INDEX.

PAGE

ABNORMAL frogs + 25) 33

ee sepa gems ` columns in

VOTAR

Abnormal tortoise 17

Adams C. Base leveling a and

its faunal significance . 839

Æthop . 226

Alce, Osteology and position ahs 541

Allen, G. M adeer . . 449

Allen, jj A: nae ane mam-

— Two > papers o on American mam-

, 21

Alligator eggs, Incubation of ~ . 193

Am un Breeding of - OII

Amphio : 320

obse T Hammar's layer 55

Vitality of eggs and sperm . 56

——- What holds blastomeres to-

ther

Annelids - í . 322

Variation and regulation i in . 563

Antarctic gs of . 139

Ant-guest ; 1007

Anthropology 397; 499, 853

1555 Fe

thropometry .

inti-bodies i in blood . 927

i : 877

43h ES 701, x

> > > > > D> > D >

Pe Ea 5

pi.

ocotyle

laatia, Spermatogenesis ‘of 919

\uks, Osteology o i $41

iust n mamm , 245

M geben y i 887

BACULITES, Larval coil of 39

Baker, F.C. Mollusca of Genesee

iver. “6

Bangs, O. - - Mamunals from qe

—— Mammals of Liu Kiu Island 36%

Banks, N. Flies as carriers of

disease

— Synopsis of North American

Phalan

— nene: i faunal distribu-

> . 839

aec t S. Moses and PF.

leder. ^3 A. Ancestry al Marsu-

pialia I17

— Origin and Maii of Aus-

tralian mars

Bigelow, M. A PK ng Whitney,

and Lucas’ "Studies of anim

ife

ood, Formation 1 of anti-bodies i in CAD

Blood relationship, ~~ ental

proo

Botany .

. 245

.- 66, 322, 41 F 503, 687, 784;

866, 944

Brain, Segmentation . 4

De D: Ameiuru - QII

Sic sinl Diptera (Pho-

iy æ) fr m Tex

Bullhead, Dresda of

. 9II

Ee pad carbonate in animals . . 687

. Protozoa from

men Yor

Cambarus, Habits of .187

Campbell, D. H. New book on fos-

sil plants

Castle, W. E. Artificial Partheno-

geneses . 233

bie mega endi 1: =. Plants used

by z " I

Chau 455

Child, c M. "Habits and natural

: 975

— of Stichostemma .

Clark, H. L. Synopsis _of North

American holothuria . 479

—— T.D. As Cotión. ameri-

. 940

-= ey el æ of Brazil 63

—— Coccidae stanfordiana . 940

Colletti: + 220

Collins, F. S. Recent t papers on

ge. . 687

Colors of c rayfis sh . -933

Commensal Öligochæta i : s

Commopte

Conant, H. S. The conchometer bós

AE 'onchometer . Rx 4108

Crayfish, Colors CON UE 933

iv INDEX.

PAGE

Daisy, Variation in ray flower 51

Dall, w. H. Hinge-teeth of bi-

valve 175

Dean, B. Figures of living Nautilus re

Notes on SEN nautilus

Death of Dr. Lutken e p

Deer, Louisiana . - 449

Desmognathus, Respiration : of 8

Didelphys 1A?

Diptera, Development of. 36

ra, New. ae Qi o Ur

DMESEOIDES- OS oie us SS EO

EASTMAN, vm e Dean's Paleonto-

logical no

adips "VE EA AA SE ies

Ecitomyia . 346

Ecitons of Texas 157

Eckel, E. C. Snakes of New York 1 51

53» 496

Editorials

Eigenmann, C. H. » md Cox, | p.o

es of saltatory variation . .. 3

Eisen, G. ichaelson's Oligo.

