

nucleoli of the polar nuclei of four micropylar and four chalazal nuclei.

which it is formed.

From each of these groups one nucleus,

the polar nucleus, moves toward the center of the sac, where

it unites with the corresponding nucleus from the opposite end.

1e three remaining apical nuclei, with their accompany ing

cytoplasm, constitute the egg apparatus ; the three basal ones,

No. 430.] THE EMBRYO SAC OF ANGIOSPERMS. 779

which usually become surrounded by definite cell membranes,

form the three antipodal cells. The polar nuclei, either before

or after fertilization, fuse to form the primary endosperm nucleus.

The most marked deviations from the typical development

observed by the earlier investigators were an increase in the

number of archesporial cells (Rosa sp., Helleborus, etc.) and an

increase in the number of antipodal cells. Of the latter varia-

tions the most marked examples were various species of grasses,

first noted by Hofmeister. Further examples of both of these

Wis na

Hi ds

| LH

SERES

P bl

(es

Fic. 2. — Ovules of Arisema triphyllum: A, B, in longitudinal section; C, D, transverse

sections of the nucellus; the archesporial cells are shaded.

deviations from the type have been recorded by later investi-

gators, and other modifications, not hitherto observed, have

been discovered.

The departures from what may be termed the typical devel-

opment may be referred to several categories. Besides those

already referred to, these are three in number: (1) an increase

in the number of nuclei before fertilization has taken place ;

(2) peculiarities in the formation of the endosperm ; (3) the

so-called double fertilization.

The increased number of archesporial cells observed by

Strasburger in Rosa livida, Helleborus cupreus, and other form Et

has been found to occur in a considerable number of forms.

lGoebel Outlines of Morphology, etc.

780 THE AMERICAN NATURALIST. [VoL. XXXVI.

Treub describes a large number of sporogenous cells in Casua-

rina, several developing into complete embryo sacs. The multi-

cellular archesporium in this case is comparable to that of the

pteridophytes and gymnosperms. This increase in the number

of sporogenous cells is seen in a lesser degree in Arvisema tri-

phyllum (Fig. 2). In the latter the original hypodermal arche-

sporial cell divides longitudinally into 2—4 (occasionally more).

One of these, as a rule, becomes at once the embryo sac, but

sometimes there are first transverse divisions. It is possible

that there may sometimes be more than one primary arche-

sporial cell in Arisaema, z.e., the whole sporogenous tissue may

not certainly be referable to the division of a single hypodermal

cell. This point, however, needs further investigation.

CASUARINA.

The very peculiar genus Casuarina, according to Treub's

investigations,! differs decidedly from the other angiosperms,

not only in the large development of the sporogenous tissue of

the ovule, but also in the structure of the developed embryo

sac. This has no antipodal cells, and all the cells of the egg

apparatus, which are variable in number, are formed from the

division of a primary cell. The egg, moreover, is surrounded

by a cellulose membrane before fertilization takes place, a con-

dition unknown among other angiosperms. Before the egg

cell divides (and Treub believes before it is fertilized) there is

a rapid increase in the number of endosperm nuclei. A demon-

stration of the actual fertilization was not made, and this point

must remain for the present in doubt. Should it be verified, it

might well be compared with the condition existing in Peperomia.

PEPEROMIA.

In the genus Peperomia the writer? found that there is

normally a doubling of the nuclei of the embryo sac before

fertilization ; Ze. instead of the eight nuclei usually found in

the mature embryo sac, there are sixteen (Fig. 3). This fact

1 Le cit.

2 Campbell. The Embryo Sac of Peperomia, Annals of Botany, vol xv

(March, 1901), pp. 103-118.

No. 430.] THE EMBRYO SAC OF ANGIOSPERMS. 781

was confirmed by Johnson,! who also discovered that the endo-

sperm nucleus was the product of the fusion of several nuclei,

instead of two, as in the typical angiosperms. In Peperomia

there is no clearly defined egg apparatus, nor are there any

proper antipodal cells, although a varying number of nuclei

become enclosed in cell membranes and form small flattened

cells applied to the wall of the

embryo sac. It is to be hoped

that a further study of the embryo

sac of Casuarina may be made,

employing carefully stained mi-

crotome sections. A comparison

with the embryo sac of Peperomia

would be of great interest.

The increased number of nuclei

in the embryo sac of Peperomia,

while intermediate in character

between that of the typical angio-

sperms and the lower vascular TR

plants, might equally well be com- Fte: ee Gh a Aad ce

pared with that of the gymmno- there were present sixteen free nuclei, of

sperms Gr She Beterospofous = er o9 PH WEN.

pteridophytes. The nearest approach to it is found in Gnetum,?

where the structure is not dissimilar, no true archegonium being

present, but the egg cell being developed from one of the free

nuclei. However, as the affinities of Gnetum are very obscure,

this resemblance does not necessarily imply any connection

between Peperomia and the typical gymnosperms.

An increased number of nuclei in the unfertilized embryo

sac has been recorded as an occasional variant in several low

monocotyledons, — e.g., Naias, Zannichellia, Sparganium, — but

in all these forms the normal embryo sac is of the ordinary

angiospermous type. :

1 Johnson. On the Endosperm and Embryo of Peperomia pellucida, Botai-

cal Gazette, vol. xxx, July, 1900.

2 Lotsy. Contributions to the Life-History of the Genus Gnetum, Annales

du Jardin Botanique de Buitenzorg, tome xvi (1899), pp. 46-114.

Karsten. Beiträge zur Entwickelungsgeschichte der Gattung Gnetum, Bota-

nische Zeitschrift, Bd. l, 1892.

782 THE AMERICAN NATURALIST. . [Vor. XXXVI.

VARIATION IN THE ANTIPODAL CELLS.

The three antipodal cells commonly found at the chalazal

end of the embryo sac are frequently looked upon as merely

vestigial structures, as in many cases they doubtless are.

That they may be of importance physiologically has been

en.

p f ferti

Fic. 4. — A, the chalazal t

lization : e. z., endosperm terra aM., two ot the three sped cells. B, section of a

young seed of the same species: em., embryo; end., endosperm; ant., the large mass of

antipodal cells. d — € cells in young endosperm: from a somewhat —— stage,

more highly magn sichiton

kamtchatcense, showi , end., and th } 3 ipodal a SRDA

repeatedly diosa and there is no question that this is oftener

the — than was formerly supposed. In many forms — e.5»

Naias, Lilaa — the antipodal cells are large, with abundant

cytoplasm, and every indication of being active cells. In other

cases, such as some Ranunculaceze,! they become very large,

- 1Mottier. Contributions to ae Embryology of the Ranunculacez, Botanical

Gazette, vol. xx, 1895.

No. 430.] THE EMBRYO SAC OF ANGIOSPERMS. 783

and become multinucleate. An increase in the number of

antipodal cells was observed as a regular phenomenon in vari-

ous grasses, by Hofmeister, and has been noted also by later

observers, — Cannon! found in Avena fatua as many as thirty-

six. There are in these cases the original three antipodals

which subsequently increase by division, this occurring before

the egg is fecundated.

The greatest number of antipodal cells yet recorded occurs

in Sparganium simplex (Fig. 4), where there may ultimately

Fic. g: dof Spatii b gitt ef oli : em. , embryo; 5., e nlarged basal

CTR EEA x 6. B, b lon of th f th more enlarged C, the

large basal endosperm nucleus, : a, and the ordinary ones, 4, from the embryo sac of er

be 150 or more. At the time of fertilization there are but

three, small and inconspicuous antipodal cells ; but after fertili-

zation these increase rapidly in size and begin to divide actively,

forming a very conspicuous hemispherical mass of cells at the.

base of the embryo sac. This large group of antipodal cells

develops while the endosperm is still rudimentary and is doubt-

less of great physiological importance.

1 Cannon. A Morphological Study of the Flower and meee of the Wild

Oat, Proc. California Acad. of Sciences, third series, vol. i, No. 10,

784 THE AMERICAN NATURALIST. [Vor. XXXVI.

Among other similar cases may be mentioned that of the

peculiar aroid, Lysichiton, which occurs upon the northern shores

of the Pacific. The original antipodal cells divide in this case also,

and their nuclei become enormously enlarged (Fig. 4 D).

From a study of the numerous instances which have been

studied, it is clear that the antipodal cells are by no means

merely vestigial structures, but are often of great physiological

importance, replacing the endosperm to some extent, or acting

as a medium for the transport of nutriment to the developing

embryo.

THE ENDOSPERM.

In the typical angiosperms the primary endosperm nucleus

resulting from the fusion of the polar nuclei divides to form

many free nuclei lying in the peripheral layer of cytoplasm.

Walls form between these nuclei, and a layer of tissue is thus

developed, lining the embryo sac. By subsequent centripetal

divisions the embryo sac becomes finally completely filled

with endosperm. When the embryo ultimately fills the whole

embryo sac the cell divisions in the endosperm may be entirely

suppressed (z.g., Naias). In other cases the first division of

the primary endosperm nucleus is followed by a cell wall

extending across the cavity of the embryo sac, which is thus

filled from the first with tissue.

A modification of the type has been noted in a number of

low monocotyledons. Of the two nuclei resulting from the

division of the primary endosperm nucleus, only the upper one

divides further. The lower one, which may be separated by a

wall from the upper part of the embryo sac,! sometimes becomes

enormously enlarged (Fig. 5 C) but does not divide further.

It is possible, although this has not been demonstrated, that in

some cases there is no fusion of the polar nuclei.

A condition somewhat intermediate between the ordinary

form and that just described was recently observed by the

writer in Spathicarpa sagittefolia, a South American aroid.

In this casé (Fig. 5) a group of a few very large cells, with

Finis The Life History of Sagittaria variabilis, Botanical Gazette, vol. xxiii,

I -

No. 430.] THE EMBRYO SAC OF ANGIOSPERMS. 785

enormous nuclei, lies at the base of the endosperm and is

sharply separated from the small-celled endosperm in the upper

part of the embryo sac. It is probable that the basal group of

large cells is the product of the lower of the two nuclei derived

from the first division of the endosperm nucleus.

In the genus Peperomia the very large endosperm nucleus

is the result of the fusion of several (usually eight) nuclei.

It then divides to form the rudimentary endosperm found in

the ripe seed.

DOUBLE FERTILIZATION.

Much interest has been aroused by the discovery of the

so-called double fertilization which has been demonstrated for

a large number of angiosperms. This double fertilization con-

sists in the fusion of the second generative nucleus of the pollen

spore with the endosperm nucleus, which thus becomes the

product of three nuclei. This has led to a theory that the endo-

sperm nucleus is sexual in its nature, and the endosperm arising

from it is an embryo. That this view can hardly be maintained

is evident from the condition found in Peperomia, which pre-

sumably represents a more primitive condition than that of

the typical angiosperms. In Peperomia, as already stated, the

endosperm nucleus is the product of several similar nuclei, and

as such a multiple fusion of sexual cells is quite unknown else-

where, it is safe to assume that this fusion is not of the nature

of a true fertilization.

That the second generative nucleus discharged into the

cavity of the embryo sac should fuse with the only available

nucleus, the endosperm nucleus, is not surprising. That its

character is impressed upon the resulting nuclei is also to be

expected. This is shown most clearly in the case of hybrid

maize. It has been clearly demonstrated that one type of maize

pollinated with another will produce ears in which the endo-

sperm of the grains shows a hybrid character, due presumably

to the fusion of one of the pollen nuclei with the endosperm

nucleus.!

1 Webber. Xenia, or the Immediate Effect of Pollen in Maize. U.S. Department

of Agriculture, Division of Vegetable Physiology and Pathology, Bulletin Av. 22, 1900.

786 , THE AMERICAN NATURALIST.

SUMMARY.

Peperomia, in regard to the embryo sac, probably represents

the most primitive form yet described among the angiosperms.

The absence of a definite egg apparatus and antipodals, and

especially the increased number of nuclei, point to this. The

single-celled archesporium of Peperomia is, however, probably

less primitive than the multicellular sporogenous complex

found in Casuarina and some other forms.

The similarity between the structure of the embryo sac in

Peperomia (and perhaps Casuarina) and Gnetum is striking ;

but as the affinities of the latter are very doubtful, this does

not throw much light upon the relationships existing between

gymnosperms and angiosperms.

The typical embryo sac may very well have been derived

from one like that of Peperomia, by the suppression of one

nuclear division. The marked polarity, and the specialization

of the egg apparatus and antipodal cells, are probably secondary

characters, and the fusion of the polar nuclei has its prototype

in the multiple fusion of the nuclei in Peperomia, to form the

endosperm nucleus.

Peperomia offers a basis for an explanation of the homolo-

gies of the embryo sac. The egg cell probably represents an

archegonium reduced to a single cell, and possibly the synergidze

may also represent potential archegonia, although it is quite as

likely that they are derived from vegetative prothallial cells.

All of the other structures, the polar nuclei (and their product,

the endosperm nucleus) and the antipodal cells, represent vege

tative prothallial tissue. The increase in number met with in

the antipodal cells of Sparganium, for instance, merely empha-

sizes their power to assume the róle of active prothallial tissue.

The fusion of the polar nuclei is in no way to be considered

as a true sexual process. The regular occurrence of a multiple

fusion in Peperomia is a strong argument against such an

assumption. It is much more probable that it is to be inter-

preted as a stimulus to further growth. The fusion of the

second pollen nucleus with the endosperm nucleus must be

considered as more or less accidental.

1 Strasburger. Einige Bemerkungen zur Frage nach der “ doppelten Befruch-

tung” bei den Angiospermen, Botanische Zeitschrift, 1899.

THE VASCULAR SYSTEM OF THE COMMON

SQUID, LOLIGO PEALII.

LEONARD W. WILLIAMS.

THE knowledge of the histology of the vascular system of the

decapod mollusks is very incomplete. The nature and extent

of the capillary system especially have been so little understood

that two contrary opinions find expression in the text-books of

zoólogy. It has been maintained by Lang that the arterial and

venous systems are connected partly by capillaries and partly

by lacunze, which are portions of the primary body cavity, while

Parker and Haswell affirm that the blood flows through a com-

plete system of capillaries. I have been unable to find any

published work which decides between these views. All agree

that the vascular system is highly developed and that the capil-

lary system is extensive, but no one seems to have investigated

the structure of the vessels or to have determined the exact

nature of the so-called lacunz or sinuses. Prof. Ulric Dahlgren

called my attention to this subject, and under his direction I

worked upon the vascular system of the squid at Woods Hole

during the summers of 1899 and 1900, and at Princeton Uni-

versity during the intervening time. The material, Loligo

£eaíii, was furnished me by the Marine Biological Laboratory

through the kindness of Professor Dahlgren, and by the United

States Fish Commission through the kindness of Dr. H. C.

Bumpus, and further material was obtained from the traps at

Elberon, N. J. ;

The extent of the capillary system was determined by inject-

ing the vessels with a saturated aqueous solution of Berlin blue.

The injected tissues were studied while fresh and transparent

or were fixed in acetic corrosive sublimate er 95 per cent

alcohol and mounted in serial sections. The structure of the

walls of the vessels was studied after impregnating them with

silver in order to show the outlines of the endothelial or other

787

788 THE AMERICAN NATURALIST. | [Vor. XXXVI.

lining cells. Before injecting the solution of silver nitrate it

was necessary to remove the chlorides from the vessels by

irrigating them with a 5 per cent solution of potassium nitrate.

This solution was followed by a ¥% per cent solution of silver

nitrate, and after a few moments this was removed and the

tissues fixed by the injection of 95 per cent alcohol Small

pieces of the several organs were dehydrated, cleared in creo-

sote, and mounted. The contraction of the intrinsic muscles

of the vessels renders complete injection very difficult, and to

obviate this difficulty by dilating the vessels the injection fluids

and the water in which the squid were kept were saturated with

amyl nitrite. This method was not uniformly successful.

The general arrangement of the vascular system of the

squid will be recalled by a glance at the diagram (Fig. 1) of

obs. t

205 cs ph ss aa A

i ao bre cd => pp indi `

RYC

] us

br's os UC

brh pe:

Fic. 1. — Diagram of the vascular system of the squid as seen from the left. aa, anterior aorta;

ork, branchial heart; őrs, brachial sinus; cs, cerebral sinus; A, heart; zs, inner buccal

sinus; zs, nephridial sinus; 04s, outer buccal sinus ; os, optic sinus ; p posterior aorta;

pc, postcava; pf, peripheral heart; ss, salivary sinus; vc, vena cava.

the system seen from the left. Arterial blood enters the one-

chambered heart from the gills through the two branchial veins

and is forced out through the anterior, posterior, and genital

aorte. The anterior aorta forks near the head, and each branch

bears a muscular enlargement, called a peripheral heart, at the

point where it breaks up into the branches which supply the

head. The two branches of the posterior aorta which pass out

of the mantle to the fins also have peripheral hearts. The

latter are vascular sphincters, which probably contract synchro-

nously with the mantle, and so prevent the transmission of an

excessive blood pressure to the extra-pallial vessels. The blood

returning from the arms through the brachial veins reaches the

brachial sinus, which is connected with five sinuses, that partially

enclose the pharynx, the eyes, and the brain. These sinuses

open into the anterior vena cava, which also receives a ven

from the sinus that encloses the salivary gland. The anterior

No. 430.] THE COMMON SQUID, LOLIGO PEALII. 789

vena Cava opens into the apex of the V-shaped nephridial sinus

which is surrounded by the nephridial chamber. The venous

blood from the dorsal end of the mantle returns through the

two postcavee which open into the ends of the arms of the

nephridial sinus. This sinus opens at each side into a bran-

chial heart which forces the blood through the gill to the

systemic heart. The sinuses are not interposed between

the veins and arteries but receive blood from the veins and

return it to the veins. The aortz, the larger arteries, the

branchial arteries and veins, the anterior vena cava, and the

postcavee are all peristaltically contractile. The wave of con-

traction passes slowly away from the heart along the arteries,

and toward the heart along the veins. The peristalsis of the

vessels continues long after the stoppage of the hearts and

the apparent death of the animal.

The Structure of the Arteries and Veins. — The systemic

or arterial heart is composed of smooth muscle fibers. The

external surface is firm, but the inner surface is formed by a

latticework of interlaced fibers. The great irregularity of this

surface interferes with the success of silver impregnations,

and although some impregnations and microtome sections indi-

cate that there is an endothelium, such a large number have

yielded only negative results that it is probable that an endo-

thelium is not present in the heart. The arterial blood which

passes to the tissues of the heart from its lumen becomes

venous and is carried to the nephridial sinus by two or three

small veins. It is probable that these veins are separated from

the lumen of the heart by capillaries. The vessels through

which the blood passes are certainly as small as capillaries,

but it was impossible to demonstrate an endothelium.

The branchial hearts are quite different from the systemic

heart in that the muscle fibers are cross striated and are sepa-

rated by numerous clusters of apparently glandular polygonal

cells, which possibly have a function in connection with the

pericardial gland that is attached to the inner edge of the

heart and receives a large blood supply from it. Silver impreg-

nation shows that an endothelium is not present and that the

muscular and glandular cells are in contact with the blood.

790 THE AMERICAN NATURALIST. [Vor. XXXVI.

A branch of the posterior aorta brings arterial blood to the

tissues of the branchial heart.

The aortæ and the larger arteries are peristaltically con-

tractile and thick walled. Their walls are formed by a thick

sheet of peculiar muscular tissue, which is enclosed by two

thin coats of connective tissue. The muscular tissue is formed

of fusiform cells whose large oval nuclei have a small quantity

of protoplasm at each end, but the greater portion of each cell

consists of radiating fibers which interlace with those of neigh-

boring cells. The muscle resembles connective tissue more

than muscle, but it is actively contractile, for stimuli can be

só applied that the peristaltic wave will move in the opposite

direction to the blood current, and the peristalsis continues long

after the final stoppage of the hearts.

god cr EOE

j \

Fic, 2. — Arterioles which show shed effect of the contraction of the scattered intrinsic

scle fibers of the vessels.

The muscular coat of the smaller arteries 1s much thinner,

and the peculiar muscle cells are replaced by simple fibers,

which are so irregularly distributed upon the smallest arteries

that their contraction makes the vessels moniliform, as is shown

in Fig. 2. Arterioles arising as lateral branches of an artery

commonly have a strong band of muscle fibers at their origin.

The aortz and their largest branches may have an endo-

thelium, but the unavoidable contraction of their muscular

walls throws the inner surface into longitudinal wrinkles,

between which the silver is deposited in streaks that obscure

the cell outlines. It is probable that an endothelium is present

in these vessels ; for, although almost all the evidence is nega-

tive, a few preparations seem to show the endothelium. The

^ intermediate arteries are lined | by an endothelium composed of

No. 430.] THE COMMON SQUID, LOLIGO PEALII. 791

flat elongate cells that encircle the vessel. This endothelium

gradually merges into the typical endothelium lining the

smaller arteries. Bergh! found a similar arrangement in the

arteries of the pulmonate gasteropods, except that he ascer-

tained definitely that the larger arteries and the heart do not

possess an endothelium but are lined by muscle fibers. He con-

cludes that the endothelial cells are undifferentiated (wngeformte)

muscle cells. . There is nothing in the squid which tends to

support such an explanation, for the larger vessels are almost

certainly lined either by connective tissue or endothelium.

The veins have the same structure as the arteries except

that their muscular coat is thinner or wanting and that all

have a typical endothelium (Fig. 3).

They may readily be distinguished

from the arteries by their thinner

walls. In many places an artery is

accompanied by a pair of veins which

anastomose frequently. Fig. 4 repre- ric, 3.— Endothelium of a vein.

sents a portion of such an artery with par

its companion veins; the narrow initial caliber of the branches

of the artery, due to the contraction of the intrinsic muscles of

the vessels, should be noted.

The nephridial sinus passes through the nephridial cavity

and is invested by the secretory nephridial epithelium. Both

surfaces of the sinus wall are increased by interdigitating

evaginations from the lumen of the vessel and from the

nephridial cavity. The consequent irregularity of the surface

of the walls causes many artifacts in silver preparations, so

that the cell outlines cannot be followed certainly ; neverthe-

less, sections indicate that an endothelium is present. The

nephridial epithelium of the sinus is columnar and is supported

by a membrane, the proper wall of the sinus, which carries the

nutrient arteries of the wall.

The Capillaries. — The Berlin-blue injections which were

intended to demonstrate the extent of the capillary system

were perfectly successful. The injection fluid passed from

the arteries through the tissues until it filled and flowed from

1 Bergh, R. S. Anat. Hefte, I. Abth., Bd. x.

792 THE AMERICAN NATURALIST. [Vor. XXXVI.

the veins, so that it is certain that the vascular system was

fully injected. Serial sections of many portions of the squid,

and gross mounts of portions of the skin, mesentery, and

intestinal wall show conclusively that there is a perfect capil-

lary network which connects the arteries and veins in all parts

of the body. Like other capillaries, these (Fig. 4) branch and

anastomose frequently without altering their diameter, and their

walls are formed by an endothelium composed of flat oval cells

with sinuous margins. The silver impregnations of the capilla-

ries, though rarely successful, leave no doubt as to the nature of

the wall.

Fic. 4. — Artery, i ins, and

Hets 3 £. ah norting

T o E rE

b f the gill. æ, artery; v, veins.

Some arterioles and capillaries are connected with lacuna-

like cavities which may be called the end sinuses of the veins.

Fig. 5 is a camera sketch of such a sinus situated in the

visceral body wall. Similar sinuses have been found in the

testis, glands, and muscles, and it is very probable that they

are more widely distributed. Each sinus is a small irregular

cavity, into which an arteriole passes and then breaks up into

branches; the latter pass out of the sinus and communicate

with the surrounding capillary plexus, some of whose branches

_ open into the sinus, so that the blood must pass through the

~ perforating arteriole and then through the capillaries into the

No. 430.] THE COMMON SQUID, LOLIGO PEALII. 793

sinus! The sinuses are connected with the veins by very small

vessels. In partially successful injections the colored arteriole

can be seen surrounded by a space filled with blood corpuscles,

but in perfect injections both the arteriole and the sinus are

filled with the injection mass. The arteriole, capillaries, and

the sinus are all lined by endothelium. It was impossible to

determine whether the endothelium of the sinus was reflected

over the surface of the perforating arteriole. It is worthy of

note that when the blood had not been driven out of the sinus

the proportion of corpuscles in the blood of the sinus was six

or eight times as great as in the other vessels, —a fact which

Ul 7

zu 7

7 7^. Y

7 LL LLL

7, ha 77 "LL OOO = 1272»

C ge UE 7

Uf 277 77^

RN a (M

Wipe

e DEA ay,

LT,

Fic. 5. — End sinus of vein, from visceral body wall.

may be due to the filtering out of the corpuscles as the blood

was driven through the vessels by the injection fluids.

The Capillaries of the Gills. —The gills are not easily injected

because a valve at the origin of the branchial artery prevents

the free flow of injection fluids in either direction. The valve

is formed by four tubercles which project in the direction of the

blood current into the lumen of the artery. A band of muscle

fibers forms a sphincter whose contraction forces the tubercles

together so that the lumen of the vessel may be closed.

1 The connection between these vessels and the veins has been inferred from

the course of the injection fluids through the sinus.

794 THE AMERICAN NATURALIST.

Notwithstanding the resistance of this valve, successful capillary

injections of Berlin blue were obtained as well as good silver

impregnations of the veins and arteries. The capillaries are

enclosed between epithelial plates which form the upper and

lower surfaces of the gill and are separated from one another

by small columns of cells which bind together the gill surfaces.

The vascular endothelium could be traced only to the beginning

of the capillaries, but microtome sections leave little doubt that

the endothelium also extends into the capillaries.

The salivary sinus and the sinuses of the head (brachial,

inner and outer buccal, optic, and cephalic), which are usually

cited as examples of lacunz, must be considered to be enormous

dilations of the veins. They are so large that it is impracti-

cable to examine every portion of their walls, but silver impreg-

nations of portions of several sinuses show that they are lined

by an endothelium. All the sinuses have essentially the same -

structure, so that the description of one sinus will convey a

true conception of all. The outer buccal sinus surrounds the

sides and dorsal surface of the oval pharynx as the pericardium

invests the heart. The outer wall of the sinus is supported by

the muscular ring at the base of the arms, and the inner wall is

supported by the pharynx and by the muscular septum which

separates the inner buccal sinus from the outer. The walls are

formed by connective tissue and by the endothelium which it

supports. Ten or fifteen small veins which arise in the peri-

stomial membrane open into the ventral end of the sinus,

and the brachial sinus opens into the dorsal end. The blood

passes from the buccal sinus through the optic sinuses to the

vena cava. Beside having definite walls, these sinuses, unlike

lacunz, do not connect arteries and veins, but receive blood

from the veins and return it to the veins. The wide dis-

tribution of the capillary vessels, the presence of an endo-

thelium around every blood-containing cavity, except possibly

the hearts, and the absence of demonstrable lacune, all lead

to the conclusion that the arterial and venous vessels of the

squid are connected by capillaries so that the vascular system

is closed.

- BROWN UNIVERSITY.

WINDS AND STORMS AS AGENTS IN THE

DIFFUSION OF INSECTS.

F. M. WEBSTER.

Bv the term * winds" I wish to include even the mildest

breezes as well as the strongest gales, the latter, either alone

or in connection with thunderstorms, such as may and ordi-

narily do occur in our latitude during the warm months of the

year. Throughout the northern states the keen, biting March

winds find comparatively few insects abroad to be caught

up, or who willingly surrender themselves to its power; and in

late autumn, when the breeding season is nearly or quite over

with the majority of species, they seem to prefer the quiet of

the Indian summer to move about. In the one case, diffusion

appears to be the one overwhelming object; while in the other,

the object seems to be to reach particular locations or con-

ditions that will best protect from the winter blasts which

are to come later on in the year.

Whatever conduces to diffusion, must, as a rule, tend to

increase fecundity — first by bringing the sexes together during

the mating seasons, and second by enabling them to secure

a greater food supply.

The effect of even very light breezes in enabling the sexes

to meet at the proper time may be well illustrated by placing

a female of some of our larger moths in a wire cage, and,

hanging this in the open, noting the number of males that will

Soon be attracted and remain hovering about the cage, making

every effort to reach the female within. The same may be

observed in many Hymenoptera. It will be noted further that

these males come to the cage against and not with the wind.

With a continual absolute quiet, how much less liable would

the sexes be to reach each other at the proper time for mating,

1 Read before Section F, Zoólogy, American Association for the Advancement

of Science, July 1, 1902.

795

796 THE AMERICAN NATURALIST. | [Vor. XXXVI.

especially such as are wingless among the females, or which,

having wings, do not readily use them? If we trap insects by

baits or by lights, we find that our catches will be largely of

males or spent females, thus showing that, generally speaking,

the male seeks the female. In the cases of some of our hyme-

nopters and especially where males do not, or rarely, occur,

and with others, like the Aphides, where the males only occur

at a single definite time, this rule will not hold good, and our

captures will be all of them females; and I think it is among

such that will be found those that give themselves most freely

to the embraces of the winds during the breeding seasons.

The influence of the winds in enabling insects to detect the

location of their food is apparent in many ways. If we place

the dead body of a bird, snake, or small animal under a bell

glass in the fields, no carrion-loving insects will come to it;

but if we substitute for the bell glass a wire cage, these will

soon find their way to it, coming against and not with the

wind. The two species of dung beetles, Aphodius inguinatus

and A. serval, which appear, the one in the fall and the other

in the spring, quickly find the fresh droppings of animals by

aid of the wind. The plum curculio, Conotrachelus nenuphar,

is said to approach a plum orchard more readily against the

wind than with it, as has been determined by painting indi-

viduals and liberating them to the windward and leeward of a

plum orchard just at the season for oviposition; and there is

little doubt but that many of our wood-boring beetles are

enabled to select a weakened tree by their acute sense of

smell. Thus do the light winds enable many insects the better

to find their food and each other, thereby greatly increasing

their numbers in any given place.

The high winds and gales, unaccompanied by rain, thunder,

and lightning, also exert a powerful influence in the diffusion

of insects. But before taking up the direct subject of influ-

ences of high winds exclusively, it may be well to call attention

to some of the effects of prevailing winds, that is, winds both

light and heavy but which come most continuously from one

direction during the period when insect diffusion is most sus-

ceptible to their influences. A most convincing illustration in

No. 430.] THE DIFFUSION OF INSECTS. 797

this direction is offered by the spread of the San José scale in

orchards, especially in the Middle West. If infested trees,

brought from the nursery and transplanted to the orchard,

happen to be set along the north or east margins of an orchard,

the spread through that orchard is comparatively slow; but

on the other hand, if the infested trees happen to be placed along

the west or south margins, the progress is much more rapid.

So also, where the introduction is into a section of country

largely devoted to tree fruits, the spread over the country is

much more rapid towards the northeast than in any other

direction. This is for the reason that our prevailing winds,

during the period when the young are carried from one place

to another by these winds, blow from the southwest or west,

more generally the former. In the case of the Hessian fly,

Cecidomyia destructor, Y have noted that, where a field of wheat

has been seriously attacked in the fall, and adjoining fields have

escaped, some of these last may be seriously attacked the fol-

lowing spring, while others seemingly equally exposed to attack

almost entirely escape injury. A careful examination into the

facts, however, will show that this is due to the direction of

the prevailing winds at the time when the spring brood of flies

were abroad, and the wind simply carried them toward those

fields that were located in one direction and away from those

located in the opposite direction, thereby to a certain extent

protecting the one from attack and causing the destruction of

the other. In the case of the San José scale, active only in the

very young larva, I have noted that, where infested trees

happen to be placed in a gully extending up a steep hill, the

spread will be much more rapid upward than downward, as the

air current is in that direction. So, then, prevailing winds

have in some cases much to do with the spread of insects in

certain directions. The influence of high winds on insects is

illustrated in one way by the great number of butterflies that

are sometimes encountered by ships at sea, long distances

from land. Indeed, the entomological fauna of very many of

the islands of the sea indicates very strongly that insects

have become established on such islands by having been blown

from the mainland, or from other islands located at considerable

798 THE AMERICAN NATURALIST. | [Vor. XXXVI.

distances away.! Dr. Henry C. McCook, in his great work on

American Spiders and their Spinning Work, has shown how

the huntsman spider, Heteropoda venatorius, on account of its

aeronautic habits, might well have circumnavigated the globe

with the aid of the trade winds, basing his calculations on the

localities where the spider is known to occur and the direction

of these winds. The late Dr. D. S. Kellicott told me of the

sudden appearance of considerable numbers of the cotton-worm

moth, Aletia argillacea, at Columbus, Ohio, immediately fol-

lowing a gale from the southwest ; and, besides, we know that

this moth has been found as far north as western Ontario,

with no evidence of its having developed there in a single

instance. Years ago, while studying the habits of the buffalo

gnat, in the Southern States, one of the most perplexing ques-

tions that confronted the planter, relative to the habits of this

bloodthirsty insect, was their sudden appearance in a locality

in such immense swarms as to kill thousands of head of cattle,

mules, and other domestic animals, before these could be gotten

toa place of safety. Early in my investigations, I found that

the adult gnats emerged from the waters of the bayous and

clustered upon the surrounding vegetation in such numbers as

to fairly cover the same, and a sudden high wind would carry

these gnats along with it and distribute them for miles over

the cotton plantations, there to carry on their bloody work.

Thus, these insects might appear suddenly in immense num-

bers, ten or twelve miles from their place of breeding, in one

direction, while in the direction opposite they might not occur

1 Since the preparation of this paper the following note has come to me:

The ‘Blue Page’ Moth. — During the gale that reached Barbados and St. Vin-

cent on August 26, 1901, numbers of a large moth were found in Barbados, of

a kind not known to breed there. They had evidently been brought by the

high southwest wind. Some were caught and identified as Urania sloanei, the

‘blue page’ of Trinidad, and they had apparently come from the mainland or,

more probably, from Trinidad itself. They were found as far north as Dominica,

direct distance from Trinidad to Barbados is about 160 miles, and to Dominica is

. Some roo miles more. — The (Barbados) Agricultural News, June 7, 1902-

No. 430.] THE DIFFUSION OF INSECTS. 799

at all, — the direction depending on that of the wind ; anda wind

prevailing from the same direction during the breeding season

one year would affect a certain territory, while the following

year the prevailing winds might be from another quarter, and

so cause these insects to terrorize an entirely different section

of country. Formerly, and before the advent of the electric

motor as a means of propelling street cars, when these gnats

developed, during some seasons, in the St. Francis bottoms,

across the Mississippi River west of Memphis, Tenn., these

insects might occur in their breeding grounds in immense

swarms, but so long as the prevailing winds were from an

easterly direction few of them would be observed in Memphis ;

but let a strong gale set in suddenly from the west and these

. bloodthirsty insects would suddenly appear in such numbers as

to prevent the running of the street cars, which were then

drawn by mules; these last would be killed in their harnesses,

and the cars were necessarily abandoned until these gnats

disappeared. Of course the introduction of electricity, and the

disappearance of the mules as a means of motive power has,

in this case, overcome the difficulty, but former conditions

offered a good illustration of the far-reaching influences of the

wind on some insects. It may be mentioned that the gnats

that were so driven about by the wind were all sterile females

and the species was not, in this particular case, permanently

diffused by this means. Somewhat similar effects of winds

are to be observed as affecting the various species of migra-

tory locusts, the exact territory devastated by them often being

dependent on the direction of the prevailing winds during the

migrating season.

Relative to the concluding point in this paper, viz., the influ-

ence of wind and thunderstorms combined on insect diffusion,

I beg to call attention to a most interesting series of papers

contributed to Prometheus, a German scientific journal much

like our Scientific American, by Prof. Karl Sajo, of Budapest,

Hungary. Professor Sajó says that it is known that “ before

thunderstorms the crayfish come out of the water into the

grass on the banks of the river or lake; many fishes act as if

they were insane, and many birds and mammalia become

800 THE AMERICAN NATURALIST. | [Vor. XXXVI.

irritated and angry. Even the micro-organisms are subject to

similar changes ; for instance, before thunderstorms in late fall,

the wine fermentation can reach so great a violence as to cause

the fermenting juice to suddenly run out of the vats. The

greater the change in the atmosphere, the greater the unrest

of the living being." If one happens to be at a farmhouse in

our own country, where ice is not freely used, and a thunder-

storm occurs, any farmer's wife will complain that the thunder

and lightning have soured the milk. Continuing, Professor Sajó

calls particular attention to the “great unrest and activity that

takes place in the insect world just in the sultry hours pre-

ceding a thunderstorm, and to the fact that insects in the air

at the time the storm bursts are driven like chaff to great

distances, — perhaps into other countries, across rivers, lakes,

and mountains; not only the species that fly but many that

do not fly may thus be transported to new homes." An

again, * Many Aphides creep to the crowns of the plants, then

drop themselves at the proper moment into the violent current

of the storm. A number of these insects land in places where

there is no food supply for them and they die. A part of them

reach places where their species is already established, and

fare no better. Part are thrown into the water, sometimes

in oceans, and perish. A proportionally small number arrive

at such places as may be called really favorable for their diffu-

sion, vzz., where the species has never established itself before,

or, having done so, died out before the arrival of newcomers,

and, therefore, natural enemies had not preceded them. Such

individuals as are thus thrown into favorable places have a

chance to multiply into large, populous colonies within a short

space of time, and continue until their enemies find them out,

or they become over-populous and devour all of their food

supply, resulting in what to them is famine.”

- There is probably not an American entomologist who

has not encountered illustrations similar to those enumerated

by the writer of the above, and, while we may not have

wholesale introductions of new things among us, there is

no doubt that localities are often first colonized by certain

kinds of insects in this manner, whereas the wind or the

No. 430.] THE DIFFUSION OF INSECTS. 801

thunderstorm acting separately would not bring about such a

condition of affairs.

I have stated that, in applying trap lights or lanterns, or edible

baits like sweetened sour beer, we, as a rule, secure males

and spent females, but the influence of weather conditions that

usually precede a thunderstorm (that is, a close, sultry con-

dition) has the effect of bringing out both sexes, — a result

due, so far as can now be determined, to some subtile action

on their sexual life. As Professor Sajó so aptly illustrates

this point, I will quote him again quite fully. ** What influence

the weather has, especially on the activity of sexual life, must

be known to every zoólogist; even man is not an exception

from these ‘living barometers.’ Not only children, not only

the female sex, but the sick ones experience the influence of

the weather on the functions, especially on the nervous system ;

and everybody without exception are thus influenced, though

not all may be aware of the fact. The same causes that in

many produce unrest and irritation render others dizzy,

stupid, or sleepy, according to the temperament of the indi-

vidual.” The effect of electricity on the nervous systems of

insects, especially as relative to their love affairs, would con-

stitute an interesting study, and one that ought to be carried

out; but even as it is, we can see that the thunderstorm, in

conjunction with the wind, may accomplish in the diffusion of

insects that which neither element alone would bring about.

THE COLORS OF FISHES.

DAVID STARR JORDAN.

Pigmentation. —The colors of fishes are in general produced

by oil sacs or pigment cells beneath the epidermis or in some

cases beneath the scales. Certain metallic shades, silvery blue

or iridescent, are produced, not by actual pigment, but, as

among insects, by the deflection of light from the polished

skin or the striated surfaces of the scales. Certain fine stria-

tions give an iridescent appearance through the interference

of light.

The pigmentary colors may be divided into two general

classes, ground coloration, and ornamentation or markings.

Of these the ground color is most subject to individual or

local variation, although usually within narrow limits, while

the markings are more subject to change with age or sex. On

the other hand, they are more distinctive of the species itself.

Protective Coloration. — The ground coloration most usual

among fishes is protective in its nature. In a majority of

fishes the back is olivaceous or gray, either plain or mottled,

and the belly white. To birds looking down into the water, the

back is colored like the water itself or like the bottom below it.

To fishes in search of prey from below, the belly is colored like

the surface of the water or the atmosphere above it.

In shallow waters or in rivers the bottom is not uniformly

colored, The fish, especially if it be one which swims close

to the bottom, is better protected if the olivaceous surface is

marked by darker cross streaks and blotches. These give the

fish a closer resemblance to the weeds about it or to the sand

and stones on which it lies. As a rule, no fish which lies on

the bottom is ever uniformly colored.

In the open seas, where the water seems very blue, blue

colors, and especially metallic shades, take the place of olivace-

ous gray or green. As we descend into deep water, especially

804 THE AMERICAN NATURALIST. [VoL. XXXVF.

in the warm seas, red pigment takes the place of olive. At

a depth of 50 to 150 fathoms in the tropics a large percentage

of the fishes are of various shades of red. Several of the large

groupers of the West Indies are represented by two color

forms, a shore form in which the prevailing shade is olive

green, and a deeper water form which is crimson. In one

case an intermediate color form also exists, which is lemon

yellow. On the coast of California is a band-shaped blenny

which appears in three colors, according to its surroundings,

blood red, grass green, and olive yellow. The red coloration is

also essentially protective, for the region inhabited by such

forms is the zone of the rose-red alga. In the arctic waters,

and in lakes where rose-red alga are not found, the red ground

coloration is almost unknown, although red may appear in

markings or in nuptial colors. It is possible that the red, both

of fishes and alga, in deeper water is related to the effect of

water on the waves of light, but whether this should make

fishes red or violet has never been clearly understood. It

is true, also, that where the red in fishes ceases violet black

begins.

In the great depths, from 500 to 4000 fathoms, the ground

color in all fishes becomes deep black or violet black. This

shade is also protective. In these depths the sun's rays

scarcely penetrate, and the fish and the water are of the same

apparent shade, for black coloration is here the mere absence

of light.

In general the markings of various sorts grow less distinct

with the increase of depth. Bright red fishes of the depths

are usually uniform red. The violet-black fishes of the oceanic

abysses show no markings whatever (luminous gland excepted),

and in deep waters there are no nuptial or sexual differences

in color.

Ground colors other than olive green, gray, brown, or silvery

rarely appear among fresh-water fishes. Marine fishes in the

tropics sometimes show as ground color bright blue, grass

green, orange yellow, or black; but these showy colors are

almost confined to fishes of the coral reefs, where they are

often associated with elaborate systems of markings.

No. 430.] THE COLORS OF FISHES. 805

Protective Markings. —The markings of fishes are of almost

every conceivable character. They may be roughly grouped

as protective coloration, sexual coloration, nuptial coloration,

recognition colors, and ornamentation, if we may use that term

for brilliant hues which serve no obvious purpose to the fish

itself.

Examples of protective markings may be seen everywhere.

The flounder which lies on the sand has its upper surface cov-

ered with sandlike blotches, and these again will vary according

to the kind of sand it imitates. It may be true sand or crushed

coral or the detritus of lava, in any case perfectly imitated.

Equally closely will the markings on a fish correspond with

rock surroundings. With granite rocks we find an elaborate

series of granitic markings, with coral rocks another series of

shades, and if red corals be present, red shades of like appear-

ance are found on the fish. Still another kind of marks indi-

cates rock pools lined with the red calcareous alge called

Corallina. Black species are found in lava masses, grass-green

ones among the fronds of ulva, and olive-green among Sargas-

sum or fucus, the markings and often the form corresponding

to the nature of the algz in which the species makes its home.

Sexual Coloration. — In many groups of fishes the sexes are

differently colored. In some cases bright red, blue, or black

markings characterize the male, the female having similar

marks, but less distinct, and the bright colors replaced by olive,

brown, or gray. In afew cases, however, the female has marks

of a totally different nature, and scarcely less bright than those

of the male. ,

Nuptial Coloration. — Nuptial colors are those which appear

on the male in the breeding season only, the pigment after-

wards vanishing, leaving the sexes essentially alike. Such

colors are found on most of the minnows and dace (Cyprinidz)

of the rivers and to a less degree in some other fresh-water

fishes, as the darters (Etheostominz) and the trout. In the

minnows of many species the male in spring has the skin

charged with bright pigment, red, black, or bright silvery, for

the most part, the black most often on the head, the red on

the head and body, and the silvery on the tips of the fins.

806 THE AMERICAN NATURALIST. | [Vor. XXXVI.

At the same time other markings are intensified, and in many

species the head and sometimes the body and fins are covered

with warty excrescences. These shades are most distinct on

the most vigorous males, and disappear with the warty excres-

cences after the fertilization of the eggs.

Nuptial colors do not appear among marine fishes, and in but

few families are the sexes distinguishable by differences in

coloration.

Recognition Marks. — Under the head of “recognition marks”

may be grouped a great variety of special markings, which may

be conceived to aid the representatives of a given species to

recognize each other. That they actually serve this purpose is

a matter of theory, but the theory is plausible, and these mark-

ings have much in common with the white tail feathers, scarlet

crests, colored wing patches, and other markings regarded as

recognition marks among birds.

Among these are ocelli, black or blue ringed with white or

yellow, on various parts of the body ; black spots on the dorsal

fin; black spots below or behind the eye; black, red, blue, or

yellow spots variously placed; crossbars of red or black or

green, with or without pale edges; a blood-red fin among pale

ones or a fin of shining blue; a white edge to the tail ; a yellow,

blue, or red streamer to the dorsal fin, a black tip to the pec-

toral or ventral; a hidden spot of emerald in the mouth or in

the axil ; an almost endless variety of sharply defined markings,

not directly protective, which serve as recognition marks, if not

to the fish itself, certainly to the naturalist who studies it.

These marks shade off into an equally great variety for which

we can devise no better name than “ornamentation.” Some

fishes are simply covered with brilliant spots or bars or reticu-

lations, their nature and variety baffling description, while no

useful purpose seems to be served by them, unless we stretch

still more widely the convenient theory of recognition marks.

In many cases the markings change with age, certain bands,

stripes, or ocelli being characteristic of the young and gradu-

ally disappearing. In such cases the same marks will be found

permanent in some related species of less differentiated colora-

tion. In such cases it is safe to regard them as ancestral.

No. 430.] THE COLORS OF FISHES. 807

In case of markings on the fins and of elaborate ornamenta-

tion in general, it is best defined in the oldest and most vigorous

individuals, becoming intensified by degrees.

Intensity of Coloration.—In general, coloration is most

intense and varied in certain families of the tropical shores,

and especially about coral reefs. But in brilliancy of individual

markings some fresh-water fishes are scarcely less notable,

especially the darters (Etheostominz) and sunfishes (Centrar-

chidz) of the streams of eastern North America. The bright

hues of these fresh-water fishes are, however, more or less

concealed in the water by the olivaceous markings and dark

blotches of the upper parts.

Coral Reef Fishes. —The brilliantly colored fishes of the

tropics seem to scorn the need of protective coloration. They

save themselves from their enemies in most cases by excessive

alertness and activity (Chatodon, Pomacentrus), or else by

burying themselves in coral sand (/w/es gaimardi), a habit

more frequent than has been suspected.

Fading of Pigments in Spirits.— In the preservation of

specimens most red and blue pigments fade to whitish, and it

requires considerable care to interpret the traces which may be

left of red bands or blue markings. Yet some blue pigments

are absolutely permanent, and occasionally blood-red pigments

persist through all conditions. Black pigment seldom changes

in spirits, and olivaceous markings simply fade a little without

material alteration. It is an important part of the work of the

systematic ichthyologist to learn to interpret the traces of the

faded pigment left on specimens he may have occasion to

examine. In such cases it is more important to trace the

markings than to restore the ground color, as the ground color

is at once more variable with individuals and more constant in

large groups. In other words, it varies a good deal, but in

almost constantly narrow limits.

Variation im Pattern. — Occasionally, however, a species is

found in which, other characters being constant, both ground

color and markings are subject to a remarkable range of varia-

tion. In such cases the actual unity of the species is open to

serious question. The most remarkable case of such variation

808 THE AMERICAN NATURALIST.

known is found in a West Indian fish, the vaca, which bears

the incongruous name of Hypoplectrus unicolor. In the typical

vaca the body is orange with black marks and blue lines, the

fins checkered with orange and blue. In a second form the

body is violet, barred with black, the head with blue spots and

bands. In another form the blue on the head is wanting. In

still another the body is yellow and black, with blue on the

head only. In others the fins are plain orange, without checks,

and the body yellow, with or without blue stripes and spots, and

sometimes with spots of black or violet. In still others the

body may be pink or brown, or violet black, the fins all yellow,

part black orallblack. Finally, there are forms deep indigo blue

in color everywhere, with cross bands of indigo black, and these

again may have bars of deeper blue on the head or may lack

these altogether. I find no difference aniong these fishes

except in color, and no way of accounting for the differences

in this regard.

A species of puffer (Tetradon setosus) shows similar remark-

able variations, being dark gray with white spots, but varying

to indigo blue, lemon yellow, or sometimes having coarse

blotches of either. Lemon-yellow varieties of several species

are known, and these may be due to a failure of pigment, a

sort of semi-albinism. True albinos, individuals wholly with-

out pigment, are rare among fishes.

FLOWERS AND INSECTS IN NEW MEXICO.

T. D. A. COCKERELL.

IN a certain sense we may say correctly that the flora of our

country is fairly well known. New species, and even genera,

are frequently described, but most of these are segregates from

the more comprehensive groups of earlier authors, and radically

new types are not often met with. Nevertheless, there is

hardly a plant upon which new observations may not be made

with ease. Our whole flora needs redescribing from living

plants, every species needs close study to determine the char-

acter and range of its variations, and the relations between

plants and insects offer a field for research which appears

practically inexhaustible. The following notes record the

results of some recent researches, and may serve as a con-

tribution to entomophytology. They are classified according

to locality.

Rio Rurposo, WHITE MOUNTAINS.

This is an alpine region in southern New Mexico, possessing

a remarkably interesting flora, with quite a number of apparently

endemic types. Prof. C. H. T. Townsend collected there a large

number of bees, together with the flowers on which they were

found. Some of these have been recorded, but the following

data are new: |

(1) Verbena macdougalii Heller. A very common species in New Mexico,

formerly regarded as V. stricta, but separated by Heller on what

seem to be perfectly valid grounds. The lilac-purple flowers are

arranged in long spikes, so that the plant looks entirely different from

the ordinary kinds of Verbena, which are adapted to butterflies. At

the foot of Baldy Mountain, near Elizabethtown, Mrs. O. St. John

found a variation (mut. nov. rosella) with pink flowers. The ordinary

form was growing at the same place, which had an altitude of about

9600 feet.

809

810 THE AMERICAN NATURALIST. [Vor. XXXVI.

Although our common types of Verbena (V. macdougalii, V. bipinnati-

fida, and V. bracteosa) look so entirely different, they all have the

throat of the corolla covered with the same peculiar moniliform hairs.

V. macdougalii is an excellent bee plant. On the Rio Ruidoso, Town-

send collected from the flowers the following bees: Anthophora

cleomis Ckl. 9, A. montana Cress. 9, Clisodon terminalis Cress.,

Anthidium maculosum Cress., Anthophora californica Cress. 8,

Megachile fortis Cress., Melissodes ruidosensis 5 t

crenulaticornis Ckll. $, Calliopsis rhodophilus Ckl. 9. It will be

noted that these are long-tongued bees.

(2) Monarda stricta Wooton. Another very common New Mexico plant,

only lately recognized as a distinct species. Its conspicuous heads

of pink flowers are attractive to certain long-tongued bees, of which

Townsend collected Anthophora cardui Ckl. $, A. cleomis Ckll. d,

Clisodon terminalis Cress., and Megachile fortis Cress. The flowers

are also visited by Bombus, as I have recorded elsewhere (Annals

and Magazine Natural History, November, 1899), and Town-

send took a single male Melissodes montana Cress. He also took

a butterfly, Epargyreus fityrus Fabr., not hitherto recorded from

New Mexico. These results accord fairly well with those of Loew,

who observed the insect visitors of Monarda in the Botanic Garden

at Berlin.

Bombus. Other long-tongued bees. Short-tongued bees. Butterflies.

M. stricta, New Mexico 9 5 o

M. sf., Berlin 2 I 2

Both lists are doubtless very incomplete.

(3) Rhus glabra L. A widely distributed shrub, with conspicuous panicles

of greenish flowers. The visitors might be expected to be much like

those of the Umbelliferz, but Townsend actually obtained a number

of bees: Bombus sonorus Say, B. prunelle@ Ckll, Colletes gilensis

Ckl. 3, Coelioxys gilensis Ckll, Heriades gracilior Ckll., Colletes

americana Cress. $. He also obtained a butterfly, Basz/arcAza weide-

meyerit Edw. In Europe Müller found the honeybee abundant on

the allied Rhus hirta. ln Illinois Robertson found fifty-eight species

of insects on x Reto of which nineteen were bees, mostly short-

tongued.

(4) Potentilla thurberi Gray. This is a beautiful species, differing from

ie ordinary species of the genus by its dark crimson flowers, on which

. Townsend took the following : ro fortis Cr., Colletes gülensis

Ckll. 8, Vespa occidentalis C

(5) Geranium atropurpureum Heller. A common New Mexico plant,

only r recently recognized as distinct. Visited by males of Co//efes

qs CklL, a short-tongued bee. -

No. 430.] FLOWERS AND INSECTS IN NEW MEXICO. 811

(6) Heliopsis scabra Dunal. Visited by Megachile fortis Cr, Anthidium

perpictum Ckll., Megachile fidelis Cr., — all laoa bees.

From Z7. APT in Illinois, Robertson collected twenty-two

bees, fourteen being long-tongued.

A few other miscellaneous observations are to be recorded :

Anthophora cleomis, 8, at Prunella z vulgaris, and 9 at Verbascum

thapsus ; Clisodon terminalis at Prunella vulgaris and Verbas-

cum thapsus ; Halictus angustior at Er rysimum asperum; Melis-

sodes ruidosensis at Erigeron macranthus,; Vespa diabolica at

Solidago trinervata and Sicyos parviflorus ; Andrena barberi at

Solidago trinervata.

La Curva, ORGAN MOUNTAINS.

This locality, also in southern New Mexico, is at the eastern

side of the Mesilla valley, and has an altitude of about 5000 feet.

The records given below are based on collections by Prof. C.

H. T. Townsend :

(1) Datura meteloides DC. A magnificent species, with large, heavily

scented white flowers. Although no doubt properly a sphinx-moth

flower, it was visited by the following bees: Xylocopa arizonensis Cr.,

Caupolicana yarrowi Cr., Anthophora montana Cr., Agapostemon

viridulus Fabr. (new to New Mexico), Augochlora neglectula Ckll.

The Caupolicana flies early in the morning, before sunrise. According

to Mr. Friese (zz Z//£.) the Chilian C. curvipes Friese (3) agrees with

C. yarrowit.

(2) Lippia wrightii Gray (Verbenacez). A shrubby plant, characteristic

of the re egion. vo y Xylocopa arizonensis Cr., Caupolicana

yarrowi Cr., hora montana Cr., Perdita albovittata Ckll.

(12), 4 score avai CklL, Syahalonia crenulaticornis

Ckll var. $ (clypeus yellow except the hind border broadly), — all

bees, five long-tongued.

(3) Tourerea multiflora (Nutt.) Visited by pase Perdita mentzeliaru

Ckil. and Anthophora californica Cr. 1 learn from Dr. iiber

that Tourerea, Eaton and Wright, takes the oig of Hesperaster ;

the combination here given will be credited to h

(4) Phacelia congesta Hooker. Visited p bees: Ceratina nanula Ckll.

Ge 9,138), Halictus ruidosensis Ckll. (19), Perdita phaceliz

These are all bees.

812 THE AMERICAN NATURALIST. [VoL. XXXVI.

Other observations are: /Vomada gutterrezig Ckll., on Ver-

besina encelioides; Naomia foxi D.T. and Exomalopsis solani

Ckll. on Solanum eleagnifolium ; Augochlora neglectula Ckll.,

at Parthenium 2ncanum.

ROMEROVILLE.

A locality about six miles from Las Vegas. In Entomological

News, 1901, p. 40, five insects are recorded from Ribes, sf. at

this place. It can now be stated that the Ribes is R. Zeptan-

thum veganum Ckll. (Proceedings of the Biological Society of

Washington, 1902, p. 99).

Las: VEGAS.

(1) yeas verticillata L. This is very attractive to flies; the following

e taken from its flowers: Syritta pipiens L. (July 3, N. Stern),

paes latifrons Loew (July 3, M. Winters), Echinomyia algens

Wied. (July 21), Peleteria iiis Fab. (July 21), Myiophasia

ænea Wied. (July 21). Lygeus reclivatus Say is also common on

the plant. Robertson collected 115 species of insects from this plant

in Illinois. -

(2) ger oct Nutt. The yellow-flowered currant, common both

and cultivated in the vicinity. On May 12 Loyola Dillon and

Olive Barnes collected on the flowers four species of bees and

one wasp: Bombus nevadensis aztecus Ckll. 9, Halictus coriaceus

Smith 9, H. armaticeps Cr. 9, H. pruinosus Rob. 9, Odynerus, sf.

On May 9 of last year a 9 Synhalonia frater, Cr. var. (new to New

. Mexico), was taken at the flowers. This and the Bombus can prob-

ably get some nectar, but the Halicti only get pollen.

(3) Anogra albicaulis is a large white evening primrose, doubtless intended

Tor moths, but on May 29 Eldon Tuttle and Leo Tipton discovered a

minute new bee of the genus Perdita visiting its flowers at Las Vegas.

— Perdita anogre, n. sp, 8. Length about 33 mm.; head and thorax

. very dark olive green; cheeks unarmed; face below antenna all

bright. yellow except the clypeal dots, yellow not extending upwards

n middle line, but at sides going a little above the antennz and then

terminating abruptly, just touching the facial fovea, making slightly

. more than a right p with the eye ; m pale green, the lower

half of P. efi

th y bordered with yellow; face

and mesothorax not conspicuously hairy ; antennae pale orange, dark

brown above as far as third joint of flagellum ; vertex and mesothorax

bcne sculptured, but rather shining ; tubercles light — pleura

No. 430.] FLOWERS AND INSECTS IN NEW MEXICO. 813

all dark; tegule pale brownish, with a yellowish spot ; wings rather

small, hyaline, brilliantly iridescent; nervures and stigma sepia brown,

third discoidal distinct ; marginal cell obliquely truncate ; second sub-

marginal narrowed to a point above ; legs brownish-orange, the femora

black behind, tibiae with a dark brown stripe on outer side, tarsi with

at least the last joint darkened ; abdomen rather broad, shining black,

with transverse wedge-shaped bluish-white marks on the sides of seg-

ments two to four; ventral surface black.

In my table in Bulletin of Laboratory of Denison University, XI, this

runs to section 30, where it is at once distinguished by its spotted

abdomen. It is closely allied to P. sexmaculata, agreeing in the

ornamentation of the face and color of the antennz, but the color

of the abdominal spots is entirely different, as also is the shape of

the second submarginal cell, and P. anogre is a erred insect than

sexmaculata.

(4) Verbena bipinnatifida Nutt. is a plant with umbel-like heads of bril-

liant t purple flowers, allied to the Verbena of gardens, and adapted to

butterflies. On June 7 I saw it visited by the butterfly Pyrameis

cardui and the day-flying hawk moth Dedlephila lineata; but to my

surprise I also saw a female Anthophora montana, a long-tongued

bee, sucking vigorously and apparently successfully. She had not

collected any pollen

(5) Phlox nana Nutt. A small species characteristic-of our region, with

brilliant pink flowers. It is adapted to butterflies, but on June 7 I saw

a short-tongued bee, Agapostemon texanus, - repeatedly trying to

suck and of course getting nothing. ias n, in his revision of `

Phlox, says the corolla of JP. nana is “red” or ** white." It is always

whitish beneath, but the üpper surface is typically bright pink, vary-

ing, however, at Las Vegas to pale mauve (forma lilacina) and white

with a large pink “eye” (forma oculata), the markings round the

throat remaining the same in each case.

(6) Sophora sericea Nutt. is a plant allied to Astragalus, with conspicuous

w l It seems to be adapted to bumblebees, and is visited

by Bombus morrisoni.

ENGLE.

Engle is a locality near the Rio Grande about fifty miles

north of Las Cruces. Nothing has hitherto been known of

its insect fauna.

Astragalus bigelovii Gray. One of the so-called **loco-

weeds " common in the southern part of New Mexico in open

ground. The leaves are densely covered with white silky hairs.

814 THE AMERICAN NATURALIST. [VoL. XXXVI.

Miss Nora Newberry, during the latter part of April and the

first few days of May, collected from the flowers a number of

long-tongued bees, which prove to be as follows :

(1) Sy#halonia lycii Ckll., both sexes ; the male was not before known.

In the male the clypeus, labrum, and a small, broadly triangular

supraclypeal area are cream-color. (In the males of .S. edwardsii

and frater the clypeus is very bright yellow, and there is no supra-

clypeal mark.)

(2) Anthophora affabilis Cr. 9. ‘The identity of this has been confirmed

by Mr. Fox, who compared it with Cresson's type. In the Mesilla

valley it visits Lycium torreyi.

(3) Anthophora portere Ckll. Several females; only two specimens of

this species were previously known. It is readily distinguished from

A. affabilis by the black hair mixed with the light on the mesothorax.

The female nearly agrees with the description of 4. zgnava Cr., but

Mr. Fox has kindly compared it with Cresson's type, and assures me

that it is different. Mr. Fox remarks that the female seems to agree

with 4. edwardsii Cr., except that it is larger; the male, however,

has not the tooth on inner side of hind joint of posterior tarsi, and the

lateral face-marks are not “lanceolate,” but are shaped like a rose-

thorn. A male 4. porteræ was taken at flowers of Ribes longiflorum,

at Las Vegas, N.M., May 15, 1902, by Eldon Tuttle and Leo Tipton.

It differs from the original type in having more yellow on the scape

and rather more black hair on the mesothorax. A character over

looked in the original description is a little pencil of black hair

overlapping the upper anterior corner of each eye.

It is to be noted that both Synhalonia and Anthophora visit

Ribes, Lycium, and Astragalus, plants of very different affinities

— but similar in their adaptation to long-tongued bees

Trout SPRING.

This is a locality in Gallinas Cafion, N.M., some miles above

Las Vegas Hot Springs. It is of interest on account of the

mixed boreal and austral elements in its fauna and flora, the

result in large measure, no doubt, of the narrowness of

the cafion, whereby some slopes get little or no sun, while

others are well warmed. The abundance of Pinus scopulorum

and Populus angustifolia indicate the Transition Zone, while

Heracleum lanatum, Dasiphora fruticosa, and Ribes irriguum are

No. 430.] FLOWERS AND INSECTS LN NEW MEXICO. 815

distinctly Canadian Zone types. A fine bush of the last men-

tioned grew on the north side of a huge rock, shaded from the

sun. An Upper Austral representative is Ribes longifiorum

(a variety, however, with the petals deep red, the calyx tube

often reddish outside, the flowers about 14 mm. long, and the

apex of the leaves obtuse, thus closely approaching the north-

ern R. aureum, as restricted by Coville), while at its flowers we

found the typically southern butterfly Epargyreus tityrus Fabr.,

which extends to South America and the West Indies. The

bright yellow flowers of the Canadian Zone Thermopsis were

visited by a female Megachile wootoni Ckll., a bee of a strictly

alpine and boreal type. A magpie (Pica pica hudsonica) was

also to be regarded as a boreal representative. The pink flowers

of the Upper Austral and Transition Phlox nana were seen in

the cafion a short distance below Trout Spring.!

The insect visitors of one species of plant were rather care-

fully studied by my wife and myself, May 24, 1902.

dris missouriensis Nutt. Visited by bees, butterflies, and

flies. The bees crawl in under the petaloid divisions of the

style, and so doubtless effect cross fertilization. The butter-

flies and flies (Bombyliide) suck from between the perianth

segments, and appear to get their meal without paying for it.

The following insects were observed :

(1) Colias eurytheme Q, Lycena, Thanaos, — all butterflies, sucking.

(2) Bombylius major L., with proboscis 8-8} mm. long, very abundant,

sucking. In Europe a syrphid (Rhingia) with a proboscis 11 mm.

long visits /ris pseudacorus, as is described by Müller.

The remaining species are bees :

(3) Megachile wootoni Ckll., Megachile latimanus Say. A male of each

found sitting on the flowers, perhaps waiting for the females.

(4) Synhalonia frater Cresson. One of each sex.

(5) Anthophora bomboides neomexicana Ckll. One &.

(6) Halictus coriaceus Smith. One 9.

! Mr. Eldon Tuttle lately found in Las Vegas a flower of Phlox nana with the

limb 8-parted, a character which, if not aberrational, would take it out of the

Polemoniacez. I have since found a flower with the limb 7-parted.

816 THE AMERICAN NATURALIST. [Vor. XXXVI.

(7) Augochlora confusa Rob. Two 9. These differ from a specimen

received from Mr. Robertson in having the basal area of metathorax

bounded by a sharp rim, but Robertson’s description indicates that

such specimens occur also in Illinois. The species is new to New

Mexico.

(8) Osmia pusilla Cresson, one $; Osmia, two new species, males. I have

sent these to Mr. Titus, who is revising the genus.

Las VEGAS Hor SPRINGS.

Verbena macdougalii was visited July 11 by bombyliid flies,

Systoechus vulgaris Lw. (det. Coq.), as observed by Miss M.

Holzman.

Ribes cereum Dougl. was visited May 24 by Bombus juxtus

Cresson.

I will take this opportunity to record from Las Vegas Hot

Springs the dragon fly Hyponeura lugens Hagen. Prof. J. A.

Needham, who kindly identified it, states that the genus is new

to the United States, but there are specimens from Arizona

in the Cornell University collection, not hitherto recorded.

APPENDIX.

Two New Species of Osmia.

The two new species of Osmia collected on flowers of Iris at

Trout Spring, N.M., were sent to Mr. Titus with the expecta-

tion that they might prove identical with species which he had

already described in MS. As this is not the case, they are

briefly diagnosed below. Some additional notes and comments

will be given by Mr. Titus when he publishes his revision of

the American species of Osmia.

Osmia iridis Cockerell and Titus, z. sf.

$. Length 9 mm., stout; head and thorax yellowish-green ; abdomen

dark blue-green; antennz long, slender, entirely black, subtruncate at

apex; pubescence of head and thorax abundant, erect, white; vertex

very broad ; mandibles entirely black ; legs black, with black hair (white

on first four femora), hind femora slightly bluish ; abdomen short and

broad, with white hair on first segment, on the others mixed black and

white ; sixth segment reflexed, entire. Hind tibial spurs black, strongly

No. 430.] FLOWERS AND INSECTS IN NEW MEXICO. 817

curved at tips; eyes perfectly black; cheeks and sides of vertex with

some long black hairs mixed with the pale; tegule black, with a

slight green tinge; wings slightly dusky, nervures black ; second sub-

marginal cell long; anterior edge of clypeus nodulose; clypeus and

front as densely punctured as is possible. Mr. Titus adds: * A very

distinct species; apical margin of second ventral segment is different

from any species heretofore seen. The antennz are crenulated slightly

beneath, and this gives them slight resemblance to males of Zgma7ia

section, but the species does not belong there. It would be well to

note the hairiness of the labial palpal joints 1 and 2

Osmia chlorops Cockerell and Titus, x. sf.

4$. Length about ro mm., brassy green, with bluish tints on abdomen

and thorax; the abdomen shining, the head and thorax densely punc-

tured; pubescence white, long and abundant on face; legs strongly

tinged with green. Head large, face almost golden; eyes (in life)

green, black anteriorly ; mandibles black; antennz long, black, crenu-

lated ; first joint of flagellum covered with a seal-brown velvety pile ;

tegulz with greenish punctured margins ; wings somewhat dusky ; hind

tarsi with the basal joint broadened distally, and covered on the inner

side with short brown-black hair; hind tibial spurs black, curved at

tips; sixth dorsal segment of abdomen notched ; apical segment deeply

and broadly notched; third ventral segment deeply and broadly emar-

ginate, the edges of the emargination fringed with short shining hairs.

NOTES AND LITERATURE.

ZOOLOGY.

Gardiner's ** Maldive and Laccadive Archipelagoes,’’ Part II.! —

This second part contains ten “reports.” The first, by F. F. Laidlaw,

deals with the Amphibia and Reptilia. Allbutone of the species are

abundant in Ceylon, and the exception, Lygosoma albopunctatum, is a

lizard that is widespread in the Oriental region. For most of the

species the region in question is near the western limit of its area of

distribution. Only two Amphibia are listed, a Rana and a Bufo ;

four lizards, two Hemidactyli, a Calotes, and the Lygosoma; bos

snakes; one species of Testudinidæ (Nicoria and two marine

turtles (Chelone).

The Lepidoptera are listed by E. Meyrick. "There are sixty-seven

species, of which practically all occur also in India or Ceylon. A

single species of Notarcha, regarded as new, replaces the widespread

LV. multilinealis.

Mr. A. E. Shipley has enumerated the six species of Echiurodea

and nineteen sipunculoidea. The former belong to the genera

Bonellia and Thalassema, characteristic of the warm and temperate

seas. A new genus of sipunculids, Lithacrosiphon, allied to Aspido-

siphon, is described and figured.

The land and fresh-water mollusks are treated by E. A. Smith.

He says: * The present collection comprises eight land and two

fresh-water forms from the Maldives, and four terrestrial species

from the Laccadives, three of which are included among those

from the Maldives. The latter group, judging from the collection at

hand, does not possess a single indigenous species, all the forms

occurring either on the Indian peninsula or in Ceylon or other locali-

ties. As many plants are introduced into the islands by foreign

vessels, some or all of the Mollusks there may be accounted for by

importation. The list includes Succinea vitrea, known elsewhere only

from Calcutta and Bombay; a new species, Sifa/a vagata, probably

1 The Fauna and Geography id the Maldive and Laccadive Archipelagoes, etc.

Edited by J. Stanley Gardiner. Cambridge, University Press, 1902. Vol. i, pt. ii,

PP- vi, 119-222, Pls. VI-XIII, fes 24-40.

819

820 THE AMERICAN NATURALIST. [Vor. XXXVI.

from India; Zuplecta indica, var. malabarica, found on the adjacent

Malabar coast; Xestina bombayana, agreeing “in all respects with

those from the mainland of India"; Rhachis punctatus and Opeas

gracilis, which are resistant forms associated with plants, with which

they may be transported; Tornatellina manilensis; Melampus casta-

neus, from the shore, a wide ranging species ; Leptopomoides halophilus ;

Melania tuberculata, found from Syria to Australia and West Africa;

and Cyrena ceylonica, of mangrove swamps.

The pigments of corals have been investigated by Dr. MacMunn,

who finds them “ either chlorophylloid or of a closely connected kind

of pigment, which latter absorbs the violet end of the spectrum and

seems generally changeable into the next kind by the agency of heat,

etc., namely, into the dark pigment which gives the coral its dark

color in the fresh condition.”

The sixteen species of Chetognatha are treated by L. Doncaster.

In his consideration of the variation and distribution of the group

he makes many interesting points. He lays stress on the fact that

since closely allied species are found living together without barrier

of habitat, they can hardly have arisen through natural selection ;

[but it must not be forgotten that there are others than spacial bar-

riers]. Remarkable is the occurrence here of a species (Sagitta

flaccida) previously described only by Conant from the Bahamas.

The six dragon flies are listed by F. F. Laidlaw. One seems to be

new, the others are well known Ceylon and Indian species, or have a

wider distribution.

The first installment of a series of papers on marine crustaceans

is given by L. A. Borradaile. He here considers the Portunidz, a

family that “is highly variable and varietal [exhibiting varieties]

and is probably undergoing rapid evolution in many directions.”

The account of each species is accompanied by valuable notes

on its variation.

The longest contribution is by Gardiner, continuing his account of

the coral formation of the islands. It is clear and interesting. He

assumes, first of all, the former existence of a continent connecting

India and Madagascar, which underwent subsidence, leaving certain

elevations where mountain peaks were. The archipelagoes occupy

the position of one or two of these mountains. Second, the moun-

tains became eroded by the action of oceanic currents to nearly level

plateaux two hundred fathoms below the surface. Third, upon these

plateaux there began to grow deep sea corals, assisted by nullipores

and other organisms. By growing on the skeletons of their ancestors

No. 430.] NOTES AND LITERATURE. 821

these corals formed banks up to forty or fifty fathoms, at which

height the true reef-building corals were able to attach themselves. In

time they gained the supremacy and built up the reef to near the sur-

face. A lagoon was formed in the center of the reef partly by the

more rapid growth of the organisms on the edge of the bank and

partly by the subsequent solution of the central parts. This article

is well illustrated by charts and diagrams. CRD.

Jordan and Heath’s ‘Animal Forms.” ! — This “second book of

zoölogy ” is the companion volume of the already well-known Anima/

Life by Jordan and Kellogg, and the two books may be obtained

separately or bound together in one volume with the title Animals.

Whereas the first volume, Animal Life, dealt with animal ecology,

the second book gives an elementary account of animal morphology.

The introductory chapters deal with the differences between animals

and plants, the characteristics of an animal as exemplified by a mam-

mal, and the cell and protoplasm. Then follows an examination of

typical representatives of the great groups, from the simplest to the

most complex. While animals are here considered primarily from

the morphological viewpoint, considerable attention is given to

functions of organs and to habits and life histories.

Animal Life was designed for the first half year in zoology in the

high school, and Animal Forms to complete a full year’s work.

However, Animal Forms may be used alone or as the first book.

With regard to the adaptability of the book to the conditions

obtaining in secondary schools, Animal Forms and its companion

volume are unquestionably the best existing books for supplementary

reading for the high-school beginners ; but as a žastis for an elementary

course they are far from practicable, for it will be difficult to follow

either book in close correlation with laboratory study. In spite of

the authors’ emphasis upon “a basis of observation," there has

already been manifested in the use of Animal Life in some schools

a tendency to drift backward towards the old-time recitation method.

The informational side of zoólogical teaching must not be allowed

to displace scientific training, and a definite laboratory course, and

not a text-book, must give the foundation, to which may be added

supplemental and closely correlated reading. As sources for such

collateral information from which high-school teachers may select

topics relating to the laboratory study, Animal Forms and Animal

1 Jordan, David Starr, and Heath, Harold. New York, Appleton, 1902. 8vo,

vi + 258 pp., 140 figs.

822 THE AMERICAN NATURALIST. [VoL. XXXVI.

Life deserve the highest commendation, and every pupil in elementary

zoology should be made familiar with them.

A laboratory manual to accompany Animal Forms is announced as

in preparation. This perhaps will throw some light upon the problem

of correlation between laboratory work and the use of the text.

Aside from the secondary schools, where undoubtedly the books

will have the greatest sale, the combined volume, Animals, will offer

a mine of information and inspiration to college students who use

it for collateral reading. Moreover, it should be especially recom-

mended to the growing body of readers who eagerly seek general

books relating to animal life. M. A. B.

BOTANY.

Notes. — Late numbers of the Botanical Gazette contain a descrip-

tive list of the plants collected by Dr. F. E. Blaisdell at Nome City,

Alaska, prepared by Miss Eastwood. One hundred and seventy

species are recorded, several of them considered as new to science.

In the Bulletin of the Torrey Club for February Miss Eastwood

describes and figures a number of new Californian anthophytes.

Dr. Rydberg's * Studies on the Rocky Mountain Flora," VII, in

the March Bulletin of the Torrey Club, contains a number of new

Ranunculacez, Papaveracez, and Fumariacez.

In No. 4 of the current volume of Transactions of the Academy of

Science of St. Louis, Professor Norton discusses the unexplored

botanical regions of the Southwest, and describes and figures a

number of new spermatophytes.

Several southwestern plants are named by Cockerell in Torreya

for March.

Several new species of trees from the Eastern and Southern States

are described by Ashe in the March number of the Botanical Gazette.

Fifty additions to the “Catalogue of Ohio Plants” are made by

Kellerman in the Ohio Naturalist for December last.

A list of the climbing plants of Ohio is published by Miss Dufour

in the Okio Naturalist for February.

Dr. Harshberger gives an account of a botanical ascent of

Mt. Katahdin, Maine, in the Plant World for February. The

article is illustrated by half tones and zonal diagrams.

No. 430.] NOTES AND LITERATURE. 823

In Vol. LIII, Part III, of the Proceedings of the Academy of Natural

Sciences of Philadelphia, Dr. Harshberger has an illustrated paper on

the ecology of the San Domingo flora.

Vol. XX of the Acta Horti Petropolitani consists of the first part

of a “Flora Manshuriz" by V. L. Komarov, occupied with about

one hundred pages of bibliography and discussion, in Russian,

followed by the pteridophytes and monocotyledons as far as the

Orchidacez.

Hemsley and Pearson publish, in No. 224 of the Journal of the

Linnean Society (Botany), an account of what is known of the flora

of Tibet.

A paper by Willis and Gardiner, on the botany of the Maldive

Islands, is published in Vol. I, Part II, of the Annals of the Royal

Botanic Gardens, Peradeniya.

In the opening number of the current volume of Nyt Magazin

Jor Naturvidenskaberne, Prof. N. Wille contrasts the present vege-

tation of the Norwegian district of Telemarken with the records for

the same district published by H. J. Wille in 1786.

A very interesting detailed study of rheotropism, by Professor

Newcombe, presented at the Chicago meetings of last winter, has

been published in the Botanical Gazette.

Mendel’s laws of inheritance in hybrids and crossbreeds are dis-

cussed by Weldon in Biometrika for January, and by Correns in

Heft 3 of the current volume of Berichte der deutschen botanischen

Gesellschaft.

An illustrated article on caprification as practiced in Algeria,

appears in the January number of the Revue horticole de l Algérie.

The self-pruning of plants forms the subject of an article by

Schaffner in the Ohio Naturalist for January.

Photosynthesis is being passed in review by André Richter in

current numbers of the Revue générale de botanique.

A portrait of the Spanish botanist Colmeiro, with biography, is

published by Lázaro e Ibiza in Vol. XXX of the Anales de la Sociedad

Española de Historia Natural.

A portrait of Alvah A. Eaton forms the frontispiece to the April

number of the Fern Bulletin.

A portrait of von Heldreich appears on the first page of the

Deutsche botanische Monatsschrift for March.

824 THE AMERICAN NATURALIST. [Vor. XXXVI.

Livraison 2 of the current volume of the Bulletin du Jardin impé-

riale de St.-Pétersbourg contains a portrait of the late Dr. J. Klinge.

In the double number (149-150) of the Bulletin de /Mcadémie

Internationale de Géographie Botanique is published an excellent

portrait of the veteran Chilian botanist, R. A. Philippi, accompanied

by a list of his publications, numbering two hundred and twenty-six

entries.

A short biographical sketch of Schweinitz, with portrait, is pub-

lished by Mr. Shear in the Plant World for March.

Vol II of Radde's Die Sammlungen des kaukasischen Museums,

published in Tiflis, contains twelve portraits and a considerable

number of plates and maps.

Lamarck's herbarium forms the subject of an article by Bonnet

in the Journal de botanique for April.

Dr. Harshberger has an article on the botanical gardens of Jamaica,

in the Plant World for March..

Vegetable pathology, and the method of teaching it as practiced

in St. Louis, are discussed by Dr. von Schrenk, of the Shaw School

of Botany, in the Bulletin of the Torrey Botanical Club for February.

. Professor Marshall-Ward considers predisposition and immunity,

in plant diseases, in Vol. XI, Part V, of the Proceedings of the

Cambridge Philosophical Society.

In the labels for his exsiccate of Ohio fungi, reprinted in current

numbers of the Oio Naturalist, Professor Kellerman gives tran-

scripts of the original descriptions of the species.

Kellerman and Jennings, in the Ohio Naturalist for April, give

details of experiments to test the comparative susceptibility of maize

and sorghum to Cintractia when taken from different hosts.

— Fusarium Lini and the “ flax-wilt” that it causes are the subject of

Bulletin No. 50 of the North Dakota Experiment Station.

Papers on fungi, by Peck, Salmon, and Long, are published in the

Bulletin of the Torrey Botanical Club for Februa ry.

The ascomycetous genera Urnula and Geopyxis are considered by

Miss Kupfer in the Buletin of the Torrey Botanical Club for March.

Current numbers of Zorreya contain keys to the species of a

number of genera of agarics, by Earle.

The Botanical Gazette for July contains the following articles:

Arthur, * Uredinez occurring upon Phragmites, Spartina, and Arun-

dinaria in America”; Nelson, “Contributions from the Rocky

No. 430. NOTES AND LITERATURE. 82

Mountain Herbarium,” III; Atkinson, “Three New Genera of the

Higher Fungi"; Berry, “ Notes on the Phylogeny of Liriodendron ”;

Cook, * Polyembryony in Ginkgo"; von Schrenk, * Root Rot at

Apple Trees caused by Zhelephora Galaria "; and Wilcox, “ Stipa

fassei not a good species.”

The Bulletin of the Torrey Botanical Club for June contains the

following articles: Kirkwood and Gies, “Chemical Studies of the

Cocoanut, with some Notes on the Changes "— Germination ”

Curtis, “Some Observations on Transpiration ” ; Peirce, * Forcible

Discharge of the Antherozoids in Æsterella Cali ifornica’’; Harper,

“ Taxodium distichum and Related Species, with Notes on some Geo-

logical Factors influencing their Distribution”; N elson, * New

Plants from Wyoming,” XIV; Wight, “The Genus Eritrichum in

North America.”

The Annals of Botany for June contains the following articles :

Yapp, * Two Malayan ‘Myrmecophilous’ Ferns”; Ward, “On the

Relations between Host and Parasite in the Bromes and their Brown

Rust, Puccinia dispersa"; Hill, “On Variation in the Flowers of Cer-

tain Species of Primula”; Copeland, * The Mechanism of Stomata Kd

Thiselton-Dyer, “ Morphological Notes,” VII ; Farmer and Hill, “ On

the Arrangement and Structure of the PESE Strands in Angiop-

teris veris and some other Marattiacez "; and Fritsch, * Algological

Notes.

Vol. XXIII of the Zransactions of the American Microscopical Society,

issued in May, contains the following botanical articles : Jackson, “A

New Species of Crenothrix (C. manganifera)” ; and Bessey, “ Structure

and Classification of the Conjugate, with a Revision of the Families

and a Rearrangement of the North American Genera.”

Heft 9 of Engler's Das Pflanzenreich, a volume of 438 pages, with

numerous figures, is a revision of Myrsinacez by Mez, and bears

date May 6, 1902.

A revision of Conocephalus, by Bargagli-Petrucci, is a prominent

feature of the Nuovo Giornale Botanico Italiano for April.

A revision of the Podostemacez of India and Ceylon, by Willis,

appears in Part III of the Annals of the Royal Botanic Gardens,

Peradeniya, issued in May.

The laws of plant distribution in alpine regions are analyzed by

Jaccard in Hora of April 3o.

An interesting addition to the botany of the Atlantic islands is a

comparative account of the mosses of the Azores, Madeira, and the

826 THE AMERICAN NATURALIST. [Vor. XXXVI.

Canaries, published, in connection with an article on some mosses

of the latter islands, by Cardot, in No. 5 of the Bulletin de 7 Herbier

Boissier for the present year.

No. 10 of Marcus E. Jones’s Contributions to Western Botany, issued

as an individual publication by the author from the Mammoth Record

Print of Robinson, Utah, under date of June 1, 1902, contains a

revision of the Alliums of the Great Basin region, with notes on the

markings of the bulbs and a key to the species based on them;

further notes on Astragalus; a study of the Nyctaginaceze, chiefly of

the region of the Great Plateau ; and descriptive or synonymic notes

on a number of miscellaneous species.

The flora of Scott and Muscatine counties, Iowa, is the subject of

a paper by Barnes, Reppert, and Miller, in the recently issued eighth

volume of Proceedings of the Davenport Academy of Sciences.

A botanical series of the University of California Publications

begins with the issuance, under date of June 7, of a paper by H. M

Hall, entitled * A Botanical Survey of San Jacinto Mountain." It

appears to be a well done piece of work, on the prevalent ecological

lines, and is well illustrated.

A catalogue of British marine algz, by Batters, is in course of publi-

cation as a supplement to current numbers of the Journal of Botany.

A large part of Vol. III, fasc. 3-6, of Ze Botaniste, dated June 10,

is occupied by a paper on Euglenia, by Dangeard.

Cladophora, as represented in the salt water of New England, —

a difficult group, — is revised by Collins, in RAodora for June.

Studies of spore germination in the common mushroom and other

agarics, by Miss Ferguson, form Buletin No. 16 of the Bureau of

Plant Industry of the national Department of Agriculture.

The Geastrez of the United States are revised in an illustrated

paper by C. G. Lloyd, forming Bulletin No. 5 (mycological series,

No. 2) of the Lloyd Library.

In the Journal of Botany for June, Arthur Lister describes and

figures a Chondrioderma, — C. asteroides, — which in aspect resem-

bles to a marked degree a small Geaster.

The Technology Quarterly for June contains an article by Whipple

on the physical properties of gelatin, with reference to its use in

culture-media.

A paper on albino phenomena in the vegetable kingdom, by

Pantanelli, is published in Nos. 10—12 of Vol. XV of Malpighia.

No. 430.] NOTES AND LITERATURE. 827

An account of double fertilization in Monotropa uniflora is pub-

lished by Shibata in “Vora of December 4 last.

In the Journal de Botanigue for May, Guignard has an article on

double fertilization in Solanacez.

The mechanism of secretion of extra-floral nectaries is the subject

of an article by Haupt in Fora, Vol. XC, Heft 1, which also con-

tains a paper by Lepeschkin on the significance of the water-secret-

ing organs of plants.

Meierhofer has a well illustrated article on the bladders of Utricu-

laria in Flora of Dec. 4, 1901, which also contains a paper b

Brenner on climate and leaf form in Quercus, and an extensive

study of Gunnera by Schnegg.

An account of the developmental history of Quercus is given by

Brenner in Vol. XC, Heft 3, of Fora.

An important addition to the literature of plant physiology, in its

economic bearings, is Dr. Whitten's thesis on * Das Verhältnis der

Farbe zur Tótung von Pfirsichknospen durch Winterfrost," recently

published in Halle.

A handbook of the economic plant products of Ceylon, by Willis

and Wright, is in course of publication as supplements to current

numbers of the Avna/s of the Peradeniya garden.

An interesting account of the fiber Agaves, by Marshall, is reprinted

from the Journal of Geography in the American Journal of Pharmacy

for July.

An illustrated account of the new botanical laboratories at Liver-

pool is contained in Mature for June 12.

Several views of the Cape Town Municipal Gardens are published

in the Gardeners’ Chronicle of June 7.

A portrait of J. B. Ellis forms the frontispiece of No. 62 of the

Journal of Mycology.

A memorial of the Italian botanist Gibelli is separately issued

from Vol. XIV of Malpighia.

A biographic sketch of Ramírez, with portrait, is published in the

Memorias y revista de la Sociedad Cientifica “ Antonio Alzate” of

October last.

In Biometrika for April, Mr. Yule has a paper on the variation of

the number of sepals in Anemone nemorosa, and Mr. Tower a paper

on the variation in the ray flowers of Chrysanthemum leucanthemum,

on which subject is added a further note by Pearson and Yule.

828 THE AMERICAN NATURALIST. [Vou. XXXVI.

An editorial in the same number of Biometrika contains some

desirable cautions as to the polymorphism of certain species which

appears to result from the frequency polygons derived from tabulated

individual measurements.

The first fascicle of Vol. III of the Contribuzioni alla biologia

vegetale of the R. Lstituto botanico di Palermo contains the following

papers of general interest: Terracciano, “ Contributo alla biologia

della propagazione agamica nelle fanerogame "; Albo, “ Sul signifi-

cato fisiologico della nicotina nelle piante di tabacco”; La Floresta,

* Formazione di radici avventizie nelle foglie di Gasteria acinact-

folia"; Borzí, * Anatomia dell apparato senso-motore dei cirri delle

Cucurbitacee " ; and La Floresta, ** Struttura ed accrescimento secon-

dario del fusto di Xanthorrhoea." The articles are well illustrated

and a French résumé is given of their contents.

Rhodora for June contains the following articles : Collins, ** Marine

Cladophoras of New England"; Nye, H. A., * Blooming of Hepat-

; Brainerd, “Two more Rare Plants from Lake St. John,

Quebec" ; Blanchard, “ Some Vermont and New Hampshire Plants

in the Middle Connecticut Valley," I; Fernald, “The Seneca

Snakeroot in Maine" ; Webster, * C/athrus columnatus in Lawrence,

Mass."; Robinson, *Two New Hypericums of the Adpressum

Group”; and Fernald, ** An Anomalous Skullcap.”

The third Report of the Michigan Academy of Science contains

the following botanical articles : Pollock, * Relation of the Fibro-Vas-

cular Bundles in the Root and Hypocotyl in Æchinocystis lobata”’;

Reed, “Ecology of a Glacial Lake”; Larsen, “Disease of the

White Birch”; Denton, “Interfoliar Scales of Monocotyledonous

Aquatics"; Clark, “ Notes on the Flora of Eaton County” ; Davis,

“Notes on Utricularia cornuta” ; Davis, * Wolffia Notes” ; Long-

year, “Notes on Michigan Saprophytic Fungi"; Longyear, “ New

Species of Michigan Fungi"; Longyear, *Sclerotium Disease of

the Huckleberry”; and Conover, “ Asparagus plumosus.”

The Botanical Gazette for May contains the following articles :

Miyaké, “On the Starch of Evergreen Leaves and its Relation to

Photosynthesis during the Winter”; Newcombe, * Rheotropism of

Roots” (conclusion) ; Overton, * Parthenogenesis in Thalictrum pur-

purascens” Leavitt, “Subterranean Plants of Epiphegus” ; Fair-

child, * Notes of Travel,” VIII; Cockerell, “A New Heliotropium

(CR. xerophilum)." ‘5

No. 430.] NOTES AND LITERATURE. 829

The Botanical Gazette for June contains the following articles :

Rimbach, “ Physiological Observations on the Subterranean Organs

of some Californian Liliacez "; Smith, * The Parasitism of Botrytis

cinerea”; Shaw, “The Development of Vegetation in the Morainal

Depressions of the Vicinity of Woods Hole"; Webb, * A Morpho-

logical Study of the Flower and Embryo of Spiræa ” ; Fairchild, D. G.,

* A Precocious Poplar Branch”; and Wilcox, * Numerical Variation

of the Ray Flowers of Compositz.”

Botanical papers in No. 7 of the Bulletin of the New York Botani-

cal Garden are: Earle, * Mycological Studies," I; Williams, “A

Preliminary List of Montana Mosses" ; and Hollick, “ year oe and

Botanical Notes, Cape Cod and Chaa t Island, Mass.

Torreya for June contains the following botanical articles : Rydberg,

“ Our Yellow Lady's-Slippers ” ; Underwood, ** The Bracket Fungi” ;

Lloyd, “ An Interesting Irregularity in a Rose Flower”; and Brit-

ton, E. G., “ Notes on a Long Island Moss.”

The Bulletin of the Torrey Botanical Club for May is entirely occu-

pied with Cryptogams, mostly fungi.

Botanical articles in the Ohio Naturalist for June are: Kellerman

and Tyler, “ Further Additions to the Catalogue of Ohio Plants” ;

Dresbach, “ Moulds Injurious to Foods”; and Tyler, “ Rosette

Plants of Ohio.”

A large part of Engler’s Botanische Jahrbücher continues to be

occupied with descriptions of African plants.

As usual with the Journal of the Royal Horticultural Society, the

number for April contains several articles of general botanical interest.

After an intermission of eight years, the Journal of Mycology begins

its eighth volume with No. 61, bearing date May, 1902.

The great multiplication of species in Crataegus is commented on

by Professor Bessey in Science for May 16.

A paper by Knuth on the geographic distribution, adaptive struc-

ture, and classification of the species of Geranium, is published in

Engler’s Botanische Jahrbücher of May 2.

A monograph of the Senecios of North and Central America, by

Dr. Greenman, is published as a contribution from the laboratory of

the Berlin Garden, in the first Heft for the current year of Engler’s

Botanische Jahrbücher. Keys are given to the sections, under each

of which are enumerated the species recognized as belonging to it.

830 THE AMERICAN NATURALIST. [VoL. XXXVI.

Dr. Holm has an article on the grass genus Arctophila Rupr. in

the Ottawa Naturalist for June. Three new species are described.

An analytical key to the ferns of the northeastern states, based on

stipe characters, by C. E. Waters, is printed in the Johns Hopkins

University Circular for June.

A lecture on the past, present, and future of vegetable pathology,

by Dr. G. Mottareale, is reprinted from Z/a//a Orticola for March

to May.

In Education for February, Dr. Harshberger discusses the question

as to what part of the varied things comprised in * botany " is of

most worth.

In Popular Science News for June, G. W. Browning has an illus-

trated article on insect-catching plants, other than those that are

carnivorous.

Spindle formation in Agave is the title of a cytological paper by

Osterhout, published as No. 8 of the current volume of Proceedings of

the California Academy of Sciences.

An interesting note on the formation of hair masses in the stomachs

of goats that have fed on the fruits of the sweetbrier rose is published

by Sir William Thiselton-Dyer in ature for May 8.

A study of galls and the insects producing them, by M. T. Cook,

appears in No. 7 of the Okio Naturalist, and is separately printed as

a bulletin of the University of Ohio.

Professor Brewer has distributed separates of an article on the

relation of forestry to public health, from the Proceedings of the

Twenty-Ninth Meeting of the American Public Health Association.

The decorative possibilities of some native climbers are shown by

Professor Nelson in Bulletin No. 50 of the Wyoming Experiment

Station.

The Revue horticole for June 1 contains an illustrated article on

the botanic garden of St. Pierre before its destruction by the eruption

of Mont Pelée.