cheta 862

Eycleshymer, A. c. Breeding h hab-

. QII

s of Ameiur

FIL à

Fishes, F ossil ; 292

Friedenthal’s Experimental proof

of blood-relationship

GEOGRAPHICAL distribution and

base lev song ng . (os 539

lcs opha 2427

Goode, ators volume - 496

Hasirs and natural history of

Stichostemma: 5-2 . —. 1.975

Hargitt, C. W. Synopsis of Hy-

dromeduse . 301, 379 575

Harris; 2}, A. Habits of Cam

barus . 187

Henshaw, s. Bentenmüller s ; Mon-

ograph of Sesiidæ 783

—— Dickerson's Moths and but-

rflies 2 382

—— Howard’s Insect book . 861

Howard’s Mos is 784

exapod thorax i357

Hinge-teeth of bivalves 175

Hoffmann, R. Du Chaillu’s Word

of the great fore 57

— — Herrick's Haie life of wild

birds Sg » B5

——— Moos fo 58

Holothurians, ‘Synopsis su. ; 475

Hrdlicka, A. Dwight's Human

spines.

—— Manouvrier’s Anthropometry reed

PAGE

Hydromedusz, Synopsis of, 301, 379,

575

Hydrozoa, New names for . . 789

INDIANS, Plants used by. . . . 1

Indian worksho 213

cotta Methods in study of. 553

357

ora ‘

Invertebrates, ‘Synopses of:

Mid be spur 301, 379, 575

"Holot hur

XVI. Pili

. 669

XVII. Rotatoria .

rires H. S. Putter on ME

29to

—— "Significance of „Spiral swim-

organ

Sings of Rotato ori . 72

rete R. H. Three polymelous

irae D. s AM e error Corrected 226

Correction 4

— Evermann and Marsh's Fishes

. 401

of P R i ico

—— Fishes "of . 9041

—— diis on dishes. of tropical

Vost

gue Fowler’ s 5 Fishe

: 941

— Gill and Smith | on morin-

guoid 225

tm d Snyder on Japa-

317

tons sna Snyder’ s Check

list of fishes of Japan . 43

—— — Notes on recent d literature 226

Recent fish liter 81

—— Recent papers on Mii ei - 599

——— Report of Scottish fisheries

DMNEO 105-0 o Le MM

—— Transplanting Californiatrout 225

KELLOGG V. L hen adeno in

development of = : 3

t, W. J. Colors of cra rayfish 33

FER JS Schmeil’s Zoblogy 54

Koenenia I5

reda CA Sea beach at -—

Kuhlia .

LAMELLIBRANCH hi : 158

u Kiu Islands, Mammals s. 2 RGt

EMI PETRY Fi of . 1029

ovell, J. H Cub of northe ern

apetalous flower S.

MACRASP $5105

Mammals, Interrelations ob :

— u Kiu Islands . s MEN

INDEX.

PAGE

Mammals of Panama . . . . . 631

Manx cats, Inheritance in . . . 52

Marsu D eiue TH Oh 6 Sow aes

—— Origin . 245

Mauck, A. V Swarming and v vari- À

ation in myriapod 477

Meek, S. E. Osburn's Fishes of

Ohio IAN . 857

Microthelyphonia dic. POLS

Mollusca of Genesee River . 659

ma 66

Morgan, T. H Regeneration in

the egg, embryo, and adul - 949

un gie NEN and Sidon

de. » 477

hvu ophaga . ANY ueris did

Papaa ena flies Cos AP gr

NAUTILUS, Notes on living. . . 819

Needham, J. G. Miall and Ham-

mond’s Harlequin fly . . . . 319

New st from Jap 2

an «9

n the PI REA Naturii.