An account of the botanico-geographic regions of North America,

as exemplified in the new botanical garden at Dahlem-Steglitz, is

given by Professor Engler in Appendix IX of the otisblatt des

K. Botanischen Gartens und Museums zu Berlin, dated May 15, 1902-

Several new or little known Southern California plants form the

subject of a note by LeRoy Abrams in No. 6 of the current volume

of the Bulletin of the Southern California Academy of Sciences.

No. 430.] NOTES AND LITERATURE. 831

Parts III and IV of Spegazzini's *Nova addenda ad floram Pata-

gonicam," separately issued from the Anales del Museo Nacional de

Buenos Aires, Vol. VII, bears date of April 16, and includes species

numbered from 474 to 977 inclusive, — a considerable number of

them being new.

Visitors to the beautiful little island of Capri, in the Gulf of Naples,

will find help in a study of its flora in a tasteful little handbook by

Cerio and Bellini, entitled Fora dell Isola di Capri, published by

Emilio Prass of Naples.

A portrait and biographical sketch of Dr. Adam Kuhn, first pro-

fessor of botany in America and at the University of Pennsylvania,

with an illustration of the genus Kuhnia, named in his honor, are

published by Dr. Harshberger in the Alumni Register, of Philadel-

phia, for April.

A portrait of Sir George King, formerly director of the Calcutta

Garden, is published in the Gardeners’ Chronicle of May 11.

A note on Parkinson, with figure of the Parkinson statue in the

palm house at Sefton Park, Liverpool, is printed in the Gardeners’

Chronicle for May 17.

CORRESPONDENCE.

Editor of the American Naturalist :

Mr. Robertson’s criticisms in the American Naturalist for July

appear to be exceedingly captious and trivial. In my paper on the

colors of flowers I say of the visitors of Vymph@a advena, “ I have col-

lected on the flowers in Maine four Diptera, two Coleoptera, and one

small bee, Za/ictus nelumbonis, which confines its visits to this flower.”

So far as possible my remarks on the visits of insects to flowers are

based on numerous observations I have made in the field. The

above remark is true of this bee, so far as I have collected it, in this

locality. —

Again, as to the number of visitors to the Umbelliferz. It is

certainly true that several species are visited by more than two hun-

dred insects, while others approach quite closely to this number.

I still believe that when a more careful and extended study of these

plants shall have been made, the number of visitors to many species

will be found to exceed two hundred. The visitors to many umbel-

lifers have, moreover, never been collected.

As regards the visits of beetles to dull yellow flowers, it certainly

never occurred to me that there was anything in my paper which

could lead any reader to believe that I considered the statement

as new. I mentioned the later observations of Müller, and also

those of Loew, Schultz, and Knuth, as well as one of my own, and

added that it did not appear necessary to carry the illustrations

further. In the brief reviews of the coloring of the different plant

families which space renders possible, I am compelled to omit

many observations both of my own and of others. Mr. Robertson

tells us that he has also made some observations on this subject,

and the real animus of his remarks seems to be that he has been

hurt because no reference was made to these. Mr. Robertson was

not long ago severely censured by a correspondent of the Canadian

Entomologist for his loose way of naming supposed new species of

bees based on single specimens or on trivial characters. Of his

regard for the rights of others a single illustration will serve.

Transactions of the Academy of Science, St. Louis, Vol. X, No. 2, P 45

after describing Andrena viciniformis, which he calls a new species,

832

CORRESPONDENCE. 833

he adds, “ This may be the same as A. dunningii.” A comparison

of the two descriptions will leave little doubt that the species are

identical. In renaming 4. dunningii an injustice is done both to

the author of this species and to the person in whose honor it was

named. Again, in the Canadian Entomologist, Vol. XXXIII, No. 8,

p. 231, he writes: “Zpeolus lectoides n. sp. 9. — Closely resembles

E. lectus Cr., and may be the same." Described from one specimen.

Additional instances might easily be given. An author who will

publish such descriptions as these certainly shows little desire for

the advancement of the interests of science.

Joun H. LovELL.

WALDOBORO, MAINE, August II, 1902.

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BEDDARD, F. E. Mammalia. The — Natural History. Vol x. 8vo,

xii 4-605 pp., 285 figs. $4.00. — BLAISDELL, A. F. Life and Health. A Text-

Book on Physiology for Ee Schools, teas, and Normal Schools. Boston,

Gin Co., 1902. 8vo, vi + 346 pp. I10 figs.— Davis, W. M. Elementary

Physical eighislit Boston, Ginn & Co., 1902. 8vo, xviii + 401 pp., 190 figs.,

9 pls. — EMERTON, J. H. The Common poma of the United States. Boston,

Ginn & Co., 1902. 8vo, xviii + 225 pp., 501 $1.50. — GARDINER, J. S. The

Fauna and Geography of the Maldive and Kredit. ue spe REM etc. Vol. i,

pt. 119-222, Pls. VI-XIII.— GILBERT, J. K., and BRIGHAM, A. P.

An Pusodacilón to Physical Geography. New York, Appleton, 1902. te xvi +

380 pp., 263 figs. — JoRDAN, D. S., and EvERMANN, B. W. American Food and

Game Fishes. A Popular Account of all the Species found in America North of

the Equator, with Keys for Ready Identification, Life Histories, and Methods of

Capture. New York, Doubleday, Page & Co., 1902. 8vo,1+ 573 PP» colored

plates, text figures, and photographs. $4.00.— JoRDAN, D. S., and HEATH, H

Animal Farms. A Second Book of Zoölogy. New York, Appleton, 1902. 8vo,

vii + 258 pp. 140 figs. — RABL, CARL. Die Entwicklung des Gesichtes. Tafeln

zur Entwicklungsgeschichte der äusseren Kórperform der Wirbelthiere. I. Heft,

Das Gesicht der Sáugethiere, I, (Kaninchen, Schwein, Mensch). Leipzig, hn

mann, 1902. vi-+ 21 pp. folio, 8 pls. 12 marks. — ROOSEVELT, THEO., VAN DY

T. S., ELLIOT, D. S., and STONE, A. G. The Deer Family. New York, ai

lan, 1902. 8vo, ix + 334 pp., 21 a is maps. $2.00.— SAGE, D., TOWNSEND,

C. H., SurrH, H. M., and Har . C. Salmon and Trout. New York,

Macmillan, 1902. 8vo, x + 417 w 20 pls, 13 figs. —SANDys, E., and VAN

` pp» 9 pls.— SMITH, EUGENE. The Ho ome Aquarium and How to Care for It.

A Guide to its Fishes, Other Animals, and Plants, etc. New York, Dutton, 1902.

8vo, ix + 3 PP., 137 figs. $1.20.— WHEELOCK, IRENE GROSVENOR. Nestlings

orest and Marsh. Chicago, McClurg, 1902. 8vo, xvi + 257 PP» 2? pls.,

text dn

ASHMEAD, W. H. Papers from the Harriman Alaska Expedition, XXVIII.

The Hymenoptera. Proc. Wash. Acad. Sci. Vol. iv, pp. 117-274, Pls. IX-XI. —

— V. Seven New Mammals ses Saee Texas. Proc. Biol. Soc., Wark-

xv. Dp cote El

BATESON, W., and SAUNDERS, E. R. Experimental Studies in the Physiology of

Heredity. Royal Society Reports to the Evolution Committee, Rept. I. — BUSK,

. A Review of the American Moths of the Genus Depressaria Haworth, with

_ Descriptions of New Species. Proc. U. S. Nat. Mus. Vol. xxiv, pp- 731-749: —

: T R. V. Henicops dolichopus, A New Chilopod from Utah. Proc.

834

PUBLICATIONS RECEIVED. 835

U. S. Nat. Mus. Vol. xxiv, pp. 797-800. — CockERELL, T. D. A. A New Ribes

from Mexico. Proc. Biol. Soc, Washi ington. pp. 99—100. — DUERDEN, J. E.

Report on the Actinians of Porto Rico. Bull. U. S. Fish Com. Sor 1900. Vol. ii,

I i

Cal. Pubs., Bull. Dept. Geol. Vol. iii, No. 2, pp. 31-50, Pls. I-IV. — Field

Columbian Museum. Annual Report of the Director for the year 1900-1901. —

GREEN, E. H., and Tower, R. W. The Organic Constituents of the Scales of

Fish. Bull. U. S. Fish Com. for rgor. Pp. 97-102. — Hay, W. P. A List of

ps Batrachians and Reptiles of the District of Columbia and Vicinity. Proc.

Soc., Washington. Vol. xv, pp. 121-145.— IMBERT, A. Mode de fonc-

vas iE. économique de Fetes nisme. Paris, C. Naud, 1902.

Scientia, Série Roo No. 14. — INGEN, J. van. The Siluric bir near

Batesville, Ark. Pt. i, Geological Relations. Columbia Univ. S. ol of Mines

wart. Vol. xxii, pp. 318-329. — JOHNSTON, J. B. An idis ga to ; Delis the

Primitive Functional Divisions of the Central Nervous System. /ourn. Comp.

Neurol. Vol. p pp- iur 2 figs. shop STON, J. B. The Brain of Petro-

myzon. Journ. Comp. Neu Vol. x i, pp. 1-86, Pls. I-VIII. — JULIEN, A. A.

A Study of the iet of po tes. d n. Geol. Vol.ix,pp.673-693. 3 figs.

— Kemp, J. F. The Rôle of the Igneus Rocks in the Formation of Veins.

Trans. Amer. Inst. Mining Engineers. Richmond Meeting, 1901. 30 pp. — KEMP,

e Deposits of Copper Ores at Ducktown, Tenn. Trans. Amer. Inst.

Mining Engineers. Richmond Meeting, 1901. 22 pp., 11 figs. — Kemp, J. F., and

Hitt, B. F. Preliminary Report on the Pre-Cambrian Formations in Parts of

Lawson, A. C. and ParacHE, C. The Berkeley Hills. A Detail of Coast

Range Geology. Univ. Cal. Pubs., Bull. Dept. Geol. Vol. ii, No. 12, pp. 349-450,

Pls. X-XVII, map. — Lyon, M. W. Description of a New Bat from Colombia.

Proc. Biol. Soc., Washington. Vol. xv, pp. 151-1 ge — MasoN, O. T. hag

for Collectors of American Baskets. Proc. U? S. Nat. Mus. Vol. xxiv, pp. 31.

44 figs. — Mearns, E. A. Descriptions of Three. si Birds from he Soo hera

United States. Proc. U. S. Nat. Mus. Vol. xxiv, pp. 915-926. — MEARNS, E. A.

Two New Species of Poisonous Sumachs from the States of Rhode Island and

Fede Prec. DiN Soc. Warbing ton. Vol. eda 147-149. — MENDELSSOHN, M.

ét Paris, C. Naud, 1902. 8vo, 99 pp.

Scientia, Biol., No. 13. — MERRILL, Gzo. ?- A Newly Found Meteorite from

Admire, Lyon County, Kan. sage y. s Nat- Mns: bis xxiv, pp. 207-313, Pls.

L-LVI. — MILLER, G. S., JR. d Nicobar Islands.

Proc. U. S. Nat. Mus. Vol. xxiv, pp. 751-795, Pls. XLI-XLII. — MILLER, G. S., JR.

Directions for Preparing Study Specimens of Small M Proc. U. S. Nat.

Mus. Vol. xxiv, pp. 25. 3 figs. — NELSON, A. shady Vines in Wyoming Homes.

Bull. Wyoming Agr. Exp. Sta., No. 50. 15 pp., 11 pls. — NEWLAND, D. H. The

Serpentines of Manhattan Island and Vicinity and as Accompanying Minerals.

Columbia School of Mines Quarterly. Vol. xxii, pp. 307-410. 1901.— North

Carolina State Horticultural Society. The Cow Pea. Southern Pines, N.S. 63

PP. 12 figs. — OBERHOLZER, H.C. A Review of the Larks of the Genus Oto-

coris. Proc. U. S. Nat. Mus. Vol. xxiv, pp. 801-884, Pls. XLIII-XLIX. —

836 THE AMERICAN NATURALIST.

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en Serge. Compt. Rend. VITI Cong. Internat. Geol. 7 pp.— OSBORN, H. F.

Des méthodes précises mises actuellement en œuvre dans l'étude des vertébrés

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ous Stimuli, and the dig of ind on Daily Depth ene ae Bull. U.S. Fish

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LXI rmudi

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The American Naturalist.

ASSOCIATE EDITORS:

~ J-A. ALLEN, Pu.D., American Museum of Natural History, New York.

E. A. ANDREWS, Pu. D., Johns Hopkins University, Baltimore.

M. DAVIS, M.E., Harvard U. bp. "ambu

om ork City.

N fi er sily.

ARLES A. KOFOID, PH.D., University of Cali PARE Berkeley.

3. G: NEEDHAM, PH. Se Lake Forest Oni Er Si

ARNOLD

, ent of. Aeneas Washington.

LEONHARD Seer SGER, LL.D., Smith institution, Washington.

Ww. TRELEAS SE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, PH.D., University of Nebraska, Lincoln.

= M. WHEELE R, BEDS University af Texas, Austin.

HE AMERICAN N. HTURALIST i is an —— Louer. mue

atural Histor and A im ' sent to its readers the

THE

AMERICAN NATURALIST

Vor. XXXVI. November, 1902. No. 431.

BIOMETRIC EVIDENCE IN THE PROBLEM

OF THE PAIRED LIMBS OF THE

VERTEBRATES.

BASHFORD DEAN.

From early development onward a fish is accurately poised

in its living medium. Its long axis ina position of rest remains

normally horizontal, in spite of the most varied changes in

the size and shape of the fish’s body and the shiftings and

differentiation of its component parts. To preserve an accu-

rate balance under conditions of rest as well as under the strain

of the most active movement implies obviously a delicate

adjustment of the morphological elements of the animal to the

physical ones. And as the latter are relatively constant it fol-

lows that the acute strain in the evolution of the fish’s body,

both in ontogeny and phylogeny, has fallen upon the mor-

phological elements. These, then, become subject to form

changes, to position changes, and, most important, to physio-

logical changes, to enable them to fulfill the mechanical require-

ments of habitat. To what degree the results are successful

in adapting the vertebrate body to aquatic living can best be

understood by comparison of the “area curves” and “entering

; 837 T

838 THE AMERICAN NATURALIST. [Vor. XXXVI.

angles,” not only among fishes of different groups but between

fishes and cetaceans; for in these practically the same “ lines "

and “entering angles ” ! obtain.

From a physical standpoint we conclude, therefore, that the

position of the gravity center of the animal during different

stages of growth plays an important róle in connection with its

ever-increasing body length, and that special organs of balancing,

t.e., fins, will in their development be subject par excellence to

the mechanical needs in shifting which the changing position

of the gravity center and increasing body length demand.

From this standpoint, then, one can attack the long-troubled

problem of the origin of the vertebrate limbs. Not, however,

with the aim of solving it indisputably, — for this I firmly

believe can be done only by the paleontologist, — but rather

for the purpose of finding a probable clue to the labyrinthine

confusion in latest morphological studies ; for the voluminous

conclusions of Braus, following Gegenbaurian teachings, and

supported by Semon in his recent work on Ceratodus, — cf.

also the related work of Klaatsch, — are vigorously denied by

the newer phases of the fin-fold theory of Dohrn, Mollier,

Rabl, and others, which in turn are in the latest time criticised

by Fürbringer. In short, there is at present a deadlock in the

discussion as to the mode of origin of the limbs of the verte-

brates which cannot yet be loosened by orthodox embryological

methods. Perhaps it will be bettered when a greater number

of forms will have been examined, or when we shall have a

clearer knowledge of the processes of accelerated and retarded

development. Perhaps, on the other hand, to take a more

somber view, it will never be adequately loosened by study of

embryonic structures, since these may have been so modified

during phylogeny as to mask hopelessly their true homologies.

For the discussion has to a large measure narrowed itself down

to the interpretation of embryological details, which if given

fixed morphological weight become of great importance in

theoretical discussion, but which are relatively valueless if

! The entering angle terminates with * wonderful uniformity" at 36 per cent

of the total length of the animal, whether teleost or cetacean. Cf. especially

H. DeB. Parsons, Trans. Amer. Soc. Engineers, vol. ix, penn 17 pp» 7 pls.

No. 431.] PAIRED LIMBS OF THE VERTEBRATES. $ 39

viewed as nothing more than adaptations to recent physiological

needs. Thus, as an example, a tangle of nerves and blood ves-

sels on the tailward side of a fin lobe of a shark may be either

an important morphological condition as vestige of an ancestral,

paddle-shaped, ceratodont fin, ¢.g., as Braus maintains, or it

may be nothing more than a newly developed condition to

enable muscles and nerves to get in better touch with an

enlarging and specializing margin of the fin, —which may

indeed in its growth be developing from stage to stage special

embryonic and larval movements. Again, the emancipation of

nerves, vessels, and muscles from the basis of an embryonic

fin may be the result either of the migration of a gill bar ele-

ment in the Gegenbaurian sense, or merely of a purely local

change in the requirements for balancing the constantly grow-

ing animal. For the center of gravity may shift as the body

grows, and the fin (in sharks functional even when the “larva "

is growing within the egg case) may thus have to move forward

and backward as mechanical needs demand.

“But,” the Gegenbaurian would maintain, “ our theory is

supported by fin migration. Of course we are willing to grant

that the record is obscured by cenogensis, but we have still

morphological evidence that the ventral fin migrates backward,

and that the pectoral is relatively stationary, its girdle resem-

bling the adjacent branchial arches. And we believe, more-

over, that the mechanical fin needs, which have been noted

above, are of the utmost importance, since it is in response to

them that the ventral fin has traveled backward and that the

pectoral fin has retained, relatively, its primitive position.

Indeed, the very perfect segmental character of the ventral fin

can be best explained in terms of a continued hindward migra-

tion, for the ventral is not the more steadfast and more prim-

itive fin."

Now, it has long seemed to me that such a line of argument.

could be invalidated if it could be shown that during the growth

of the individual the fins responded to their mechanical require-

ments in just the opposite way ; that is, that the ventral fin

Was zot, and that the pectoral fin was, the migrating element.

And the present paper has grown out of an attempt to trace,

840 THE AMERICAN NATURALIST. [Vor. XXXVI.

and to this end by biometric methods, the actual stages in the

development of the paired fins in accordance with their physi-

cal requirements. Assuming, for example, a complete series

of a form in which the limb characters can be favorably exam-

ined, say a shark,’ in stages ranging from adult to embryos in

which the paired limbs are just appearing as distinct structures,

we can then determine by measurement what change occurs

from stage to stage in the position of the gravity center (with

reference to the total length of the animal), and by similar

measurement of the fins in similar terms we can ascertain what

ratio these measurements bear to the possible shifting of this

center. For all will admit that the fins themselves, as light

and delicate structures, can in their growth play little direct

part as a cause of the changing position of the center of gravity,

and that their migration, forward or backward, is rather

an outcome of the shifting of bulkier organs, — muscles,

brain, viscera, sense organs. We can therefore conclude that

by such a system of measurements one can obtain evidence as

to whether the ventral fins migrate backward, z.e., in terms

of the total length of the individual, and show the correla-

tion in growth with the pectorals which the Gegenbaurian

theory demands; or whether, on the other hand, it is the

ventrals which are constant, and resemble the dorsal fins,

while pectorals play the part of the mobile and specializing

members.

The following notes are based upon a cestraciont shark,? a

form particularly interesting from the standpoint of morpho-

logical conservatism, since there is reason to believe that it

has been little modified since Carboniferous times. And the

measurements are recorded of sixty individuals ranging in

length from 330 mm. to 14.5 mm. For this careful and labo-

rious computation I am much indebted to my former student,

Prof. C. H. Brookover, who made it the beginning of a research

1 Better, of course, would be sets of many genera and species; but as long as

these are lacking, we may base conclusions as to essential characters upon the

NEUF of a single form.

? Heterodontus japonicus Macleay. The embryos were collected during the

writer’s visit to Japan, 1901, thanks to the many — which he enjoyed at

Misaki as guest of the Im mperial University of Tokyo

No. 431.) PAIRED LIMBS OF THE VERTEBRATES. 841

upon the general problem of limb migration. He was, how-

ever, unable to carry out his plans, and he generously turned

over to me his measurements for use in the present paper.

DETAILS OF DATA.

Straight specimens were selected for measurement. Gravity

center was determined in terms of the total length ! by balancing

the specimen either on a knife-edge or within the delicate tips

of curved forceps. By this simple method the limit of error was

found to be surprisingly small, estimated by repeated tests at

about r per cent. In the measurements individual variations

are taken into account, since it was found that a more advanced

embryo is sometimes smaller in size and variable slightly in

proportions, and in preparing the table of averages as plotted

in Fig. 1 this source of error has been largely counterbalanced

by the use of guide points representing the average of several

(usually four) successive individuals. All measurements are

computed in percentage of the total length of the specimen, and

are estimated from the snout tip. The landmarks whose rela-

tive position is computed are indicated graphically in the present

Fig. 1,2.¢., anterior margins of pectoral, of ventral, and of dorsal

fins, and posterior margins of pectorals and ventrals. Of the

latter margin the functional (dermal) one is alone considered.

THE RESULT OF THIS ANALYSIS.

The center of gravity is found to shift forward (about

5 per cent of the total length) in the earlier embryos (embryos

from 27 mm. to about 6mm.), in later stages (up to embryos

of 200 mm.) backward to the extent of about 8 per cent of the

total length. In well-grown specimens the position of the

gravity center remains practically unchanged.

The pectoral fin undergoes conspicuous changes: from the

time it can be distinctly differentiated from the ventrals it

1 The transverse plane in which the center occurs was alone determined; its

more accurate position within this plane has evidently no bearing upon the present

problem.

842 THE AMERICAN NATURALIST. [Vor. XXXVI.

TABULATED MEASUREMENTS OF FINS AND GRAVITY CENTER

OF CESTRACION EMBRYOS.

(By C. H. BROOKOVER.)

From Snout Tip To

OTAL ANT. Post ANT. OST.

GRaviry.| PerEct. PECT. VENT. Vent. | Domsar. | DORSAL.

Fin. Fin. Fin. Fin.

mm %o % % % % % %o

M 14.5 m 314 38— cm a au ste

2 15 — 32 40 — — — —

3 M I 27 a m" i s

no spine no spine

4 20.5 ses 26 38+ 42+ 49+ =

no spine no spine

5 21.5 inq: 25— 324 41— 484- 37+ 6

no spine

6 24— mae 24+ a= wt 44— 5 58+

7 24 = 28— 37— 45— 52+ 62+

8 26.5 — 26+ 324 40 4- 48— eges

9 26.5 31 -— 33— 414 48+ 3 57

s 27 28 24+ 30 37 44+ S 53

i 26 — 25— 31— 39— 45+ 3 53

A T 5 Sra 24+ = + 38+ 46— 32 57

2 o o

: : a vent tent

14 31 — 23 31 9 45 32 55

5 3r mee 25 31 7 43 32

16 32.5 — 24 30 8 4I 29 50

17 34 = 22 28 4 po aes 31 50

18 3455 ene 22 29 4 40 28

19 35 ae 24 31 8 45 30 2r

20 35 — 20 26 o 38 29 48

21 35 — 22 28 4 40 29 50

22 2 v" 23 29 I 45 29 50

.23 3 T: e 22 28 9 46 30 50

24 39 seme 22 29 9 44 29 51

25 39 = 21 27 5 43 27 50

26 39 ke 21 27 4 40 27 48

27 40 — 20 26 2 28 48

28 4° — 21 29 6 42 28 50

29 40 26 21 27 $ 41 27 48

o 40.5 = 20 26 5 40 28 48

I 4r — 22 27 5 43 26 47

4 4r mM 21 29 4 41 29 50

J 41.5 — 20 26 o 39 26 47

+ 42.5 vex 22 28 6 42 28 5o

] 43 Mem 22 29 29 49

J 43 26 21 26 : 40 25 45

44-5 26 20 26 4 39 29 50

3 44-5 — 21 27 7 43 27 49

) 45 — 22 27 6 43 27

> 46 — 20 26 4 38 27 48

4 46.5 a 22 27 35 aE +

42 50 25 20 26 7 41 27 46

43 ae 24 38 24

4 50 j 40 as 47

51 — 40 25 50

$. : E ar 26

V STRE 39 23

E : 40 25 47

TE 1 4t 25 48

1A CA 4t 27 47

R E 40 24 46

me 41 24 5I

ru 42 27 5

E ) 43 26 52

82 , 43 27 52

105 7 46 27 53

187 7 50 27 57

s 53. 30 57

41 53 31 59

41 S Ri "o

No. 431.] PAIRED LIMBS OF THE VERTEBRATES. 843

increases greatly in size, measured antero-posteriorly, from

6 per cent of length to about 24 per cent, its anterior and

posterior margins undergoing somewhat different fates. The

anterior rim of the fin passes forward, suddenly in earlier stages,

slowly in later ones, altogether a distance of over 12 per cent

of the total length, a distance considerable enough, I take it,

to entail important adaptive changes in the structures of the

encroached-upon region, especially in view of the fact that this

fin margin grows forward as the gravity center passes back-

ward. (Cf. also infra, re the continuous hindward growth of

the outer gill slits.) Interesting in this connection is the

slight variant in embryos of about 80 mm. ; here, the gravity

center shifting a little forward, the anterior fin margin passes

backward, thus indicating a nice adjustment in the mechanical

relations between them. Ina general way the advance of the

anterior rim of the pectoral is at the same pace as the retreat

of the center of gravity. The behavior of the hinder rim of

the pectoral shows a less close adjustment to the gravity center.

In early stages this fin rim passes forward (about 11 per cent

between stages of 15 mm. to 56 mm.) to about the transverse

plane of the center of gravity ; thereafter, however, it grows

hindward, gradually increasing the distance from the gravity

center (11 per cent) (and from the anterior fin rim (total, 24 per

cent)). By this noteworthy hindward growth it comes in the

older stages to overlap somewhat the anterior rim of the

ventral fin.

The ventral fin is conservative in its position during growth:

measuring about 6 per cent of the total length in earlier stages,

it increases to but 12 per cent in latest ones, a gain in per-

centage of but 6 per cent as against 18 per cent in the pec-

toral. Like the pectoral, the entire fin passes slightly forward

in younger stages, then slightly backward. The anterior rim

remains almost fixed with reference to the body length, the

posterior passing slightly backward to a degree corresponding

almost exactly with the shifting of the gravity center. Thus,

in an embryo of about 80 mm. the center of gravity is 18 per

cent in advance of this rim, in one of 115 mm. 19 per cent,

in one of 145 mm. 19 per cent, of 187 mm. Ig per cent, a

844 THE AMERICAN NATURALIST. | [Vor. XXXVI.

correspondence so close that we can interpret it only in terms

of cause and effect.

The dorsal fins are even more conservative in position than

the ventrals. In the first place they maintain practically the

same relative distance from one another, measured from the

anterior rim of the fins! : thus, ina specimen of 21 mm. the inter-

val measures about 23 per cent, of 40 mm. about 22 per cent, of

60 mm. 23 per cent, of 82 mm. 25 per cent, of 105 mm. 26 per

cent, of 197 mm. 27 per cent; in other words, between the two

fins there is but a slight variation in the interval (say, 5 per cent

of the entire length) in stages widely different in size (e.g.,

measuring between 20 and 200 mm.). In general, however, we

note that the growth of the anterior rims of first dorsal and

ventral are indicated by parallel lines, and similarly the growth

of the anterior rim of the second dorsal and the posterior rim

of the ventral.

SUMMARY.

The foregoing characters yield what seems to me convin-

cing evidence that the pectoral fin is subject to changes with

respect to the gravity center, z.e., physical changes, which cause

it to become more highly specialized than the ventral fin,

and that the latter is conservative, after the fashion of the

unpaired fins. The result of biometric data, it will be seen,

confirms strikingly the views of the fin-fold theorists, most

clearly expressed in this particular regard by Wiedersheim in

his *Gliedmassenskelet." On the other hand, the present study

yields no evidence that there has ever been a migration of the fins

in the Gegenbaurian sense; thus, we find no reason to believe

that the ventral fin is a structure which has shifted its position

from in front hindward ; it is indeed in about the same position

fore and aft in the adult as in early embryos (20 mm.). It is

important, furthermore, that the only fin (leaving anal and caudal

out of discussion) whose anterior margin passes forward is the

pectoral, since with this condition is correlated the fact that this

*In the younger stages the measurement was made from the actual most

anterior point of the fin; in the later, when a spine appears, from the point where

the anterior fin rim, if produced, would intersect the dorsal-most line of the body.

No. 431.] PAIRED LIMBS OF THE VERTEBRATES. 845

rim alone is in advance of the gravity center. Thus, as this

center passes backward, the posterior margin of the ventral

fin is found to follow it pari passu, and, as effecting a counter

balance, like equal weights placed in opposite scale pans, the

anterior rim of the pectoral passes headward.

Again, contrasting the embryos shown drawn to scale in

Fig. 1, and referring now to the position of the external gill

openings, we note that there is a marked tendency for these

openings to pass backward. In the earlier specimens all are

widely interspaced and the last is well in front of the fore rim

of the pectoral; in the next embryo the openings are drawn

together, altogether somewhat tailward ; and in the latest, as

far at least as to the second opening, the series have taken a

position hindward of the rim of the pectoral.

' More concretely (measurements taken from the specimens) :

PERCENTAGE OF LENGTH FROM SNovT Tip. |

| PERCENTAGE OF TAIL-

WARD MIGRATION.

GILL OPENING.

Stage Ir. Stage 2. Stage 3.

I 11% 13% 16 % 5 %

y 19 % 19 % 23.5 % 4-5 %

In other words, as far as these openings are concerned, and

with them the outer part at least of the gill canal, it is evident

that in the older stages a definite migration has taken place, to

a degree that, in terms of the total length of the animal, the gill

Openings come to occupy a position in the body wall which is

largely behind that of the earlier stages. Numerically consid-

ered, the interval between gill openings I and V is about 8 per

cent of total length in stages 1 and 3; but in the latter the

region of the gill openings is nearly 5 per cent farther tailward ;

therefore, in this stage the openings occupy a position in the

body surface which is about five-eighths new, z.e., never before

occupied by gills. From this it follows that the gill region, at

least in its outer part, shows no affinity during proportional

&rowth with the neighboring region of the pectoral fin. In fact,

from an early stage onward, they are evidently growing in oppo-

Site directions.

846 THE AMERICAN NATURALIST.

If, now, we figure the matter up, we find that the distance

the gill openings pass tailward (contrasting embryos vary-

ing from about 20 to 300 mm.), added to the distance which

the anterior rim of the pectoral passes headward, makes the

considerable sum of 20 per cent of the length of the animal.

And from this it follows that there occurs in this region a

process of “shortening up," which is surely enough to account

for the presence of anastomosing vessels, blended muscle bands,

plexus of nerves, and “collectors.” The presence of such com-

plicated structures,! especially in the region of the anterior or

posterior rim of a (pectoral) fin, is therefore to no little degree a

product of the mechanical needs of fin migration.

COLUMBIA UNIVERSITY, October 17, 1902.

! Another complicating feature to be considered in this connection is the dispo-

sition of segments arising in the caudal region and growing cephalad. In the

earlier embryos (15 to 20 mm.) the total number of segments behind the last gill arch

is enclosed within a space measuring about 80 per cent of the entire length; later

specimens (300 mm.) include the segments within a space of 76 percent. The size

of the embryo at which the formation of new segments ceases has not been deter-

mined in the present form, nor is it as yet ascertained in kindred forms.

cis 70 80 90 100 %

ESAL 2 Dorsal

PrcrORAU y | VexmaL ^

$$ à ^

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FET EE Ed

I3 9990

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measurements of si

“indicating the ofthe center of gra ity and of the fins. I

i the NOM [prn Lachs Spee UE mae Ina srecimers

cis ic Kao aaoi a DACAAR. The leata of Eng aie QUUM ue ini.

cated (in millimeters) in the vertical columns at the left. oe te oa s

EIECTUS arms hee

TEES T aiuto

IIPPURUPPTTILLLLLERA

— j

Ao à 138 mm. are T na

NOTICE OF INTERESTING NEW FORMS OF

CARBONIFEROUS FISH REMAINS.

C. R. EASTMAN.

TuRoucH the kindness of Professor G. Hambach, of Wash-

ington University, St. Louis, a number of highly instructive

Carboniferous fish remains have recently been brought to the

writer's attention, two of which seem to possess considerable

interest, and are therefore made the subject of the following

notice. The first to which attention is invited is a remarkable

specimen of Erismacanthus from the vicinity of St. Louis, now

preserved in the private collection of Dr. Hambach. It is

unquestionably the largest and at the same time most perfect

spine belonging to this genus yet brought to light, and appears

to be distinct from other described species. The second speci-

men noticed in the present paper yields new information regard-

ing the dentition of Campodus corrugatus, and is from the Coal

Measures of Winchester, Illinois.

Genus Erismacanthus M'Coy.

The two European and one American species of this genus

that have been described are evidently closely related to Physo-

nemus, but differ in that the spines are divaricated, the two

branches extending in opposite directions in the same vertical

plane. The imperfect ichthyodorulites known as Gampsacan-

thus, Lecracanthus, and Dipriacanthus are also of the same

general nature, and all these bodies are interpreted by most

authors as lateral head spines of presumably cochliodont or

petalodont sharks. The occurrence of spines on either side of

the head in sharks and chimæroids has been observed in a

number of forms, such as in Ovacanthus armigerus Traquair,

from the Calciferous sandstone of Eskdale, Scotland, in the Per-

mian JZezaspis armata, and in Mesozoic cestracionts (Hybodus).

849

850 THE AMERICAN NATURALIST. (VoL. XXXVI.

The Physonemus group of spines has been theoretically associ-

ated by Jaekel with the teeth of petalodonts, but this conjecture

has not been corroborated as yet by any direct evidence.

Very interesting stages of modification are displayed by the

group of Physonemus-like spines throughout their existence in

the Lower Carboniferous. The earliest and most primitive

forms of the typical genus are found in the Kinderhook, in

accompaniment with small forms of Stethacanthus. The two

or more species of Kinderhook Physonemi that are known are

of small size, attenuated, and quite destitute of surface ornamen-

tation. Erismacanthus is also represented in the Kinderhook

by two small, comparatively unornamented species, and it is

noteworthy that the branched condition is here rudimentary.

The Burlington species of Physonemus and Stethacanthus

display a marked increase in size, but the ornamentation is

feeble, and remains so throughout the stage represented by the

Keokuk Limestone. Stethacanthus seems to have attained its

maximum size in the Keokuk, as Physonemus did in the Bur-

lington (P. gigas N. and W.), a considerable falling off in this

respect being true of both genera in the St. Louis division.

The spines of Stethacanthus remain unornamented from their

first appearance in the Berea Grit of Ohio until their extinction

in the St. Louis Limestone, but those of Physonemus and Erisma-

canthus increase in complexity of ornamentation throughout the

Mississippian series, ultimately displaying great elaboration.

An inspection of the forms illustrated in Plate XXII of the

sixth volume of the 7//inois Paleontology, or of the large spine

immediately to be described, will, we think, satisfy any one as

to the correctness of this latter statement and of the above

generalizations.

Erismacanthus formosus sp. nov. (Fig. 1).

DEFINITION. — A very large species, the spines attaining a total length

of at least 22 cm. Anterior branch stout, much produced, gently arched,

one side more or less flattened and provided with a double row of obtusely

conical denticles, the other smooth and convex, appearing as if imbedded.

Posterior spine considerably arched, prominently keeled, laterally com-

: and bearing a double series of closely set denticles along the distal

; half of the posterior margin. The posterior spine is relatively wide in its

No. 431.] CARBONIFEROUS FISH REMAINS. 851

basal portion, and tapers gradually toward the acuminate distal end ; its

surface is ornamented with numerous longitudinal costae which increase by

bifurcation, and by rows of small, rounded or spiniform tubercles occupying

the intercostal spaces, these being especially numerous on the broad basal

portion of the spine, and extending also over the proximal portion of the

anterior branch.

The above description is based upon the remarkable spine

shown in Fig. 1, together with two or three imperfect specimens,

all collected by Dr. Hambach from the St. Louis Limestone in

the vicinity of St. Louis, Missouri. The large specimen has

a total length of 21.5 cm., of which the anterior branch

forms about four-sevenths and the posterior spine three-

sevenths. It is nearly seven times the size of Æ. maccoyanus

vitare are "p x both

Fic. n AES Jermens p, ngr EM Toar Limestone; St. Louis, Mo. Late

x i. fram

hath aaa i

ats ar than the natural sos.

St. J. and W., which accompanies it in the same formation, and

twice that of E. jonesi M'Coy, the largest known European

species. From both of these the present species differs not

only in size, but also in the form of cross section and denticula-

tions of the anterior branch, as well as in the general surface

ornamentation. It appears improbable that the anterior branch

was terminated by a cluster of denticles, but there is a double

row of about twenty robust conical enameled bosses along the

flattened face of this arm, which have evidently become obtuse

through wear. This flattened face was probably entirely exposed,

but there is reason to believe that the opposite, or more convex

face, was buried in the integument. Its smooth surface pre-

sents the same appearance as the inserted portion of most fin

spines, and betrays none of the usual indications of wear.

852 THE AMERICAN NATURALIST. . [Vor. XXXVI.

Obviously if this portion of the spine had projected freely, and

were not anchored by insertion in the soft parts, it would have

been not only a useless appendage but a positive hindrance to

the creature, besides being constantly exposed to injury; but

if we conceive of it as inserted in the integument in the same

manner as the anterior prolongation of Oracanthus vetustus,}

its function as a defensive organ and as a support for the pos-

terior spine is readily understood. Evidence that this was a

paired spine, and therefore not occupying a position in the

median line of the back or of the head, is afforded by the

fact that the exposed face of the anterior branch is turned so

as to present the denticles toward the left-hand side; and the

organ may therefore be interpreted as a lateral head spine

belonging to the left side of the body.

The posterior spine is very similar to the single defenses of

Physonemus, but is more strongly keeled and broader at the

base. The distal half of the posterior margin bears a double

row of closely set denticles resembling those of E. maccoyanus.

The sides of the spine are occupied by numerous fine, smooth

longitudinal costz, of which about fifteen are to be counted

along the line where the cross section shown in the figure is

taken. The intercostal spaces, more particularly in the basal

portion of the spine, are studded with numerous small tubercles,

which are stellate in the unworn condition, and vary in shape

between conical and spiniform. In many cases their summits

are mucronate with the points reflected toward the basal por-

tion of the spine, and the resemblance of these tubercles to

certain types of placoid scales is very marked. It is probable

that the imperfect spine illustrated in Pl. XXII, Fig. 3, of the

sixth volume of the Minois Paleontology, represents a young

individual of this species.

Genus Campodus de Koninck.

This genus of cestraciont sharks is represented in the Coal

Measures of North America by three species, one of which —

C. variabilis (N. and W.)— is of importance as affording a

. 1 6f. J. S. Newberry, Trans. N. Y. Acad. Sci., vol. xvi (1897), p. 287, Pl. XXIL

Fig. 3.

No. 431.] CARBONIFEROUS FISH REMAINS. 853

complete insight into the dentition of Orodus-like forms. The

species described by Newberry and Worthen as C. corrugatus

has been known hitherto only by a few of the lateral teeth,

Fic. 2. — C. (N. und W.). Aim Measures ; Winchester, Ill.

Oral aspect of a i portion of t ial dentition, x 34.

and we have been without information regarding the symphysial

series. One specimen, however, of the symphysial dentition

has recently come to light, and seems worthy of particular

description.

854 THE AMERICAN NATURALIST.

Campodus corrugatus (Newberry and Worthen) (Fig. 2).

The symphysial dentition of this species differs from that of

C. variabilis in that the individual teeth are much more robust,

less intimately fused with one another, and less differentiated

in form from the lateral series; they are also apparently fewer

in number than in the corresponding series of C. variadclis.

All these characters point to the conclusion that as between

the two species the latter is much more highly specialized

and stands nearer to the line which gave off Edestus and other

extremely modified cestracionts.!

The unique specimen of C. corrugatus shown in Fig. 2

exhibits five very robust teeth, which are recognizable by their

peculiar form as having pertained to the symphysial series.

Of how many teeth the entire series was constituted cannot be

told, but owing to the large size of the teeth it is probable that

the number was less than in C. variabilis, which has as many

as thirteen. The apical extremities of the crowns have unfor-

tunately all been broken away, and the bilateral symmetry of

the series has been disturbed by mechanical agencies since the

death of the creature, so that the teeth appear to succeed one

another in spiral fashion, instead of being arched in a single

plane. The apical portions of the crowns differ from those of

C. variabilis, also, in that they are more elevated, and are

inclined backward at a slight angle.

The coronal surface of each tooth is marked by a prominent

transverse crest, from which numerous branches are given off

approximately at right angles on either side. The anterior

margins of the crowns are very prominently buttressed, an

especially large projection occurring in the median line on both

the anterior and posterior faces. Fig. 2 is photographed from

a cast of the original specimen, which was obtained from Bates

and Company's coal mine at Winchester, Illinois.

1 A detailed account of the dentition of Edestus and related forms is given in

Bull. Mus. Comp. Zoöl., vol. xl, No. 3, 1902.

THE MICROCOSM OF THE DRIFT LINE.

LAETITIA M. SNOW.

IN the spring of 1902 my attention was called to the

extremely interesting life relations of insects along and

around the line of drift thrown up by the waters of Lake

Michigan, and a series of collections and observations were

made between April 14 and May 31. The collections included

one hundred and fourteen species, only fifty-one of which it

was possible, with the means at hand, to identify. The speci-

mens have, however, been preserved.

PuvsicaL FEATURES.

As with life everywhere, the physical features of the habitat

are of great importance to the population. For instance, on

Windsor Park beach, including the region north to Seventy-

Second Street and south to the Steel Works, the forms were,

as a rule, much more abundant than on the two beaches in

Jackson Park. On the former beach the gradient is low and

the sand smooth ; the water action is rather gentle, the insects

stay when the water retreats, and the fine sand keeps them

on the surface. At Jackson Park beaches, on the other hand,

unless the water has recently been high and beyond the pebble

zone, the collecting is poor. In the pebble zone the gradient

is greater, the water action greater, and there is the possi-

bility of the insects being washed out again into the lake.

That the physical conditions of the beach account for the

greater number of forms, and not the fact that collections at

the two places were made at different times, may be proved

by the fact that on the same day a pebble region of beach

north of Windsor Park gave no results, while a smooth region

yielded a number of forms.

855

856 THE AMERICAN NATURALIST. [VoL. XXXVI.

CAUSES OF DISTRIBUTION.

The waves of the lake wash up great quantities of refuse,

among which are numbers of insects. These animals may be

either land or water forms; if the former, they have either

come directly upon the shore, or have flown lakeward or been

blown by an off-shore wind, and then been washed in, dead or

dying. On account of the lake current moving southward

along this region, specimens collected at Chicago may have

come from points farther north (13). To this group of

stranded land forms belong the ladybirds (Coccinellidz),

Chrysomelidze, leaf-eating Scarabzidz and Carabide, possibly

the Elateride, Lucanide, and the Rhyncophora among the

Coleoptera, the Lygæidæ and Pentatomide among the Hemip-

tera, the Hymenoptera, Neuroptera, Trichoptera, and Lepi-

doptera, — all of which are herbivorous. Also to this group

belong some predaceous carabids and one roach. To the

stranded water forms belong Benacus griseus, Hydrophilus

triangularis, and two other water beetles.

These stranded forms may revive and depart, may serve as

food for predaceous forms, or, if dead, fall to the share of the

scavenger insects. The flies and ants are undoubtedly scav-

enger forms, and inhabit this region on account of the food

supply. In this class I would also place a small carabid beetle

(undetermined) which occurred for a while in great numbers

under stones and débris, so as to preclude the idea of its hav-

ing been stranded during migration. Several small beetles

seem to have this scavenger habit ; for example, three Staphy-

linidze, some Scarabzeidze, and several Carabidae. In this latter

family we find some predaceous forms which seem to inhabit

this region normally, to feed on the dying insects or on their

destroyers ; for example, several black forms (undetermined)

and Galerita janus. The spiders are also predaceous, and feed

on the flies, ants, and other small living insects. I might

add that birds come in as a final factor and eat these various

forms.

No. 431.) MICROCOSM OF THE DRIFT LINE. 857

ORDER OF SUCCESSION.

These insects do not come all at once, nor “hit or miss,"

but follow a fairly regular order of succession. It was seldom

that the same form was dominant at two visits. Between

April 14 and 21 very few insects were found, and no regular

record was kept. The temperature was cool and the winds

variable during that period. On April 25, after four days of S.,

S.W., and W. winds, followed by E. and S.E. winds, more forms

appeared. Two days of strong (thirty to forty miles per hour)

W. wind, followed by a S.E. wind, brought in a few forms on

WIND PREVIOUS TO COLL. T Temp. F. SPECIES.

Bow Dav or Corr.

Days. Direction. Av. Vel.|Direction.| Av. Vel. | Max. | Min. | New. | Old. | Total.

Apr. 25 2 W.to E. 19 S.E. 14 69 42 IO | I II

28 | 2 W: 374 S.E. 15 61 48 | 3 | o 3

May 2 2 E. to N.E. 20 S.W. 18 82 53 37 5 42

12 3. |N.E.toE.to N.E| 21% N.E. 10 56 43 27 10 37

16 7 N.E. t E. 16$ E. 10 61 48 | 6 19 25

23 2 S.W. to S. 12] S. E. 10 77 6r | 13 19 32

27 2 W. to N.W. 15 NES 18 50 40 8 7 15

a 3 S.W. 17) S.E. 15 75 60 | 2 14 16

(Data kindly furnished by the U.S. Weather Bureau, Chicago, Ill.)

April 28. All this time the temperature was cool, only once

rising over 69° F. On May 2 a N.E. wind (twenty miles),

preceded by a W. wind, brought in a good many forms, thirty-

seven new species being found, besides several previously col-

lected ; the rising temperature (53°-82°) probably influenced

the numbers. On May 12 twenty-five new species were

found, besides an abundance of old ones. This rich supply

was probably due to three days of W. and S.W. wind, fol-

lowed by four days of N.E. and E. wind (averaging eighteen

and three-fourths miles) which continued until May 16 with

decreased velocity (thirteen and one-fourth miles), the tem-

perature never exceeding 61? F. On this day only five new

858 THE AMERICAN NATURALIST. (VoL. XXXVI.

species appeared, but numbers of old ones were found. On

May 23, after a warm period with S., S.W., and S.E. winds

of low velocity, fourteen new species were collected, a few old

ones appearing. A N.E. wind on May 27, preceded by W.

and N.W: winds of moderate velocity, brought in countless

Colorado potato beetles and ten new species, the temperature

dropping twenty degrees in two days. On May 31 a S.W.,

followed by a S.E., wind brought only two new species, but a

good many old ones occurred. These results are placed on the

preceding page in tabular form.

DomINANT Forms.

The first form to predominate notably was Chrysomela ele-

gans (May 2), while. Hippodamia parenthesis came in a good

second. The little Coccinella sanguinea occurred throughout

the season in about the same numbers, but absence of other

forms made it at times the dominant species. Black carabids

occur also throughout, but never as dominant forms. The

next visit, May 12, showed Diabrotica vittata as the charac-

teristic species, while of Chrysomela only a few specimens

were found. The small brown and black scavenger carabid

was present in great numbers, and should probably be classed

as the dominant form. Coccinella sanguinea was numerous,

as were also black carabids. The following collection showed

the Diabrotica, although not the characteristic species, still

occurring in fair numbers; the scavenger carabid seemed to

be the most numerous form, with quantities of Coccinella san-

guinea. Of the larger forms Bombus sp.? and Hydrophilus

triangularis were the most important. This time (May 16)

marked the first occurrence of Galerita janus, and the flies

were first noted. The various forms of Hemiptera occurred

throughout the collecting trips, but were never dominant forms.

On May 23 the Colorado potato beetle (Leptinotarsa decem-

lineata) was the dominant form at Jackson Park. This appears

to have been the vanguard of the great “migration” which

took place on, or just before, May 27; for on that date the

. beach at Cheltenham was- nearly covered with the beetles,

No. 431.) MICROCOSM OF THE DRIFT LINE. 859

dead, dying, or active. In the latter case they had in many

instances crawled up on stones and driftwood, until the sur-

faces of such articles were, in places, covered with them, some-

times two layers deep. On May 31, however, very few live

ones remained, and the total number of individuals was

greatly diminished. The form that approached the potato

beetle in number on May 27 was the May beetle, or June bug

(Lachnosterna sp. ?).

Besides those above mentioned, other forms occurred in

greater or less abundance at different times, some running

through the whole period, some starting late and continuing

to the end, some starting early and disappearing.

MIGRATIONS.

Would the fact that certain forms culminated at certain

times indicate that they simply reached the adult stage at that

time, laid their eggs and disappeared, or that they were per-

forming mass movements, broadly called ** migrations"? The

great swarm of potato beetles on May 27 closely resembled

mass movements recorded by many authors, in that the num-

ber was large and that the animals were found on the 27th

and by the 31st had almost entirely disappeared, no new ones

appearing. If it were merely a general emergence, it is not

likely they would have been found only along the lake front.

This great washing up of one species recalls the predominance

of crickets (Memobius fasciatus) described by Needham (1).

Much has been written on this subject, both descriptive and

theoretical, but the problems as to the * why " and “ whither ”

of insect flights have never been solved. The search for food

has been suggested as the cause, and the suggestion has been

accepted by many as at least a secondary factor. In some

cases over-production, followed by a scarcity of food, would

seem a sufficient reason for the movement ; as, for instance,

the migrations of locusts, both in this and other countries

(2, 3). Certain examples of the flights of dragon flies point

to the same cause (3, p. 509; 4; 10). Kobelt (3) quotes

Gatke as saying: “Es ist schon darauf hingewiesen worden,

860 THE AMERICAN NATURALIST. [Vor. XXXVI.

dass auch die Bewegung der Nachtschmetterlinge meteorolo-

gischen Beeinflussungen unterworfen sind. Diese Ansicht

stiitzt sich auf wiederholte Beobachtungen, nach welchen die-

selben unter gleichen Bedingungen wie die Vögel, und fast

immer zusammen mit diesen in ostwestlicher Richtung hier

vorbeiziehen "; which may be the key to the whole question.

As far as I am aware, no barometric observations have been

made directly on the subject of insect migration (15).

That a feeling of hunger is not alone a sufficient cause for

some of the direct flights recorded, is shown by the fact that

butterflies often continue in their path, over the very flowers

they use for food, only a chance one stopping on the way

(3, P. 509). Also, if they simply spread out to find food,

the progression would be diffuse, directions inclined to be

random, and the advance a gradual one, as is the case with the

spring “ dispersal through isolated individuals" of Danais (5)

which go north as the spring advances, various generations

taking part in the advance.

In many instances the direction of flight has not been noted,

but in Europe the general tendency seems to be from east to

west. In our country, however, no such law appears to hold.

More attention has been paid to the consideration of whether the

animals, chiefly butterflies, flew against or with the wind, some

holding that the latter feat was impossible. That it is at least

possible, has been shown by some direct observations (6, 7).

It has been suggested that, as in some cases the swarm was

composed of individuals of one sex (males), the flight origi-

nated in a search for mates; but as the absence of females

can be explained in some instances by their wingless condition

(8, 1900, p. 13), while in most observations the sexes were nearly

equal in number, this cause may apparently be ruled out.

If they are then not necessarily merely drifted by the wind,

or impelled by immediate hunger or by the sexual instinct,

why do they follow such direct routes, fly with such apparent

aim, and often repeat the flight at intervals? Walker attrib-

utes it to *a propensity to migrate” (8, 1901, p. 353), thus

throwing it back upon hereditary tendencies, while Kefer-

stein (4) adds to over-production and some aid from wind,

No. 431.] MICROCOSM OF THE DRIFT LINE. 861

an unknown impulse to take a common migration. Riley (5)

makes the statement that “all insects acquire the migratory

instinct when crowded together through excessive multiplica-

tion." The probable response to meteorological influences has

been noted.

Hancock (9) distinguishes between dispersal and migration

in the statement, * Individuals of a species which effect a

more or less regular periodical change in their habitat are

truly migratory. Migrations may be primary, consisting of

local flights, such as movements by insects hatched in tem-

porary regions, to which they confine themselves to passing

to and fro, from point to point; or secondary, as the repeated

periodical changes of residence covering foreign fields, which

naturally establishes a nomadic habit." In addition to these

distinctions it is suggested that the term “migrations” be

confined to periodical changes of habitat resulting from the

normal sequence of temperature and season, while such abnor-

mal occurrences as the devastating swarms of Rocky Moun-

tain locusts or irregular and unusual swarms, as in certain

instances of butterfly, moth, and dragon-fly flights, be given,

temporarily at least, the term “immigration.” Mr. Hancock's

*primary migrations or local flights" would be simply dis-

persal flights of individuals or groups of individuals within

their area of distribution.

Was the flight of Colorado potato beetles on May 27 a dis-

persal (or local) flight, diverted out of its course and driven

lakeward by west winds, or was it a periodic migration? The

predominance of other insects at various times was probably

due to the spreading of groups of individuals by local flights :

may not these local flights give some evidence of the dispersal

paths of various species ?

CONCLUSION.

By whatever means the animals reached the region, — by

migration or dispersal movement, — the life relations of the

beach are not altered. We have here a little community of

food providers and food obtainers, whose population varies

with the season, the wind (probably), the beach conditions,

862 THE AMERICAN NATURALIST. [Vor. XXXVI.

and the relative abundance of the various forms. For example,

we found (1) the occurrence was in succession ; (2) the popu-

lation increased apparently at times when an offshore wind

was followed by a lake breeze; (3) it also increased with the

temperature; (4) the greatest numbers occurred on beaches

of low gradient and smooth, fine sand; (5) the abundance of

scavenger forms depended upon the abundance of dead her-

bivorous and predaceous forms and other refuse; (6) the abun-

dance of predaceous forms depended upon the presence of the

active scavenger and herbivorous and smaller predaceous forms.

I gratefully acknowledge the kindness of Dr. Charles B.

Davenport and Mr. Charles C. Adams, and present my thanks

for their assistance in identification and for valuable advice

and suggestions.

HULL ZOOLOGICAL LABORATORY,

July 18, 1902.

List.

Of the one hundred and fourteen species collected fifty-one

were identified, the whole collection being placed in the follow-

ing orders and (where possible) families : —

Coleoptera, 75: Carabida, 33; Scarabaida, 10; Chryso-

melidz, 9; Coccinellide, 7; Staphylinide, 3; Hydrophilidz,

3; Cerambycide, 2; Elateride, 2; Silphidz, 1; Histeridz, 1;

Halipide, 1; Lucanidz, 1; Curculionide, 1; Calandridz, 1.

Hemiptera, 11: Pentatomide, 5; Lygæidæ, 2; Reduviide,

I; unknown, I.

Diptera, 10: Muscidae, 5; Syrphidz, 2; Phoridæ, 1; Tach-

inida, 1 (?); unknown, 1.

Hymenoptera, 9: Apidæ, 4; Formicide, 1; Vespide, 1;

Andrenidz, 2; Tenthredinide, 1.

Lepidoptera, 3: Arctiidz, 2; Noctuide, 1.

Trichoptera, 2: Phryganeide, 2.

Neuroptera, 2 : Chrysopide, 2.

Orthoptera, 1: Blattidz, 1.

Arachnida, 1 : (Lycosidz ?)

No. 431.] MICROCOSM OF THE DRIFT LINE.

IDENTIFIED SPECIES.

863

First Last coL-

NAME. coul irn: DOMINANT. *

COLEOPTERA

ADAoditt Hmetariur > oa counts dua April 14-21 -— May

Hippodamia 13-punct. : 25 = I2

Hippodamia parenthesis 25 "od 2

ippodamia convergens 2 ine -

Coccinella san, MEE 25 dics 3!

Cyllene pictus . . (^ UNE UM May 2 Bugs js

Necrophorus marginatus 2 a n;

Coccinella g-no i 2 e 3t

Chrysomela aan 2 2 31

Anatis 15-punctata x um

Agonoderus pallipes . 3 ar 23

Casnonia AEE 2 RS 16

Drasterias elega 2 un Hi

Disonycha Miia - 2 gz ei

Megilla maculat. 12 27

Diabrottica vittata 12 I2 16

Diabrottica 12-punctata . 12 Oc I

co - ro-lineata 12 27 3l

osoma scrutator «T 12 3t

si wilcoxii ái pien 12 = x

Elaphrus ruscarius arg 3 12 cut oe

Platynus | ees nx 12 — 16

Galerita 7 i ; 16 = 23

MTE A triangulari ux y 16 ike 3t

Lucanus dama . . 23 n g

Calosoma calidum 2s > 23 qu 3t

a ind. i wo poe 23 He VETE

Phymatodes sp. ? 3 See kid 23 — vum

opinus incrassatus. . . y Sirs 23 exe e

ZLachnos. Spo cru. ww ie 27 aK x

Cotalpa CAME oe, os UR 27 — xum

ss utter deigdae tt, 1. 3. ok co 27 side a

: s POO ORME c0. soene e 4M .31 Ser ee

Coptocycla aurichaleea . . . o p s. 3I pagea mec

YMENOPTERA l

ND runnen S ee ee April 15-21+ = e"

Polistes variatus NE i var . May 2 ncn 16

PPM Bie? oe Se a N 2 = ues

Augochlora sp.? . . i ic. OR 2 un ea

SOME WT Uu uo. i RO nio 12 vom 27

GAME ERI S o. oso o 27 — eM

Nomada sp. ? jo 4d 4 4 v s 27 — —

864 THE AMERICAN NATURALIST.

IDENTIFIED SPECIES (Continued).

First LAST COL-

NAME. Y COLLECTED. Dominant. LECTED.

DIPTERA

nao di torvos d a TER eco ER UP April 25 — May 16

Lucilia Se Ua ce ie 28 — 3t

rota ZUGFENEEMAS ER dex May 23 on? -— ==

HEMIPTERA

Oncopeltus O° ogg i May 2 I Be

Benacus griseu. s 16 Eus 2x

LEPIDOPTERA :

Estigmene acre ( "I i ENE EE a

Eyprepia(?) . AA a ay Mare aa x

ARACHNIDA

Present throughout

REFERENCES.

1. NEEDHAM, J. G. Occ. Mem. of Chicago Ent. Soc. Vol. i, No. 1,

. RiLEY, C. V. Locust Plague in the United States. Chicago, 1877.

3. KoBELT, W. Verbreitung der Thierwelt. Lieferung Io, pp. 499-509.

4. KRFRESTEIM, A. Zeitg. f. ges. Naturwiss. Bd. xxii (1863),

PP. 249-275.

5. RILEY, C. V. Sci. Amer. Vol. xxxviii, N.S., 1877, p. 215.

8. Tutt, J]. W. Ent. Rec. and Journ. Var. 1900, 1901, 1902.

9. Hancock, J. L. Amer. Nat. 1894, pp. 483-487.

10. CALVERT, P. P. Trans. Amer. Ent. Soc. Vol. xx, pp. 272 ff.

II. RiLEY, C. V. Amer. Nat. Vol. xv,

12. WEBSTER, F. M. The Trend of TEM Diffusion i in North America.

Ann. Rep. Ent. Soc. of Ontario. 1901, pp. 63-67.

13. United States Department of Agriculture, Weather Bureau Bulle-

14. HAGEN. Insectenzüge. Svett. Ent. Zeitg. Bd. xxii-xxiii (1861),

(apparently an important reference, which I was not able to obtain).

Ik PACKARD, A. S., JR. Amer. Nat. Vol. xi, pp. 22 ff.

STUDIES OF LOCALIZED STAGES OF GROWTH

IN SOME COMMON NEW ENGLAND

PLANTS.

JOSEPH A. CUSHMAN.

In the following are given some of the stages in detail found

in a few of our common plants, and some general notes which

may be safely drawn, and which may be useful in studying

plants from this point of view. The cases given are not

chosen as particularly striking ones, but are such as would come

naturally under the notice of anyone. In a following paper

the writer hopes to present more striking examples from some

of the less generally known plants now in the Botanic Gardens

of Harvard University. The few given, however, will show

the truth of the principle involved as well as more striking

ones.

In a paper by Robert T. Jackson, “ Localized Stages in

Development of Plants and Animals," Memoirs Boston Society

of Natural History, Vol. V, No. 4, 1899, this principle was

worked out and applied to plants. It was there applied to

shoots and various growths of trees almost exclusively. It is

in the spirit of that paper that I have attempted this one,

giving the results of my study of plants from this point of

view. My thanks are due to Dr. Jackson for reading this in

manuscript and for many suggestions.

The new point intended to be brought out here is the appli-

cation of the principle of localized stages to the early spring

growth of herbaceous plants. These are adult plants, not

young plants or seedlings. The principle is also applied to

the stages immediately preceding the flower.

The principle of localized stages in development, to quote

from the above paper, is that, “throughout the life of the

865

866 THE AMERICAN NATURALIST. |. (Vor. XXXVI.

individual, stages may be found in localized parts, which are

similar to stages found in the young and the equivalent of

which are to be sought in the adults of ancestral groups.”

This principle may be well demonstrated in a greater or less

completeness in each of the following examples.

Viola pedata L. (Bird’s-Foot Violet).

Under this species Gray’s Manual gives, “leaves all 3—5-

divided, or at the earliest only parted, the lateral divisions

2—3-parted, all linear or narrowly spatulate, sometimes 2-3-

toothed or cut at the apex," thus recognizing the fact of the

differences in the leaves but with no significance of the various

forms or connection between them.

On April 18, 1902, a study was made of the leaves of this

species in adult plants which were then well started. Among

the larger and stronger plants it was found that the earliest

leaves went back in the accompanying series usually to Pl. I,

Figs. I0 or 11. After further search plants were found with

the leaves on the outside of the rosette, similar to Figs. 9, 8,

or7. Noticing that the smaller plants seemed to be simpler or

more primitive, attention was turned to them, and some small

ones were found with the outside leaves going back to Fig. 6,

and the smallest to Figs. 5 or 4. These were along a woods

road in a somewhat dry, sandy place. In the middle of this

road, where there was very little vegetation, were found plants

evidently very young and weak, with the result that some were

obtained with the simple leaves shown in Figs. 3, 2, and I.

That these were not seedlings was shown by the rootstock.

They were probably weak plants of the preceding year.

In nearly all the stages shown many specimens were obtained

of each stage, and in all cases enough to warrant the form as

not abnormal Where the outer leaves were most primitive

the more complete were the stages before the typical leaves

were produced in the center. Seedlings of this plant were not

obtained, but the earliest of the stages given are very similar

. to corresponding ones of Viola tricolor L. in both young and

adult. Later stages would naturally be specific. The seedling

No. 431.] STAGES IN PLANT GROWTH. 867

would probably not repeat all of the stages given here, but the

majority of them. The data here being fairly complete will be

referred to again later.

Chelidonium majus L. (Celandine).

Seedlings of this plant were found at Arlington Heights in

May, among the adult plants, and afforded a very convenient

opportunity to study the relations of the two.

The leaves of the seedling after the cotyledons are developed

as follows (Pl. II, Figs. 1, 2): First leaf usually slightly tri-

lobed and trinerved; second leaf usually decidedly five-lobed

and five-nerved ; the third leaf is decidedly trilobed, with the

median lobe notched, lateral ones entire or with one notch.

From the third on, the trilobed character persists for a short

period, the leaves becoming, later, three-pinnate, five-pinnate,

and so on.

In comparing this with the first leaves of the adult plant

when it starts from the ground in the spring, a striking like-

ness is at once noted. In the plant shown in PI. II, Fig. 3, the

first leaf is exactly comparable to the second leaf of the seed-

ling, and the second to the third of the seedling. The

third leaf is like the three-parted one of the seedling, the

fourth similar. The fifth is five-pinnate, and so on. In all

cases where the first formed leaves were found on the plant,

these comparisons were found to be true. It was especially

noticeable in the plants found at Arlington Heights, which

were on a sloping gravel bank with poorer nourishment than

in other cases noticed in Cambridge. In this plant the early

development in spring growth representing localized stages,

and in the seedling representing direct development, not only

show great similarity but may be exactly compared stage by

stage.

Aquilegia canadensis L. (Wild Columbine).

The seedlings of this plant have the first nepionic leaves typi-

cally trifoliolate, with the terminal leaflet having two well-marked

notches and each of the lateral leaflets one less-marked notch

868 THE AMERICAN NATURALIST. |. [Vor. XXXVI.

on the lower side. By raising a number of the seedlings of

this plant there was found a marked variation from the typical

form. In two cases the first nepionic leaves were trifoliolate,

but both the terminal and lateral leaflets were entire, no sign

of notches being present. The second leaves in the two cases

given were typical of the first of the other seedlings, thus

showing that these two individuals were less accelerated in

development than the others. These two repeated the char-

acter of a trifoliolate leaf with entire edges, which is usually

dropped out in the development of the seedling.

By comparison with the early spring growth of adult plants,

the first leaves of some weaker individuals were trifoliolate and

entire, as are those of the two seedlings noted above. This

is, then, not an accidental variation, but a true stage, which in

the seedling is usually skipped by acceleration of development,

and rarely appears there. In several cases the first leaves were

like the typical first leaves of the seedling. This, then, shows

two rates of acceleration, as it takes one longer to arrive at

a certain stage than it does another.

A striking similarity is noticed in the leaves immediately

preceding the flower. At the base of the flower stalk the

leaves are normal, but higher up the number of leaflets is

reduced until, a short distance below the flower, there appears

a trifoliolate leaf with notches varying considerably, some being

like the typical first nepionic leaf of the seedling, others vari-

ously notched. The majority, however, are like the seedling.

The next leaf above usually drops out the notches and has an

entire outline. This can be directly compared to the earlier

stage noted in two seedlings and in the average simple leaves

of spring growth of the adult plant. Just beneath the flower

is often found a single leaf, entire and exactly comparable to

the terminal leaflet of the trifoliolate leaf. Thus at the flower

the leaves retrace the steps which they went through in the

early growth of the plant and in the reverse order. This last

stage — the simple leaf below the flower — is more primitive

than anything that is found in the seedling, showing that local-

ized stages may be used to fill in the steps which are crowded

out by the acceleration of development in the seedling.

No. 431.] STAGES IN PLANT GROWTH. 869

Ranunculus acris L. (Buttercup).

This species has the typical leaves three-divided, and the

divisions considerably cleft and toothed, but the seedling starts

off with fairly simple nepionic leaves (Pl. II, Fig. 4). Compar-

ing the localized stages shown in the early spring growth with

the direct development of the seedling, a great similarity is

again noted. The plants were well started when observed,

and weak growths had to be largely depended upon to show

the stages. Pl. II, Fig. 5, shows a leaf of which one-half is

comparable to the first nepionic leaf of the seedling. The

other half is like the second nepionic. Here, then, is a leaf

showing an acceleration of development in its two halves, —

the half with two lobes being more accelerated than the other.

This is often noted in compound leaves which are pinnate, —

there being fewer leaflets on one side than on the other, — but

is less often definitely shown in simple leaves. A first leaf of

spring growth is figured, Pl. II, Fig. 6, which is comparable to

the second nepionic leaf. The steps in complication of the

lobes and nervation are shown in Figs. 7—9, and the seedling

in its later development passes through similar stages.

Beneath the flower the leaves become more simple usually

by four distinctly reverse, but often varied, steps. The lower

one of the four has about the same number of divisions as

Fig. 8, but is drawn out into linear divisions. Higher up is a

leaf that may be compared to the second nepionic of the seed-

ling, or to the leaf figured in localized development (Fig. 5), in

that it has five lobes. It is, like the preceding, deeply cut and

elongated. Above this is a three-lobed leaf of the same char-

acter and in its lobing comparable to the first nepionic leaf of

the seedling. Just beneath the flower is usually a simple, elon-

gated leaf, showing once more a localized stage in the adult

plant more primitive than any of the stages shown in the

direct development of the seedling.

This species is one which is usually well developed in the

early spring, and these earlier and weaker forms are hard to

find without careful searching. The first formed leaves quickly

dry, curl, and then drop off.

870 THE AMERICAN NATURALIST. [VoL. XXXVI.

Spirea salicifolia L. (Common Meadowsweet).

This is not herbaceous, and the stem, unlike the others before

noted, is woody, and the early spring growth is from the buds

at the sides as well as new shoots from the ground. Sucha

lateral growth is figured, Pl. III, Fig. 3. The first leaf indica-

tions are mere bracts. The first of what may be called the true

leaves is entire, lanceolate, with a simple median vein. This is

the first in the series and the simplest. This seems to be a case

like that noted under Aguzlegia canadensis, where a stage is

found simpler than that of the ordinary seedling first stage, as

the first nepionic leaves of two species of Spirzea as figured by

Lubbock are like No. 2 of Fig. 3. The second leaf is cuneate

and tridentate, trinerved, the broadest part near the tip. The

third leaf is five-dentate like the second nepionic leaf figured

by Lubbock. The complexity of dentations increases and the

broadest portion of the leaf moves from near the tip to near

the base. The leaves become doubly serrate, but the tip holds

the form of the second leaf given, throughout.

The spire formed by the flowers usually has one or more of

the lower branches with more leaves than flowers, and here the

reversal of the stages is well shown. Near the main stalk the

leaves are typical, but moving distally along the branch they

will be seen to lose little by little in the number of dentations.

This keeps on until there are but three, as in the second leaf

figured, and then there are several simple ones. The wedge

shape, with its broadest part toward the tip, is not reassumed,

but in dentations the stages are exactly reversed beneath the

flowers.

Tanacetum vulgare L. (Tansy).

As far as was found, the seedling of this species has not been

published, and therefore it is given in a little more detail than

the others (Pl. III, Figs. 4 and 5).

Cotyledons glabrous, subfleshy ; lamina oblong, obtuse, ses-

sile. The first nepionic leaves are paired and vary considerably.

. As a whole they are spatulate, obtuse, and may be distally

No. 431.] STAGES IN PLANT GROWTH. 871

toothed. Ten seedlings in a single pot were compared as to

their first nepionic leaves as follows:

Both first leaves simple . :

One simple, one one- hod ‘

One simple, one two-notched

Both one-notche

One one-notched, one Showin

Both two-notched . Sr

De bee BK wu

This crude table is enough to show the individual variation

of seedlings under the laws of acceleration of development

and the consequent necessity of making sure of the typical

stages.

The second pair of nepionic leaves have typically two lateral

teeth on each side in some cases, with more complicated forms

than this, but even in this case, so far as seen, only one of the

pair has the more complicated form, the other being typical.

The midrib begins to be distinct.

The next leaves are quite dissected and begin to decidedly

resemble the typical adult leaf. It may have in the third pair

of nepionic leaves from three to seven or more lateral teeth on

each side, and these may also be divided or notched.

Among the adult plants at the very base in the first spring

growth, leaves may be found which repeat many of the simpler

conditions. Fig. 6 is decidedly three-parted, much as the first

nepionic leaf in many ways, but less strikingly so than Fig. 7,

which is quite like the second nepionic leaf of the seedling.

Figs. 8 and gare like later seedling stages. All of these leaves

are from the base of fresh spring growths.

The flowers of this plant are in a dense corymb, each portion

of which comes from the axil of a leaf. The leaves in the

upper part become inconspicuous, but repeat, however, many of

the characters seen in the seedling and in the reverse order,

those immediately below the upper heads being like the first

and second nepionic leaves in the number of teeth and general

character. This is exactly comparable to the principles of

senescence of the late Professor Hyatt, stages occurring in the

old age of the individual corresponding to stages passed through

872 THE AMERICAN NATURALIST. [Vor. XXXVI.

in nepionic development, but in the reverse order of succession.

He has shown this at length in his “ Genesis of the Arietidae,"

published in part as a paper of the Museum of Comparative

Zoólogy of Cambridge, and as No. 673 of the .Swthsontan

Contributions. It is given more simply and concisely in a

paper, * Cycle in the Life of the Individual (Ontogeny) and in

the Evolution of its Own Group (Phylogeny)," Proceedings of

the American Academy of Arts and Sciences, Vol. XXXII,

No. 10, as well as in other papers. This case seems to be

comparable in that it is a failure to develop the full characters

of an earlier adult condition, due to the strength of the plant

going into the production of flower and seed instead of develop-

ing leaves. The tip of the leaf in all cases repeats the triden-

tate character of the first nepionic leaf, both in the tip of the

leaf proper and in the tips of all the divisions.

Viola tricolor L. (Pansy).

This plant is very seldom a perennial, but often comes up

from the rootstock the second year. Such plants are easily

obtained, and so exactly do the stages compare with the seed-

ling that were it not for the rootstock and the absence of

cotyledons the two might be easily mixed. The seedling is

figured by Lubbock, Seedlings (1892), and in that and in

seedlings examined the stages were exceedingly like those of

the early spring growth from a rootstock. The reversal of these

stages beneath the flower is seen with varying definiteness.

The old single form known as heart's-ease seems to show this

best. The number of notches decreases and the leaves return

to the forms seen in the beginning, those directly beneath the

flower being exactly like the first of the spring growth of the

adult and the first nepionic (Pl. III, Figs. 1 and 2).

Rosa rubiginosa L. (Sweet Brier).

In this plant, perhaps, of those given here, the stages are most

easily and completely seen. The seedling varies, having the first

nepionic leaf single as in Pl. IV, Fig. 1, or it may be trifoliolate

like the second leaf of Fig. r, as in Fig. 2; in each case the

No. 431.] STAGES IN PLANT GROWTH. 873

second nepionic leaf is trifoliolate, as well as several following

ones. Then the number of leaflets increases to five, and in the

full-grown plant becomes seven. In the first spring growth the

first leaf, especially upon the lower or weaker parts of the plant,

is very often trifoliolate, like the second nepionic leaf in all

cases and like the first in some. In some cases, however, a

simple leaf may be found, but rarely.. When it does appear,

it may be compared to the simple leaf seen in the first nepionic

leaf of Fig. 1. After this trifoliolate leaf in the spring growth

of the adult is one of five leaflets, then usually five leaflets until

the flowering stage. Then the number drops by the same steps

that it increases, but in the reverse order. Going toward the

flower, leaves are encountered with five, then three leaflets, until

finally just below the flower there is a simple one. Here, then,

are repeated the exact steps by actual count of the stages of the

seedling in the early spring growth, and as exact a reversal

of the same stages beneath the flower. The same is true of

Rosa lucida and other species (Pl. IV, Fig. 3).

Although these examples given might be multiplied almost

without number, they will do to illustrate several general prin-

ciples which may be drawn from them. The various conditions

are more or less combined in any actual case, as they often

are in such problems, but the effect of each may be noted

nevertheless.

First, the relation between the occurrence of the earliest

stages and the age of the plant. The plant coming up the

second year, other conditions being the same, seems to be more

primitive in its first characters than older and stronger plants.

It therefore repeats more stages in arriving at the typical form

than older individuals. The case of Viola pedata illustrated

this, the young plants of the preceding year being the ones in

which the earliest stages were noticed. In some plants, how-

ever, the typical leaf of the species is not reached for several

years. The young trees of Carya alba Nutt. the shagbark

hickory, have when very young — the second year, for example

— only single and trifoliolate leaves, and it is some time before

the number rises to five leaflets. The typical one of seven

leaflets does not appear until quite a late period.

874 THE AMERICAN NATURALIST. [VoL. XXXVI.

Weak individuals are more apt to show earlier stages than

are more vigorous plants. They represent plants which are

less accelerated in their development, not attaining the charac-

ters usual in plants of their age and other conditions. This is

true whether the whole is weak or whether the growth comes

from adventitious or weaker buds. The same is true of plants

injured by outside agencies; for example, when the leaves are

stripped by insects and a later growth put out from the weaker

buds, before undeveloped.

Soil and moisture conditions come into play here. The poor

sandy soil and lack of moisture mentioned in the first two exam-

ples given, shows its effect by producing weak individuals.

Plants in poor, dry soil often repeat stages not seen in strong,

vigorous plants in other places.

In studying these localized stages, especially in the early

spring growth of herbaceous plants, some things must be espe-

cially looked out for. The first leaves which appear are often

very small and would be overlooked unless they were being

sought for especially. Many of these plants which start very

early in the spring adopt the rosette form, and these first early

leaves are hidden under the later ones. These small early

leaves are usually delicate and soon wither and drop off, espe-

cially in rapid growth or where the rosette form is adopted.

For these reasons many plants, even in early spring, may show

no traces of the very early stages which they would have shown

if examined a short time before. The surest way to find these

first leaves is to notice where the plant is growing the year

before and mark it in some way so that it may be looked for

the following spring.

As shown, seedlings may vary considerably, and also these

stages, so that to avoid errors there is necessary a knowledge

of the seedling or allied ones, and of ancestral forms. By 4

close study of these localized stages and their variations — within

limits —in many individuals, steps in the phylogeny may be

determined which it would be impossible to fix in any other

way. Many stages may be found in this way that in the accel-

eration of the seedling are crowded out of the record preserved

there.

No. 431.] STAGES IN PLANT GROWTH. 875

Coupled with a knowledge of the seedlings and of ancestral

forms these localized stages may be used to great advantage in

determining not only the phylogeny in a broad way, but the

relations, especially to plants in the same genus or family.

By comparing the character of the stages repeated and the

acceleration with which they are gone through with, something

may be learned of the position of the species in relation to

others within the genus, according to the general principle that

greater acceleration of development means a higher form.

Comparison of seedlings of different species may also deter-

mine this to a considerable extent, but many seedlings of each

species should be compared to get the relation of the variation.

Thus from the seedlings of various species of Aquilegia raised

and compared, a definite relation was noticed in regard to the

relative difference in time of appearance of the same stage in

different species. In this way localized stages, together with

a comparative study of seedlings, may be made to yield profit-

able results in a field where there is otherwise very little data

to depend on.

876 THE AMERICAN NATURALIST. | [Vor. XXXVI.

EXPLANATION OF PLATES.

Leaves are numbered in order of their development, except below the flower, —

there, in the reverse order. c — cotyledons.

Figures drawn from nature by the writer.

PLATE I.

Viola pedata L.

IG. I. First leaf of spring growth of plant one year old. Found in middle of

old woods road-dry soil.

Fics. 2, 3. Second and third leaves of same plant. XI.

S. 4, 5. First leaves of spring growth of plants found at side of same road

in slightly more favorable conditions. Plants small and evidently weak. XI.

Fic. 6. First leaf of slightly stronger plant in same place. I.

a 7-9. First leaves of decidedly stronger individuals in better situa-

xt.

me 10, r1. First leaves of thrifty ub mo more than one year's growth. X I.

Fics. 12-14. Later leaves of same plan

Fic. 15. Most complex leaf noted in vides jm best developed plant. XI.

No. 431.] STAGES LN PLANT GROW TH. 877

PLATE I.

878 THE AMERICAN NATURALIST.

[VoL. XXXVI

PLATE II.

(All natural size.)

Chelidonium majus L

FIG. 1 Seedling showing cotyledons and first and second nepionic leaves.

Fic. 2. Seedling as preceding, but showing also third nepionic

Mass.

preceding.

leaf.

Fic. 3. Leaves of early spring growth. In sandy soil at Arlington Heights,

4th, 6th, and 8th leaves not shown—each like the one immediately

Ranunculus acris L.

FIG. 4. Seedlingshowing cotyledons and Ist, 2d, and 3d nepionic leaves.

IG. 5. Simplest leaf found i

weak plant.

in an

ult plant; first leaf of spring growth of

wo halves of leaf show difference in acceleration of development.

Compare right half with r of Fig. 4; left half with 2 of same figure.

veins

Compare with 2, Fig. 4.

Compare with 2, 3, Fi

Fics. 8,9. Later stages in adult plant and complication of the network of

Fic. 10, Typical adult leaf.

No. 431.] STAGES IN PLANT GROW TH. 879

PLATE II.

880 THE AMERICAN NATURALIST. (VoL. XXXVI.

PLATE III.

(All natural size.)

Viola tricolor L.

Fic. 1. Stages in early spring growth from rootstock of preceding year.

Fic. 2. Stages below flower. Numbered in reverse order of development for

comparison with preceding.

Spirea salicifolia L.

FIG. 3. Growth from lateral bud on plant of previous year. Bract-like leaves

at base not numbered

Tanacetum vuigare L.

Fic. 4. Seedling with cotyledons and typical pair of first a leaves.

Fic. 5. Seedling with second pair of nepionic leaves als

Fics. 6-10. Leaves found near the base of adult aa in spring growth

Compare 6 and 7 with 1 and 2 of Figs. 4 5:

Fic. 11. Simplest leaf to be found ier early spring growth has withered.

6-9.

Very e when compared with Figs.

No. 431.] STAGES IN PLANT GROWTH. 881

PLATE III.

882 THE AMERICAN NATURALIST. [Vot XXXVI. —

No. 431.] STAGES IN PLANT GROWTH. 883

884 THE AMERICAN NATURALIST. . [Vor. XXXVI.

PLATE V.

Aguilegia canadensis L.

Fic. 1. Seedling with first nepionic leaf simple in outline, second normal. X 1.

Fic. 2. Seedling with normal form of first and second nepionic leaves. X I.

Fic. 3. Stages in early spring growth from ground. Compare with seedlings

and Fig. 4. x 4.

FIG. 4. Stages below flower, partly vimus in apparent crowding, but

each leaf in its relative position and shape. Numbered in reverse order for com-

parison with Fig. 3 and seedlings.

No. 431.] STAGES IN PLANT GROWTH. 885

e. 28 V 1

AON Se —

h e

|. `

PLATE V.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XII. THE TREMATODES.

PART I[1— THE ASPIDOCOTYLEA AND THE MALA-

COCOTYLEA, OR DIGENETIC FORMS.

H. S ERATI.

Tne following are the families, subfamilies, and genera of the

suborders Aspidocotylea and Malacocotylea, as arranged by

Braun, Fischoeder, Looss, Liihe, Monticelli, Stiles and Hassall,

and others :

Order. — Trematoda Rud.

Suborder II. Aspidocotylea Mont.

Family. Aspidobothride Burm.

Genera: Macraspis, Stichocotyle, Aspidocotylus, Platyaspis, Cotylaspis,

Cotylogaster, Aspidogaster, Lophotaspis.

Suborder III. Malacocotylea Mont.

Family I. Paramphistomidz Fisch.

Subfamily I. Paramphistominz Fisch.

Genera: Gastrothylax, Paramphistomum, Stephanopharynx.

Subfamily II. Cladorchinz Fisch.

Genera: Gastrodiscus, Homologaster, Diplodiscus, Cladorchis, Chiorchis.

Related genus: Balanorchis.

Family II. Fasciolida Rail.

Subfamily I. Fascioline S. et H.

Genera : Fasciola, Fasciolopsis, Campula.

Related genera: Pleorchis, Paragonimus.

Subfamily II. Omphalometrinz Lss.

Genera: Omphalometra, Cathzemasia.

l Part I appeared in the American Naturalist, vol. xxxiv (August, 1900),

P- 645. The figures illustrating Part II will appear in the continuation, in the

‘December number of the American Naturalist.

887

888 THE AMERICAN NATURALIST. [VOL XXXVI.

Subfamily III. Opisthorchiine Lss.

Genera: Opisthorchis, Holometra, Metorchis.

Related genus: Podocotyle.

Subfamily IV. Telorchiinz Lss.

Genera: Telorchis, Orchidasmus.

Related genus : Deropristis.

Subfamily V. Echinostomine Lss.

Genera: Echinostoma, Stephanochasmus, Dihemistephanus, Stephano-

prora.

Related genus: Rhopalias.

Subfamily VI. Psilostomine Pratt.

Genera: Psilostomum, Crepidastomum, Rhytidodes, Allocreadium, Caly-

codes, Azygia, Halicometra, Cotylotretus

Related genera : Ptychogonimus, Orchipedum.

Subfamily VII. Anisocceline Lss.

Genera: Anisoccelium, Anisogaster.

Subfamily VIII. Centrocestinz Lss.

Genera: Centrocestus, Ascocotyle.

Related genera: Acanthochasmus, Anoiktastoma.

Subfamily IX. Cotylogoniminz Pratt.

Genera: Cotylogonimus, Cryptocotyle.

Subfamily X. Philophthalminz Lss.

Genera: Philophthalmus, Pygorchis.

Subfamily XI. Seung Pratt.

Genera: Opisthioglyphe, Plagiorch

Related genera: Glossidium, ‘eee Pachypsolus.

Subfamily XII. Reniferinz Pratt.

Genera: Styphlodora, Ochetosoma, Renifer, Oistosomum, Astiotrema.

Subfamily XIII. Bunoderinz Pratt.

Genera : Bunodera, Tergestia.

Subfamily XIV. Haplometrinz Pratt.

Genera: Haplometra, Hzmatoleechus, Ostiolum, Macrodera.

Related genera: Opisthogonimus, Asymphylodora.

Subfamily XV. Anaporrhutinz Lss.

Genera: Anaporrhutum, Plesiorchorus.

Related genus: Callodistom

889

No. 431.] MORTH-AMERICAN INVERTEBRATES.

Subfamily XVI. Gorgoderinz Lss.

Genera: Phyllodistomum, Gorgodera.

Subfamily XVII. Microphallinze Ward.

Genera: Microphallus, Levinsenella.

Subfamily XVIII. Brachyceeliine Lss.

Genera: Phaneropsolus, Lecithodendrium, Pycnoporus, Brachyccelium.

Related genera: Cymatocarpus, Brandesia.

Subfamily XIX. Pleurogenetina Lss.

Genera: Prosotocus, Pleurogenes, Gymnophallus, Lepidophyllum.

Subfamily XX. Cephalogoniminz Lss.

Genera: Cephalogonimus, Emoleptalea, Prosthogonimus.

Related genus: Stromylotrema.

Subfamily XXI. Dicroceeliine Lss.

Genera: Dicroceelium, Lyperosomum, Athesmia.

Related genera: Eumegacetes, Anchitrema.

Subfamily XXII. Hemiurinz Lss.

Genera: Hemiurus, Lecithocladium, Pronopyge, Lecithochirium, Lecithas-

ter, Liopyge, Derogenes.

Subfamily XXIII. Syncceliine Lss.

Genera : Progonus, Synceelium, Otiotrema.

Related genus: Halipegus, Accaccelium, Eurycoelum.

Subfamily XXIV. Harmostominz Lss.

Genera: Harmostomum, Ityogonimus, Glaphyrostomium, Scaphiostomum.

Subfamily XXV. Urogoniminz Lss.

Genera: Urorygma, Urogonimus, Urotocus, Urotrema.

Subfamily XXVI. Zoógoninz Odh.

Genera: Zoógonus, Zoógonoides.

Subfamily XXVII. Clinostomine Pratt.

Genera: Clinostomum, Nephrocephalus.

Additional genera: Sperostoma, Microlistrum, Mesotretes, Hapalometra,

istomum.

Family III. Schistosomidae Lss.

Genera : Schistosoma, Bilharziella, Keellikeria.

Family IV. Holostomide Brds.

Subfamily I. Cyathocotylinz Pratt.

Genus: Cyathocotyle.

890 THE AMERICAN NATURALIST. [VoL. XXXVI.

Subfamily II. Diplostominz Brds.

Genera: Diplostomum, Polycotyle.

Subfamily III. Hemistominz Brds.

Genus: Hemistomum.

Subfamily IV. Holostominz Brds.

Genus: Holostomum.

Family V. Gasterostomide Brn.

Genus: Gasterostomum.

Family VI. Didymozoénide Mont.

Genera: Didymozoón, Nematobothrium.

Family VII. Monostomidze Mont.

Subfamily I. Microscaphidiinz Lss.

Genera: Microscaphidium, Deuterobaris.

Subfamily II. Pronocephalinz Lss.

Genera: Pronocephalus, Pleurogonius, Glyphicephalus, Adenogaster, Crico-

cephalus, Pyelosomum.

Related genus: Charaxicephalus.

Subfamily III. Haplorchidinz Lss.

Genera: Haplorchis, Galactosomum.

Additional genera: Opisthotrema, Cycloccelum, Notocotylus, Ogmogaster,

Stictodora, Mesometra, Monostomum.

ORDER. — TREMATODA RUD.

Small parasitic flatworms, with unsegmented, flattened or cylindrical,

unciliated bodies, with usually anterior mouth opening and bifurcate intes-

tine, and without anal opening, which attach themselves to their hosts by

means of suckers, or hooks, or both

KEY TO THE SUBORDERS.

A,. Usually ectoparasitic trematodes living upon the external surface or

the gills, or in the mouth or cloaca of aquatic animals (except genus

Polystoma), to which they attach themselves by means of suckers,

or hooks, or both ; suckers, when present, are usually near either

one or both ends of the body ; when at the anterior end, in most

cases, a single pair is present; when at the posterior end, in most

cases, one or more pairs are present, or, in their place, a sucking

disk . . Heterocotylea Mont.

A, Endoparasitic PRSE viti ‘wes heic to their hosts either

by means of one or more median (unpaired) suckers or a large

ventral sucking disk; hooks never present.

No. 431.] NWORTH-AMERICAN INVERTEBRATES. 891

B, Either a large ventral sucking disk or a mid-ventral row of suck-

ers present; no oral sucker; intestine not bifurcate (except

possibly Aspidocotylus) . . . Aspidocotylea Mont.

B,. Either one or two or, in a few cases, more than two median

suckers present; an oral sucker invariably present (except

Gasterostomum) ; intestine, except in rare cases, bifurcate

Malacocotylea Mont.

KEY TO THE GENERA OF THE SUBORDER ASPIDOCOTYLEA AND THE

FAMILY ASPIDOBOTHRID/E.

4,. A single mid-ventral row of suckers present, extending nearly or quite

the length of the body, which is elongate and cylindrical.

&,. Suckers contiguous and confluent, on a distinct ridge of the body ;

one testis: in gall bladder of Chimera

Macraspis Olss. (Fig. 1) (55, 66)'

b, Suckers not contiguous and not on a ridge ; two testes : in gall pas-

sages of skates (as adults) and in cysts in the end-intestine of

lobsters (as larva) . Stichocotyle Cunn. (Fig. 2) (62, 68, 66)

a, Very large circular or oval sucking disk present.

b,. Sucking disk circular, containing a number of small round sucking

pits; intestine possibly bifurcate: in the intestine of fresh-water

fsh . . . . . . . Aspidocotylus Dies. (Fig. 3) (58, 66)

5, Sucking disk circular or elliptical and containing three or four lon-

gitudinal rows of more or less rectangular depressions separated

from one another by ridges.

¢,. Sucking disk with three rows of depressions.

d,. Sucking disk irregularly circular or elliptical, with crenulate

border and with transversely elongated depressions; one

testis.

e, Number of depressions about twenty-five; marginal

sense organs and eyes absent : in intestine of cheloni-

ans . . . . Platyaspis Mont. (Fig. 4) (58, 66)

e,. Number of depressions about twenty-nine; twenty

marginal sense organs and two eyes present: in

mantle chamber of fresh-water mussels

Cotylaspis Leidy (Fig. 5) (22. 17; 66)

d, Sucking disk elliptical, with very long transversely elongated

median depressions and small round lateral ones; two

testes; marginal sense organs: in the intestine of fish

Cotylogaster Mont. (Fig. 6) (58, 66)

1 The numbers in parentheses following the name of a genus or a species refer

to the publications in the bibliography in which descriptions of it may be

found.

892 THE AMERICAN NATURALIST. [VoL. XXXVI.

cĉ» Sucking disk with four rows of depressions; one testis; mar-

ginal sense organs.

d. Number of depressions seventy-seven; small protrusile

tentacle-ike sacks at the corners of the ridges: in the

stomach of sea turtles . . . . Lophotaspis Lss. -

d, Number of depressions 64-120; no protrusile sacs: in

fresh-water mussels

Aspidogaster v. Baer (Fig. 7) (58, 66)

KEY TO THE FAMILIES, SUBFAMILIES, AND GENERA OF THE

SUBORDER MALACOCOTYLEA.

. But one sucker present (see p. 908).

s Two suckers present, the oral sucker and the acetabulum, together with

a large variously constructed ventral projection or disk of use in

attachment (see p. ).

4,. Two suckers present, the oral sucker and the acetabulum, the latter

either at the posterior end or in the mid-ventral surface; no other

organ of attachment present.

6,. Acetabulum at posterior end, either terminal or subterminal

Family I. Paramphistomide

c,. Lateral pharyngeal pockets not present; cirrus sac absent

Subfamily I. Paramphistomine

4,. Genital pore opens near oral sucker into a large pouch;

acetabulum terminal; body cylindrical: in herbivores

Gastrothylax Poir. (Fig. 8) (11)

4, Genital pouch not present; testes near center of body;

ovary behind them: usually in herbivores.

£, Pharynx with ring-shaped projection near its hinder

end; intestine long and serpentine

Stephanopharynx Fisch. (11)

€» Pharynx without ring-shaped projection ; body conical ;

acetabulum subterminal

Paramphistomum Fisch. (Fig. 9) (1!)

c» Lateral pharyngeal pockets present; cirrus sac present

ubfamily II. Cladorchinz

d,. Ventral surface with numerous papilla.

e,- Acetabulum minute and subterminal; body made up of

two portions, a slender anterior portion and a very

broad, disklike posterior portion, which contains numer-

ous papilla on its concave ventral surface : in intestines

of herbivores ^ Gastrodiscus Leuck. (Fig. 10) (11)

ey. — large and subterminal; entire ventral sur-

ace covered with papillae : in intestine of mammals

Homologaster Poir. (Fig. 11) (11)

No. 431.] MWORTH-AMERICAN INVERTEBRATES. 893

d,. No ventral papilla present.

e€, Testes simple, not lobate or dendritic.

f, Acetabulum terminal with excretory pore in its

center; testes often fused to form a single body:

in the rectum of amphibians

Diplodiscus Dies. (Fig. 12) (41)

Ja- Excretory pore not in acetabulum ; testes in extreme

hinder part of body with ovary in front of one of

them; genital pore with a muscular fold around

it; very small worms: in the stomach of herbi-

n o . . Balanorchis Fisch. (11)

Testes dendritic, near center of body.