New York, Snakes

Nichols, L. Spermatogenesis of

sellus

Heller s "Catg-

No nh American Indian

yoJ« BE

ids of North American plants 503

vs Spee i

Nuttall, G. H. F cific anti-

bodies in bloo 7

Nutting, £3. rrections. in

nomenclature x yendo. . 781

OLIGOCHATA, Parasitic or com-

mensa oe ay E 905

Ophidia of New York . 151, 42

Ortmann, A. E. Origin of Ant-

arctic flora and fauna . 139

Osborn, H. L. Anatomy of Axo-

lotls (ovr 087

— Case of polydactylism cree 81

PADAMADRÀ . 5 3 tous BS

obotany . . 73, 509, 606

Paleontology . . 327, 418

ama, Mammals of... . . . 631

araponyx, Variation in . . . . 52

Patani Oligochaeta . - 905

Parker, €: H. Abnormalities in

tortoi 17

— Cope's Reptiles of North

i . 228

— yraa of pigment s 938

-Excretion in annelids 39

—— Hopkins’s Hospital bulletin . 779

— — Hydra and electric - 938

—— Korschelt and Heiders 1 Em.

bryology . - 403

Parker, xd e Loweg on skin abs

porcu « £48

iic DAN girdle of reptiles Ot

— dés lum the crustacean

—— Reighard and Jennings - on

—— Ribmann and Teiler's Human

physiology . «220

—— Schenk and Gürber's Physi-

ology . $

Schultze’s "Heart bets: in

DMBA ER oe aye AO

Sixtus’s Reptilian affinities of

ee ae eee

—— — on the intestine of

Cet (7

——— The neurone 62

— gs Strassen on ‘the tenira

Parypha, Daninn of 64

Peirce, G-J. ormal and intra-

dics. respiration 3

Penhallow, D.-P. Se ward" s Juras-

flo 3 tain

ora of Great Bri . 606

—— White on fossil plants . 509

Zeiller’s — € any - 509

Peters, A. W. ethods in study

In Mt: 3

Petrogra 423, 946

hy 235,

Phagocytosis | in dipteran "—

erde i Synopsis les

PME, SS 685, 939

Plants aed by Tadians : I

tylis

C. "X. Duplication of

parts and regulation in Nereis . 563

ne -— icd ics ONE

Pseudosc |a usa 4

Psyllo oh jd. a ze tS

Pterocles , Position or. I

Publications received, 85, 242, 429, Sil,

698, 873

ND, H. Friedent hal’s Ex-

perimental proof of blood-rela-

1017

fisco of gift s, appointments , re-

vit ri and deaths . 78, 332, Mo

Reese, A. M. Incubation of alli-

gator eg (uw T9

Regenerati Pk va 949

Regulation in Nereis . . . 563

Rehn, J. A. G. Application of

Didelphys marsupialis . $ 147

grenienn normal and intramo-

lecular . 46

INDEX.

PAGE

Review

Arnold’ s Sea d at ebb tide ie

Atkinson's American fungi 66

deccm of plant life . . 866

. 869

Bentenmüller on Sesiidz . 783

ergen's s Foundatio inak botany 784

Bittner’s Triassic fossils of

Siberi «330

Piaciiman’ s Primitive Algae and

Fdoeladteé 27250 2 v ee

Bohn's Evolution of pigment 938

xe Pg on vingi ater Algæ

of the Faer . 691

Pieds on ds characters « 235

Buller’s Spermatozoa of ferns 413

aullery on social Ascidians . 230

Check list of fishes of Japan . 943

Coleman's Redwood mealy-

"940

Comere’s Desmids of France. 687

pac North American rep-

228

Coulter and Rose's Umbel-

Ca us d ded ' Adirondack gneisses xd

Cyclope ag n hor

ticultu . 602

Dean's PUR ntological notes FT

oe North Am

cans of yesterday Sot

Dickerson's Moths and butter-

ies . 782

Diener’s Triassic of ‘Armenia in

Du worst World of the

great for ica ay

Dwight's Heiss spine oy

Evermann and Marsh, Fishes

P Pd Rico . à i

Fishes of Japan . 941, 942, 943

Fowler on tropical Pacific

shes » $17

— Fishes in the Philadel-

phia aeea my . + 941

Fraser's 8

Fürbringer "Pettortd region

of re 61

Gemmi | on vitality ‘of eggs

Gill and S Smith « on 'moringuoid ;