J, Intestine long and serpentine; no bulblike widen-

ing of cesophagus; ovary near acetabulum: in

mammals . . . . . Cladorchis Fisch. (11)

Ja Intestine straight; esophagus with bulblike widen-

ing; ovary behind testes, which lie one behind

the other and are each cross shaped

Chiorchis Fisch. (11)

Š. Acetabulum in the mid-ventral surface, but not at posterior end.

orms hermaphroditic . Family II. Fasciolidæ

d,. Ovary between testes ns p. 6):

d@,. Ovary behind testes (see p. 903).

d,» Ovary in front of testes

é,. Uterus does not extend back of testes.

f, Intestinal ceca long and with lateral projections ;

yolk glands very voluminous: usually in the liver

or lungs of mammals Subfamily I. Fascioline

Worms of large size, broad and leaf-like ;

acetabulum near anterior end; intestinal

. coeca, testes, and ovary profusely branched.

h,. Body elongate; anterior end conical and

more or less set off from remainder of

worm . Fasciola L. (Fig. 13) (44,83)

ñ Body circular

Fasciolopsis Lss. (Fig. 14) w

g+ Worms not of large size; cem ranches

intestinal coeca short an

h,. Testes two, one behind ie other: in liver

of Cetacea

Campula Cob. (Fig. 15) (4, =

A, Testes numerous, in two longitudinal rows

intestinal coeca with e two leinchés,

which pass anteriad : in

Pleorchis eg (Fig. 16) (86)

894 THE AMERICAN NATURALIST. [VoL. XXXVI.

f» Intestinal ceca without lateral projections.

4,. Genital pore immediately behind acetabulum (but not on a papilla) ;

body thick and egg-shaped ; intestinal coeca long; testes lobate,

in same transverse plane; yolk glands very large; uterus very

small, near acetabulum: in lungs of mammals

Paragonimus Brn. (Fig. 17) (87)

Zə Genital pore in front or at left side of acetabulum or, if behind,

on a papilla.

h,. Yolk glands dendritic, z.z., diffuse and not compact.

z. Either a ring of spines or two spiny, retractile tentacles at

anterior end.

Jy- Testes in same transverse plane behind acetabulum;

mouth surrounded by spines.

k,. Testes and yolk glands near middle of body, which

is broad Anoiktastoma Stoss. (Fig. 18) (44)

&, Testes and ovary at extreme hinder end of body:

in small intestine of birds and mammals

Subfamily VIII. Centrocestine

Z. Intestinal coca long; «esophagus wanting;

testes and yolk gland large

Centrocestus Lss. (Fig. 19) (44)

4. Intestinal coeca short, not reaching acetabulum ;

esophagus long; a dorsal lip over mouth;

oral sucker prolonged posteriorly into a long,

blind sac Ascocotyle Lss. (Fig. 20) (44)

J» Testes one behind the other, not in same transverse

plane.

£,. Two spiny, retractile tentacles present, one on

either side of mouth: in the opossum

Rhopalias S. et H. (Fig. 21) (86, 7)

&, Tentacles not present; body usually elongate and

cylindrical; mouth surrounded by spines.

4. Testes and ovary close together in extreme

hinder end of body; oral sucker large and

surrounded by a single row of spines: in rep-

tiles and fish

Acanthochasmus Lss. (Fig. 22) (44; 48)

A. Testes and ovary usually not all close together

and not in extreme hinder part of body,

although in posterior half.

m, Intestinal coeca do not reach to end of

body and not of equal length; testes

elongate ; mouth surrounded by a single

row of spines: in intestine of fish

Subfamily VII. Anisocceline

No. 431.] MORTH-AMERICAN INVERTEBRATES. 895

7",. Testes very small and elongate and in center of

body; ovary lobate; uterus does not extend to

end of intestinal coeca

Anisoceelium Lühe (Fig. 23) (52)

7. Testes elliptical, in hinder half of body; ovary

spherical; uterus extends to end of bod

Anisogaster Lss. (Fig. 24) (49)

. Intestinal cceca extend to end of body, or nearly so, and

are of equal length Subfamily V. Echinostomine

7#,- Mouth surrounded by a single row of spines.

o,. Oral spines set in a more or less reniform

ridge and interrupted mid-ventrally

Echinostoma R. (Fig. 25) (44)

0,. Oral spines not set in a ridge or interrupted

ventrally; a number of unusually large

spines just back of mouth: in crocodiles

Stephanoprora Odh. (71)

m,. Mouth surrounded by a double row of spines.

o, Spines entirely surround mouth; long pre-

pharynx present: in fish

Stephanochasmus Lss. (Fig. 26) (44, 48)

o,. No spines on ventral side of mouth, but a little

back of this is a group of short spines

Dihemistephanus Lss. (Fig. 27) (48)

fî». Neither a ring of spines nor tentacles present at anterior end.

J, Testes two in number, one behind the other, often obliquely, but

never in same transverse plane; body usually elongate

k,. Ovary separated from one or both testes by half the length of

the body, the space being filled by the uterus; cirrus sac

. Subfamily IV. Telorchiine

very long: in turtles .

/,. Both testes at extreme hiid end of body

Telorchis Lühe (Fig. 28) (50, 10)

L. One testis at extreme hinder end and one near ovary

Orchidasmus Lss. (Fig. 29) (44. 45)

&,. Ovary not widely separated from testes; uterus usually in front

of ovary.

4. Excretory vesicle Y-shaped, with very long stem, which winds

between the testes ; receptaculum seminis large ; no cirrus

sac: in the liver of mammals, birds, and reptiles

Subfamily III. Opisthorchiinze

m,. Yolk glands do not extend forward of acetabulum ;

uterus confined between intestinal coeca

Opisthorchis R. BL (Fig. 30) (44)

ma Yolk glands partly or wholly in front of acetabulum ;

uterus usually overlapping intestinal coeca.

896

THE AMERICAN NATURALIST. [Vor. XXXVI.

7;. Yolk glands entirely in front of acetabulum;

uterus extends back of ovary to testes

Holometra Lss. (Fig. 31) (44)

7. Yolk glands partly behind acetabulum

Metorchis Lss. (Fig. 32) (44)

/,. Stem of excretory vesicle usually entirely posterior to testes ;

cirrus sac present.

m,. Testes lobate; prepharynx present

Subfamily II. Omphalometrinz

nı (Esophagus absent; intestinal cceca do not extend

into extremity of body; yolk glands meet in

median line: in Ciconia

Omphalometra Lss. (Fig. 33) (44)

7 (Esophagus present.

o. Acetabulum pedunculate; ovary lobate; ute-

rus extends in zigzags to genital pore; yolk

glands meet in median line

Podocotyle Stoss. (51)

9, Ovary spherical; acetabulum sessile; uterus

massed between them; yolk glands laterad of

intestinal cceca

Cathaemasia Lss. (Fig. 34) (44)

m,. Testes not lobate.

#,. Yolk glands extend in front of acetabulum.

o, Anterior end with projecting ridges or with six

papilla.

f, Body subcylindrical; anterior end with

four projecting, radial ridges : in turtles

Calycodes Lss. (Fig. 35) (49)

f. Anterior end with six papilla, two being

ventral and four dorsal: in fish

Crepidastomum Brn. (8)

o, Anterior end without ridges or papille; body

road and flat.

£2, Yolk glands very extensive, overlapping

intestinal coeca; ovary near testes an

not near acetabulum: in fish

Halicometra Odh. (Fig. 104) (7°)

pe Yolk glands not — intestinal

Opisthioglyphe Lss. (Fig. 47) (44)

a,. Yolk glands do not extend in front of acetabulum

Subfamily VI. Facute

o, A pair of earlike papillae present near mo

body elongate.

No. 431.] MORTH-AMERICAN INVERTEBRATES. 897

f, Acetabulum small; testes ovoid: in turtles

Rhytidodes Lss. (Fig. 36) (49)

f; Acetabulum very large; testes reniform:

in Spilotes pullatus, a Brazilian snake

Cotylotretus Odh. (Fig. 130) (71)

0, Anterior papillz absent.

f, Anterior portion of body covered with

largespines and often inflated ; remainder

of body with small spines: in fish

Deropristis Odh. (Fig. 105) (70)

f, No large anterior spines.

g,- CEsophagus long; uterus short; recep-

taculum seminis and testes large: in

fish

Allocreadium Lss. (Fig. 37) (44. 45)

gy. Œsophagus very short or wanting;

receptaculum seminis absent or

small.

r, Yolk glands extend back of

testes.

s, Large suckerlike genital pa-

pila present in front of

acetabulum ; uterus extends

to end of body

Ptychogonimus Lühe (Fig. 38) (51)

5, No geni papilla present;

uterus short, in front of

testes; prepharynx present;

in birds and fish

Psilostomum Lss. (Fig. 39) (44)

7, Yolk glands do not extend back

of testes; ovary far from ace-

tabulum, the space between

filled by the uterus; body elon-

gate: in fish

Azygia Lss. (Fig. 40) (44)

Ja- Testes either two in number, in same transverse plane in hinder end

of body, or many in number.

k. Testes numerous, in median field of body; no pharynx present;

ovary just back of acetabulum: in ducks

rip Brn. (9)

£, Testes two in number, in same semet

sparen IX Colonin

4. Genital pore in e ; yolk glands very voluminous :

in birds . Hoike Lühe (Fig. 41) (50, 10)

898 THE AMERICAN NATURALIST. [Vou. XXXVI.

4. Genital pore on a muscular suckerlike

papilla at posterio-lateral side of acetab-

ulum; œsophagus long; yolk glands

small: in mammals

Cotylogonimus Lühe (Fig. 42) (50, 10)

ña. Yolk glands compact, and lobate or tubular; worms oval

or elliptical in shape; suckers large and oft projectinge

testes in extreme hinder end of body ; cirrus sac long:

Th OUR o v Subfamily X. Philophthalminze

4,. Cirrus sac extends back of acetabulum; yolk glands

tubular, laterad of intestinal coeca: under the eyelid

of birds . . Philophthalmus Lss. (Fig. 43) (44)

2, Cirrus sac does not extend back of acetabulum ; yolk

glands lobate, not laterad of intestinal ceca : in

cloaca of birds . . Pygorchis Lss. (Fig. 44) (44)

é,- Uterus extends back of testes, usually to end of body.

fi. Intestinal cceca long, extending more than half the length of body.

&1- Ovary immediately behind or at side of acetabulum, or when

not near it the intestinal ceca do not extend into hinder

extremity of body.

h,. Genital pore near the acetabulum or somewhere in front

of it, but not at the extreme anterior or posterior end of

body.

4. Mouth surrounded by papilla-like or long spinelike pro-

jections . . Subfamily XII. Bunoderine

Jj; Mouth anomna by six contractile projections ;

no spines present; uterus consists of a descend-

ing and an ascending branch, without spinal

windings : in fish

Bunodera Rail. (Fig. 45) (44, 86)

Ja Mouth surrounded by long spinelike projections ;

uterus winds spirally: in Belone

Tergestia Stoss. (Fig. 46) (44)

Z4. No projections surrounding mouth.

Jy Intestinal cæca usually do not reach the end of

the body (longest in Opisthioglyphe) ; body usu-

ally spiny; excretory canal usually Y-shaped;

cirrus sac present

Subfamily XI. Plagiorchiine

£,. Uterus does not extend back of testes; oesoph-

| ids ovary at side of acetabulum: in

amphibian.

Opistiogyphe ay (Fig. 47) (44)

E, Uterus extends back of

fae (Esophagus ree ey

No. 431.] MORTH-AMERICAN INVERTEBRATES. 899

m,. Yolk glands extend forward of the acetabulum.

7. Testes not in same transverse plane; cirrus sac ex-

tends back of acetabulum : inamphibians, reptiles,

and birds — Plagiorchis Lühe (Fig. 48) (50, 10)

a Testes in same transverse plane.

o. Cirrus sac extends far back of acetabulum:

in turtles Pachypsolus Lss. (Fig. 49) (49)

0, Cirrus sac extends to middle of acetabulum;

body divided into two parts : in crocodiles

Oistosomum Oldh. (71)

m,. Yolk glands do not extend in front of acetabulum;

body elongate, tapering to posterior end, which is

truncated: in fish — Glossidium Lss. (Fig. 50) (44)

4. CEsophagus present.

74. Body covered with minute spines or scales.

7. Testes not in same transverse plane.

o,. One intestinal coecum much longer than the

other; testes elongate ; a single row of

spines around mouth: in fish

Subfamily VII. Anisocceline

f, Testes very small, in center of body ; ovary

lobate; uterus does not extend to end of

intestinal coeca

Anisoceelium Lühe (Fig. 23) (52)

f, Testes in hinder half of body; ovary

sp ean uterus extends to end of body

nisogaster Lss. (Fig. 24) (48)

2,. Intestinal coeca $^ equal length.

Pi Oral un mper m et im

, stem of CAT

cretoty vesicle very long, winding between

testes ; receptaculum seminis minute ; cir-

rus sac long : in aquatic vertebrates

Astiotrema Lss. (Fig. 51) (44. 45)

f, Oral sucker smaller than acetabulum; yolk

glands do not extend in front of latter ;

stem of excretory vesicle short : inreptiles

typhlodora Lss. (Fig. 52) (44, 77)

t Testes in nearly or quite the same transverse plane

and lobate ; oral sucker smaller than acetabulum ;

yolk glands extend about to acetabulum ; recep-

taculum seminis absent or minute ; ; stem of

emm vesicle very long: i

Renifer Pratt it (Fig. 53) (77)

THE AMERICAN NATURALIST. [Vor. XXXVI.

m Body not spiny; cirrus sac very wide in front of

acetabulum ; yolk glands composed of a single row

of follicles on each side: in turtles

Endiotrema Lss. (Fig. 54) (44, 45)

. Intestinal ceca reach into extremity of body; testes usually in

oblique plane or one behind the other.

k,. Large suckerlike genital papilla present in front of acetabulum ;

suckers large; excretory crura join at anterior end

Ptychogonimus Lühe (Fig. 38) (51)

k,. No genital papilla present . Subfamily XIV. Haplometrinz

4. Genital pore just behind pharynx; cirrus sac very long and

rrow.

m. Acetabulum larger than and near oral sucker; body

elongate; testes and ovary in forward half of body;

receptaculum seminis wanting or minute

Macrodera Lss. (44)

m,. Acetabulum smaller than oral sucker and not near it;

uterus very long and deeply colored ; receptaculum

seminis very large; Laurer's canal wanting ; cesoph-

agus wanting or short.

7,. Uterus in long longitudinal folds; ovary beside

acetabulum : in lungs of amphibians

Hæmatolæchus Lss. (Fig. 55) (44)

7, Uterus in transverse folds; ovary behind acetabu-

lum, which is minute; testes very large: in frogs

Ostiolum Pratt (Fig. 56) (77)

4. Genital pore near acetabulum.

m. Genital pore just in front of acetabulum.

7,. (Esophagus present.

f, Yolk glands extensive and dendritic ; body

elongate: in amphibians

Haplometra Lss. (Fig. 57) (44)

f. Yolk glands small on each side of

acetabulum ; testes in same transverse

plane just behind them; ovary to right

of and just behind acetabulum; body

broad: in fish

Callodistomum Odh. (70)

o,. Pharynx absent; body broad; yolk glands very

small lobate bodies just behind acetabulum

Phyllodistomum Brn. (Fig. 63) (8)

7, (Œsophagus absent; body very broad; testes 4

large number of small round bodies

Anaporrhutum Ofen. (Fig. 64) (67)

No. 431.] NORTH-AMERICAN INVERTEBRATES. gol

m,. Genital pore behind and to left of acetabulum;

cirrus sac U-shaped: in snakes

Opisthogonimus Liihe (Fig. 58) (50, 10)

A, Genital pore at the extreme anterior or posterior end of the body.

4. Genital pore in front of or at side of oral sucker

Subfamily XX. Cephalogoniminz

Jı Testes one behind the other; cirrus sac very long.

&,. Excretory vesicle large, with long stem and crura, both

of which throw out side branches; genital pore in

front of oral sucker

Cephalogonimus Poir. (Fig. 59) (44)

&,. Excretory vesicle with long stem, short crura, and

without side branches ; genital pore at right of oral

sucker: in fish Emoleptalea Lss. (Fig. 60) (44, 45)

J». Testes in nearly same transverse plane and lobate; genital

pesi at left of oral sucker ; ovary lobate: in birds

Prosthogonimus Lühe (Fig. 61) (50, 10)

fə Genital pore at extreme hinder end of body; testes and ovary

in median line, the latter just behind acetabulum, the former

just in front of cirrus sac with the uterus between: in bats

Urotrema Brn. (9)

£». Ovary not immediately behind or at side of acetabulum; intestinal

coeca extend to end of body.

Jı. Genital pore in front of acetabulum and median; yolk glands

small, being spherical, lobate, or dendritic bodies near the middle

of the body.

í Yolk glands spherical ; excretory crura join near the pharynx;

acetabulum large . Liopyge Lss. (Fig. 88) (44)

7,. Yolk glands lobate or dendditic:

Jı- Pharynx absent; yolk glands us = lobate : in urinary

bladder of cold-blooded vertebra

Shamil XVI. Gorgoderinz

kı- Body elongate; either one pair of testes or four testes

on one side and five on the other

Gorgodera Lss. (Fig. 62) (44)

k,. Body broad and leaflike; one pair of testes

Phyllodistomum Brn. (Fig. 63) (8)

Ja Pharynx present ; body broad and leaflike; testes either nu-

merous or two in number Subfamily XV. Anaporrhutinz

A,. Testes made up of a large number of small bodies; yolk

glands small and dendritic; uterus not laterad of

intestinal ceca Anaporrhutum Ofen. (Fig. 64) (67)

E, Testes two large, lobate bodies; yolk glands lobate;

uterus extends laterad of intestinal coeca: in turtles

Plesiorchorus Lss. (Fig. 65) (46)

THE AMERICAN NATURALIST. [VoL. XXXVI.

A, Genital pore on edge of body.

Genital pore on right edge of body opposite

pharynx; suckers very large; testes in same

transverse plane in front of acetabulum: in birds

Stromylotrema Lss. (Fig. 66) (44. 45)

i, Genital pore on left edge of body opposite acetab-

ulum ; but one testis present: in fish

Asymphylodora Lss. (Fig. 67) (44)

J» Intestinal coeca very short, extending but little, if any, beyond

the acetabulum, and often not to it; excretory vesicle usually

wide, long, and V- or Y-shaped ; testes usually in same trans-

verse plane.

£y. Intestinal coeca reach about to acetabulum or a little beyond,

and are S longer than the cesophagus ; yolk glands

small soa . . Subfamily XIX. Pleurogenetinæ

h,. Genital pore at left side of body near forward end;

cirrus sac large.

i, Testes small, lobate, just behind yolk glands and

ovary, all being behind acetabulum: in fish

Lepidophyllum Odh. (Fig. 106) (79)

i, Yolk glands and ovary in front of testes: in amphi-

bians and reptiles.

jı- Testes one on each side of acetabulum ; cesoph-

agus long; body elliptic

Pleurogenes Lss. (Fig. 68) (43. 44)

Jo. Testes far in front of acetabulum and ovary;

body oval or round; cesophagus short

Prosotocus Lss. (Fig. 69) (44)

Ža Genital pore in or very near acetabulum

i,. Genital pore in anterior border T acetabulum ;

excretory vesicle Y-shaped and very long

Gymnophallus Odhner (Fig. p" (69)

i, Genital pore not in acetabulum

Jj. Genital pore at left side at acetabulum ; yolk

glands behind testes

Levinsenella S. et H. (Fig. 71) (44 15)

Ja Genital pore in front of acetabulum ; yolk glan

in front of testes: in Corone

Ochetosoma Brn. (9)

£y Intestinal ceca very short, usually falling short of acetabu-

lum; ovary usually near acetabulum; Ld glands small ;

testes usually in same transverse plan

Subfamily XVH: Brachycoeliinz

Ay. Body shi ca and spiny. .—

No. 431.] NORTH-AMERICAN INVERTEBRATES. 903

z,. Acetabulum small, about size of oral sucker ; cirrus

sac large and extending back of acetabulum;

yolk glands dendritic, extending in front of ace-

tabulum : in turtles

Cymatocarpus Lss. (Fig. 72) (44)

łą Acetabulum very large; cirrus sac small, in front

of acetabulum; yolk glands lobate, behind

acetabulum: in amphibians and reptiles

Brachyceelium Duj. (Fig. 73) (44)

A» Body short, oval, usually spiny.

4,. Testes behind and ovary to right of acetabulum.

Jı» Genital pore median, in front of acetabulum;

yolk glands, one on each side of acetabu-

lum: in bats

Pycnoporus Lss. JÁ i 74) (44)

Ja Genital pore at left side of acetabu

Subfamily XVII. Mod aiti

&,. Yolk glands lobate, in hinder part of body ;

vesicula seminalis large, in front of

acetabulum ; nospines: infresh-water fish

Microphallus Ward (Fig. 75) (92, 94)

&,. Intestinal coeca reach a ai: beyond ace-

um; spines presen

kovilendlié 3. et H. its: 71) (44. 15)

f, Testes in front of or on each side of acetab-

ulum.

Genital pore median, between oral sucker and

acetabulum; testes one on each side of and

ovary to left of acetabulum.

£,. Genital pore near acetabulum; aesophagus

long; yolk glands behind acetabulum;

no cirrus sac: in bats and chameleons

iu E n Lss. (Fig. 76) (44)

£, Genital pore n ; esophagus

short ; yolk gasis ri in iot of acetab-

ulum; long cirrus sac present: in birds

and mammals

Phaneropsolus Lss. (Fig. 77) (44)

Ja Genital pore at left side of body; acetabulu

in hinder part of body ; all the genital glands

in front of them; cesophagus short: in frogs

Brandesia Stoss. (Fig. 78) (44)

4,. Ovary behind testes. :

e, Testes in front of acetabulum, in same transverse plane.

904 THE AMERICAN NATURALIST. [VoL. XXXVI.

f, Ovary behind acetabulum; suckers large; genital pore near

pharynx ; no cesophagus present: in cloaca of birds

Eumegacetes Lss. (Fig. 79) (44, 45)

J. Ovary in front of acetabulum ; suckers not large

£j. Genital pore at extreme hinder end: in birds

Urorygma Brn. (9)

£2 Genital pore in front of acetabulum ; intestinal coeca not

reaching acetabulum.

A, Genital pore median.

z,. Cirrus sac long; yolk glands in front of acetabu-

m: in birds and mammals

Phaneropsolus Lss. (Fig. 77) (44)

i,. No cirrus sac; yolk glands behind acetabulum: in

bats and chameleons

Lecithodendrium Lss. (Fig. 76) (44)

h,. Genital pore on left edge of body; cirrus sac long;

genital glands all in front of acetabulum: in am-

phibians and reptiles

Prosotocus Lss. (Fig. 69) (44)

€,. Testes behind acetabulum.

J; Yolk glands dendritic and along the sides of the body.

gı- Testes one behind the other; body elongate; intestinal

ceeca extending to end of body.

h,. Yolk glands consist of a series of branched tubular

follicles on each side.

i. CEsophagus very long ; each intestinal coecum sends

a long projection forward, giving the digestive

tract the form of an H; acetabulum often

pedunculate : in fish

Accacelium Mont. (Fig. 80) (44 59)

i, Acetabulum not pedunculate; digestive tract

not in form of an H; acetabulum in middle o:

body, the anterior half of which is flat, the

posterior half cylindrical

Eurycoelum Brock. (55)

l, Yolk glands not of tubular follicles, but of spheroidal

masses; cesophagus short; suckers near together:

usually in the liver Subfamily XXI. Dicroceeliine

í» But one yolk gland, the left one, present: in liver

of birds . . . Athesmia Lss. (Fig. 81) (44)

z,. A pair of yolk glands present.

Jı- Body flat and leaf-like

coat Duj. (Fig. 82) (44 86)

a» Body cyli

g due Rue Lss. (Fig. 83) (44)

No.431.] MORTH-AMERICAN INVERTEBRATES. 905

Testes in same transverse plane or nearly so; intestinal

ceeca extend to end of body; excretory canal Y-shaped ;

cirrus sac wanting: in bats and chameleons

Anchitrema Lss. (44)

J», Yolk glands compact, or if dendritic, of small extent and not

along the sides of the body.

£j. Genital pore in front of acetabulum.

f. Yolk glands in hinder end of body.

ñ} Distal ends of intestinal ceca joined; distal ends

of excretory crura also joined: in marine fish

Subfamily XXIII. Syncoeliinze

J,. Yolk glands compact, spherical ; testes in same

transverse plane: in the stomach

Progonus Lss. (Fig. 84) (44)

Ja Yolk glands dendritic: on gills of sharks.

&,. Intestinal coeca have side branches; testes

and ovary dentritic ; acetabulum pedun-

culate; anterior portion of body cylin-

drical, posterior flat and folded

Otiotrema Setti (Fig. 85) (44)

£, Intestinal ceca without side branches;

ovary deeply lobate; testes in two longi-

tudinal rows of isolated lobes

ceelium Lss. (Fig. 86) (44)

Distal ends of intestinal coeca not joined; testes

usually in same transverse plane in hinder end

of body; uterus in transverse folds, passing to

anterior end of body: in amphibians

Hal indie Lss. (Fig. 87) (44)

A,. Yolk glands near or towards the middle of the body

(except Derogenes) ; excretory crura usually joining

over the pharynx; small cylindrical or elliptical

forms, in most of which the hinder end of the body

is telescopic : in marine fish

Subfamily XXII. Hemiurine

z,. Hinder end of body not telescopic (no appendix) ;

genital pore near branching of intestine.

J, Testes behind ovary and diagonally behind

each other; yolk glands in front of ovary;

all genital glands spherical

Liopyge Lss. is r 88) ao

29.

Ja Testes in front of ovary, in nearly

transverse plane; yolk poke: bake pe

behind ovary

Derogenes Lühe (Fig. 89) (52, 55)

906 THE AMERICAN NATURALIST. [Vor. XXXVI.

7, Hinder end of body telescopic (with appendix).

Jı Body elongate, finely ringed.

&,. Yolk glands spherical, behind ovary; ap-

pendix either long, medium, or short;

receptaculum seminis large

Hemiurus R. (Fig. 90) (44, 55, 57, 76)

kə Yolk glands composed of three or four long,

involved tubes on each side; appen-

dix long

Lecithocladium Lühe (Fig. 91) (55)

J, Body fusiform or ovate; not ringed.

kı. Yolk glands spherical, near acetabulum ;

excretory crura do not join anteriorly ;

testes in same transverse plane; appen-

dix very short

Pronopyge Lss. (Fig. 92) (44)

k,. Yolk glands deeply lobate or tubular, be-

hind testes.

Z4. Yolk glands apparently joined to form

a single star-shaped body

Lecithaster Liihe (Fig. 93) (55)

l» Yolk glands distinctly separate from

each other, and each composed of

three or four tubular lobes

^ Lecithochirium Lühe (Fig. 94) (55)

£» Genital pore on left edge of body nearly opposite acetabu-

lum ; but one yolk gland present, a small ovoid structure

in center of the body; testes in same transverse plane:

in fish . Subfamily XXVI. Zoógoninz

h,. Testes jum back of E ARU yolk gland just in

front of ovary . . Zoógonus Lss. (Fig. 95) (47)

ñ, Testes one on each side of acetabulum; yolk glan

just behind ovary

Zoógonoides Odh. (Fig. 96) (79)

d, Ovary between the testes, which are often one behind the other.

4,. Genital pore in front of acetabulum; suckers large; cesopha

and intestinal coca long; testes and cirrus sac large; yolk

glands extensive, along entire sides of body :

Spzrostoma R. yos ig. 97) (44, 86)

*,. Genital pore behind acetabulum.

Jı Genital pore at hinder extremity of body; cesophagus absent; -

yolk glands voluminous: in birds and bats

Subfamily XXV. Urogonimine

£i. Suckers very gi testes obliquely behind each other;

_ body broad rogonimus Mont. e ig. 98) (44)

No. 431.] MWORTH-AMERICAN INVERTEBRATES. 907

g> Suckers not large; testes directly behind one another ;

body elongate . . Urotocus Lss. (Fig. 99) (44)

Ja Genital pore not at hinder extremity of body but usually

near testes.

gı: Genital pore between testes, which are very large and

broken up into a large number of distinct parts; no

pharynx, a long cesophagus, and long intestinal coeca

present: in sea turtles

Hapalotrema Lss. (Fig. 100) (44)

g, Testes not broken into small parts.

A. Mouth surrounded by circular ridge; intestine

usually with short lateral projections

Subfamily XXVII. Clinostominz

í. Genital pore just in front of testes; pharynx

absent: in mouth of birds

Clinostomum Leidy (Fig. 101) (21, 6)

i, Genital pore a short distance in front of pos-

terior end; pharynx present: in crocodiles

Noaha Odh. (71)

. Mouth not surrounded by ridge

Subfamily XXIV. Harmostominz

i,. Genital pore in front of anterior testis.

j, Body linguiform ; mouth a slit; ovary and

testes in extreme hinder end : in mammals

Harmostomum Brn. (Fig. 102) (10)

Jo Body very elongate and cylindrical; genital

glands not in extreme hinder end: in

birds . . . Scaphistomum Brn. (9)

Z, Genital pore between the testes.

j, Body very elongate, tapelike; acetabulum

very small; oral sucker large; genital

pore near — end of hinder testis ;

testes far apart: in Talpa

Ityogonimus Lie (Fig. 103) (50, 10)

. Body elongate, linguiform ; genital pore just

behind anterior testis ; mouth circular: in

Mycotheria from Brazi

Glaphyrostomum Brn. (9)

€». Worms diecious . » . Family III. Schistosomide

@,. Hinder portion of bódy pida in both sexes; female shorter than

male: in the blood and liver of birds Bilharziella Lss. (44)

d,. Body cylindrical in both sexes or expanded in female.

e, Female longer than male and filiform, and enclosed in the

gynzcophoric canal of the latter: in the blood of mammals

Schistosoma Wein. (Fig. 107) (44)

908 THE AMERICAN NATURALIST. | [Vor. XXXVI.

€» Male and anterior portion of female filiform; posterior

portion of female swollen and reniform: in the mouth

and gill cleftsof fish, living in cysts, a male and a female

being in a cyst . Keellikeria Cob. (Fig. 108) (86)

4,. Two suckers present, the oral sucker and the ventral acetabulum,

together with a large, a constructed ventral projection, or disk,

of use in attachment š i amily IV. Holostomidz _

6,. Body circular, not made up of two portions; acetabulum usually

„covered by a large disklike structure which possesses a large

cavity extending itself into a longitudinal groove: in intestine of

birds. . Cyathocotyle Mühl. (Fig. 109) (60)

4, Body head acd seid up of two distinct portions, the anterior

portion being usually flattened and containing the acetabulum

and the special organ of attachment.

cı» Lateral edges of anterior portion not bent ventrally ; special

organ of attachment an elongated depression lined with

papilla. ... , («1a Babfamily H. BORET

d,. A row of subdere suckers on hinder part of body: i

intestine of the alligator

Polycotyle Will.-S. (Fig. 110) (95)

d,. No dorsal suckers: in intestine of reptiles and birds

Diplostomum v. Nord. (Fig. 111) (3)

Cy Lateral edges of anterior portion bent ventrally.

d,. Anterior portion trough-shaped ; special organ of attach-

ment an elongated elevation, which may project over the

acetabulum.

é,- In birds and land mammals

Hemistomum Dies. (Fig. 112) (3)

£, In Delphinus .... . . Braunina Heider (12)

. Lateral edges of anterior " natis meet and fuse mid-

ventrally, making this part of the body cup-shaped; spe

cial organ of attachment a conical projection: usually

in birds . . . . . Holostomum Nit. (Fig. 113) (3)

a, But one sucker present and no other organ of attachment.

ó, Mouth in the middle of the ventral body surface; sucker at

anterior end; intestine not bifurcate; genital and excretory

pores at posterior end: in the intestine of fish

Gasterostomum v. Sieb. (Fig. 114) (33)

6,. Mouth at anterior end, sucker oral. 3

€, Worms found in pairs in cysts on the gills, outer surface, or 1n

the mouth of fish; intestine often rudimentary or wanting

Family VI. Didymozoónidze

d,. Anterior portion of body slender and cylindrical, posterior

portion thick and cylindrical, or reniform

Didymozoón Tasch. (Fig. 115) (3)

No. 431.] MWORTH-AMERICAN INVERTEBRATES. 909

d,. Body very elongate (up to a meter in length) and filiform ; intestine

wanting but mouth present . . ematobothrium v. Ben. (3)

€» Worms not found in pairs enclosed in cysts Family VII. Monostomide

d,. Testes directly behind one another in middle of body ; ovary behind

them; body usually elongate, anterior end not set off; genital

pore near anterior end; cirrus sac wanting ; uterus not volumi-

nous ; intestinal coeca do not extend to end of body : in intestine

of reptiles . . SN Microscaphidiinze

é, Uterus canna bete ioteetinal caec

Microscaphidium x (Fig. 116) (44, 45)

£,. Uterus overlaps intestinal coeca

Deuterobaris Lss. (Fig. 117) (44, 45)

Testes either in same transverse plane or obliquely behind one

another; ovary usually in front of testes or between them.

e, Genital pore at hinder end of body, which is broad; pharynx

wanting: in Halicore Opisthotrema Leuck. (Fig. 118) (3)

€» Genital pore in anterior half of body.

` fy Intestinal coeca join at their hinder ends; testes obliquely

behind one another: in water birds

Cycloceelum Brds. (Fig. 1 19) (44)

f. Intestinal coca not thus joined.

g, Anterior end more or less triangular, being set off by a

circular muscular ridge from rest of body ; testes and

ovary often lobate in hinder end of body: in marine

Subfamily II. Pronocephalinz

turtles

E. pes dds with a deep indentation on the

ventral side.

i, Ovary in front of testes, of which two are

resent.

Intestinal coeca without side projections and

with their posterior ends laterad of the

testes ; latter not in same transverse plane

Pronocephalus Lss. (Fig. 120) (44)

Intestinal ceca with side projections and

with their posterior ends mediad of "ege

which are in same transverse plan

k,- Ventral surface with four pe

rows of groups of glands

Adenogaster Lss. (49)

k,. No ventral glands.

Anterior circular ridge very high;

vesicula seminalis very long and

winding skeinlike outside the

cirrus sac

Glyphicephalus Lss. (49)

QIO THE AMERICAN NATURALIST.

, Anterior ridge not noticeably high; vesicula seminalis

not skeinlike Pleurogonius Lss. (Fig. 121) (49)

Ovary behind testes, of which about fifteen are present, arranged

in two longitudinal parallel rows Charaxicephalus Lss. (49)

h,. Anterior circular ridge without ventral indentation ; intestinal caeca

with numerous side projections; testes and ovary deeply lobate

in hinder end of

i, Hinder end of body truncated or concave ; cirrus sac long and

with vesicular seminalis extending the greater part of the

length of the body . Cricocephalus Lss. (Fig. 122) (44)

i. Hinder end of body rounded ; cirrus sac in transverse plane

Pyelosomum Lss. (Fig. 123) (44)

4» Anterior end not set off from rest of body.

h,. Longitudinal ridges or rows of glands on ventral surface; testes in

same transverse plane in hinder part of body.

i. Three rows of glands on ventral surface ; intestinal coeca long

and without projections; ovary between testes: in birds

Notocotylus Dies. (Fig. 124) (44)

i,. About fifteen longitudinal ridges on ventral surface; cirrus sac

very large: in intestine of cetaceans

Ogmogaster Jag. (Fig. 125) (3)

A,. Ventral surface without ridges or rows of glands

i,. Yolk glands laterad of intestinal cceca in hinder portion of body;

testes in oblique plane.

Jı. Ovary between testes ; intestinal coeca very narrow ; uterus

extends to end of body : in birds

Stictodora Lss. (Fig. 126) (44)

Jo Ovary in front of peri intestinal cceca broad ; uterus in

front of ovary : h

Galactosomum Lss. (Fig. 127) (44)

i, Yolk glands on both sides of intestinal coeca in middle and hinder

part of body.

jı But one testis present; ovary in front of it: in Bagrus

Haplorchis Lss. (Fig. 128) (44)

ja Testes in same transverse plane: ovary behind them ; intes-

tinal coeca enclose both ; body circular or oval: in Box

Mesometra Lühe (Fig. 129) (54)

(To be continued )

NOTES AND LITERATURE.

ZOOLOGY.

Beddard's ‘‘ Mammalia.” !— Through the publication of this

excellent manual of the class Mammalia Mr. Beddard has placed the

general student under lasting obligations. The subject could hardly

be more judiciously treated in the limited space of a single con-

venient volume of six hundred pages. In scope and general char-

acter it occupies nearly the same field as Flower and Lydekker's

An Introduction to the Study of Mammals, Living and Extinct, pub-

lished in 1891, and now necessarily in some respects a little out of

date. The two works are, however, naturally constructed on practically

the same plan.

The “ Scheme of Classification ” (pp. ix-xii) recognizes only two

subclasses, the Prototheria (Allotheria, Marsh; Multituberculata,

Cope), including the echidnas and the duckbill, and the Eutheria,

comprising all the other members of the class. These latter are

divided among the following thirteen orders: .

Marsupialia, Creodonta,

Edentata, Rodentia,

Ganodonta, Tillodontia,

Ungulata, Insectivora,

Sirenia, Chiroptera,

Cetacea, Primates.

Carnivora,

The orders are further subdivided into 28 suborders (of which six

are extinct) and 109 families. The number of species is stated to be

3000, but this is obviously far too low an estimate, even for the

existing species. Trouessart, in his Catalogus Mammalium (1897,

1898), listed upward of 7300, and hundreds have since been added.

Doubtless 8000 to 10,000 species would not be too high an estimate

for both fossil and recent.

1 Beddard, F. E. Mammalia. The Cambridge Natural History, vol. x.

London, Macmillan, 1902. 8vo, xii + 605 pp» 285

git

912 THE AMERICAN NATURALIST. [VoL. XXXVI.

In the brief introduction (pp. 1-4) the class Mammalia is very

satisfactorily defined; and in reference to their position as the

“highest of the Vertebrates,” the term “highest,” in comparison

with “ specialization," is considered, and its use in such a connection

is amply defended, since “ specialization " does not necessarily imply

the perfection and complexity of structure that is involved in the

term “highest.” “Most specialized" and “highest,” and “least

specialized” and “lowest” are therefore not the equivalent alter-

natives some writers have considered them to be, and in preferring the

former to the latter as more definite and exact expressions they have

not thereby secured greater accuracy of statement.

An exposition of the structure of mammals occupies pages 5-78,

and deals concisely with their external form, skeleton, skull, teeth,

the organs of digestion, respiration, reproduction, the brain, etc.,

with numerous well-chosen illustrations. The present distribution of

the class and the subject of zoólogical regions is treated less satis-

factorily (pp. 78-89): because the tiger “is as much at home in a

tropical jungle as on the icy plains of northern Asia" is no proof

that *the world cannot be mapped out into areas bounded by

parallels of temperature "; because there are a few wide-ranging

types it does not follow that temperature does not constitute one of

the most powerful controlling influences in the distribution of life

over the globe. The primary regions and their subdivisions here

adopted are, however, those now most generally accepted.

Chapter III (pp. 90-95) considers * The Possible Forerunners of

the Mammalia,” and Chapter IV (pp. 96-104) is entitled “The

Dawn of Mammalian Life "; both subjects are carefully and conserva-

tively treated. The rest of the volume takes up the orders of

mammals in their systematic sequence, from monotremes to primates.

Respecting this part of the work the author says: *I have given, I

hope, adequate treatment from the standpoint of a necessarily

limited treatise to the majority of the more important genera of mam-

mals both living and extinct." Such groups as the Edentata and

Marsupialia are given relatively, and very appropriately, much more

space than is accorded to many of the other orders. On the other

hand, the Rodentia and Chiroptera, with their multitude of forms, are

treated so briefly that one almost regrets that the plan of the work

did not permit of an additional hundred pages or so, in order to

more fully notice these and a few other groups now very briefly

noticed or quite omitted. But the author has given us so much that

is good in this excellent manual that it is rather ungracious to make

No. 431.] NOTES AND LITERATURE. 913

such omissions, doubtless seemingly necessary from the publishers’

standpoint, a matter of criticism.

Notwithstanding the general excellence of Mr. Beddard's Mam-

malia, there are a few slips and misstatements that should hardly

pass without notice. While generally free from typographical errors,

the reference on p. 415 to “Wortman and Malkens” might puzzle

readers unprepared to recognize that * Malkens" is a misprint for

Matthew. As a general criticism, it may be stated that the author is

averse to the adoption of most of the recent changes in nomenclature,

due to the enforcement of the foundation principle of nomenclature,

the rule of priority. A few of them — and among them some of the

most unwelcome— have been adopted, but in t inst the author

is content to say, as in the notorious case of Trichechus for the wal-

ruses: “This family contains but one genus, Trichechus, . . . Or

Odobenus, as the more correct term seems to be.” And so on in

many other instances, temporary convenience, at the cost of a

vitiated nomenclature, being preferred to the slight inconvenience

of introducing to the general public, and particularly to the rising

generation of students, names recently established as the correct

names, and which will therefore, sooner or later, become the familiar

names in the literature of the subject. The conservatism of habit

is thus allowed to retard progress in the attainment of a correct and

stable nomenclature. Incidentally it may be added that the author

recognizes only one species of walrus instead of two ; which is only

to be explained on the ground of inertia or conservatism, and lack of

actual investigation of the point in question; for although Mr. Bed-

dard is almost ultra-conservative in the matter of species and genera,

it is hardly possible to believe that he could say, ** There is but one

species of walrus, though it has been attempted to show that the

Pacific and Eastern forms are different," if he had actually compared

the skulls of the two forms. :

In his comment on the elephants he notes that (p. 22 1) the African

elephant *has been sometimes referred to a distinct genus or sub-

genus, Loxodon,” and later (p. 223) speaks of the Indian elephant as

“ E-uelephas indicus, if the genus Loxodon is to be accept : "; whereas

in reality the Indian elephant would in such case retain the name

Elephas indicus, Euelephas being a synonym of Elephas. J udging from

his treatment of the rhinoceroses (to pass over many other similar

cases), the author’s conception of genera is open to revision, "e

such groups (as well as the higher groups) are supposed to indicate

degree of differentiation regardless of whether they consist of a

914 THE AMERICAN NATURALIST.

single species or many ; and not that diverse types are to be thrown

together, if they happen to have some striking features in common

and are each represented by single species.

The enumeration of Reithrodontomys as a characteristic South

American genus (p. 480) is probably an inadvertence, as is perhaps

the statement that Capromys “ is found only in the islands of Cuba

and Jamaica," whereas there is a species in the Bahamas, another on

Swan Island, coast of Honduras, and still another in Venezuela. It is,

however, surprising to find currency again given, especially in a work

of high scientific standing, to the long since exploded myth of the

" happy-family" relations of the prairie dog, owl, and rattlesnake.

It is also rather surprising to find the statement (p. 518) that the

common mole of the eastern United States (Scalops aguaticus) is

“a creature of largely aquatic habits," — a wholly false inference,

derived apparently from the animal's unfortunate specific name. It

is.also not quite true that the arctic fox (Canis lagopus) “is known by

its bluish summer coat and pure white winter dress as ‘blue fox’ and

‘white fox’ respectively.” This has been so often stated in books of

natural history that Mr. Beddard cannot be blamed for repeating it ;

but the truth is that the blue and white phases are not altogether

seasonal conditions, since at some localities, as the Aleutian and

Pribilof Islands, only blue foxes are found. Indeed the blue foxes

of these islands are regarded by the latest authorities on the group

as specifically distinct from the white fox.

The foregoing criticisms relate for the most part to minor matters,

which do not seriously detract from the value of the work. As would

be expected, Mr. Beddard’s treatment of his subject is thorough, and

fairly represents the latest generally accepted views and results in

Systematic mammalogy, a field in which he has long been an ener-

getic worker and a recognized authority. The publishers have done

well their part in placing before the public a needed work of great

merit in pleasing form. The paper and typography are all that could

be desired, and the illustrations, abundant and well-selected (those

of structural characters are mainly after Flower, Wiederscheim,

De Blainville, Osborn, and others), are printed with admirable clear-

ness. Mr. Dixon’s numerous full-length drawings are of unequal

merit, but most of them are both satisfactorily artistic and effective,

while some are admirable. Í A

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

AND DEATHS.

EDUCATIONAL GIFTS.

Ohio State University, $5000, from Prof. S. W. Robinson, Vassar College ;

10,000, by the will of Adolph Sutro.

APPOINTMENTS.

Jean Bréthes, custodian of insects in the museum at Buenos Aires. —

Dr. Oskar Ebert, custodian of the collections of the Prussian Geological

Survey. — T. W. Galloway, professor of biology in Milliken University,

Decatur, Ill. — Professor Griffon, professor of botany in the N ational

School for Agriculture at Grignon, France. — Dr. Maurice Jaquet, con-

servator of the oceanological museum at Monaco. — Dr. Fridolin Kasser,

professor of botany in the Vienna Agricultural School. — Dr. Adolf

Klautsch, district geologist of the Prussian Geological Survey. — Dr. Benno

Kiihn, geologist of the Prussian Geological Survey.— Dr. R. S. Lillie,

instructor in physiology and histology in the University of Nebraska. —

Dr. Lorenz v. Liburnau, docent for zoólogy in the Vienna Agricultural

School.— Dr. Raymond Pearl, instructor in zoólogy in the University of

Michigan. — Dr. Henry F. Perkins, assistant professor of biology in the

University of Vermont. — A. R. Ruggles, assistant to the state entomologist

of Minnesota. — Dr. Sommerfeldt, docent for mineralogy in the university at

Tübingen. — Dr. Paul Sorauer, docent for plant pathology in the university

at Berlin. — Dr. F. L. Stevens, professor of biology in the Agricultural

College at Raleigh, N.C. — Dr. Robert H. Wolcott, assistant professor of

zoülogy in the University of Nebraska.— Dr. Wilhelm Wolff, district

geologist of the Prussian Geological Survey.— Dr. Oskar Zoth, professor

of physiology in the university at Innsbruck. — Prof. R. Albrecht Zimmer-

mann, botanist of the biological station Amani, in German East Africa.

DEATHS.

Alfred Blavy, entomologist, in Montpellier, France. — Oliver Collett,

student of the Mollusks of Ceylon, at Colombo, June 13, aged 35.— John

Edward Fletcher, British entomologist, at St. Johns, F ebruary 26, aged 66.

a — $5 14

916 THE AMERICAN NATURALIST.

— Petrus Heude, missionary and naturalist, near Shanghai, China, Janu-

ary 3, aged 66. — George Samuel Jenman, government botanist, in George-

town, British Guiana, February 28, aged 56.— Prof. Dr. A. Ludwig G.

Leimbach, editor of the Deutsche botanische Monatsschrift, in June, aged 53.

— William Neale Lockington, a student of Crustacea and fishes, and at

one time a considerable contributor to this journal, at Worthing, Sussex,

England, August 3, aged about 60.— John William May, entomologist, in

London, June 17, aged 78. — Antonio Mori, professor of botany, at Modena,

April 6. — Felix Nawrocki, emeritus professor of physiology, in Warsaw,

June 3, aged 64. — David Pacher, author of a flora of Carnathia, aged 86. —

Dr. P. Plósz, professor of physiology, at Budapest, aged 57. — Dr. Bernard

Schmidt, docent for botany in Tübingen, May 28, aged 35. — Dr. Michael

Tichomiroff, professor of anatomy in the university at Kieff, May 30,

aged 54.— Charles M. Wakefield, student of the insects of New Zealand,

May 11, aged 64.

(Wo. 430 was mailed October 21.)

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THE

AMERICAN NATURALIST

Vor. XXXVI. December, 1902. No. 432.

STUDIES OF GASTROPODA.

AMADEUS W. GRABAU.

MARINE gastropods appear to have reached their acme of

development in the present geologic period. What the

Jurassic and early Cretaceous time was to the cephalopods,

the Tertiary and present periods are to the gastropods.

This is indicated not only by the great number of species,

but also by the fact that so many series have branched out

into bizarre types, in which excessive development of spines

and tubercles suggests that the limit of variation is approached.?

Phylogerontic types are furtl e to be found in the majority

of series, while some groups, such as Strombus, Cyprza, etc., are

represented only by phylogerontic forms in the modern seas.

SO EDITOS

3*9

! The author has for the past five years been engaged in the study of the

Phylogeny of gastropods, particularly the Fuside and some related types.

monograph on Fusus and some of its allies is ready for the press, while another,

dealing especially with Fulgur and its allies, is in preparation. The present paper

is intended as a contribution to the principles of study of the molluscan shell as

applied to Gastropoda, principles first worked out for the Cephalopoda by the

~ late Prof. Alpheus Hyatt and some of his contemporaries, and for the Pelecypoda

_ by Prof. R. T. Jackson.

. 2 See Beecher, Origin and Development of Spines, Amer. Journ. of Sci., Ser. 4,

vol. vi (1898), pp. 329-359; also Studies in Evolution., pP 93 et seg. Scribner, Igol.

O17 :

918 THE AMERICAN NATURALIST. [Vor. XXXVI.

The Protoconch of Gastropods. — The apical whorl of a gas-

tropod shell has come to be generally known as the protoconch,

though this term had been preoccupied for the corresponding

apex of cephalopod shells. The protoconch of gastropods is

essentially different from that of cephalopods in that it is (at

least in the majority of types) twisted into one or more coils

of a spiral nature, whereas that of cephalopods, so far as

known, is a mere swollen bulb. The earliest portion of the

gastropod protoconch agrees essentially in form with that of

the cephalopods (Spirula, Ammonoidea) and pteropods, but

spiral coiling appears very early in the majority of forms.

Thus the protoconch of the Gastropoda may on the whole be

considered as more specialized than that of the Cephalopoda.

It might perhaps be thought desirable to dignify this specialized

type of protoconch by a distinct name, as has been done by

Jackson in the case of the pelecypod protoconch (* prodisso-

conch"). If so, the name “ protorteconch," suggested to me

by the late Professor Hyatt, would be most applicable. The

shell or conch of gastropods may be specifically described by

the term ** torteconch," a name also used for the spirally coiled

shell of cephalopods (Turrilites, Trochoceras).

The characteristics of the protoconch of most living gas-

tropods are very variable, as might perhaps be expected in a

class whose living members are on the whole highly specialized.

Even in Tertiary times this specialization

of the protoconch is noticeable. It is not

always possible to determine the precise

line. of demarcation between protoconch

Fic. 1.— Rhopalithes rugoides and conch, since in a large proportion of

fosoid (bulbow) premens types the two grade into each other imper-

with riblets on last portion ceptibly. In certain types (Buccinum, etc.)

the conch. Eocene. Paris à line and a slight change in the growth

of the shell indicate the place where we

may most reasonably make the separation between protoconch

and conch. In a number of cases (Fusus, Hemifusus, certain

Murexes, etc.) the end of the protoconch is strongly marked by

the existence of a pronounced varix and an abrupt change of

ornamentation (Fig. 5). In the majority of cases, however, no

No. 432.] STUDIES OF GASTROPODA. 919

such definite line of demarcation exists. In general, the proto-

conch coils in the same direction as the conch, though con-

Spicuous exceptions to this rule are found in Pyramidellidae

and other groups.

The early whorls of the protoconch of

gastropods (except such ovoviviparous

types as Cymbium) are smooth rounded

coils of the type found in adult Natica,

and best exemplified by Lunatia heros of

our northern coast. In the majority of

cases the initial whorl is minute, while

the succeeding ones enlarge gradually and

regularly. In some types the initial whorl

is large and swollen (Fusus, Hemifusus,

Fulgur, etc., Figs. 1, 7), when it is gener-

ally more or less elevated and oblique, a early conch whorls.

feature carried to excess in certain volutes.

This type of protoconch has been termed “ bulbous” by Dall.

Again the initial whorl may be compressed so as to produce

a conical form, the sloping sides of which are flat. This type,

observable in so-called Clavilithes from the

British and American Eocene and in some

other genera, may be denominated a “ tro-

choid" type of protoconch.? The flatten-

ing may be confined to the apical whorl

or may be continued over one or more of

the succeeding whorls. The normal nati-

coid type of whorl may continue, with

regular increase, thus giving rise to the

large ** Melo " type of protoconch (Dall) so

"s i prominent in Melo and some other genera.

G. 3. — Protoconch of Syco-

typus canaliculatus. pena When the whorls become depressed so as

| epar um, umbi- practically to lie in one plane, a “ planor-

boid” protoconch is produced (Dall).

Again the naticoid initial whorl may be succeeded by a number

1 Trans. Wagner aiu Znst., vol. iii (1890), p P.

? 'The * trochiform " type of protoconch of ae is more comprehensive, includ-

ing the naticoid type.

920 THE AMERICAN NATURALIST. [Vor. XXXVI.

of whorls which increase slightly or not at all in size, thus giving

a long and slender type of protoconch which Dall has called

*pupiform." This type of protoconch is characteristic of

Clavilithes of the Paris Eocene (Fig. 2), of many Eocene and

recent species of Turbinella, and of other forms.

The trochoid type of protoconch is apparently a case of

specialization, and is confined to a comparatively small number

of gastropods.

The naticoid initial whorl of the protoconch, so far as has

been observed is umbilicated. This is well shown in Fulgur,

Fie. 4. TE canaliculatus in the phylembryonic stage. a h, heart ; mo, mouth;

eye; sh, shell (protoconch); ž, tentacle. x 3

Sycotypus, Fasciolaria, Buccinum, and other types in which

the initial whorls are large enough to be readily examined.

The umbilicus is best shown in young individuals which have

just completed the early protoconch stage (Fig. 3). It is well

marked in Fulgur at the end of the protoconch (phylembryonic)

stage, before the velum is fully developed (Fig. 4). This latter

persists until just before the animal leaves the egg capsule, by

which time the young conch has begun to develop and the

umbilicus is closed. I believe that we are warranted in assum-

ing that the umbilicus exists at some stage in the protoconch

of the majority of gastropods, though it is conceivable that in

extremely accelerated types, even this — apparently the most

No. 432.] STUDIES OF GASTROPODA. 921

primitive condition of a coiled protoconch — is lost. From

the characters of the initial whorls of the protoconch we may

argue that the radicle of the coiled gastropods must have been

a naticoid type with a well-marked umbilicus. Such a type

is found in Straparollina remota Billings, one of the earliest

coiled gastropods of the Etcheminian or Lower Cambrian of

the Atlantic border province of North America. That it is

probably not the most primitive type of gastropod is suggested

by the consideration that the earliest stage (ana-phylembryonic)

of the protoconch is not coiled, but rather cap-shaped like

modern Patella. Such primitive types are found in Lower

Cambrian species which have variously

been referred to Platyceras, Scenella,

or Stenotheca, owing to the want of

sufficient characteristics to define their

exact relations.

Our modern patelliform shells are

probably not primitive types, as shown

by the protoconchs of Acmza and

Crepidula, which are coiled. Tem :

Ornamentation of the Protoconch.— Pe s Midi diu X4. med

In a number of species of gastropods vs icee ep esc A aid

the later whorls of the protoconch are the abruptly beginning, accelerated

ornamented by riblets (Figs. 1, 5), and Sue | developes

more rarely by true revolving lines or

spirals. In others a carina appears on the whorls of the

. protoconch.! These structures normally belong to the conch,

Where their strength and variation constitute the chief features

by which the whorls are characterized. Their occurrence on

the protoconch is due to a backward pushing of the normal

conch characters, until they appear on the final whorls of the

protoconch. This is in conformity with the law of tachygenesis,

or acceleration in development, which is the key to the under- :

standing of the taxonomic values of shell characters in

1 For an account of the characteristics of the protoconchs of a pep of

gastropods, see Baker, F. C., On the Modification T wo r IX

alio Prec PE Ami pm (1890), pp. 66-72; (1894), pp. 223-224; Roch. Acad.

Sez. (1891), pp. 130-132.

922 THE AMERICAN NATURALIST. (Vor. XXXVI.

gastropods. Such accelerated type of protoconch is particularly

characteristic of the Fusidze, though not confined to them. In

some members of this group, Ze, Fusus caloosaensis and its

congeners, the ornamentation of the protoconch begins very

early, showing extreme acceleration. This is also true of Fusus

apicalis Johnson. Neither of these species is a true Fusus.

The classificatory value of the protoconch of gastropods has

not generally been recognized, Dall, so far as I know, being

the only author who has made extensive use of it. It is, how-

ever, apparent that if our classification is to express genetic

relationship, this, the earliest formed portion of the shell,

demands a careful consideration. It is generally conceded

that organisms of a common ancestry are more nearly alike

in their earliest stages of development, differentiation becoming

more and more marked inlater stages. This is to be expected

if the law of recapitulation of ancestral characters is accepted,

for then the earlier stages must more nearly represent the

features of the immediate ancestor in any particular restricted

group. Furthermore, the smaller the taxonomic group, the

more nearly identical must be the earlier stages, since in a

small group the members have not become widely separated

from their common ancestor. Thus, while the initial whorl

of the protoconch does not differ widely, except in size, in the

majority of gastropods, indicating the characters of the more

distal radicle of the class, — ż.e., the naticoid ancestor, — the

whole character of the protoconch must be considered in the

determination of the more immediate relationships. There-

fore, I believe it is not too much to say that the protoconchs

of all the species within a given genus should agree as to their

essential characteristics, and that no species can be considered

congeneric in which the protoconchs show a radical difference.

Thus, as will be shown in a forthcoming memoir on Fusus and

some of its allies, the American Eocene species of Fusus do not

show the typical Fusus protoconch, but some have a protoconch

like that of many species of Pleurotoma and like Levifusus

trabeatus (i.e, Fusus meyeri, etc.). Others again have the pro-

. toconch of Fulgur (Fusus guercollis, etc.), of which group they

are the Eocene ancestors. The species referred to have adult

No. 432.] STUDIES OF GASTROPODA. 923

conch features closely similar to those of Fusus, and the con-

sideration of these alone has caused them to be placed in that

genus. (Compare Fig. 6.) These examples, however, are due

to parallelism rather than close genetic relationships. Great

similarity exists between the protoconchs of Fusus and Hemi-

fusus, thus showing a close genetic relationship between these

two genera, while on the other hand Fasciolaria appears to be

more widely removed.

Septa in the Apex of Gastropods. — Gastropoda in which the

spire of the shell is long, generally develop septa near the apex.

These septa partition off the protoconch and earliest whorls of

the conch, and they may generally be seen

in specimens with broken apices. Turri-

tella, Cerithium and its allies, Fusus, and

in fact turreted shells in general show this

septum. It is generally a curved, more or

less funnel-shaped element, though often

only meniscus-shaped. The septum is invari-

ably convex backward, as might of course be

expected. The apical portion is generally

uniformly curved and lies freely in the cav-

ity of the protoconch or early conch. Septa , 6 — Fuss asper Sow

are sometimes very numerous as observ- Atypical primitive Fusus.

able in longitudinal sections (Triton, etc.), Kowss, Paget. ~ ıb

but generally the number is comparatively smal]. The septa

are entirely imperforate and mark the withdrawal of the apical

portion of the coiled visceral hump from the apex of the shell

and the cutting off of the useless space by a partition wall.

In a number of types the portion of the shell thus divided

off becomes invariably broken away. An interesting type of

structure which belongs here is found in the recent Scaphella

magellanica Sby. and some related forms. In this, according

to Dall,! « the larva is clothed with a cuticular or horny proto-

conch, probably similar in form to that which when shelly

results in the ‘bulbous nucleus.’ Later on, but while still in

1 Dall, W. H. Blake Gastropoda, Bull. Mus. Comp. Zool, vol. xviii (1889),

P- 452; Proc. U.S. Nat. Mus., vol. xii (1890), p. 311, Pl. IX, 6; Trans.

Wagner Free Inst. of Sciences, vol. iii (1890), p. 67, Pl. VI, Fig. 5-

924 THE AMERICAN NATURALIST. [VoL. XXXVI.

the ovicapsule, the deposition of limy matter begins as a slender

cone or elevated point along the line of the axis of the proto-

conch, and as the larva grows the posterior part of the mantle

secretes a shelly dome. Being thus cut off from the horny

protoconch, the latter falls into shreds and is lost. The nucleus

of the larva, still in the ovicapsule, then presents a slightly

irregular dome, with a slender point rising from the apical

To this apical point Dall has applied the term “ calca-

In this case the protoconch proper was horny and decid-

uous, while the first formed septum was calcareous and hence

came to take the place of the protoconch when this was lost. Prob-

ably the types of nuclear whorls named by Dall “ Caricella type "'

and ** Scaphella type " from characteristic genera are secondary

septal deposits of this type, as already suggested by that author.

Characteristics of the Conch.— The simplest type of whorl of

the gastropod conch (torteconch) is round and smooth, showing

only the lines of growth. This type, which is virtually only a

continuation of the primitive protoconch whorls, is seen in

Natica and similar types, and in these the umbilication is gen-

erally retained. Omitting Pleurotomaria !

and its allies, this type of gastropod is

most characteristic of the Paleozoic strata,

the chief modification being the elonga-

tion of the spire, thus producing a lox-

onemoid type. |

One of the earliest modifications of the

smooth shell is the appearance of spirally

revolving, more or less elevated lines and

xvii inet the formation of transverse elevated folds

whorls x 10, showing bulbous OF ribs? (Fig. 7). Our observations are

Tea weenie * still insufficient to settle the question as

to the order of appearance of these two

types of ornamentation. Where both are present the ribs in

most cases are the first to appear to the unaided eye, the spirals

1 The development of Pleurotomariidz and Bellerophontidz, and of Euompha-

lus and other related genera, has been well described by Koken (Mewes Jahrb. für

Mineralogie, Beilage, Bd. vi, pp. 305-483).

2 These ribs must be distinguished from varices, such as occur in Scalaria,

Harpa, and Murex.

No. 432.] STUDIES OF GASTROPODA. 925

not appearing visible until some time after. On the other hand,

fine radiating lines — the precursors of the visible spirals — are

seen in the embryonic hyaline shell of Fulgur, etc., before it

has become opaque by the deposition of secondary calcareous

material (Fig. 8). It is most probable, however, that these

fine thread-like markings are

merely due to a change in the

texture of the hyaline shell,

corresponding in that respect

to the primitive lines of

growth, and are not equivalent

to the elevated spirals, whose

existence is due to a regular,

though slight, plication of the

mantle edge which builds the

shell If we adopt this view as

the most probable one, the ribs, ' rs. iy hyaline shell of Fulgar (Syco-

so far as my observations go, Wid fia h m

must be considered the first

modification of the shell in the ribbed and spiraled forms.

It is, however, by no means true that spirals do not appear until

after the ribs have been formed. In a great many Paleozoic

genera ribs never occur, while spirals are well developed. This

is well exemplified in the Ordovician genera Cyclonema and

Trochonema, the most primitive types of which (ż.e., T. (Gyro-

nema) livatum U. and S., etc.) are umbilicated shells of a round-

whorled or naticoid outline, with spirals more or less strongly

developed. Among the earliest ribbed shells of naticoid form

is Holopea pyrene Billings from the Middle and Upper Trenton

group of Canada and central United States. The ribs are scarcely

anything more than coarse undulations, very like the dying

stages of the ribbed condition in more highly specialized genera

of later geological periods. They have, however, the chief

qualities of the ribs in their most accentuated development,

and must be regarded as such. As nearly as can be deter-

mined from the illustrations given by Ulrich,! the earliest

Stages are free from undulations, thus indicating that, as we

1 Paleontology of Minnesota, vol. ii, Pl. LXXIX, Figs. 13-18.

926 THE AMERICAN NATURALIST. [Vor. XXXVI.

should naturally expect, the ribbed were derived from the

smooth species which precede them immediately in time. The

immature specimens of H. pyrene illustrated by Ulrich show

well the faint beginning of undulations which become pro-

nounced only in the adult.

A somewhat more specialized ribbed shell from another

phylum is found in Nazca nexicosta Phillips of the Devonian.

This in form is truly naticoid, with the ribs sharp, narrow, and

uniform, separated by uniform intervals which are much wider

than the ribs. The earliest stages, as far as can be judged

from the illustration,! are ribless. The whole aspect of the

last whorl of this shell is strikingly like the final whorls of the

protoconch of highly specialized Tertiary and recent gastro-

pods. Natica subcostata Schl. of the Devonian (Stringo-

cephalen Kalk) of Paffrath, Germany, is another example, in

which the costz or ribs have become compound, secondary

shorter ones being intercalated between the longer ones. Natica

costata from the Lower Trias of St. Cassian carries the simple

type of costz into the Mezozoic era, while JV. armata from the

Upper Trias of that region shows the further complication of

revolving spirals which are cancellated by the sharp ribs. In

the later forms generally the umbilicus is closed. A similar

succession of smooth, simple-ribbed, and cancellated-ribbed

shells appears in Jurassic Nerita and Neritopsis, as illustrated

by Hudleston in his monograph of the “ Inferior Oólite Gastro-

poda."? Among the turreted types simple ribs appear in some

of the Devonian Loxonemas, some species of this genus show-

ing the further specialization of cancellating spiral lines.

Similar successions may be found in a great many other series,

and it is perhaps not too much to say that in the majority of the

larger phyletic series, except those highly specialized, the radicle

is a smooth, round-whorled form, succeeded by types in which

the adults are ribbed, and later cancellated, after which pro-

gressive modification may be carried further. The index to the

history of the phylum is in general to be found in the life

history of the individual member under consideration as revealed

1 Whidborne. Devonian Fauna of the South of England, Pa/eontogr. Soc.

Mon., Pl. XIX, Fig. r. 2 Palzontogr. Soc. Mon., Pl. XXVII.

No. 432.] STUDIES OF GASTROPODA. 927

by the succession of characters in the shell whorls, from the

earliest (nepionic) conch stage to the adult (ephebic) stage.

The revolving elevated lines, or “ spirals,” may in general be

considered as primary and post-primary. The primary ones

are the first to appear, and they increase in number by the

exogenous appearance of new ones on the upper and lower

portions of the whorl, outside of those which appeared first.

Secondary spirals appear between the primary ones as these

diverge, owing to the uniform increase in size of the whorls.

Tertiary spirals and spirals of a higher cycle appear in special-

ized forms, but all such forms begin with only primary spirals,

and generally only a few — sometimes even only one of these.

The higher cycles come in later progressively, being most

numerous in the adult stage. This teaches us that in the

primitive type of a series we may expect to find primary spirals

only, even in the adult, and experience shows that these are

characteristic of the earlier members of any series, and that

they generally accompany simple ribs and simple rounded

whorls. (See Figs. 6 and 7.)

The first modification in the form of the whorls in gastropod

shells is generally a change from the primitive rounded outline

to an angular one, which causes the division of the whorl by a

median or submedian keel, or carina, into an upper, more or

less flattened “ shoulder” portion and a lower, generally larger

body portion (Fig. 5). Sometimes more than one carina

arises, in which case all except the lower portion of the whorl

becomes. flattened, the section assuming the appearance of a

portion of a polygon. The angulation, which finally develops

into a carina, and often into a more or less accentuated “ keel,"

is generally due to the strengthening of one of the primary

spirals above the others. When the shell is ribless the angu-

lation becomes a smooth keel. This feature appears early in

the pleurotomarioid shells, in which the angulation is generally

accompanied by an emargination or sinus of greater or less

depth, which occupies the place of the carina. This group of

shells, which appears to be a very heterogeneous one, proba-

bly had its beginning in the Lower Cambrian species of

Raphistoma, of which Æ. attleborensis Shaler and Foerste is

928 THE AMERICAN NATURALIST. . [Vor. XXXVI.

the representative in the Etcheminian limestones of the Atlan-

tic coast region. That this species is derived from the round-

whorled Straparollina remota of the same horizon seems probable

on comparison of the species, and this is in line with the theo-

retic consideration which derives the angular-whorled forms

from more primitive round-whorled species.

When the angulation appears far down on the whorl, so that

the suture of the next whorl touches it, a trochoid shell is pro-

duced, which varies in outline from the gently tapering form of

many Cerithiums, etc., where the basal angle is very obtuse,

through the flat-based Trochus, where the angle is strongly

acute, down to Xenophora with concave base and overhanging

lateral margins, where the acuteness of the angle is extreme.

When the angle is relatively far up on the whorl, so that the

body portion is the largest, we have again two types of spires

produced. When the succeeding whorl joins the earlier one

below the angle, a turreted or terraciform spire is produced, in

which the length and slenderness of the spire depends on the

amount of the embracing of the whorls and the obtuseness of

the shoulder angle. The long graceful spire of Fusus toreuma

and its near allies is a result of a pronounced slope of the

shoulder, together with a very moderate degree of embracing

of the whorls, or what might be called a slender coiling. The

strong contraction of the body of the whorl below the angle

accentuates the slenderness of the spire, producing what is

generally called a depressed suture. In some other species of

Fusus all these features are less accentuated, and the spire as

a result becomes more condensed and relatively stouter, thus

losing some of its gracefulness. In other gastropod shells we

can study all degrees of condensation of the spire owing to the

increased embracing of the whorls, the flattening of the shoulder

even to right angles with the axis of the spire, and the assump-

tion of a cylindrical form by the body of the whorl. When

the later whorls embrace the preceding ones up to the shoulder

angle, a uniformly sloping trochoid spire is produced, the degree

of slope of which depends on the angle which the shoulder

makes with the longitudinal axis of the spire. Thus we may

have every gradation from the long slender spire of certain

No. 432.] STUDIES OF GASTROPODA. 929

pleurotomoid shells to the perfectly flattened or even slightly

sunken spire of certain species of Conus. In this latter genus

it is perceptible that even in the most flattened species the

young whorls form an elevated spire, which varies in intensity

of slope in different individuals. The flattened or sunken

condition appears only in the later stages.

A type of modification of the whorl, which at least in appear-

ance belongs here, is due to the formation of a notch in the

Fic. 9. — Mel long Ant ture individual, showing

growth and the spines. Slightly reduced.

posterior margin of the aperture of the shell, where the body

whorl joins the preceding one. This feature has so far been

found only in the old-age stage of the individual, or in the

adult or even earlier stages of phylogerontic types, t.e., such

as belong to a declining group of gastropods. This posterior

notch may cause a transgression of the final portion of the last

‘whorl on the spire, thus covering up a portion of the preced-

ing whorls. This is generally accompanied by a resorption of

previously formed spines or other ornamentation which would

930 THE AMERICAN NATURALIST. [Vor. XXXVI.

interfere with the comfort of the animal. Accompanying the

formation of this notch is generally a change in the outline

and ornamentation of the last whorls, as will be discussed

further beyond. This type of structure is well shown in Melon-

gena melongena (Fig. 9) and M. patula of the modern trop-

ical seas, and may well be termed a * melongenoid" type of

growth. A striking modification of this type is seen in Strom-

bus, where it appears only in the adult and is accompanied by

the formation of a lip, and by fingers in pteroceroids.! This

may be termed a ** stromboid " type of growth. It appears peri-

odically in Cassis, where it has the value of varices. An extreme

type, the “cyprzoid,” is seen in Cyprza and Ovulum, where

the spire is entirely covered in the adult. Still another type,

characteristic of the Eocene Clavilithes and related genera, is

produced by the flattening of the top of the posterior canal and

the production of a flat sutural shelf, which however is unaccom-

panied by an increase in the amount of embracing of the whorls.

This shelf thus runs like a regular spiral path around the spire

part way, in rare cases almost all the way, up to the apex.

This type deserves to be specified as the *clavilithoid " type

of growth. In extreme cases it spreads out laterally into a

flange, which in certain species is broken up into blunt,

vertically flattened spines (Fig. 18).

One of the most pronounced modifications of the aperture

is the formation of an anterior notch at the point most distant

from the apex. This notch which lodges, and is due to the

formation of an anterior mantle fold which serves the purpose

of a 'siphon, i is in many specialized types drawn out into a long

slender canal, which finds its most perfect development in

Fusus and in certain Murexes, where it is occasionally trans-

formed into an almost closed tube, from the apposition of the

upper borders. This type of modification makes its appear-

ance in the Ordovician genera Subulites and Fusispira, where,

however, the canal is more suggested than actually developed.

Ornamentation of the Whorls. — The simplest types of what

for want of a better term we may call ornamentation of the

1 From unpublished studies on Strombidz, by Miss I. H. Ogilvie, it appears

that the Pterocera has a polyphyletic origin.

No. 432.] STUDIES OF GASTROPODA. 931

whorls are, as already suggested, simple ribs, or spirals, or both.

When the whorls remain round throughout the life of the indi-

vidual no other regular ornamentation is as a rule produced,

except that in the final (gerontic) portion of the last whorl the

ribs, and in some cases the spirals, may disappear, leaving the

shell smooth. If, however, a shoulder is produced through

the formation of an angulation, a new succession of ornamental

types will arise, which generally appears in the same sequence

in widely separated groups of gastropods. The first effect of

the angulation seems to be to concentrate the growth force on

it. The ribs become accentuated on the angle, and correspond-

: ingly fainter away from it. Finally, they disappear above and

below the angle, being represented on the latter, however, as

a series of tubercles. These tubercles, though generally blunt,

may at times assume a somewhat spinous form ; but they appear

to be distinct from true spines, which, so far as observed,

are a feature of later growth. In the cases in which actual

succession has been traced out in series (Fulgur, Fusus, Hemi-

fusus, etc.) it was found that the knobbed or tuberculated stage

is succeeded by a stage in which the tubercles become con-

fluent, producing a pronounced ridge or keel on the angulation.

This after a time disappears, and with it the angulation, the

outline of the whorl becoming rounded again, with only the

spirals persisting. These in some cases may disappear also,

or at least become very faint. After the whorl has returned

to its primitive rounded outline a new type of modifying ele-

ment arises in the form of spines. These begin as notches in

the margin, generally in the spiral zone of the angulation of

the earlier whorls, but sometimes in a second or even a third

zone lower down on the body of the whorl (e.g. Melongena).

They mark a distinct and periodic fold in the outer lobe of

the mantle, which sometimes is accompanied by a temporary

cessation in shell building, as indicated by more or less strongly

marked resting lines in the lines of growth of the shell (Fulgur

carica, etc.). They thus have all the characters of varices of

ounced in Murex, where their multiple devel-

the type so pron

the most

opment and frequent compound character produce

striking feature of the group of shells generally associated

932 THE AMERICAN NATURALIST. (Vor. XXXVI.

under that generic name. When the spine has been fully

developed, the fold in the mantle which caused it may gradu-

ally diminish in size, with the active resumption of the shell-

building process, until it finally disappears. In this case the

spine is gradually closed on the apertural side, thus becoming

symmetrically developed (Fulgur, Hemifusus). In some cases,

however, the fold of the mantle appears to be lost abruptly

when the shell-building process recommences. In such cases

the apertural side of the spine remains open, becoming entirely

external by the flooring over of the emargination produced by

the spine in the shell margin. This type is most characteristic

of the spines forming the varices of Murex.

Through a process in acceleration in development, which will

be more fully discussed below, the spines may be crowded back-

wards, z.e., appear earlier and earlier, thus condensing the pre-

ceding stages more and more until finally some of them are `

dropped out altogether. Thus it will eventually happen that as

in Fulgur carica of the recent fauna the keeled and smooth stages

are dropped and the spines follow immediately upon the tuber-

cles, and to some extent even encroach on these. A gradual

passage from the true tubercles to the true spines is thus pro-

duced, and it becomes practically impossible to determine where

one type ends and the other begins. This feature may be readily

observed in the various species of Hemifusus, in most of which

varieties occur showing all stages from the widely separated

tuberculated and spinous stages to those in which spines and

tubercles grade into each other without allowing a line of

demarcation to be drawn. In trochoid and other shells in

which the whorls embrace up to the angulation, the spines,

if present, are either imbedded in the succeeding whorl and

more or less covered up, or else they are progressively

resorbed as the new whorl increases. In Melo and some

other genera the spines project upwards and are generally

unclosed on the apertural side. This produces the spiral

“corona,” so striking a feature in some shells. Finally, the

remarkable apical character of Vetus (Cymbium) proboscidialts

uld be mentioned. In this a depressed, smooth, central,

apical area occurs, due to a secondary deposit which covered

No. 432.] STUDIES OF GASTROPODA. 933

up all the preceding whorls. This is margined by a sharp

elevated and thin edge.

Varices. — Varices are the periodic rows of spines, or the

reflexed or bent-over lips which mark recurrent stationary

periods in growth in many gastropod shells. Several types may

be recognized, — the spinous, e.g., Murex, the simple reflexed

lip, best shown in Scalaria, and the periodically accentuated

normal rib, perhaps the most abundantly represented type.

The simplest type of the spinous varix is that in which each

stopping place is marked by only one simple spine. Such a

condition, found in Fulgur and a number of other genera, is

generally not considered as falling under this category ; but it is

clear that the multispinous varix is a modification of the uni-

spinous one. This becomes more apparent when we note that

in every multispinous varix one of the spines predominates over

the others to a greater or less extent. On tracing back the spiral

from which this spine arises, which is generally possible, even

though the lower spines are progressively resorbed by the

advancing inner lip of the new whorl, it appears that in every

varix it bears the largest spine. Finally, on tracing back far

enough, only one spine is seen on the whorl, —the primary

spine, which throughout is recognizable by its predominant

size. In Murex brassica Lam. of the west Mexican coast we

have a type in which only a few additional small spines occur

besides the primary large spine. This is but slightly advanced

beyond the state found in Fulgur. In M. bicolor the shell con-

tinues to grow somewhat after the chief varix is formed, the

principal spine alone remaining unclosed, so that the lip at the

resting stage differs little from that of Fulgur. The multispi-

nous type of varix, with simple spines, is best illustrated in the

beautiful Murer tenuispira Lam., the most striking in form of

all the Murexes. Each spine arises from a spiral line, of which

it forms the apertural prolongation. The shorter spines arise

from the secondary spirals, and on the canal they are bent at

about 120? forward from the primary ones, thus lying parallel

to the primary series of the next but one varix preceding.

Only in the neighborhood of the large spine do the tertiary and

even later cycles of spirals terminate in small spines.

934 THE AMERICAN NATURALIST. [Vor. XXXVI.

If the spine from a primary spiral does not increase in length

rapidly, but increases in width, the secondary and later spirals,

instead of developing independent spines, encroach upon the

primary one and modify it by the production of lateral crenula-

tions, which, as the spine grows, diverge more and more and

become more and more pronounced, until the striking multi-

lobed character of the spine of M. pa/ma-rose@ (Lam.) and other

highly ornate species is attained. If the principal spine of such

a species is traced backward through preceding varices (which

can generally be done, since resorption does not reach up to it),

it will be found to be less and less complex in earlier and earlier

varices, and ultimately may perhaps — in a very perfect speci-

men — be resolved into a single spine. (See Fig. 10.) In the

growth of the spine itself, from its beginning on the final or

first varix (counting backward) to its full development, as

already noted, it progressively increases in complexity. A stage

somewhat earlier than the adult stage in this spine shows the

same degree of complexity as the adult. of the representative

of this spine in the next preceding or second varix. A still

earlier stage in the principal spine of the first varix corresponds

to the stage just preceding the adult in the principal spine of

the second varix, and to the adult stage in the corresponding

spine of the next earlier or third varix, and so on. Inthe same

manner, though less perfectly developed in most cases, the

spine next below (anterior of) the principal spine on the final

or first varix has the characters of the adult of the principal

spine in the next preceding or some earlier varix, and the third

spine has the adult characters of the principal one in a still

earlier varix. The last spine of the final varix — presumably

the last formed one — has the characters of the principal spines

in one of the earliest varices, when the shell was still very

young (Fig. 10, where 4-7o corresponds to K-z, and -ro to

L-r). Thus each spine passes through a succession of stages,

and in its reappearance in a new varix it has made marked

advances. The life history, therefore, of a single spine may be

read by noting the characters of all the spines of that varix

progressively from the smallest to the largest. This also indi-

cates in general the life history of the group to which the

No. 432.] STUDIES OF GASTROPODA. 935

Species under consideration belongs. Thus in an ancestor of

M. palma-rose we might expect that the principal spine on the

last formed varix of the adult shell would have the characters

of this same spine in an earlier varix in pa/ma-ros@ or those of

a more anterior spine in the adult varix of that species. Such

correspondence of characters in localized areas (e.g., a single

varix), with the changes characteristic of the life history of the

group, has been called by Jackson the formation of localized

234 nE

Bee, ag:

fu Kits

An LIT Ve r 123 4

7273 3 92

423 ZO

y 8 4 rot

Fic. ro. — Dia of the varices of Murex brevifrons Lam., from the West Indies.

last or most recent varix, Z the earliest recognizable one. The corresponding spines oe

numbered alike. The dotted line indi hat th ining p

Stages in development, and he has found this phenomenon in

a great many types of animals and plants.!

One of the commonest types of varix is that due to accentua-

tion of normal ribs. Even in the spinous Murexes the bases of

the spines are merged in such an accentuated rib, which recurs

with greater or less frequency, though with regular periodicity,

in the various species. This accentuated rib, whether spinous

or not, appears to mark the periodic culmination of constructive

vitality, after the expenditure of which a rest is necessary

before building again commences.

The varices of Harpa are due to a periodic reflection of a

smooth lip, which has the appearance of a margin rolled back.

1 Jackson, R. T. Localized Stages in Development in Plants and Animals,

Mem. Boston Soc. Nat. Hist., vol. v, No. 4.

936 THE AMERICAN NATURALIST. [Vor. XXXVI.

In Scalaria the varices are simple labial flanges, which stand

out as sharp ridges, with the appearance of ribs. In He/ix

albolabris and other types the varix is formed only in the

adult. In all types with successive varices it may be observed

that the varices of the same longitudinal

zone do not form a line parallel to the

axis of the shell but that this line runs

spirally backward, in the direction opposite

to that in which the whorls coil (Fig. 12).

This is due to the fact that there is a

progressive diminution in the proportional

amount of building which was performed

between the resting stages. If theamount

of building were proportional to the size

aide EU of the shell the varices would run in

with " proportional Straight, constantly diverging lines (Fig.

TET II) instead of spiral ones. If the propor-

tional increase were too great for the size of the shell, and

progressively increasing, the lines of varices formed by the

corresponding ones of all the whorls would pass spirally forward,

l.e., iv the direction of coiling of the shell Neither this nor

the case of increase proportional to the

size of the shell has been noted, and it

hence appears that we have here an

illustration of Minot's law of senescence,

the animal progressively growing old from

the beginning. This feature is best shown

in Scalaria, where in large specimens an

additional feature, first pointed out to the

writer by Prof. R. T. Jackson, is also seen.

This is the more rapid shortening of the

building stages, in the old age of the indi- Fic. 12.— Diagram

vidual, so that the varices of the last— — teristic retard ed growth

old age — whorl are not continuous with

those of the preceding ones but fall between them, so that

sometimes a varix may be the whole length of a period

behind what it should be (Fig. 13). In phylogerontic species

of Scalaria this feature appears in the adult or earlier whorls,

No. 432.] STUDIES OF GASTROPODA. 937

which whorls will then have a larger number of varices than

those preceding them.

Other Ornamental Fezatures.—Dall! has demonstrated the

purely mechanical origin of the columellar plication in gastro-

pod shells to be due to the sliding inward and outward over the

columellar surface of a mantle lobe folded by being crowded

into too small a space. In Cypraa these plications are most

numerous and developed in the adult on both sides of the aper-

ture. The intensity of the plications varies with the nearness

or remoteness of the muscle of fixation. In a like manner the

liræ of the outer lip may be explained. |

We must, however, be careful not to

mistake the strong spirals, which some-

times appear through the thin covering

of the inner lip, for plications, or the

interspiral spaces appearing prominently

in the thin outer lip, for lire. Countess

von Linden has discussed at length the

development of the color pattern in gas-

tropods,? and the reader is referred to

her paper.

Individual Old Age, and PAylogerontic

Characters in Gastropods. — Gerontism,

or old age, is marked in its earlier stages Fric. 13. — Diagram illustrating

. . gerontic characters in the var-

in gastropod shells by the disappear- ices of the last whorl.

ance of features characterizing the adult.

Such disappearance of features is generally in the reverse

order in which they were formed. In the fusoid types the

shoulder and the spines are the first to be lost, if they have

been developed; or if there were only spinous tubercles, as

in the Fusidz, these disappear together with the angulation,

and the gerontic lip becomes smooth except for the spirals.

The normal outline is next more or less modified by an attempt

at straightening out the whorl and making it more cylindrical,

1 Loc. cit., p. 58. |

2 von Linden, Maria. Die Entwicklung der Skulptur und der Zeichnung bei

den Gehiuseschnecken des Meers, Zeitschr. für wissensch. Zoologie, Bd. lxi,

pp. 261-317. 1 pl.

938 THE AMERICAN NATURALIST. [Vow. XXXVI.

producing a consequent obscurity of the canal. The last stage

in gerontism is observed in the loss of the power to coil, which

is at first expressed in the separation of the inner lip from the

columella ; and if any further coiling occurs, in the loose appo-

sition of the whorls and the consequent formation of an

umbilicus. These features. never go very far in the normal

gerontic individual, for death intervenes and ends the process.

Among the highly specialized types of gastropods uncoiled

phylogerontic species are relatively uncommon. Perhaps the

most striking example in a highly specialized group in which

such a want of regular coiling does exist in the adult is Ver-

metus. This, as is well’ known, has all the characters of a

Turritella in its young stages, showing its derivation to be

from that group. In the adult, however, it loses the power to

form a regular spiral, turning and twisting in all directions and

completely uncoiling at times, so as to make these portions of

the shell almost indistinguishable from worm tubes such as

Serpula. In primitive gastropods non-coiling, through the loss

of the power to coil, appears to be the normal expression

of old age, or gerontism, since in most cases the coil is the

only feature which can be modified.

Shells with the final portion of the coil unrolled are well

represented in the Ordovician. They even occur in the Cam-

brian, as indicated by Platyceras primevum Billings from

the Lower Cambrian. From what is known of this little shell

it appears that it represents a strongly umbilicated, low-spired

naticoid type, with a portion of the last whorl free. If, as

appears to be the case, the early whorls are normally coiled,

and only the later coil loosely, this little shell represents

the first phylogerontic type appearing side by side with the

naticoid radicle of the entire group of gastropodous mollusks.

Leaving out the loose coiling Eccyliomphalus and Eccyliop-

terus as of questionable affinities, the first well-pronounced

example of a phylogerontic type with the final whorls not coiled

is found in the Ordovician pleurotomarioid genus Lophospira.

This genus, which comprises mainly normally coiled umbili-

cated forms, contains a few species which have become phylo-

gerontic and show various degrees of non-coiling (Fig. 14).

No. 432.] STUDIES OF GASTROPODA. 939

Trochonema vagrans Ulrich and Scofield (Fig. 15), from the

Stones River group, represents a phylogerontic type among

the carinated naticoid gastropods, while Dyeria costata (James),

from the Upper Lorraine of Cincinnati,

represents a phylogerontic type of the

group of simple naticoid shells with

depressed spire. In the Silurian, P/atyo-

stoma (Diaphorostoma) niagarensis fre-

quently shows individual gerontism by a

loose outer lip. Platyceras miagarense

represents a phylogerontic type, possibly

derived from the former. In the Devonian

the phylogerontic non-coiling Platycera

abound. We find all degrees of coiling,

from the close-coiled non-umbilicate Dia- "9 '* RA gore

phorostoma, which appears with slight showin Paley of Mi

vol. Fig.

modifications throughout most of the xesota, pl. Ixxii,

26.)

series, to the straight * Orthonychia,”

which appears as a terminal member in most groups. Very

often a number of species of “ Platyceras” can be traced to a

species of Diaphorostoma or Strophostylus occurring with them,

the gradations being perfect. From such evidence it appears

that the numerous species classed together as Platyceras must

be split up into groups, each of which

has been derived from a close-coiled

ancestral species, probably within the

same geological horizon. If so, the

name Platyceras becomes meaningless

for generic purposes. The great diffi-

culty which besets the proper breaking

up of what appears most certainly to be

PN Sec Twice eeut A OOD of polyphyletic origin lies in the

U. and S., illustrating geron

final whorl. (After Ulrich, 7. small number of ornamental characters

læontology of Minnesota, vol. which can be made use of in tracing out

ii, pl. ioiii. Fig. 13.) i

relationships. In many Platycera spines

appear, but these cannot be regarded as clues to affinity, since

spinous types may and do arise in any group. What seems to

be the right method of procedure was made use of in the

940 THE AMERICAN NATURALIST. [VoL. XXXVI.

determination of the genetic relation of PJatyceras arkonense

Shimer and Grabau, from the Hamilton of Ontario.

Natica neriteformis, from the European Muschelkalk (Queen-

stedt), as far as can be judged from the internal molds which

are alone represented, is a phylogerontic member of a simple

naticoid group of shells, Fissurella, Acmza, and other patel-

loid types, so common in the Mezozoic and modern sea, have,

as already noted, a coiled protoconch. They therefore repre-

sent phylogerontic types, in which the power of coiling has

been lost after the completion of the protoconch stage. In

Crepidula the power to coil is still retained in a slight degree.

In the majority of specialized gastropods phylogerontism is

expressed, not in the non-coiling of the last portion of the

spire, but in its expansion and wrapping about the earlier

whorls so that these become more or less concealed. This

is generally accompanied by the loss of all ornamental char-

acters except, in some cases, the coloration. Sometimes it

happens, however, that spines arise independently on this

portion of the shell. This is most marked in Melongena and

can be readily explained by the peculiar manner in which the

shell develops. |

In the young stage every typical Melongena has the char-

acters of Hemifusus. In accelerated species of this genus

spines follow the tubercles, the two grading into each other as

in Fulgur carica. In other species, however, notably Z. colosseus

Lam. and H. (Melongena) pugilinus, a keeled or even smoothly

rounded stage intervenes between the tubercled and spinous

stages. (See ante.) This enables us to understand Melongena.?

The two typical species of this genus appear to have been

derived from the same ancestral species, which coexists with

1Shimer, H. W., and Grabau, A. W. The Hamilton Group of Thedford,

Ontario, Bull. Geol. Soc. Am., vol. xiii, p. 176.

? Only the two typical species, Melongena melongena and M. patula, are referred

to here. Most of the other species generally referred to this genus belong else-

. where. Such species as M. morio and M. Pugilina are transitional from typical

Hemifusus, eg., Æ. colosseus Lam., being produced chiefly by a condensation

of the typical Hemifusus characters. They are generally classed with Melongena,

. but have not the typical phylogerontic growth of that genus. ‘They will be

referred to under both names.

No. 432.] STUDIES OF GASTROPODA. 941

them, — namely, Hemifusus (Melongena) morio. Representa-

tives of this species occur in both west African and West

Indian waters. From the west African type appears to have

been derived the Mazatlan species Melongena patula, while M.

melongena, at home in West Indian waters, was derived from

the West Indian representative of H. (M.) morio. In both

species the phylogerontic melongenoid form of whorl appears

immediately after the tubercled and before the spinous stage,

this latter coming in somewhat later. Thus, while the mode of

coiling has become senile, the formation of the spines indicates

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Fic, 16. — Diagram a Hyatt’s Law of Serm in Development.

or description see tex

the persistence of one portion of the vigor of the adult.

Such form of acceleration may be called partial or incomplete,

differing in that respect from Fulgur, in which the acceletation

is complete. A reason for this may perhaps be found in

the following consideration: Fulgur is an accelerated type.

in the present fauna. The species in which a separation

existed between the tubercles and spines all lived in Miocene

and Pliocene times, and hence in the modern descendant (F.

carica) spines have become firmly established. We may further

suppose that the power to form spines has persisted so long

that in the adult it is on the wane; and hence if any phylo-

gerontic types of Fulgur should arise, the spines would first

disappear and probably would never be formed again. In

Hemifusus, on the other hand, the spine-forming power has

942 THE AMERICAN NATURALIST. [VOL XXXVI.

apparently been but recently acquired, judging from the fre-

quent lateness in appearance of spines in individuals. Thus

there would be more vitality or spine-forming power left, which

would be able to overcome any weakening effect of senility

appearing in other features of the group. Hence, though

a gerontic form of coiling is assumed by the whorls of Melon-

gena, the power to form spines is still retained, and these

appear after an interval in the senile portion of the whorl.

Lllustrations of the Law of Ti achygenesis and of Parallelism

among Gastropoda. — Hyatt’s Law of Tachygenesis, or accelera-

tion in development, teaches us that the adult characters of

a species are inherited at an earlier stage in the members of a

succeeding generation. This of course implies a condensation

of preceding characters, which in some cases may even be

dropped entirely. The diagram on page 941 (Fig. 16) will give

a clearer representation of the working of this law.

Let each of the lines A to / represent the life history of an

individual gastropod in a phylogenetic series, beginning with A.

Let the life history be divided into stages numbered from 1 up,

each stage being characterized by some definite feature in the

shell.

No. 1 is the protoconch stage, which persists throughout ;

No. 2, the smooth, round, whorled naticoid stage; No. 3, the

primitive ribbed and spiraled stage; No. 4, the angular stage,

with formation of nodes at the crossing of the ribs; No. 5,

the keeled stage ; No. 6, the second round-whorled stage; No. 7,

the spinous stage; No. 8, the spineless smooth stage.