LI PRIN ui e el

Guaita’s Pedigree mouse-

breeding AL E . 682

Hammar's layer s <: $4

acies on rock diagrams . 236

Heller North poter HN

pies 2: 1 - 3 4. mM

PAGE

Reviews:

Herrick's Home life of wild

birds . . 854

Hirn's Monograph of. Œdi-

M in A

Hobie 6 rock diagram 236

Holt. and Lee's Phiototactic

respon - 685

Ho pe sg Hospital bulletin F790

pror cs Sedimentar ty-

ocks 2

Howard’ s isect bee E . 861

of human excre-

men fu

—— Mosquit 784

Irving's Geology of the Black

: vun

Upper Paleozoic

fossils of Russia . . - 330

apan, Fishes of . , 942, 943.

jenen and Kellogg’ : "Animal

Jordan’ and Snyder’ s Japanese

Kemp's Calculation of rock

analyses + +» 947

—— Ha ndboo k of rocks foe MI

Kjellman on Galaxaura . .

Koorder's Flora of Celebes . 414

i :

orni

é scale in. Japan 940

Laneand riot rd’s olea)

of the superior region - 424

Looss’ Trematode fanha of

SYP - 407

Loweg on | skin of ‘porcupine . 248

Seri Indians

anonn on Anthropome-

Miall and Hammond’s Harle:

quin uy *

Michaelson’ s ; Oligochas ta 2

o land 39

of Alabama 944

and Parsons’s (€

ogy -

T€ s Graphic representa: |

on of rock analyses 427

Neumann s Revision of Txodi-

Nicholson’ s - Dictionary of

gpudesehip o < o 5 «7.9368

Nuttall on Anopheles . . . 860

Osburn's Fishes of Ohio "

—Ó Revision of ju cket

mice i

Reviews

INDEX.

PAGR

S: ; Review

Patterson’s Notes on Cero- What Me blastomeres to-

WOE coe ty qe

diss s Hydra and electric cur-

vele: Mitchell and Max-

well's ‘Plant - 324

Pieri and Winkler’ on. par-

thenogenesis 233

Pound and Clements’ s Phyto-

geography of Neb 600

Pratt's Origin of corundum . 946

Prentiss’ Crustacean M 95]

. deem on thigmotaxis

Quaintance and Scott’s s Coac-

ricanæ 40

Reighatds and Jennings s Anat-

597

— of Poe IR fisheries

boa 224

Retinána and Sellers’: s Human

physiology . 229

Rosenbusch on origin of Glau-

cophane schists 20429

Sargent's Trees and shrubs . 867

Scharff’s History of the Euro- |

pean fauna 87

Scheack es Gürber's s Human

physio . - 403

Scheels. By rox 854

Schultze's Heart beatsin Salpa 405

Scott's Studies in fossil 73

rd’s Jurassic flora of Great

Bub. — 2 3 v lod

Sixtus's Studies of Piae and

mammals . «279

Tbr rufi Brackish He

Spur a Ve Volcanic or of the”

Great Basin < 48$

Süssbach on intestine of Ce-

tac i81

Svedelius on Alge of the

Bal go

Tabley' s Flora of Chesh ire. 602

cse r's Mere. ad-

dre 37

Viewer n on n the Neuron 62

[EY of Magnet Cove,

+ 425

Watson's: Origin “of pheno-

crysts . '

Weathe ering of granites i

eisman parthenogene-

947

231

White c on fossil plants

m and Minne’s Esco

tion in ann

: Wolfthiigel’ s Perd helminths

Wright’s Flowers am ferns

Zeiller’s Paleobot

Zur Strassen on “posit

centrosome in — M

n H. M. Blackman

Ritter, W. E. As to social Ascid-

Rack composition, Representation

of .

ks, Sebemes of classification of

Roc

erage age

Rucker xan Eit

PME

— McGee’: s | ew Indian

—— North Americans of vircs

SAND grouse, Position o:

Sargent, F. L. Atkinson" s ` First

studies of plant life

— Bene E Foundations of bot-

any

—— Pepoon, ‘Mitchell, and Max-

t life

Scre steology of .

Ps gelben Sui measuring .

hrikes, Variation

Shufeldt, Ri W. ‘Osteology and

position of Alcz

T1 ology and position of the

creamers uu pu

Position of sand

Smith, z s Arthaber's Permian

of A *

— Bittner’ s ‘Triassic fossils oy

—— Jakolew’s Paleozoic fauna of

ussia .

—— Larval coil of Baculites

—— Pompeckj’s tod fossils

from Al

Snakes of New York

Stejneger, L. Eckel’s Snakes of

New York

ene io, d

er's Spermatozoa of ferns

and Clements’ s

ska . 60

PAGE

| B53.

sol

vill

Stejneger, I —— ade Ads

the Eur opean fau

Stethopathus ERIT 4 54

= eig 5

: 97

trong, R Va riationi in shrikes 271

n ofi inverte

XIV. Hydro Sdn: 301, 379, z 5

XV. Holothurians .

XVI. Phalan ngida ; $69

XVII. Rotatoria . : Nod

TAPE worms, Nervous system of . 321

akui on 4354

Thomas, O. Myrmecophaga and

Didel phys 3

Torsion, Instrument for measuring 299

Trelease, W. Bailey's Botan . 869

rad n of American Hor-

cultu : = + 602

— Flora of Ches hir . 602

— Koorder’s Flora of Cetebe es .

——Nicholson’s Dictionary of

gardening . . . 868

—— North American Pterido-

hytes 7

—— Plant life of Al abam : 04d

— Recent forestry subi alice 323

—— Sa hg emis d shrubs . 867

— Two mushro

oks «66

Toe s Arieta and ferns . 868

Tunicata 320

Me est bec in a myriapod : 477

—— in human spinal column :

land

Willoughby, = E.

INDEX.

PAGE

Vatlation in. Nereis < 5" . 563

—— in shrikes hay 271

51, 681

WALTON, L. B. Thorax - insects 357

Ward, H. B. Avian helminths

Looss’ Tiaa kina of

- 407

Recent work on n Anopheles . 860

—— Revision of Ixodidz

er, W. M. An n extraordinary

gest ...

—— Nests of nts

Parasitic origin

gates among a

Wheeler, W. M., and um w. 'H.

Males of some Texan ants . . 19

Wilder, aryngeo

phageal | dung id Desmognathus 183

Willcox, sitic or com-

mensal Oligichatai in New Eng-

431, E 701, 73!

. 877

Cii a a

/ Préhistoric

d Ad

workshop in Main

MEN ICHTHYS 270/127. 25»25 BA.

YERKES, R. M. Holt and vail

Phototactic response . . 68

ZOOLOGY 55, 221, 317, 401, 597, 779,

854, 940

=. AMERICAN —

=~ NATURALS]

E 6 _ A MONTHLY JOURNAL

: .DEVOTED TO THE NATURAL SCIENCES A

IN THEIR WIDEST SENSE | rd

DE. CONTENTS Pris

I. Plants Used by the Indians of Eastern North America : x 5

LUCIA 8. CHAMBERLAIN EET

II. On the Systematic Position 6 Mx tous THEIN es; T

: R. W. SHUFELDT

IH. orrelael Abnormalitis in tho Ste and Boy Platos ofthe et

of Recent poe € ur e ee ar' Tagen Ras

Wo he Great Forest, At and Others ie the Bou

x Correspondence: Identity at Xenichthys x xenurus mi Kuhlia malo

Professor D. $.

The American Naturalist.

ASSOCIATE EDITORS:

i A. ALLEN, Pu.D., American Museum of Natural History, cw York.

. A. ANDREWS, PH.D., Johns Hopkins University, Baltim

Se S. BAYLEY, PH.D., Colby University, Waterville.

CHARLES E. BEECHER, Pu.D., Yale University, New Haven.