It becomes apparent that as new characters appear, the older

ones persist for a shorter and shorter time. This is neces-

sary, for otherwise each succeeding species must be larger

to accommodate the new characters. This is acceleration by

condensation. A character newly acquired in one member

of the series appears earlier and earlier in the succeeding

members of that series, thus changing from an adult to an

infantile character. Some characters are less resistant than

others, and these will be more quickly condensed. Such is

the case with those of stages 5 and 6, the keel and the second

Tround-whorled stage. The nodes are very persistent, and the

No. 432.] STUDIES OF GASTROPODA. 943

spines (of stage 7) are.very vigorous. Thus it will happen,

as in H, that stages 5 and 6 are dropped out altogether,

stage 7, the spinous one, following upon stage 4, the tubercled

Fics. 17 and 17 a. — Cyr£ulus serotinus Hinds. Recent. A phylogerontic type of

the Fusus series. Natural size.

Fic. 18. — Clavilithes chamberlaini Johnson and Grabau. Eocene. Texas. Two stages; the

adult showing a serrated flange and clavilithoid form. (Proc. Phil. Acad. Nat. Sci.,

mber, 19or.)

one, and even encroaching upon it. This is acceleration by

elimination (example: Fulgur carica). Finally, even the vigorous

Spines disappear, and the final smooth gerontic stage is formed.

944 THE AMERICAN NATURALIST. [VoL. XXXVI.

Parallelism. — There are perhaps few if any classes of inver-

tebrates which equal that of the Gastropoda in illustrating the

law of parallelism of development. In the Gastropoda similar

features constantly arise independently in entirely distinct

series ; and since these features are generally used as a basis of

classification, it follows that our present system of classification

is in great need of revision. If we represent the natural sys-

tem of classification by the customary tree and its branches,

and then pass a more or less warped plane through the upper

part of this tree, intersecting each branch at the same relative

stage of development, we obtain a number of scattered points

where the plane and the branches intersect. If we consider

these points species, and group all the neighboring and perhaps

Some distant points into genera and so on, we will get very

much what we have to-day in the classification of marine

gastropods. In point of fact, we have only one episode in

each of the large number of distinct phyletic series, and what

we mistook for relationship is only a resemblance in a parallel

developing series which has been cut at the same stage in

development. Thus it is clear why the customary mistake of

placing the recent Cyrtulus serotinus (Figs. 17, 17 2) with its

parallel, the Eocene Clavilithes, is so generally made. Both are

phylogerontic members of entirely distinct phyletic series, but

they have reached the same stage in development. Again

Fusus, known only in the Eocene of Europe, and in this country

only from the Miocene on, has its close parallel in Pseudofusus

and other genera in the American Eocene. Here the same

type of form is developed, — a very simple matter, — but, as

shown by the life history, the two types are widely apart

genetically. Again, all gastropods with stromboid lips are placed

with Strombus, a proceeding which has no warrant from a

phyletic point of view. The common proceeding of classing all

loose coiled or non-coiled Paleozoic gastropods under Platyceras

has likewise no warrant from a phyletic view point, though it

must be confessed that this is a convenient method when we

do not know what else to call them. Thus, without multiply-

ing examples, we may sum up the result of our studies in the words

of Hyatt's Law of Morphogenesis: «A natural classification

No. 432.] STUDIES OF GASTROPODA. 945

may be made by means of a system of analysis in which

the individual is the unit of comparison, because its life in all

its phases, morphological and physiological, healthy or patho-

logical, embryo, larva, adolescent, adult, and old (ontogeny),

correlates with the morphological and physiological history

of the group to which it belongs (phylogeny)."

PALZEONTOLOGICAL LABORATORY,

COLUMBIA UNIVERSITY.

THE OCCURRENCE OF FORMICA CINEREA

MAYR AND FORMICA RUFIBARBIS

FABRICIUS IN AMERICA.!

WILLIAM MORTON WHEELER.

IN a valuable paper published in 1886? Dr. Gustav Mayr

recorded both Formica cinerea and F. rufibarbis, two well-

known European ants, as occurring in the United States.

The former was cited from California and New Mexico, the

latter from Colorado, Nebraska, California, and Montana.

Seven years later, when Professor Emery published his impor-

tant revision of the North American Formicidz,? he was so

doubtful of the occurrence of these forms in the United States

that he did not include them in his synoptic table. Concern-

ing the former species he wrote : “ F. cinerea does not occur in

North America; the form identified as such by Mayr will

be described below as F. pilicornis n. sp." In regard to F. rufi-

barbis his statements are less positive: ‘For the present I

am inclined to doubt whether forms belonging to the true

Jusco-rufibarbis series are actually indigenous to America. I

am really unable to distinguish from rather pale and very pilose

European fusca (fusco-rufibarbis) only three workers which

were received from Colorado through Mr. Pergande. The

precise locality of these specimens is not given."

More than a year ago Dr. Harold Heath of the Leland Stan-

ford University sent me numerous specimens both of F. cinerea

and F. rufibarbis which he had kindly collected for me near

San Jose, California. The F. rufibarbis was compared with

1 Contributions from the inii Laboratory of the University of Texas, No. 38.

2 Die Formiciden der Verei Staaten von Nordamerika,, Verhand. Zool.-

Bot. Ges. Wien., Bd. xxxvi (1886), pp. 419-464.

8 Beiträge zur Kenntniss der nordamerikanischen Ameisenfauna, Zool. Jahrb.,

Abth. f. gale ric - oo pp. 633-682, Taf. XXII, and Bd. viii (1894),

PP- 257-360, T:

947

948 THE AMERICAN NATURALIST. (VoL. XXXVI.

European specimens by Professor Emery, who reported as

follows (zz Zferis): ‘It is the true European form, differing

from our common type only in the total absence of erect hairs

on the thorax. In this respect it approaches var. glauca

Kusgky from Oriental Russia." It is possible that Dr. Mayr

may have seen specimens of this same ant, but it is more probable

that he had specimens of zeo-rufibarbis Emery, a common form

throughout the more western and southwestern states as far

.. as the Pacific coast. The specimens of cinerea received from

Dr. Heath were compared with European cinerea given me by

Professor Emery, Professor Forel, and Dr. Mayr. The speci-

mens from Professor Emery were collected near Bologna, Italy.

They are decidedly smaller and have a darker ground color

than the Californian specimens. These, however, agree very

closely in their larger size and somewhat reddish coloration

with the Austrian and Swiss specimens from Dr. Mayr and

Professor Forel. I believe, therefore, that there can be no

doubt concerning the occurrence in California of two species

of Formica almost or quite identical with the European cinerea

and rufibarbis. It is not so easy to decide whether one or both

of these species are imported or indigenous to the American con-

tinent, but I know of no cogent reasons for accepting the former

alternative. Certainly the occurrence of these species on the

Pacific coast and their apparent absence from the eastern states

of our Union are extremely suggestive in connection with

the like geographical distribution of many other Pacific coast

arthropods (Astacus, e.g., many Diptera, and other insects)

which are known to be more closely related to European genera

and species than to those of the Atlantic states.!

After accustoming myself to view the distribution of the

two species of Formica in this light, I was much surprised

during August of the past summer to find cinerea very abun-

dant in the vicinity of Rockford, Winnebago County, Ill. For

several weeks of three successive summers I had collected very

diligently without finding any such species in this locality. It

1 See,zg., Osten Sacken's “ Western Diptera.” Cases in point are also the

Californian ants of the subgenus Messor, and Myrmica mutica Emery, which is

hardly more than a subspecies of the European M. rubida.

No. 432.] FORMICA CINEREA MAYR. 949

occurs most commonly, however, in such peculiar situations

that there is little difficulty in understanding why it has been

so long overlooked. The formicaries are so large and populous

that it can hardly be regarded as an imported species unless

it resembles some of the European weeds which have found

the American soil so very favorable to their growth and

expansion,

The following account of the localities in which I have taken

F. cinerea, together with some notes on the structure of its

formicaries, may prove to be of interest to students of insect

distribution in general and of our American Formicida in par-

ticular. There are three of these localities some ten miles

apart, in different directions and at least three to five miles

from the town of Rockford, and in each of these localities,

which are. all open and exposed to the full heat of the sun, the

nests are of a different type. August 20, I found a single

nest, the first I had seen, under a small log in a meadow.

This nest was not very populous and contained neither larvae

nor pupa. It consisted of several inosculating galleries of the

type usually made by species of Formica and extended down at

least to a distance of 20 cm. into the black, waxy soil. The ants

were timid, like the inhabitants of all small nests of Formica,

and made no attempt to attack me. August 22, I found two

very large nests side by side at the edge of a turnpike not far

from a meadow. Each of these covered an area of somewhat

more than a square meter, and each consisted of a flat mound

of earth about 10 cm. high, strewn with little straws and sticks

brought together by the ants. This débris concealed numerous

openings from which the ants rushed forth as soon as the nest

was disturbed. Excavation was difficult on account of the

hardness of the soil, but it was easy to make out that the

formicary consisted of a honeycomb of galleries 1-2 cm. in

diameter and extended down into the soil to a depth of more

than 30 cm. It was filled with worker larve and pupæ,

together with thousands of ants, which attacked me furiously,

using their jaws and formic acid batteries to good purpose.

August 25, I discovered a locality where there are hundreds of

cinerea nests. This is a meadow about a mile and a half long

950 : THE AMERICAN NATURALIST. [Vor. XXXVI.

and a quarter of a mile wide, surrounded by woods and corn

fields. It is traversed by a cool stream, the banks of which

for some distance on either side are boggy and thickly studded

with large grass-covered hummocks. The F. cinerea have con-

structed their formicaries in these hummocks, which range

from 30 cm. to 60 cm. in diameter at the base and from 20 cm.

to 30 cm. in height. There are nests in nearly all stages

of growth, but for the most part well-established and extremely

populous, being, with the exception of the two nests above

described, the most populous nests of Formica I have seen during

the entire summer. The formicary is started in the summit of

the hummock, but ultimately invades its whole earthy sub-

stance and extends to a depth of at least 30 cm. to 60 cm. into

the black soil from which the hummock arises. In small or

moderately large nests all the grass which originally covered

‘the hummock remains intact and in excellent condition, but in

the largest formicaries the grass on the summit is partly

cut away by the ants and partly buried under the earth brought

up from the galleries and the little straws, bits of twigs, leaves,

etc., collected by the insects in obedience to an instinct which

appears to be shared to a greater or less extent by all the

species of Formica. This makes the large nests very conspic-

uous, although the numerous openings, all in the flattened

or somewhat convex summit of the hummock, are hidden

under the outermost layer of vegetable débris. The living

grass forming the sides of the hummock gives the nest great

stability and very efficiently protects it from being injured

by the feet of the pasturing cattle. Excavation of larger nests

shows that the hummocks are honeycombed throughout with a

network of inosculating galleries abruptly terminating at the

level of the moist, black meadow soil, into which only a very

few long and more or less perpendicular galleries and chambers

penetrate to a depth of 60 cm. and possibly farther.’

! During September, after this paper had been sent to the Naturalist, Y

happened on a fourth locality abounding in cinerea nests. This was 4 large

meadow almost within the city limits of Rockford. It contained formicaries of

all three types : under logs and stones, in the form of flat, irregular mounds a:

in modified hummocks.

No. 432.] FORMICA CINEREA MAYR. 951

These peculiar hummock formicaries occupy a zone on either

side of the stream midway between the dryer and more boggy

portions of the meadow, although a few of them reach quite to

the edge of the stream and are even perforated by the burrows

of frogs. The nests are so numerous as to be often within a

meter’s distance of one another. Along the outer edges of

these zones, and mingled with the dryer cinerea nests, there are

occasional nests of F. subsericea of precisely the same structure.

The main zone of this species, however, lies on higher ground,

where the hummock nests are replaced by true mound nests

entirely constructed by the ants.!

There were some slight variations in size and coloration

among the F. cinerea found in different nests in this locality,

but these are all comparable to similar variations in European

specimens. On the whole, the specimens from Illinois have

the ground color of the head and thorax more or less reddish

like the Californian and Austrian specimens. All the indi-

viduals examined have a number of hairs on the lower surface

of the head. According to Emery this is the distinguishing

trait of cinerea among all the European Formicidz. In the

United States F. schaufusst and F. subpolita and its varieties

agree with cinerea in possessing such hairs, but they may be

1 As Father Muckermann has shown in a recent paper (The Structure of the

Nests of Some North American Species of Formica, Psyche, June, 1902, pp. 355-

360), F. subsericea makes nests of at least four different styles: sm mall flat mound-

as those recorded by Father Muckermann for obscuripes.

nection that, like Father Muckermann, I do not altogether agree with Forel, who

believes that the American are inferior to the Euro ilding.

parts of the United States, to the species of assess and Ischnomyrmex

in the West and d and to the es of Lasius (LZ. aphidicola,

DUM and interjectus) in Illinois. During the men summer I saw near Rockford

a dom ped formicary of Z. interjectus 1.5 meters in diameter at the base and

60 centimeters high, and I have seen many nests of n and the other yellow

species of Lasius that were fully one-half to two-thirds as

952 THE AMERICAN NATURALIST.

readily distinguished by their coloration, which is never ashy or

silvery gray. At first sight F. subsericea resembles cinerea, but

the former never has hairs on the lower surface of the head.

This character definitely separates the two forms, notwith-

standing the fact that subsericea presents color variations in the

direction of czzerea.!

The cinerea nests were not seen til it was too late in

the year to secure the winged sexes, which, like the males and

females of our other species of Formica, probably make their

appearance during June and July. Even the dealated mother

queens were found in but one of the smaller nests. All the

nests, however, were full of worker larvae and pupa. The

latter were generally enclosed in cocoons, but quite a number

of nude pupz were also seen in many of the nests. In this

respect cinerea resembles the Formicidz of the pallide-fulva,

fusca, and subpolita groups, the worker larvae of all of which,

in contradistinction to F. rufa and its varieties, have a very

pronounced tendency to omit spinning a cocoon just before

pupation. At Rockford during the past summer many of the

nests of these species contained only nude pupa. This may

have been due to the great amount of moisture in the nests, as

June and July were unusually rainy. At any rate, I observed

that the cocoons were relatively much more abundant during

the dry weather late in August.

In its habits F. cinerea is very similar to the ants of the

fusca group. It was seen in great numbers visiting the flowers

in the meadow and attending great droves of Aphide on

the willows along the stream. The walls of the galleries

in some of the formicaries were tenanted by teeming colonies

of the minute lestobiotic, or thief ant, Solenopsis molesta Say.

In one cinerea nest I took a myrmecophilous histerid beetle

(Heterius brunneipennis Randall).

ROCKFORD, ILL, September 1, 1902.

1 At Rockford I discovered two rather large nests of a form which should, per-

haps, rank as a distinct variety of Z. fusca allied to subsericea. The ants from

these nests are smaller and more graceful in stature than the common subsericea,

the legs and antennz are red like those of cinerea, and the body is so thickly

overlaid with silvery white, appressed pubescence that the black ground color

is hardly visible. This form may be called Formica fusca var. argentata var. nov.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XII. Tue TREMATODES.

PART II.— THE ASPIDOCOTYLEA AND THE MALACOCOTYLEA,

OR DIGENETIC FORMS (continued).

H. S. PRATT.

KEY ro THE SPECIES OF NORTH—AMERICAN ASPIDOCOTYLEA AND

MALACOCOTYLEA, SO FAR AS DESCRIBED,

GENUS STICHOCOTYLE.

Larva found in Nephrops norwegicus and Homarus americanus : length

3-7 mm., with 7-22 suckers. Adult in Raja clavata: length 17-105 mm.,

with 20-27 suckers . S. nephropis Cunn. (Fig. 2) (62, 68)

GENUS COTYLASPIS.

Body white or pinkish: in various species of Unio and Anodonta

C. insignis Leidy (Fig. 5) (22, 23, 73, 16, 17)

GENUS COTYLOGASTER.

Body elongate; anterior portion of body terminated by a five-lobed disk

surrounding the mouth; sucking disk with 132-144 depressions; marginal

sense organs present; from dorsal surface near posterior end rises a broad

conical elevation, at base of which is excretory pore; length 8 mm.: in

gut of sheepshead in Minnesota . . . . - C. occidentalis Nick. (66)

GENUS ASPIDOGASTER.

Number of depressions about 64; sucking disk elliptical, covering two-

thirds of the body : in pericardium, liver, and kidney of Unio and Anodonta

A. conchicola v. Baer (Fig. 7) (20, 22, 23, 16, 17, 78)

GENUS PARAMPHISTOMUM.

a,. Body ovoid or elliptical.

4,. Acetabulum large; testes 2, lobate, in median line; 12 mm. long,

5 mm. wide: in intestine of Manatus :

P. fabaceum Dies. (32)

953

954 THE AMERICAN NATURALIST. [Vor. XXXVI.

ő, Acetabulum emarginate; 4-24 mm. long, 2-12 mm. wide : in intes-

tine of terrapin . dé. ane P. grande Dies. (29)

Body conical or subconical.

6,. Acetabulum large and subterminal; testes lobate, median: in

stomach of sheep and cattle . P. cervi Zed. (Fig. 9) (85, 87)

Č, Acetabulum in ventral surface not quite at posterior end, emargi-

nate; genital pore near anterior end often in a papilla or retracted

in a depression ; body triangular in cross section : in the muskrat

P. subtriquetrum R. (45)

GENUS DIPLopiscus.

Acetabulum large; body tapering anteriorly : in intestine of frogs

D. subclavatus R. (Fig. 12) (21, 80, 79, 41)

GENUS FASCIOLA.

. In mammals; worms of large size.

6,. Worms 18-50 mm. in length, 4-13 mm. in breadth; anterior end

distinctly set off from remainder of body; broadest part of worm

in front of its middle: in cattle, sheep, deer, man, ef a7.

F. hepatica L. (Fig. 13) (83, 85, 93)

à, Worms 20-100 mm. long, r1-26 mm. broad; anterior end indis-

tinctly set off; broadest part of worm back of its middle ; body

flesh colored: in cattle . . Dist. hepaticum Curtice (83)

Dist. carnosum Hassall (83)

Dist. americanum Hassell (83)

Dist. texicanum Francis (83)

Dist. crassum Leidy (83)

= F. magna Bassi (83, 85, 93)

4, In birds; worms not of large size; body piriform; 14 mm. long: in

lungs of kingfisher . . . . Dist. trapezium Leidy (32, 81)

= F. reticulata R. Wr. (97, 44)

GENUS FASCIOLOPsis.

Body nearly circular ; 15 mm. long, 12 mm. wide; in liver of an Indian

elephant (in Boston). . . . . . F., jacksoni Cob. (Fig. 14) (83, 44)

GENUS PLEORCHIS.

Body linguiform; 14} or I5 pairs of testes present; 5.25 mm. long,

1.5 mm. wide: in lung of a turtle (Sternotherus odoratus)

P. mollis (Leidy) S. et H. (82)

GENUS PARAGONIMUS.

Body red in life; suckers small ; ovary to left of acetabulum; 8-20 mm.

long, 4-8 mm. wide: in lungs of cat and dog, e£ al., and possibly encysted

in the muscles of swine (Agamodistomum sp.) in a larval condition

ae P. westermanni Kerb. (Fig. 17) (93, 87)

No. 432.] MORTH-AMERICAN INVERTEBRATES. 955

GENUS OPISTHORCHIS

4,. Body not spinose, lanceolate; testes not at extreme hinder end of

body; ro-21 mm. long, 1-2.5 mm. wide: in the liver of cats and

the cayote . . . O. pseudofelineus Ward (Fig. 30) (94, 81, 93)

4,. Body spinose, very elongate; testes at extreme hinder end of body;

12-29 mm. long, I mm. wide: in liver of crow

O. spestosus S. et H. (81, 84)

GENUS METORCHIS.

Uterus massed around acetabulum ; body linguiform ; testes large and

lobate; 6 mm. long, 2 mm. wide : in cats

M. complexus S. et H. (Fig. 32) (81)

GENUS TELORCHIS.

Yolk glands extend in front of acetabulum; intestinal ceca extend

beyond testes; 3.15 mm. long: in intestine of painted turtle

T. angustus Stafford (79)

GENUS DEROPRISTIS.

Anterior end usually attenuated and covered by large, coarse spines,

remainder of body by short spines; acetabulum much larger than oral

suckers; 3-6 mm. long, 1.2 mm. wide : in the intestine of Phycis tenuis

D. hispidus Abil. (40, 70)

GENUS ECHINOSTOMA.

a,. Number of oral spines "r 37; 18 mm. long, 1.5 mm. wide : in

intestine of muskrat . E. echinatum Zed. (Fig. 25) (28)

. Number of oral spines 45; 2. xà mm. E in intestine of GZde»ia

americana E. flexum Linton (36)

a,. Number of oral rien ve body elliptical; 8. mm. long: in intestine of

Botaurus minor. . . E. aspersum R. Wr. (97)

GENUS STEPHANOCHASMUS.

Number of oral spines 32, 16 being in each row ; body cylindrical

34.

anteriorly, globose posteriorly ; 2.5-3 mm. long : in capsules in peri-

toneum of filefish (A/utera dit et al.

S. valdeniflatus Stoss. i m

a, Number of oral spines 42, 21 being in each row ; body sl

lanceolate ; pharynx very large ; prepharynx long; intestinal cceca

bowe 3 mm. long, 0.6 mm. wide : in rectum of striped bass (occus

. S. tenuis Linton (38, 40)

afus) .

az. "eam elongate ; heri dps pre ohair long ; 3-15-4.5 mm. long,

0.5 mm. wide: in peritoneum of odi giam (Morone americana)

enuis tenuissimus Linton (38)

956 THE AMERICAN NATURALIST. [Vor. XXXVI.

a, Body elongate ; 4.5 mm. long: in intestine of Cottus scorpius

S. sobrinus Lev. (33)

a, Number of oral spines about 48, 24 being in each row ; body broad,

lanceolate; testes and ovary in contact with each other; 1.14-

1.85 mm. long, 0.37-0.64 mm. wide: in intestine of flounder (Para-

lichthys dentatus) Rance ha a S. dentatus Linton (39, 4°)

GENUS RHOPALIAS.

Body elongate, tapering towards the hinder end; tentacles long, being

up to 1.3 mm. in length; prepharynx and cesophagus of about same length ;

6-9 mm. long, o.8-1.16 mm. wide: in intestine and stomach of opossum

(Didelphys virginiana). . + + + + + ss R. coronatus R. (80, 7)

SUBFAMILY PSILOSTOMIN.

. Body subcylindrical, anterior end cylindrical and very contractile,

without spines; acetabulum larger than oral sucker; prepharynx,

„pharynx, and cesophagus present; yolk glands very voluminous and

extend from acetabulum to posterior end, filling the body behind

the testes, but not in front of acetabulum; 0.88-1.42 mm. long,

0.25 mm. wide: in intestine of hake (Merluccius biliniaris) et al.

Distomum vitellosum Linton (39, 4°)

a, Body elliptical, depressed, anterior end cylindrical, contractile, without

spines; yolk glands extend from pharynx in front of acetabulum

to hinder end, usually not overlapping the intestinal coeca; pre-

pharynx, pharynx, and «esophagus present; 1.6-2.74 mm. long,

0.57 mm. wide: in intestine of flounder (Paralichthys dentatus)

Distomum pudens Linton (39. 4°)

. Body elongate, linear, spinose; yolk glands extend from acetabulum

to hinder end of body; prepharynx, pharynx, and short cesophagus

present; 3 mm. long, 0.55 mm. wide: in intestine of cod

Distomum rachion (Cob.) Linton (38)

a, Body elliptical or piriform, covered with flat scalelike spines; acetab-

ulum of same size as oral sucker in middle of body; ovary to right

of and near acetabulum; yolk glands voluminous, extending from

pharynx -in front of acetabulum to hinder end of body, filling it

back of testes; prepharynx, pharynx, and cesophagus present;

0.26-0.36 mm. long, 0.12-0.18 mm. wide: in pyloric coeca of rud-

der fish (Palinurichthys perciformis) et al.

Distomum pyriforme Linton (39, 4°)

a, Body elliptical; acetabulum larger than oral sucker ; anterior dorsal

surface papillose; prepharynx, pharynx, and «esophagus present ;

yolk glands voluminous, occupying hinder half of body; 3.10 mm.

long, 0.86 mm. wide: in intestine of filefish (A/utera schoepfit)

ELE. Distomum pallens (R.) Linton (38, 40)

No.432.] MORTH-AMERICAN INVERTEBRATES. 957

. Body elongate, without spines; acetabulum prominent, larger than

oral sucker; yolk glands voluminous, back of acetabulum; 18 mm.

long, 0.34 mm. wide: in intestine of tomcod (Microgadus tomcod)

Distomum simplex (R.) Linton (38, 40, 33)

. Body elliptical, spiny ; a pair of groups of ocelli near mouth; acetab-

ulum of same size as oral sucker; testes in oblique plane; pre.

pharynx, pharynx, and cesophagus present; yolk glands voluminous,

filling body back of acetabulum ; 2-2.5 mm. long: in pyloric cceca

of Cottus scorpius . . . + + Distomum oculatum Lev. (33)

Body elongate, without spines, anteriorly cylindrical, posteriorly

depressed; acetabulum larger than oral sucker, near middle of

body; prepharynx, pharynx, and oesophagus present; testes and

ovary in median line, close together; yolk glands voluminous,

extending from acetabulum to hinder end ; 1.78 mm. long, 0.33 mm.

wide : in intestine of Mola mola | Distomum fragile Linton (39, 40)

4,. Body elongate, linguiform ; prepharynx, pharynx, and cesophagus

present; yolk glands voluminous, filling body back of testes ;

4.35 mm. long, r mm. wide: in intestine of Pseudopleuronectes

Distomum globiporum (R.) Linton (40)

americanus . s +» +

GENUS CREPIDASTOMUM.

Papille small; body elliptical ; acetabulum smaller than oral sucker;

pharynx and short cesophagus present; yolk glands voluminous,

extending from pharynx to hinder end of body and overlapping

intestinal coeca; 1.5-4 mm. long, 0.5-1.5 mm. wide: in rectum of

dep eee OU) Im UD

. Papille large; body linear, tapering towards hinder end; oral sucker

larger than acetabulum, but both large; esophagus absent; yolk

glands extend entire length of body; 1.7-2.2 mm. long, o.6 mm.

wide: in intestine of lake sturgeon (Acipenser rubicundus)

C. auriculatum (Wedl) Linton (38)

GENUS AZYGIA.

Body subcylindrical, light flesh color, posterior rounded. Ventral ace-

tabulum 3 line behind oral, } line in diameter. Oral acetabulum } line.

Length 8 lines; breadth posterior } line, anterior 4 line. i

before acetabulum: in stomach of Esox . 4. tereticollis (R.) Leidy (21)

GENUS RENIFER.

a, Body elliptical; genital pore near left edge of body; 3-4.2 mm. long,

1.1-1.6 mm. wide: in mouth of Heterodon platyrhinus

R. ellipticus Pratt (Fig. 53) (77)

pore near median line, but a little to left of

in mouth of Heterodon platyrhinus

R. elongatus Pratt (77).

a, Body elongate ; genital

it; 3 mm. long, 0.68 mm. wide :

958 THE AMERICAN NATURALIST. [VoL. XXXVI.

a, Body broad, piriform; genital pore near median line; 3-4 mm. long ;

1.5-2 mm. wide: in lung of 7ropidonotus sipedon

R. variabilis (Leidy) Lühe (21, 53, 77)

SUBFAMILY RENIFERINÆ.

. Body elliptical; acetabulum smaller than oral sucker; testes small,

spherical, in same transverse plane; ovary small, spherical, all three

being near acetabulum in center of body; genital pore median, in

front of acetabulum ; 2.8 mm. long: in intestine of fro

Distomum quietum Stafford (79)

. Body elongate, linear ; acetabulum near anterior end; genital pore

in front of it; testes spherical, in oblique plane, near middle of

body; uterus voluminous, filling hinder part of body; yolk glands

between testes and acetabulum; 6 mm. long: in ureter and kidney

of Boa constrictor . . . . Distomum horridum Leidy (18, 21)

, Body elliptical, without spines ; acetabulum near middle of body ; geni-

tal pore close to oral sucker ; testes immediately behind acetabulum ;

1.8-3 mm. long, o.8-1 mm. wide: in mouth of 7ropidonotus sipedon

Distomum aniorum Leidy (32)

a, Body elliptical; testes ovoid, in same transverse plane; ovary imme-

diately behind acetabulum, which is larger than oral sucker wes in

middle of body; excretory vesicle wide and long; 3 mm. lon

intestine of Cottus scorpius . . Distomum furcigerum Olss. dem

. SUBFAMILY PLAGIORCHIINE.

. Body elongate, with minute spines; uterus very voluminous; yolk

glands a number of roundish masses distributed laterally along the middle

region of the body ; 2.8—5.5 mm. long, 0.6-1.12 mm. wide : in intestine of

Tylosurus carribeus . . . . . . . » Distomum nitens Linton (38)

GENUS BUNODERA.

Intermediate host the crayfish (Cambarus), in which it is found in cysts

in ovary and other organs; 1 mm. long

B. nodulosa Zed. (Fig. 45) (98, 35)

GENUS HA&MATOLCCHUS.

Acetabulum very small : in lung of frogs

H. variegatus R. (Fig. 55) (20, 21, 97, 79)

: GENUS OSTIOLUM.

Acetabulum minute; body elongate; meus limb of excretory vesicle

very long ; 7-11 mm. long, 1.5 mm. wide: gs

0. oiii Pratt (Fig. 56) (77)

No. 432.] MORTH-AMERICAN INVERTEBRATES. 959

GENUS GORGODERA.

. In urinary bladder of frogs and salamanders; number of testes, 9;

form of yolk gland, veriform

G. cygnoides Zed. (Fig. 62) (20, 21, 13, 22, 44)

4,. Number of testes, 9; yolk glands deeply lobate, in form of a rosette ;

ovary lobate; 4.6 mm. long: in urinary bladder of Rana catesbiana,

R. baie R. virescens + + + + G. amplicava Lss. (2, 44)

45. Number of testes, 2; ovary veriform; yolk glands elongate and not

lobate masses; 10 mm. long: in bladder of Rana catesbiana,

R. clamitans, R. virescens. . . . « G. simplex Lss. (2, 44)

GENUS MICROPHALLUS.

Body elliptical; cesophagus very long, intestinal coeca very short; 1.7

mm. long, 1 mm. wide: in intestine of Ama calva et al.; intermediate host

Cambarus bropinguus . . . . M. opacus Ward (Fig. 75) (92, 94, 74)

GENUS LEVINSENELLA.

Body elliptical; aesophagus very long; yolk glands consist of several

distinct bodies; 0.5 mm. long: in intestine of Somateria mollissima

Z. fygmea Lev. (Fig. 71) (33, 15, 94)

GENUS CYMATOCARPUS.

Body elliptical; intestinal coeca extend about to acetabulum; testes in

oblique plane; 4 mm. long: in intestine of newt

C. hospitalis Stafford (79)

GENUS BRANDESIA.

Body elliptical; acetabulum in hinder part; genital pore on left side in

anterior half; intestinal coeca short and wide; ovary lobate between them ;

testes in nearly same transverse plane in hinder half of body ; 2.5 mm. long,

1.75 mm. wide: in capsules in pyloris of frog B. arcana Nick. (64)

GENUS PLEUROGENES.

Body circular; 3-4 mm. long: in cysts in liver of bullfr

P. medians (Olen) Stafford (79)

GENUS GYMNOPHALLUS.

Body elliptical ; excretory crura E to pharynx; o.5 mm. long: in

intestine of Somateria mollissima . . G. somateria Lev. (33, 69)

GENUS CEPHALOGONIMUS.

Body elliptical ; testes and ovary very near the acetabulum ;

very short; yolk glands extend in front of acetabulum ; excretory vesicle a

960 THE AMERICAN NATURALIST. [VoL. XXXVI.

rounded sac into which radial canals extend; 2 mm. long: in intestine of

soft-shell turtles (Aspidonectes and Amyda) . C. vesicaudus Nick. (65)

GENUS PROSTHOGONIMUS.

Body elliptical; acetabulum larger than oral sucker; 7 mm. long, 2. 8 mm.

wide: in the white of hen's egg, also in Buteo lineatus and Corvus ameri-

CONUS o oos eas . . P. ovatus R. (Fig. 61) (34, 80)

GENUS DICROC(ELIUM.

Body elongate, lanceolate; suckers near together; testes lobate, close

behind acetabulum; 4-10 mm. long, 1-2.5 mm. wide. “Stated to be fre-

quent in sheep in several western states” (Leidy). ‘“ Apparently not in

North America” (Stiles and Hassall): in cattle, sheep, man, etc.

D. lanceatum S. et H. (Fig. 82) (21, 84, 85)

GENUS LYPEROSTOMUM.

In Corvus americanus . >» . . . . L. plesiostomum v. Linst. (80)

GENUS HEMIURUS.

a,. Body cylindrical; appendix from a third the length to twice the length

of the body; acetabulum much larger than oral sucker; testes not

near acetabulum; yolk glands compact or slightly lobate ; o. 5-6 mm.

long: in stomach and pyloric coeca of marine fishes; intermediate

host, copepods D. pelagicum Stafford (78)

= H. appendiculatus R. (Fig. 90) (39, 33: 55; ei 24)

a» Body fusiform; appendix about a third the length of f body; acetab-

ulum larger than oral sucker ; testes close to acetabulum ; 1.5-3.5 mm.

long: in Macrourus bairdii. . . H. levis Linton (39, 55)

a, Body slender, cylindrical; appendix ded. a third the length of body ;

acetabulum larger than oral sucker; testes distant from acetabulum ;

yolk glands lobate ; 5.12-5.4 mm. long, 0.75-1 mm. wide: in stomach

of Remora remora. . . H. monticelli Linton (39)

. Body slender; appendix About a gem the length of sone A 3.2 mm.

long, 0.42 mm. wide: in intestine of Pomatomus saltatri.

H. ocreatus (Mol.) tds (39. 55)

^»

GENUS LECITHOCLADIUM.

a,. Body slender; appendix long; 8-15 mm. long, o.7 mm. wide: in Cory-

phena hippurus et al. . . L. tornatum (R.) Linton (38, 40, 5 5)

4, Body slender; appendix long and slender; acetabulum of same size as

oral sucker; cesophagus wanting; testes just behind vesicula semi-

nalis; the tubular yolk glands extending into appendix; 10 mm.

long, 1.14 mm. wide, length of cie 3.6 mm.: in butterfish

` (Rhombus jrienthen) ea . L.gulosum Linton (40, 55)

No. 432.] MORTH-AMERICAN INVERTEBRATES. 961

GENUS LECITHOCHIRIUM.

Body cylindrical; 2-6 mm. long: in stomach of Anguilla chrysypa

L. grandiporum (R.) Linton (38, 55)

GENUS LECITHASTER.

Body fusiform; testes immediately behind acetabulum; 1-1.5 mm. long:

in intestine of Cottus scorpius Dist. mollissimum Lev. (33

= L. bothryophorus Olss. (Fig. 93) (44, 55)

GENUS PRONOPYGE.

Body elongated, elliptical, flattened; 2-3 mm. long: in stomach of

| Trichinous lepturus. . «. . Dist. gastrocolum Leidy (32)

= P. ocreata Mol. et R. (Fig. 92) (81, 55)

GENUS DEROGENES.

Body elongate; acetabulum very large, in middle of body; testes close

to acetabulum; 2 mm. long: in stomach of Cottus scorpius et al.

D. varicus O. F. Mil. (33, 52)

GENUS PROGONUS.

Body elliptical; acetabulum very large, in middle of body; 2 mm. long:

in stomach of Cottus scorpius et al. P. miilleri Lev. (Fig. 84) (33)

GENUS HALIPEGUS. ;

Body elongated; acetabulum larger than oral sucker, in center of body ;

10 mm. long: in Eustachian recesses and mouth of frog

H. ovocaudatus Vul. (Fig. 87) (63, 79, 53)

GENUS ACCAC(LIUM.

Body cylindrical, *dorsal surface and lateral margins armed with

spheroidal tuberculate spines”; “twenty flat spines, more or less,

on the inner margin of the oral sucker”; 8 mm. long, 0.72 mm.

B 3 A. contortum (R.) Linton (Fig. 80) (38)

a, Body cylindrical and slender; acetabulum larger than oral sucker and

ona long peduncle; testes near center o body; no spines present ;

yolk glands ** in slender, threadlike folds, centrally situated ” ; 35 mm.

long, 1.12 mm. wide: in intestine of Mola mola

$ A. nigroflavum (R.) Linton (38, 39)

a, Body cylindrical, without spines ; acetabulum twice as large as ora

sucker and pedunculate; 14 mm. long, 1.7 mm. wide: in intestine of

Mola mola . . . A. macrocotyle (Dies.) Linton (38, 39, 40)

a, Body cylindrical, slender ; acetabulum pedunculate, *: consisting of four

foliate flaps " ; 6-8 suckerlike structures in mid-dorsal line at anterior

end of body; 12 mm. long, 1.28 mm. wide: in intestine of Mola

dE ee que Amor tote no A. foliatum Linton (38)

wide: host?. .

962 THE AMERICAN NATURALIST. [Vor. XXXVI.

ALLIED TO GENUS ACCACCLIUM.

* Body cylindrical, narrowest in fore part and obtuse behind; ventral

bothria larger than mouth and projecting in advance in an extent equal to

the body; skin smooth and transparent; yellow intestines and white and

brown genitals shining through”; 20-45 mm. long, 0.5-1.5 mm. wide: in

intestine of Mola rotunda . . . . . Distomum pedocotyle Leidy (31)

GENUS CLINOSTOMUM.

Body elongate; intestine with short side projections; yolk glands fill

body back of acetabulum ; cirrus sac before testes ; 6-10 mm. long, 1-2 mm.

wide: in mouth of Ardea herodias ef al.; as young form in cysts in

Lepomis auritus et al.

C. heterostomum (R.) R. Wr. et MacCallum (97, 56, 6)

Dist. gracile (Dies.) Linton (38, 6)

Dist. aquile Leidy (27, 81, 6, 85)

C. gracile Leidy (21, 81, 6)

Dist. galactosomum Leidy (30, 56, 6)

= C. marginatum R. (Fig. 101) (6)

ALLIED TO GENUS CLINOSTOMUM.

* Body elongated, elliptical, moderately wider and thicker posteriorly,

unarmed.” * Mouth subterminal and enclosed with a reniform lip, suc-

ceeded by a linear annulus. Acetabulum large, at anterior fourth of body.”

ae orifice at posterior fourth of med " Length 15-20 mm., breadth

3 mm.: in mouth of Alligator . . . . Distoma oricola Leidy (25):

DisroMiDs OF UNCERTAIN LOCATION IN THE SYSTEM.

PARASITIC IN MAMMALS.

Body elliptical and spinose ; acetabulum in anterior half of body ; genital

pore halfway between it and posterior end of body ; pharynx, short cesopha-

gus, and long intestinal cceca present; testes large, obliquely behind one

another in hinder part of body; ovary to right of them; 0.65-1.2 mm.

long, 0.35-0.64 mm. wide: in intestine of Lepus

Distomum tricolor S. et H. (82)

PARASITIC IN REPTILES.

nr elongate; cesophagus wanting ; testes obliquely behind one another

in mi : y; ovary near acetabulum at anterior fourth of body;

e uterus between. testes and ovary ; yolk glands behind testes; 6 mm. long:

in intestine of snapping turtle . . . Distomum chelydre Stafford (79)

No. 432.] MORTH-AMERICAN INVERTEBRATES. 963

PARASITIC IN AMPHIBIANS.

Encapsuled under skin of Stren lacertina Distomum sirenis Vail. (90)

Body elliptical; intestinal coeca short, reaching to acetabulum ; excretory

vesicle large, the crura reaching to acetabulum; 0.7 mm. long, 0.45 mm.

wide: in thoracic cavity of grass frog and encysted in anterior part of

bullfrog . . . . . . . . Jistomum tetracystis (Gal.) Stafford (79)

Body whitish, oblong, sublinear, narrowed anteriorly ; oral sucker larger

than acetabulum; 3 mm. long, 0.5 mm. wide: in Rana halecina

Distomum retusum (Duj.) Leidy (20, 21)

PARASITIC IN FISH.

Body elongate; suckers near each other, acetabulum somewhat pedun-

culate ; genital pore in front of latter; 15 lines to 3 inches long, $ line

wide: in mouth of Esox . . . . . . Déstomum longum Leidy (20)

* Body long and narrow, band-like, with ventral disk at anterior fourth ;

smooth.” “Ventral acetabulum about the width of body; oral disk

smaller”; 8 mm. long, 0.87 mm. wide: in mouth, throat, and gills of sand

pike (Saurus fatens) «. « + + + +: Distomum ischnum Leidy (32)

* Body flat, elongated elliptical ; echinated between oral and ventral

acetabula, which are equal, and the latter one is hemispherical and sessile.

Length 21 to 3 lines, breadth 3-5ths to 4-5ths of a line.” In intestine of

Leiostomus obliquus... + + * ? Distomum incivile Leidy (21)

* Body ovoid, anteriorly compressed, conical and incurved, posteriorly

robust and obtuse.” **Acetabulum much larger than mouth.” “Genital

pore a little in advance and to left of acetabulum, provided with prominent

circular lip”; length 1-2} lines; breadth 4-1 line: in gall bladder of

e. o sn Distomum biliosum Leidy (23

oval; 20 mm. long, 6 mm. wide: in body cavity of

uq En Distomum lageniforme Linton (38)

Body elliptical, without spines ; acetabulum larger than mouth ; testes in

same transverse plane nearit; ovary in front of testes; intestinal ceca

long; 1.25-6 mm. long, 0.7-2.6 mm. wide : in intestine of Spheroides macu-

E | 2 1 5 4 a ve vc o» y Distomum vibex Linton (39)

Body wide and thick, without spines, bluntly rounded at both ends;

acetabulum very large ; cesophagus short, intestinal coeca long; testes and

ovary close together in hinder part of body, the latter in front of former ;

uterus voluminous, filling body back of acetabulum ; genital pore to right

of pharynx; 2.75 mm. long, 1.4 mm. wide: in intestine of tilefish

Distomum fecundum Linton (39)

Body broad, spiny ; acetabulum small, in anterior half of body ; cesopha-

gus short, intestinal caeca long; testes behind ovary, side by side; genital

pore to left of pharynx; o.7-1.3 mm. long, 0.4-0.64 mm. wide: in intestine

of white perch >.. . . istoemum areolatum (R.) Linton (39)

unknown fish. .

Body elongate,

Remora remora . .

964 THE AMERICAN NATURALIST. [Vor. XXXVI.

“Head compressed oval, convex anteriorly; mouth minute, not bor-

dered. Body compressed oblong oval. Ventral acetabulum immersed

between body and head. Length 2} lines, breadth } of a line.” From

the intestine of Rusticola minor

Distomum (Clinostomum) dubium Leidy (21, 6)

* Body elongated, sublinear, depressed, unarmed, transversely rugose,

posteriorly attenuate. For about 6.5 mm. back of the ventral sucker the

body was filled with the voluminous folds of the uterus”; 20 mm. long,

3.2 mm. wide: in stomach of Raja levis

Distomum veliporum (Creplin) Linton (38)

Body cylindrical, posterior portion larger than anterior; testes side by

side, immediately behind acetabulum; 18 mm. long, 4.5 mm. wide: in

stomach of swordfish . . . . . . Jéstomum clavatum (RÀ (21, 38)

PARASITIC IN MOLLUSKS.

Body oval; acetabulum large, in center of body; intestine convoluted,

passing to extremity of body; genital orifice halfway between acetabulum

and hinder extremity of body; length } line, breadth 1 line: in pericar-

dium of Helix alternata and H. albolabris üfierinediate- host)

Distomum helicis Leidy (19, 21)

— Distomum vagans Leidy (18)

Body fusiform ; suckers large ; acetabulum near the middle of the body ;

oral sucker “ with a style inserted in the upper lip "; o.2 mm. long, .o8 mm.

wide: in liver of //yanassa obsolata . . . Distomum lasium Leidy (32)

PARASITIC IN CRUSTACEANS.

Body cylindrical; acetabulum larger than oral sucker; r.25 mm. long;

o.4 mm. wide; in Apus /ucasanus . . Distomum apodis Packard (75)

Body clavate, with a long, narrow tail (a cercaria); 0.25-0.4 mm. long:

in muscles, liver, and intestine of Planorbis and Limnza

Distomum ascoideum Leidy (24)

GENUS SCHISTOSOMA.

Male 4-14 mm. long, 1 mm. wide; female 13-20 mm. long, o.28 mm.

wide: in blood of man and cattle (?)

S. haematobium Bilh. (Fig. 107) (85)

GENUS KCELLIKERIA.

Longer diameter of reniform pr 1.74 mm., shorter diameter

1.09 mm., diameter of neck 0.13 mm. : in cysts in intestinal wall of Spanish

mackerel. uo ^ 2... . K filicolle R. (Fig. 108) (40)

No. 432.] WORTH-AMERICAN INVERTEBRATES. 965

GENUS DIPLOSTOMUM.

4,. Acetabulum and oral sucker small ; special organ of attachment of a

large cavity at bottom of a conical projection just back of acetabu-

lum, in the center of which is a short canal leading to the cavity,

3-4 mm. long: in intestine of Strix nivea et al.

D. grande Dies. (23, 80)

. Special organ of attachment open; acetabulum and oral sucker small ;

1 mm. long, 0.35 mm. wide; a larval form found in cysts in liver of

Pomotis vulgaris etal. . D. cuticola Dies, (21, 38, 80)

@,. Acetabulum large; oral sicker very small ; anterior portion of body

longer than hinder portion and not sharply separated from it; special

organ of attachment an elliptical depression; 3.5-4.5 mm. long: in

intestine of alligator D. pseudostomum Will.-S. (96)

GENUS POLYCOTYLE.

Oral and ventral suckers small; 14 mid-dorsal suckers ; 4.5 mm. long:

in intestine of alligator. . . P. ornata Will.-S. (Fig. 110) (96)

GENUS HOLOSTOMUM.

Body spinose, “divided by a constriction at the anterior third."

* Head ovoidal, mouth terminal, round, opening into a cup-shaped

pharynx"'; 3.2 mm. long, 0.8 mm. wide: in intestine of Rana pipiens

H. nitidum Leidy (21)

Body not spinose ; ventral sucker twice as large as oral sucker; ventral

projection slit longitudinally into several parts and not extending

beyond cup; 4-5 mm. long: in intestine of Ardea her

H. cornu Nitzsch (21, 80)

Anterior part of body subglobose, variable, aperture transverse; pos-

terior cylindrical; reproductive aperture terminal surrounded by a

muscular border and with a central conical protractile part; 3 mm.

long, o. 8 mm. wide: in intestine of Circus cyaneus

H. variabile Nitzsch (Fig. 113) (36)

GENUS GASTEROSTOMUM.

Body elliptical; mouth in center of body; 4-6 mm. long: in pyloric

appendages of Cottus scorpius . G. armatum Mol. (Fig. 114) (33)

a, Body ovate, being wider anteriorly ; mouth towards anterior end ; : »

2.7 mm. long, 0.92-1 mm. wide: in intestine of Zobofes su

Monostomum orbiculare (R.) aes ati

— G. ovatum Linton (39)

1.28-2.7 mm. long,

. Body elongate; mouth towards anterior end;

0.21 mm. wide: in intestine of Sarda sarda

G. arcuatum Linton (39, 40)

966 THE AMERICAN NATURALIST. [Vor. XXXVI.

SUBFAMILY PRONOCEPHALIN4E.

Body elongate, lateral margins parallel; reniform projection at anterior

‘end ; intestinal cceca long and serpentine ; testes lobate at hinder end; 18-

25 mm. long, 3 mm. wide: in intestine of Spargus coriacea

Monostomum renicapite Leidy (21, 8)

GENUS OPISTHOTREMA.

Body broad, sagittate; testes small, lobate, in same transverse plane,

laterad of intestinal coeca; ovary small, in front of testes; 10 mm. long; in

Manatus YO aare Leuck. (Fig. 118) (96)

GENUS CYCLOC@LUM.

Body elongate, attenuate gsm ; 18 mm. long: in thoracic cavity of

Gallinago eż a7. oo . . . C. mutabile Zed. (Fig. 119) (26, 80)

GENUS NOTOCOTYLUS.

Body elongate, elliptical ; testes in hinder end of body; 7 mm. long: in

intestine of Somateria mollissima et al.

N. verrucosus Fróh. (Fig. 124) (33)

MonosToMIDS OF UNCERTAIN LOCATION IN THE SYSTEM.

Body spatulate, broader posteriorly; oral sucker small ; intestinal coeca

long; 12 mm. long, 2 mm. wide: in bile ducts of muskrat

Monostomum affine Leidy (23)

Body elongate, elliptical, obtusely angular in front, obtusely rounded

behind; 12 mm ig 2 mm. wide: in stomach of jewfish (Megalops

thrissotdes) . . . . . Monostomum obscurum Leidy (27)

Body flat, oblong ovate, narrowing anteriorly, obtuse posteriorly ; 6 mm.

long, 1 mm. wide: in fresh-water fish

Monostomum spatulatum Leidy (23)

Body ovoidal, loger broad; 2 mm. long, 1 mm. wide: in body cavity

of Rana pipiens, . . . Monostomum ornatum Leidy (21)

Body elongate, posterior quac slightly broader ; cesophagus very long ;

intestinal cceca short; genital pore at end of esophagus in hinder half of

body ; 1.4 mm. long, .24 mm. wide : encapsuled under skin of Séven lacertina

Monostomum aspersum Vail. (80)

. Body ovate, spiny; 8 testes, 4 on each side; ovary lobate in front of

them; uterus voluminous; 2.3 mm. long: in Opsanus tau

Monostomum vinal-edwardsii Linton (40)

: Body Maius: ; hinder end truncated or concave; testes side by side, at

e MEM. hinder end; ovary lobate, near center of body; 5 mm. long: in

i ng bladder of catfish ‘onostomum amiuri Safford (79)

kaal

Laal

-432.] NORTH-AMERICAN INVERTEBRATES. 967

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968

THE AMERICAN NATURALIST. [VoL. XXXVI.

; LRIDY, F hapa on Entozoa in Naiades. Proc. Acad. Nat.

Sci. Phila. 1857, p

LEIDY, J. Carbio 4 to Helminthology. Proc. Acad. Nat. Sci.

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LEIDY, ] On fite infesting Mollusks. Proc. Acad. Nat. Sct.

Phila. 1877, p. 200.

Leipy, J. Distoma and Filarie. Proc. Acad. Nat. Seci. Phila.

LEIDY, J. On Some Parasitic Worms of Birds. Proc. Acad. Nat.

Sez. Phila. 1885, p. 9.

LEIDY, J. Notes on Some Parasitic Worms. Proc. Acad. Nat. Sci.

Phila. 1887, p.

LEIpy, J. 2 of the Muskrat. Proc. Acad. Nat. Sct. Phila.

1888, p. 126.

LEIpy, J. ios of the Terrapin. Proc. Acad. Nat. Sci. Phila.

1888, p.

c LEIDY, J. se of the Rock Fish. Proc. Acad. Nat. Sci. Phila.

166.

1888, p. 1

. Letpy, J. Parasites of Mola rotunda. Proc. Acad. Nat. Sci. Phila.

1890, p. 282.

. LEIDY,J. Notes of Entozoa. Proc. Acad. Nat. Sci. Phila. 1890, p. 413.

LEvINSEN, G. M. R. Bidrag til Kundskab om Grönlands Trematod-

fauna. Oversigt Danske Vidensk. Selsk. Forhandl. No. 1, p. 52.

881.

Linton, Epwin. Notes on a Trematode from the White of a Newly

Laid Hen’s Egg. Proc. U.S. Nat. Mus. 1887, p. 387.

- Linton, EpwiN. Notes of Trematode Parasites in the Crayfish.

Amer. Nat. Vol. xxvi, p. 69. 1892

. Linton, Epwin. Notes on Avian Entozoa. Proc. U. S: Nat. Mus.

Vol. xv, p. 87. 1892.

. Linton, Epwin. On Fish Entozoa from Yellowstone National Park.

ep. of U.S. Com. of Fish and Fisheries for 1889 to 1891. 1893,

p. 545.

Linton, EDWIN. Notes on Trematode Parasites of Fishes. Proc. of

U. S. Nat. Mus. .Vol. xx, p. 507. 1898.

Linton, Epwin. Fish Parasites collected at Woods Hole in 1898.

U.S. Fish Com. Bull. for 1899. 900, p. 267.

Linton, Epwin. Parasites of Fishes of the Woods Hole Region.

U.S. Fish Com. Bull. for 1899. 1901, p. 405.

. Looss, A. Ueber Amphistomum subclavatum R. Festschrift R.

Leuckart’s. 1892, p. 147.

Looss, A. Die Distomen unserer Fische und Frösche. Bibl. Zool.

= Hefti6. 1894.

Looss, NC Heec sor la faune parasitaire de l'Egypte. Prem-

dere rte Mém. Inst. Egyptien. Tome iii. Caire, 1

60.

.432.]] .VORTH-AMERICAN INVERTEBRATES. 969

Looss, A. Weitere Beitráge zur Kenntniss der Trematoden-Fauna

Aegyptens, etc. Zool. Jahrb., Abth. f. Syst, etc. Bd. xii, p. 521.

. Looss, * Nachträgliche Bemerkungen zu den Namen der von mir

vorgeschlagenen Distomidengattungen. Zool. Anzeiger. Bd. xxiii,

p. 6o1. 1900.

Looss, A. Natura doceri, etc. Centrali f. Bak., etc. I. Abth.,

Bd. xxix, p. 191.

gol.

. Looss, A. Ueber einige Distomen der Labriden des Triester Hafens.

CentrallJ. f. Bak., etc. 1. Abth., Bd. xxix, p. 3

. Looss, A. Ueber die F ácia iA Stokka a. Acantho-

chasmus und einige andere. Centralbl. f. Bak., etc. I. Abth.,

Bd. xxix, p. 595. 19or.

Looss, A. Notizen zur Helminthologie Aegyptens. IV. Ueber Tre-

matoden aus Seeschildkróten der aegyptischen Küsten. Centra/d/.

f. Bak., etc. E Abth., Bd. xxx, p. 555. 1901.

. LüHE, M. Zur Kenntniss einiger Distomen. Zool. Anzeiger. Bd. xxii,

p. 524. 1899.

. Line, M. Ueber die Gattung Podocotyle (Duj.) Stoss. Zoo.

Anzeiger. Bd. xxiii, p. 487. 1900.

Line, M. Ueber Distomen aus der Gallenblase von Mittelmeerfischen.

Zool. Anzeiger. | Bd. xxiii, p. 584.

LünE, M. Ueber einige Distomen aus cba und Eidechsen.

Centralbl. f. Bak., etc. I. Abth., Bd. xxviii, p. 555. 1900

Line, M. Ueber Monostomum orbiculare. Cenfralbl. f. Bak., etc.

I. Abth., Bd. xxix, p. 49. 19ot.

Line, M. Ueber Hemiuriden. Zool. Anzeiger. Bd. xxiv, pp. 394,

473. 190I.

. MacCarLLuM, W. G. On the Species Clinostomum heterostomum.

Journ. of Morph. Vol. xv, p.697. 1899.

MONTICELLI, F. S. Osservazioni intorno ad alcune forme del genus

Apoblema Duj. Azti Accad. Sc. Torino. Vol. xxvi, p. 495. 1891.

MONTICELLI, F. S. Cotylogaster Michaelis n. g., n. s., e rivisione degli

Aspidobothride. Festschrift R. Leuckart’s. 1892, p. 168.

MONTICELLI, F. S. Studii sui Trematodi endoparassiti. Primo con-

tributo di osservazioni sui Distomidi. — Jahrb. Suppl. Bd. iii,

p.139. 1893.

MünLING, P. Beiträge, etc. Arch. f. Naturg. 62. Jahrg. 1896,

P- 244-

MünriNG, P. Die — qn d. Wirbelt. Ostpr. Arch. f.

Naturg. 64. Jahrg. 1898, p

Im On ak nephropis Cunn., etc. Zool.

Jahrb., Abth. f. Syst, etc. Bd. viii, p. 447- 1894.

On the Occurrence of Distomum ovocaudatum

NICKERSON, W. S.

| Vidpien ds American Frogs. Zool. Bull. Vol. i, p. 261. 1898.

970 THE AMERICAN NATURALIST. [Vor. XXXVI.

64.

65.

66.

67.

68.

69.

70.

NICKERSON, W. S. Note on Distomum arcanum (n. sp.) in American

rogs. Amer. Nat. Vol. xxxiv, p. "m 1900.

NICKERSON, W. S. Description of Cephalogoni icaudus sp. nov

Science. Vol. xv, p. 581. 1902.

NICKERSON, W. S. Cotylogaster occidentalis n. sp. and a revision of

the Family Aspidobothride. Zool. Jahrb., Abth. f. Morph., etc.

Bd. xvi, p. 597. 1902.

OFENHEIM, E. von. Ueber eine neue Distomidengattung. Zeitschr.

f. Naturwissensch. Bd. lxxiii. 1900.

OpHNER, TH. Ueber die geschlechtsreife Form von Stichocotyle

nephropis Cunn. Zool. Anzeiger. Bd. xxi, p. 500. 1898.

OpHNER, TH. Gymnophallus, eine neue Gattung von Vogeldistomen.

Centralbl. f. Bak., etc. I. Abth., Bd. xxviii, p. 13. o.

ODHNER, TH. Mitteilungen zur Kenntnis -— Distomen. Centra/lbl.

f. Bak., etc. 1. Abth., Bd. xxxi, Nos. 2

Sid 2.

. ODHNER, TH. Téematodet aus Reptilien, etc. Ofver-af Kongil.

Vetenskaps-Acad. Fórhand. No. 1, p. 19. 1902.

. OrssoN, P. Bidrag till Skandinaviens Helminthfauna, II. Kongl.

Sven. Vet. Akad. Hand. Bd. xxv, p. 3.

1893.

. OSBORN, H. L. Observations on the Anatomy of a Species of

Platyaspis found Parasitic on the Unionidz of Lake Chautauqua.

Zool. Bull. Vol. ii, p. 55.

1898.

. Osporn, H. L. Notes on the Trematodes of Lake Chautauqua, N.Y.

Sczence. Vol. xv, p. 573.

PACKARD, A. S. A New Distomum in the Egg Sacs of Apus. Amer.

Nat. Vol. xvi, p. 142.

1882.

. Pratt, H. S. A Contribution to the Life-History and Anatomy of

the Appendiculate Distomes. Zool. Jahrb., Abth. f. Anat., etc.

Bd.xi 18 :

Pratt, H. S. Descriptions of Four Distomes. Mark Anniversary

Volume. 1902.

STAFFORD, J. Anatomical Structure of Aspidogaster conchicola.

Zool. Jahrb., Abth. f. Anat., etc. Bd. ix.

. STAFFORD, J. Some Undescribed Trematodes. uet Salil: Abth.

f. Syst, etc. Bd. xiii, p. 399. 1900.

STILES, C. W., and HassALL, A. A Preliminary Celene of the

Parasites contained in the United States Army Medical Museum,

Biological Department of the University of Pennsylvania (Coll.

Leidy) and in Coll. Stiles and Coll. Hassall. Veterinary Magazine.

April, 1894, p. 245; and May, 1894, p. 331.

. Stiles, C. W., and HassALL, A. Notes on Parasites, 21. A New

Species of Fluke (Distoma complexum), etc. Veterinary Magazine.

June, 1894, p. 413.

STILEs, C. W., and Hassatt, A. Notes on Parasites, 29 and 30:

_ A New spicis of Intestinal Fluke (Distoma tricolor), etc. Kiers

mary Magazine. November, 1894, p. 729.

. Warp, H. B. On the Parasites of the Lake Fish.

. WRIGHT, R. R.

.432.] NORTH-AMERICAN INVERTEBRATES. 971

. STILES, C. W., and HassALL, A. The Anatomy of the Large Ameri-

can Fluke, etc. The Journal of Comparative Medicine and Veteri-

nary Archives. Vol. xv (1894), pp. 161, 225, 299, 407, 457-

vol. xvi (1895), pp. 139, 213, 277.

. STILES, C. W., and HaAssALL, A. Notes on Parasites, 42-46. Com-

parison of the Type of A ES longissimum, etc. Veterinary

Magazine. March, 1896, p

s SLES Ce Wo aag Payal ies e The Inspection of Meats for

Animal Parasites. Bull. 19, Bureau of An. Industry, Dep't of

Agr. 1898.

STILES, C. W., and HassALL, A. Notes on Parasites, 48. An Inven-

tory of the Genera and Subgenera of the Trematode Family Fascio-

lide. Arch. d. Paras. Vol. i, p. 81.

98.

. STILES, C. W., and HassALL, A. Notes on Parasites, 50, 51, 52.

The Lung Fluke (Paragonimus westermanni) in Swine, etc. .Szx-

teenth An. Rep. Bur. An. Industry. 1899, p

«5

. SrOssicH, M. Branidi elmintol. tergest, II. Boll. poe. Adriat. Vol. ix

Estr., p. 2.

. Srossicn, M. Elminti trovati in un Orthagoricus mola. Boll. Soc.

Adriat. Vol. xvii, Estr, p. 4. 1

VAILLANT, L. Sur quelques béliinties de la sirène lacertine. Annal

d. Sci. nat., Sér. 4, Zool. Tome xix, p. 347-

. WAGENER, G. Ueber Distomum appendiculatum R. T rch. f. Naturg.

26. Jahrg. Bd.i, p. 165. 1860.

I. Notes on the

Structure and Life History of Distoma opacum n. sp. Proc. Amer.

Micr. Soc. Vol. xv,p. 173. 1894.

Warp, H. B. The Parasitic Worms of Man and the AOR Ani-

mals. ep. d tag Neb. State Board Agr. 1895, p.

Warp, H. B. n the Parasites of the Lake E n I. On

the Structure a the piste Organs in Microphallus nov. gen.

Studies from the Zovl. Lab. Univ. of Neb. 1901, p. 175.

. West, G. S. A New Species of Distomum. Journ. Linn. Soc.

Vol. xxv, p. 322. 1895.

Zeitschr.

WiLLEMOEs-SuHM, R. vox. Ueber einige Trematoden, etc.

f. wiss. Zool. Bd. xxi, p. 177. 1871

Contributions to Alterican Helminthology. Journ.

Proc. Canad. Instit. Vol. i N-S., p. 54- 1879.

Wricut, R. R. Trematode Parasites in American Crayfish. Amer.

Vol. xviii, p. 429. 1884.

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NOTES AND LITERATURE.

BOTANY.

Recent Literature on Germination.— The germination of the seed

of Carapa guianensis Aubl. is described by Harshberger in Proceed-

ings of the Academy of Natural Sciences of Philadelphia, Vol. LIV

(1902), pp. 122-125, Pl. VIII. The seeds are without endosperm,

the reserve material of the cotyledons being oil, proteid bodies, and

protoplasm. The seeds would seem to lose their vitality more or

less quickly ; the period of germination is gone through quite rapidly,

the plants reaching a height of about a foot and having produced

several well-developed pinnate leaves in a month's time. They start

to germinate before falling out of the capsule; all may germinate 7z

situ and soon be so closely bound together by the intricately woven

mass of secondary roots that it is hard to separate them. The ger-

mination of Carapa guianensis and Xylocarpus granatum (= Corapa

maluccensis) is somewhat similar, the greatest difference being the

formation of a tuber-like radicle and pneumatodes in C. ma/uccensis,

a tree of mangrove habit.

Elenora Armitage finds (Mew Phytologist, Vol. I (June, 1902),

pp. 127—128), that seeds of Euphorbia peplus germinate after being

kept in a dry place for nearly nine years. She further shows that

the seeds of this species, an annual, will germinate as soon as sown.

The results are of interest in comparison with those of Winkler

(Berichte der deutschen botanischen Gesellschaft, Vol. T), who found

that seeds of Euphorbia cyparissias, a perennial, germinated first

from four to seven years after they ripened, and those of Æ. exigua,

an annual, nine years after ripening. She suggests that the so-called

resting period is usually a delay due to the presence of a hard testa

which requires moistening. It may be mentioned here that the seeds

of Nelumbium luteum, for example, germinate at once when an open-

ing is filed in the hard shell, but only after long delay when this is

left intact.

In the Journal of the Linnean Society, Vol. XXXV (July, 1902),

pp. 396—402, Prof. John Percival considers the origin of calcium

oxalate crystals in seedlings of Z+ifolium hybridum Linn., giving the

981

“oa

982 THE AMERICAN NATURALIST. [Vor. XXXVI.

results of a considerable series of experiments. In the same num-

ber of this journal Francis Darwin makes use of germinating Setaria,

Sorghum, and Vicia faba in experiments on gravitational sensitiveness

of the root tip.

A very interesting paper by Ethel Sargent, * Thé Origin of the

Seed-Leaf in Monocotyledons ” (Wew Phytologist, Vol. I, pp. 107—113,

Pl. II), gives some of the results of five years’ work on the vascular

system in monocotyledons. She points out the fact that theoretically

taxonomic characters drawn from the embryo ate entitled to special

attention in plants as in animals, but that so far the only important

strictly embryonic character of acknowledged value is that employed

in the fundamental division of the apparently well-defined group of

angiosperms. Concluding that a detailed study of the seedlings of

one family would yield results of greater value than a general study,

work for the last four years has been confined to the Liliacez, of

which seedlings of sixty genera have been collected and preserved,

but the detailed examination of which is not yet completed. She con-

siders that there is now no doubt of the systematic value of the indi-

cations given by the vascular structure of the cotyledon, hypocotyl,

and primaryroot. At the same time, these indications she considers

not easy of interpretation, since they are often wholly or in part

concealed by adaptive characteristics which are the effect of environ-

ment upon an organism at a time when it is so little differentiated as

to be extraordinarily plastic. In the hypocotyl she found the effects

of this influence the least marked, but even here the structure must

be affected by alternations in that of the cotyledon and root. In

Anemarrhena she found a type of vascular arrangement from which

others have clearly been derived, and that this form is truly symmet-

rical throughout. This type seems to be the starting point of at

least four central tribes in the Liliacez, and must therefore be a

form of some antiquity among monocotyledons. A comparison of

some Ranunculacez, especially Eranthis, Nigella, and Ranunculus

ficaria, of well-known monocotyledonous affinities, strengthened her

opinion that the Anemarrhena type is a dicotyledonous one, and to

conceive of steps by which two separate cotyledons may gradually

unite, is, she thinks, easier than to conceive of a single cotyledon

splitting into two similar members, as suggested by Lyon for Nelum-

bium. In conclusion, she says that there is no evidence of weight

for the superior antiquity of monocotyledons, and that in this group

the complete union of two cotyledons may possibly be due to their

AMI xs as an absorbing g organ. i A. Harris.

No. 432.] NOTES AND LITERATURE. 983

Leaves of aquatics. — McCallum! gives the results of experiments

designed to discover the factors determining the production of the

well-known air and water forms of leaves in Proserpinaca palustris.

After describing the form and anatomy of the two types of leaves,

he gives the results of his experiments, which seem to show that the

production of the submerged, much-dissected type, or the aérial,

simply serrate type, is not determined by difference in illumination,

nutrition, depth of water, temperature, stimulating influence of salts,

variation of concentration of CO, or O, or contact stimulus. When,

however, the plants were grown in a saturated atmosphere, a type of

leaf very close to the regular submerged type is produced, and when

plants grown under water are subjected to artificial transpiration

induced by a change in osmotic pressure secured by a solution of

mineral salts in the water, the air type of leaf was produced.

Further experiments, the results of which will be presented later,

are in progress. J. A. Harris.

Notes. — The American Journal of Pharmacy for September con-

tains an article on ZZyoseyamus muticus, by Nagelvoort, and an

account of the drug and medicinal-plant investigations in the

Department of Agriculture, by True.

In the American Journal of Science for September, Fernald dis-

cusses the relationship of some American and Old-World birches;

Sellards describes the fertile fronds of Crossotheca and Myriotheca,

and the spores of other Carboniferous ferns, and also the validity

of Jdiophyllum rotundifolium, a fossil plant from the Coal Measures.

The Bulletin of the Torrey Botanical Club for August contains the

following articles: Slosson, “ The Origin of Asplenium ebenoides ”

(including the results of artificial hybridization of Camptosorus rizo-

phyllus and Asplenium platyneuron, and confirming the assumed

hybrid nature of 4. ebenoides, except that the artificial hybrids have ¢

not yet been made to fruit); Evans, “ Hepatice of Puerto Rico"

White, * The Saltatory Origin of Species"; and Eastwood, “ New

Western Plants."

.In Engler's Botanische Jahrbücher of August 29 are published the

conclusion of Miss Perkins's “ Beiträge zur Kenntnis der Stryraca-

ceæ”; a monographic synopsis of the genus Lisianthus, by the same

1 McCallum, W. B. On the Nature of the Stimulus causing the Change of

Form and Structure i i palustris, Botanical Gazette, vol xxxiv

(August, 1902), pp. 93-108, Figs. 1-10.

984 pur fo AMERICAN NATURALIST. [Vor. XXXVI.

author; a paper on Nectria moschata, by Glück; one on the anatomy

of Anonaceez, by Beyer; a study of Berberidacem, by Tischler;

a supplement to Miss Perkins’s monograph of Monimiacez; a paper

by Hennings, on the fungi of Japan ; a discussion of the wind as

a factor in plant geography, by Warming ; and a paper on the geo-

graphical distribution of mosses in middle Europe.

The Beiblatt, No. 70, of the same number, contains Dr. Clements's

paper on a system of nomenclature for phytogeography, read in

Denver last year, with concise comments by Professor Engler, and

No. 4 of Urban's “ Plantze nove americanz, imprimis Glaziovianze,"

dealing with palms, and written by Dammer.

The Ottawa Naturalist for September contains a paper on the

arboretum and botanic garden at Ottawa, and a short account of

the plants at Norway Bay, Bristol, both by W. T. Macoun.

Professor Greene, who had thought of discontinuing /7ttonia at

the conclusion of the fourth volume, has decided differently, and

under dates of September 9 and September 18 issues the first fifty-

six pages of the fifth volume. These two installments include the

following papers: “New or Critical Species of Acer," “A New

Study of Microseris"" ‘Some Phacelia Segregates," ‘ Segregates

of Viola canadensis,” “Some New Acaulescent Violets,” * Revision

of Romanzoffia," * Revision of Capnorea," * New Species of Cryp-

tanthe," and * A Fascicle of New Compositz.”’

Torreya for September contains the following articles: Kirkwood,

“The Vegetation of Northwestern Oregon”; Peirce, “ Extrusion of

the Gametes in Fucus”; Lloyd, * Mutual Irregularities in Opposite

Leaves"; Earle, *A Key to the North American Species of Lac-

tarius " kai, “The Pith Cells of Phytolacca decandra” ; and

Wobei: “A New Station for Zsofria affinis."

K. K. Mackenzie and B. F. Bush have issued a manual of the

flora of Jackson County, Missouri, which includes descriptions of

1141 species and 5o additional varieties, contained in 500 genera.

This differs from the usual local floras in being a descriptive hand-

book, with keys to the higher groups and to the species of the

larger genera.

In the Sierra Club Bulletin for January Miss Eastwood gives an

account of Trinity County, California, with a list of the trees and

shrubs found in that county.

The shrubs of Wyoming form the subject of Bulletin 54 of the

Wyoming Experiment Station, written by Elias E. Nelson.

No. 432.] NOTES AND LITERATURE. 985

The Bulletin of the Geographical Society of Philadelphia for April

contains a report of the Brown-Harvard expedition to Labrador in

1900, and includes an extensive account of the botany of the expe-

dition, with lists of the plants collected.

The West American Scientist for August, selling for ten cents, and

Mr. Orcutt’s Review of the Cactacee for the same month, selling for

twenty-five cents, consists of the same signature of cactus notes.

A photogram of Echinocactus cylindraceus is contained in the

August number of the Monatsschrift fur Kakteenkunde.

Die Gartenwelt of August 23 contains an article on noteworthy

trees of Hanover, illustrated by a number of pictures of trees grow-

ing under unusual conditions and displacing objects in their way.

Fasciation in the leaves of Auonymus japonicus is described by

De Camps in the Memorias de la Real Academia de Ciencias y Artes

de Barcelona, Vol. IV, No. 20.

The exhibition of botanical objects under the microscope in

museums is the subject of an illustrated article by Howe in the

Journal of the New York Botanical Garden for September.

The Yearbook of the Department of Agriculture for 1901, in addi-

tion to the usual administrative and statistical portions, contains

a large number of economic articles, many of them of botanical

interest.

The many publications of the U. S. Department of Agriculture,

including those issued by the Patent Office, have been listed by the

division of publications of the department in a recently issued bul-

letin, which will prove very serviceable to libraries possessing sets

of these publications. A catalogue of the botanical publications

in the library of the same department, by Miss Clarke, constitutes

Library Bulletin No. 22 of the department.

An interesting item in Advance Sheets of Consular Reports of

August 30 shows that during July one hundred and eighty thousand

bunches of bananas were shipped from Puerto Cortes, Honduras,

to New Orleans and Mobile, and one hundred thousand additional

bunches were paid for and destroyed by the contractors.

An economic study of berseem (Trifolium alexandrinum), much

cultivated in the Nile valley, forms the subject of Bulletin No. 23

of the Bureau of Plant Industry of the U.S. Department of Agri-

culture, written by Fairchild.

986 THE AMERICAN NATURALIST.

Nos. so and 51 of the Contributions from the Cryptogamic Labo-

ratory of Harvard, published respectively as Nos. 2 and 3 of

Vol. XXXVIII of the Proceedings of the American Academy of

Arts and Sciences, are: * Preliminary Diagnoses of New Species of

Laboulbeniacez,” V, by Thaxter; and a paper on Cawuloglossum

transversarium, by Johnston.

A monograph of the Acrasiez, by E. W. Olive, forms No. 6 of

the current volume of the Proceedings of the Boston Society of Natural

History. 'This is another of the foundation-laying studies from the

cryptogamic laboratory of Harvard.

A biographical sketch of Marc Micheli, with an excellent portrait,

has been distributed by M. de Candolle, from the Archives des Sci-

ences Physiques et Naturelles for July.

CORRESPONDENCE.

Editor American Naturalist :

In the July number of the American Naturalist, among his “ Notes

on the Ccelenterate Fauna of Woods Hole," Dr. C. W. Hargitt has

defined: what he considers to be a new species of tubularian hydroid,

under the name of Tubularia parasitica, During the past summer

I have been enabled to observe the development of the western rep-

resentative of the genus Corymorpha, C. palma, a few facts concerning

which will show, I think, that 7: parasitica is but a young form of the

pendula on which it was growing. It has seemed best that attention

should be called to this at once to prevent future inconvenience to

taxonomists.

As with C. pendula, the meduse of C. palma are permanently

attached. The eggs arise on the manubrium, break through the

ectoderm when ripe, and, in quiet water, settle at once to the

bottom. The egg case is adhesive, fastening to the first object

with which it comes in contact. Many eggs are dropped at the

same time, and often cling to the rootlets of the parent hydroid,

where 7: parasitica was found. There is no free larval stage. The

embryo is able, however, to change its location very slowly, leaving

a trail of perisarc behind it. Such a movement accounts for the

clusters of six or eight individuals which are commonly found adher-

ing to each other near their bases. The stem has a single central

canal at first, as in Tubularia. The peduncles: which support the

medusz appear very early. When there are not more than eight

proximal tentacles, buds arise on the base of the hydroid and develop

into rootlets. They are the structures, I suspect, which Dr. Hargitt

has taken for “absorbent organs."

The young Corymorpha further agrees with 7: parasitica in size,

number of tentacles, and general aspect.

I shall describe the development of C. palma more fully in another

conpanhia, Harry Beat Torrey.

ZOOLOGICAL LABORATORY, UNIVERSITY OF CALIFORNIA,

BERKELEY, CAL, Sept. 5, 1902.

PUBLICATIONS RECEIVED.

AUERBACH, FELIX. Die holdin n ihr Schatten. Jena, Fischer, See

8vo, 56 pp. 1.20 marks. — BASTIA .C. Studies in Heterogenesis.

London, Williams & Norgate, or i 63-147, xi-xvii, Pls. VI-X. ec

CUNNINGHAM, J. T. Sexual Dimorphism in the Animal Kingdom. A Theory of

the Evolution of Secondary Sexual Characters. London, Adam & Charles Black,

1900. Svo, xi+ 317 pp» 32 figs. — ELroT, IDa M., and SOULE, CAROLINE G

Clodi Dll id their Moths. New York, Century Company, 1902. 8vo, xiii +

300 pp. 80 figs. $2.50. — GELERT, O., and OSTENFELD, C. H. Flora Arctica.

Containing Descriptions of the Pacak Plants and Ferns found in the Arctic

Regions, with their Distribution in these Countries. Pt. i, Pteridophyta, Gym-

nosperma, and Monocotyledons. Copenhagen, 1902. 8vo, xi + 134 pp- 95 figs. —

International Catalogue of Scientific Literature. First Annual Issue, vol. i, M,

Botany, pt. i, 378 pp.; vol. ii, D, Chemistry, pt. i, 468 pp. London, Harrison

& Sons, 1902. 21 shillings each vol. — LAUNER, A. Naturlehre. Wien, Jos. Roth,

1902. ded ii + 377 pp» 377 figs. — WILLEY, ARTHUR. Zoólogical Results based

on Material from New Britain, New Guinea, Loyalty Islands, and elsew here, col-

lected during the years 1895, 1896, and 1897. Pt. VI. Cambridge, University

Press, 1892. Pp. 691-830, Pls. LXXVI-LXX XIII, and text-figs.

ALLEN, E. T. The Western Hemlock. U. S. Dept. Agr., Bull. Bureau For-

estry, No. 33- 55 pp.» 13 pls.— ig ei E. W. Notes on the Phylogeny of Lirio-

dendron. Bot. Gaz. Vol. xxxiv, pp. 43-63. — BERRY, E. W. Liriodendron cel-

akovskii Velen. Bull. Torrey Bot. C. Vol. sana pp- pr — BERTRAND,

C. E., and CORNAILLE, F. Les t asmondéennes

et Urbain Proc. Verb. Soc. d’ Hist. Nat. d'Autun. 1902. 15--8 pp.2pls.—

BEUTENMULLER, W. Description of Some Larvz of the Genus Catocala. Bull.

Amer. Mus. Nat. Hist. Vol. xvi, pp. 381-394. — CARD, F. W. Improving an

Mosses of Alaska. Proc. Wash. Acad. Sci. Vol. iv, pp. 293-372, Pls. XIII-

XXIII.— CarkiNs, F.C. A Contribution to the Er of the John Day

Basin. Univ. Cal. Pubs., Bull. Dept. Geol. Vol. iii, No. 5, pp. 109-172, Pl. XVI.

— CoKER, R. E. Notes on a Species of Barnacle pies Parasitic on the

Gills of Edible Crabs. Bull. U. S. Fish Com. for rgoo. Pp. 399-412. 14 figs. —

CoquiLLETT, D. W. New Diptera from North P perum Proc. U. S. Nat. Mus.

Vol. xxv, pp. 83-126. — DELLABARRE, E. B. Report of the Brown-Harvard Expe-

dition to Nachvak, Labrador, in the Year 1900. Bull. Geograph. Soc., Phila. Vol.

iii, No. 4, pp. 65-212. Plates and maps. — Dyar, H. G. Descriptions of the

Larva of some Moths from Colorado. Proc. U.S. Nat. Mus. Vol. xxv, pP- 369-

412. — ECKEL, E. C. The Utilization of Iron and Steel Slags. U. S. Geol. Sur-

vey (Mineral Resources of the United e Mani 17 pp. — EtceN MANN, C. H.

The Solution of the Eel Question. 7) Amer. Mir: Soc. Vol. xxii

pP. 1-18. 4 pls. — EIGENMANN, C. Mi The Eyes of Rhincura floridana.

PUBLICATIONS RECEIVED. 989

Proc. Ind. Acad. Sci. for 1901. Pp. 106-107. TIP H. a maa

of the Eye of Amblyopsis. Proc. Ind. Acad. Sci. for 1901. Pp. -IO

EvERMANN, B. E. Description of a New Species of Shad (Alosa SER,

with Naa on other Food-Fishes of the Ohio River. U.S. Fish Com. Rep. for

rgor. Pp. 273-288. 6 figs. — FARRINGTON; O. C. Meteorite Studies, I. Publi-

cations px Columb. Mus., Geol. Ser. Vol. 1, No. 11, pp. 283-323, Pls. XLIII-

XLVI. 6 text-figs. — FELT, E. P. Elm Leaf Beetle in New York State. Budi.

N. Y. State Mus., No. 57. Pp. 1-43. 8 pls., 2 text-figs. — FoLEv, J. A. Working

Plan for Southern Hardwoods and A Results. Year Book U. S. Dept. Agr. for

Igor. Pp. 471-476, Pls. LX V-LXVII. — Fox, W. F. A History of the Lumber

Industry in the State of New ro di S. Dept. Agr., Bull. Dept. of Forestry,

No. 34. 59 pp. 19 pls. — FLEMING, B. P. The Measurement of Water for Irri-

gation. Bull. Wyo. Agr. Exp. Sta., No. 53. Pp. 59-117. 7 figs. — FLEMING, B P.

The Measurement of Water for Irrigation. Bull. Wyo. Agr. Exp. Sta., No. 53.

Pp. 58-117. 8 figs. — GARCIA, F. Spraying Orchards for the Codling Moth. Bul.

New Mex. Agr. Exp. Sta., No. 41. 26 pp, 1 p. — GEAN, R. S. A List of the

Publications of the United States Museum (1875-1900), including the Annual

Reports, plate Bulletins, Special Bulletins, aps Circulars with Index to

Titles. 7. U..S. Nat. Mus. N . 168 pp.— Hatt, H.M. A Botanical

Survey 2 vse a Univ. Cal. Pubs. Botany. Vol. i, pp. 1-140, Pls. I-XIV.

— HaLL, H. M. A Botanical Survey of San Jacinto Mountain. Univ. Cal. Pubs.,

Botany. Vol. i, pp. 1-140, Pls. I-XIV. — HarL, W. The Timber Resources

Pp. 207-216, Pls. IX-XIV. —

moth Cave, Kentucky. Proc. U. S. Nat. Mus. Vol. xxv, pp. 223-236. — HEIL-

PRIN, Defense of the Panama Route. Philadelphia, Franklin Press, 1902.

8vo, 12 pp. — HERRERA, A. L. Le rotoplasma de métaphosphate de chaux.

Mém. et Rev. Soc. Sci. Antonio Alzate. Tone xvii, pp. 201-213. 8 figs. — HERRICK,

E.C.. The psc cuti Period in the Lobster. Bull. U. S. Fish Com. for 19or.

pea 161-166. 5 figs. — x, B.H. Fertilizers, Part I. Bull. W. Va. Agr. Exp.

„ No. 80. Pp. EPEN AE I. Studies on the Hexactinellidæ. Contri-

rtm IL. (Ihe Genera Carlitella and . Heterotella.) Journ. Coll. Sci. Imp.

Univ. Tokyo. Vol. xvii 34 Pp» ! pl. — IKEDA, S. Contributions to the

Embryology of Amphibia: The Mode of Blastopore Closure a e i

of the Embryonic Body. Journ. Coll. Sct. Imp. Univ., Tokyo. Vol. xvii. 9° pp.

4 pls. — JorDAN, D. S., and FowLER, H. W. A pui of the Meier pg

M of Japan. Proc. U. S. Nat. Mus. Vol. xxv, pp- 75-79. — JORDAN, D. S.,

and Fow.er, H. W. A Review of the Cling Fishes (Goblescidn) of the Waters

of Japan. Proc. U. S. Nat. Mus. Vol. xxv, pp- 413-416. — JORDAN, D. S., and

FowLER, H. W. A Review of the Trigger-Fishes, File-Fishes, and Trunk-Fishes

of Japan. Proc. U. S. Nat. Mus. Vol. xxv, pp- 251-286. 6 figs. — Jorpan, D. S.,

H. W. Notes on Little Known Japanese Fishes, with Description

of a New Species of Aboma. Proc. U. S. Nat. Mus. Vol. xxv, pp- 573-576-—

Jorpan, D. S., and FOWLER, H. W. A Review of the Chætodontidæ and Related

Families of Fishes found in the Waters of Japan. Proc. U. S. Nat. Mus. Vol.

xxv, pp. 513-563. 6 figs. — Jorpan, D. S., and SNYDER, J. O. Descriptions of

Two New Species of Squaloid Sharks from Japan. EA

: 5 met rS icta D. S., and SNYDER, jJ.

of Fishes confused with Bryostemma Polyactocephalum. FS U. S. Nat. Mus.

Vol. xxv, pp- et jie. D. S., and SNYDER, J. O. A Review of the

990 THE AMERICAN NATURALIST. [Vor. XXXVI.

Blennoid Fishes of Japan. Proc. U..S. Nat. Mus. Vol. xxv, pp. 441-504. 28 figs.—

KiNGSLEY, J. S. The Cranial Nerves of Amphiuma. Tufts College Studies, No. 7.

(Scientific Series.) Pp. 293-321. — KINGSLEY, J. S. The Systematic Position of

the Cecilians. Z7u/zs College Studii, No. 7. (Scientific Series.) Pp. 323-344.

2 pls. — KOEHLER, R., and BATHER, F. A. .Gephyrocrinus grimaldii. Crinoide

nouvreau provenant des Ls de la Princesse Alice. Mém. Soc. Zool., France.

Tome xv, pp. 68-79. 4figs. — LAMB, A. B. The Development of the Eye Mus-

cles in Acanthias. Tufts a Studies, No. 7. (Scientific Series.) Pp. 275-

292. gs.— MASTERMAN, A. T. The Early Development of Cribrella oculata

(Forbes). With Remarks on Echinoderm Development. Zyans. ecl Soc., Edin-

sme Vol. xl, pt. ii, pp. 373-418. 5 pls. — MEARNs, E. A o Qialoi Cats.

Proc. U. S. Nat. Mus. Vol. xxv, pp. 237-249.— MEEK, S. E. c Contribution

to ita Ichthyology of Mexico. Pubs. Field Columb. Mus. (Zool Ser.) Vol. iii,

No. 6, pp. 63-128, Pls. XIV-XXXI. — MrenRIAM, C: H. Six New Skunks of the

Genus Conepatus. Proc. Biol. Soc., Wash. Vol. xv, pp. 161—165. — MERRIAM,

J.C. Triassic me por from California and Nevada. Univ. Cal. Pubs.,

Bull. Dept. Geol. Vol. iii, No. 4, pp. 63-108, Pls. V-X VIII. — MERRIAM, C. H.

Four New ir Foxes. pole Biol. Soc., Wash. Vol. xv, pp. 167-172. — MILLER,

G. S., Jk. Two New Malayan Mouse Deer. Proc. Biol. Soc. Wash. Vol. xv,

pP. 173-175. — NEHER, E. M. The Eye of osse antrorum. Proc. Ind.

Acad. Sci. for 1gor. Pp.96-101. 6figs.— NELSON, A. Native Vines in Wyoming

Homes. Bull. Wyoming Agr. Exp. Sta, No. e I5 pp., 11 pls. — NELSON, A.

The Genus Hedysarum in the Rocky Mountains. Proc. Biol. Soc., Wash. Vol. xv,

pp. 183-186. — NELSON, E. E. The Shrubs of Wyoming. Bull. Wyo. Agr. Exp.

Sta., No. 54. 47 pp, 15 figs. — NELsoN, E. W. A New Subspecies of the Cuban

Cliff Swallow. Proc. Biol. Soc., Wash. Vol. xv, p. 211. — NEWCOMBE, F. C.

The Rheotropism of Roots. Zot. Gaz. Vol. xxxiii, pp. 177-362. 15 figs. — OBER-

HOLSER, H. C. Some New South American Birds. Proc. U. S. Nat. Mus. Vol.

XXV, pp. 59-68. OBERHOLSER, H. C. List of Birds collected by William T.

Foster in Paraguay. Proc. U. S. Nat. Mus. Vol. xxv, pp. 127-147.— OLMSTED,

A A Working Plan for Forest Lands near Pine Bluff, Arkansas. U. S. Dept.

Agr., Bull. Bureau Forestry, No. 32. 48 pp., 9 pls. — PILSBRY, H. A., and Va-

NATTA, E. G. Papers from the Hopkins Stator Galapagos Expedition. XIII.

Marine Mollusca. Proc. Wash, Acad. Sci. Vol. iv, pp. 549-560, Pl. XXV.—

PoLLARD, C. L. Two New Violets from ab Eastern United States. Proc. Biol.

Soc., Wash. Vol. xv, pp. 201-203. — POLLARD, C. L., and CocKERELL, T. D. A.

Four New Plants from New Mexico. Proc. Biol. Soc, Wash. Vol. xv, pp. 177-

179. — PREBLE, E. A. Descriptions of New Species of Synaptomys and Phena-

comys from Mackensie, Canada. Proc. Biol. Soc., Wash. Vol. xv, pp- 181-182.

— RATHBUN, Mary J. Papers from the Hopkins-Stanford Galapagos E

tion (1898-1899). VIII. Brachyma and Macrura. Proc. Wash. Acad. Sci. Vol. iv,

Pp. 275-292, Pl. XIL — REESE, A. M. Structure and Development of the

. RICHMOND, C. W. Descriptions of Eight New Birds from Islands off the West

Coast of RESET Proc. Biol. Soc., Wash. Vol. x, pp. 187-190. — RICHMOND,

CW. | ns m Tuong, Lower Siam. Proc. Biol. Soc.

Wash. Vol. xv, pp. 157, 158. — Ripcaway, C. B. Experiments in Evaporation.

No. 432.] PUBLICATIONS RECEIVED. 99I

Bull. Wyo. Agr. Exp. Sta., No. 52. Pp. 45-55. 1 pl— ROTH jd Grazing

in the Forest Reserves. Year Book U.S. Dept. Agr. Jer Igor. Pp. 333- oe

Pls. XXXII-XXXIX. — RUSSELL, FRANK. Know Then Thyself. bos, Am

: - XV, pp.

graph of the Lepidopterous Family Noctuide of Boreal North America. A

' Revision of the Moths referred to the Genus Leucania, with Descriptions of New

Species. Proc. U. S. Nat. Mus. Vol. xxv, pp. 159-209, Pls. V-VI. — SMIT TH,

H Notes on Five Food Fishes of Lake Buhi, Luzon, Philippine Islands.

Bull. U. S. Fish ue for rgor. Pp. 167-171, Pl. XXII. — SNoncnass, R. E.

and HELLER, E. Papers from the Hopkins-Stanford Gohida pen

(1898-1899). XI. The Birds of Clipperton and Cocos Islands. Proc. Wash.

Acad. Sci. Vol. iv, pp. 501-520. — STARKS, E. C. The menor and Oste-

ology of the Caproid je or KAFES Proc. U.S. Nat. Mus. Vol. xxv,

pP. 565-572. 3 figs. — STE Rr The Reptiles of the unie Mountains,

Arizona. Proc. U. SS. pes jm Vol. xxv, pp. 149-158. — TassiN, W. The

Casas Grandes Meteorite. Proc. U. S. Nat. Mus. Vol. xxv, pp. 69-74, Pls. I-IV.

— TiNsLEY, J. D.. Alkali. Bull. New Mex. Agr. Exp. Sta., No. 42. 31 pp.—

TINSLEY, J. D. Drainage and Flooding for the Removal of Alkali. Bull. New

Mex. Agr. Exp. Sta., No. 43. 29 pp. — Torrey, H. B. Papers from the Harri-

man Alaska shit on. XXX. Ane emones, with Discussion of Variation in

Metridium. Proc. Wash. Acad. Sci. Vol. iv, pp. 373-410, Pls. XXIV-XXV.

16 text-igs. — ULKE, H. A List of the bates of the District of Columbia.

Proc. U. S. Nat. Mus. Vol. xxv, pp. 1-57. =~ VENDO, K. m linze Verz Japon-

icze. Journ. Coll. Sct. Imp. Univ., Tokyo. Vol. xvi. 36pp.,7 pls. — MATSUMURA,

J. Revisio Alni Specierum Japonicum. /ourn. Coll. Sci. poA ym 4, Tokyo. Vol.

xvi. 19 pp. 3 pls. — YOSHIWARA, S., and IWASAKI, J. Notes on a New Fossil

Mam Tarn Coll. Sct. Imp. Univ., Tokyo. Vol. xvi. 12 pp., 3 pls.

fais e la Comision de Parasitologia Agricola (Mexico). Tome i, Nos. 1-8.

1900-1902. gcn del Instituto Geologico de n No. 15, pt. ii. — Boletin de

la Sociedad Aragonesa de Sciencias Naturales. Tomei, No.2. Febr — Buli.

Ans Hopkins Hospital. Vol. xiii, Nos. 136-139. — Ann. Rep. Essex Institute,

year = hig $, 1902. — rag of Geography. Vol. i, Nos. 5-6. May-June.

— La C Popular. Año ii, No. r. (Mexico) April.— 27th Ann. Rep.

Mo. Bot. pnus 133 pP» ipe egi nes Ann. Rep. Mo. Horticultural Soc., 1901.

407 pp, illustrated. — 7he Vol. v, Nos. 24-51. — Buli. N. Y.

State Mus., No. 55, a ^ report of the State Entomologist. Pp. 699-925.

i xii. October, 1901. — Proc. Nat. Seci.

Revista Chilena de Historia Natu No. 2. — Science Gossip. (New

) a

Vol. viii, No. 95.

(No. 431 was mailed November 18.)

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