DOUGLAS H. CAMPBELL, Pu.D., Stanford University.

J. H. COMSTOCK, S.B., Cornell University, Ithaca.

WILLIAM M. DAVIS, M. E., Zarvard — Cambridge.

ALES HRDLICKA, M.D., New York 2

D. S. JORDAN, LL.D., Stanford Univ.

CHARLES A. KOFOID, PH.D. d sedie of Illinois, Urbana.

J. G. NEEDHAM, Pu.D., Lake Forest University.

" ARNOLD E. ORTMANN, PH.D., Princeton University.

D. P. PENHALLOW, S.B., F.R.M. S., McGill ee Montreal.

H. M. RICHARDS, S.D., Columbia University, Ne

W. E. RITTER, PH.D, Unitor hity of California, (ege

FRANK RUSSELL, PH.D., Harvard University, Cambridge.

ISRAEL C. RUSSELL, LL. D., University of Michigan, Ann Arbor.

ERWIN F. SMITH, S.D., U. S. Department of Agriculture, Washington.

LEONHARD STEJNEGER, Smithsonian Institution, Washington.

W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, PH.D., University of Nebraska, Lincoin.

WILLIAM M. WHEELER, Px.D., University of Texas, Austin.

points of interest, editorial comments on scientific questions of the

day, reviews of recent literature, and a final nat eee for

scientific news and personal notices.

5e | who have anything eni to say are invited

ce send i in | their contributions, but the editors will endeavor to select

for publication. only that which i is of ee scientific vae and at the

same time written so as to be inte d interesting

to the general scientific reader. _ -

_ All manuscripts, books for 1 review, exchanges, = E should be

sent to THE AMERICAN NATURALIST, Cambri

Pe Feen communications should be sent direct to the

iption, $4.00, xet, ix advanos. Single copios, 35 conta.

C NEQU ME

THE

AMERICAN NATURALIST

Vor. XXXV. January, IQOI. No. 409.

PLANTS USED BY THE INDIANS OF EASTERN

NORTH AMERICA.

LUCIA SARAH CHAMBERLAIN.

THE following list of plants used by the North-American

Indians inhabiting the country east of the Mississippi River was

compiled during a course given to students of Radcliffe College

in 1899-1900, at the Peabody Museum, by Dr. Frank Russell

of the Department of American Archaeology and Ethnology of

Harvard University.

The linguistic groups of Indians included in this article are

two: the Algonquian, the greatest family: group and most

prominent at the time of the settlement of the country by the

whites; and the Iroquoian, who by their efficient organization

became very powerful in the midst of the Algonquian country.

The arrangement of tribes under linguistic groups is that

followed by the Report of the Bureau of Ethnology at Wash-

ington, D.C., for 1885-1886. The material was gleaned from

the Harvard College Library, the Boston Public Library,

the Boston Athenzeum, the Cambridge Public Library, and the

libraries of the Museum of Comparative Zoólogy and of the

Peabody Museum.

2 THE AMERICAN NATURALIST. [Vor. XXXV.

ALGONQUIAN FAMILY.

Algonquin. — Derivation: contracted from A/gomeguin, an Algon-

quian word signifying * those on the other side of the river," ż.e., the

St. Lawrence River.

TRIBES REPRESENTED.

Abnaki Menomine Pequot

Algonquin Miami Pottawotomi

Blackfeet Micmac Savannah

Delaware Narragansett Sax and Fox

Kickapoo Ojibway Shawnee

Abnaki.

Bayberry (18, p. 33) ! : the wax obtained was used with tallow for

candles. — Bean (9, p. 25): used as food.— Birch bark (9, p. 29):

the Abnakis used it to write upon. — Maize (9, p. 25): used as

food. — Reeds (18, p. 10): mats made of reeds served as chairs and

beds. — Squash (9, p. 25) : used as food.

Algonquin.

Birch (17, p. 310): bark used for sheathing the frame of canoes. —

Elm (17, p. 310): bark used for sheathing the frame of canoes

and the filaments of the bark used to sew the sheathing of canoes

together. — Fir (17, p. 310): the gum used to cover seams of

canoes. — Flag (11, p. 77): the leaves of sweet flag used to thatch

huts. — Maize (40, p. 58): used as food and the leaves used to

thatch huts. — Pine (17, p. 310): wood sometimes used in build-

ing frames of canoes. — Rushes (11, p. 77): used in making mats. —

Squash (11, p. 77): used as food. — Tamarack (17, p. 310): roots

used to sew the sheathing of canoes together. — Tobacco (11, p. 77):

chewed by the Algonquins. — Wild rice (26, p. 205): used as food.

Blackfeet.

Berries (17, p. 278): esteemed a delicacy when boiled in buffalo

blood. — Bull berries, Shepherdia argentea (16, p. 203): used as food.

— Camass root (4, p. 534): roasted bulbs used as food. — Choke-

cherries (16, p. 203): pounded up before eating. — Maize, Psoralea

esculenta (5, p. 205): used as food. — Red willow (4, p. 534): berries

! The figures in bold type refer to the bibliography at the end of the article.

No. 409.] PLANTS USED BY AMERICAN INDIANS. 3

used as food. — Sarvis berries, Amelanchier alnifolia (16, p. 203):

dried and stored for food. — Stork’s bill Hrodium cicutarium (17,

p. 422).

Delaware.

Dogekumak (7, p. 96): smoked with tobacco. — Maize (40, p. 58):

used as food; leaves used to thatch huts. — Rushes (11, p. 77):

used for mats, — Squash (11, p. 77): used as food. — Sweet flag

(11, p. 77) : leaves used to thatch huts. — Sumac (4, p. 534): leaves

smoked with tobacco. — Tobacco (11, p. 77): smoked. — Willow

(7, p. 96) : the bark of the red willow is mixed with tobacco.

Kickapoo.

Acorns (21, p. 265): used as food. — Beans (21, p. 265): used as

food. — Birch (21, p. 301): bark used for canoes. — Blackberries (21,

p. 265): used as food. — Corn (21, p. 265): used as food. — Cotton-

wood (21, p. 301): used for canoes. — Crab apple (21, p. 265): used

as food. — Dewberries (21, p. 265): used as food. — Gooseberries (21,

p. 265): used as food. — Gourds (21, p. 265): used as food. — Grapes

(21, p. 265) : many varieties used. — Groundnuts (21, p. 265): used

as food. — Hazelnut (21, p. 265): used as food. — May apple (21,

p. 265): used as food. — Melons (21, p. 265): used as food. — Osage

oranges (21, p. 265): used as food. — Peas (21, p. 265): used as

food. — Pecans (21, p. 265): used as food. — Plums (21, p. 265):

used as food. — Potatoes (21, p. 265): used as food. — Pumpkins

(21, p. 265): used as food. — Strawberries (21, p. 265): used as

food. — Sweet myrrh or anise root (21, p. 265). — Tobacco (21,

p. 265): smoked. — Walnuts (21, p. 265): used as food. — Whortle-

berries (21, p. 265): used as food. — Wild licorice (21, p. 265): used

as food.

Menomine.

Ash (32, p. 274): used in making bows, — Basswood (32, p. 258) :

fibre used in making mats; the inner bark is used in making string

and ropes; wood (32, p. 254) used in building houses. — Birch

(32, p. 254): bark used to cover the top and side of houses; also

for canoes. — Blueberries (32, p. 291): used as food. — Cat-tail flags

(32, p. 258): used in making mats. — Cedar (32, p. 258): bark used

in making mats; white cedar (32, p. 293) used in building canoes.

— Cherry (29, p. 52): wild cherry used as food. — Corn (29, p. 48):

used as food, — Crab apple (29, p. 52): used as food. — Currants

(29, p. 52): used as food. — Gooseberries (29, p. 52): used as food. —

4 THE AMERICAN NATURALIST. [Vor. XXXV.

Grapes (29, p. 52) : used as food. — Hazelnut (29, p. 52): used as

food. — Hickory (32, p. 274): used in making bows. — Hops (29,

P. 52): wild hops used as food. — Maple, Acer saccharinum (32,

p. 237): maple sugar made from. — Oak (32, p. 253): used in build-

ing houses. — Pine (32, p. 254): bark used in building houses. —

Plums (29, p. 52): used as food. — Pumpkins (29, p. 48): used as

food. — Raspberries (32, p. 291): used as food. — Rushes (32, p. 258):

used in making mats. — Snakeroot, Polygala senega (32, p. 292). —

Strawberries (29, p. 52) : used as food. — Tobacco (32, p. 253) : used

as food. — Whortleberries (29, p. 52): used as food. — Wild rice (29,

p- 47): used as food.

Miamoi.

Yellow lily, Zz/ium canadense (3, p. 312): roots used.

Micmac.

Apios tuberosa, Saa-ga-ban (4, p. 534): pear-shaped roots used as

food.

Narragansett.

Chestnut (6, p. 46): used for canoes. — Pine (6, p. 46): used for

canoes. — Whitewood (6, p. 46): used for canoes.

Ojibway.

Basswood (35, p. 236): used in making nets. — Birch (1, p. 9):

bark used for the exterior of canoes; characters traced upon the

inner surface of bark; these characters usually pertain to personal

exploits (27, p. 59). — Cedar (9, p. 116): bark used in making rope

or twine; used in making nets (35, p. 236). White cedar used to

make the hoops for canoes ; the roots used in sewing canoes (1, p. 9).

— Cherry (1, p. 9): gum used in putting canoes together. — Corn

(35, p. 236) : used as food. — Groundnut (24, p. 55): used as food.

— Pine (24, p. 73): wood used in making fire by friction. — Potatoes

(35, p. 236): used as food. — Red willow (24, p. 135): bark smoked.

— Root, Oduhpin (24, p. 55): used as food sometimes. — Spunk (24,

p. 73): wood used in making fire by friction. — Sumac (24, p. 135):

leaves smoked. — Swan potato, Wahbeziepin (24, p. 55): boiled and

eaten. — Tobacco (24, p. 56): leaves smoked. — Wild rice (3, p. 120):

used for food.

Pequot.

Corn (39, p. 4): used as food. — Indian hemp (39, p. 2): used to

make twine.

No. 409.] PLANTS USED BY AMERICAN. INDIANS. 5

Pottawotomi.

Beans (17, p. 82): used as food. — Maize (17, p. 82): used as

food. — Melons (17, p. 82): used as food. — Tobacco (17, p. 82):

leaves smoked.

Savannan.

Chinkapin nuts x p. 53): used as food. — Hickory nuts (25,

P. 53): used as foo

Sax and Fox.

Basswood bark (29, p. 126): twine obtained from it to bind rushes,

— Beans (29, p. 126): used as food. — Cane (14, p. 43): flageolet

made from it or of two pieces of soft wood hollowed out and fastened

together by strips of leather. — Corn (29, p. 126): used as food. —

Gooseberries (14, p. 29) : used as food. — Grapes (14, p. 29) : used as

food. — Melons (14, p. 44): used as food. — Nettle (29, p. 126):

twine obtained from the bark. — Onions (4, p. 534): used as food. —

Pecans (15, p. 20): used as food. — Plums (14, p. 29): used as

food. — Potatoes (4, p. 534): sweet and white potatoes used as food.

— Pumpkins (14, p. 41): used as food. — Rushes (29, p. 126): mats

made from. — Sap pine, Kee-chi-heyja-ka (22, p. 419): a healing gum

which the Sax and Fox always take with them when they travel.

Shawnee.

Apple, meshemenake (2, p. 291): used as food. — Beans, mzsoochethake

(2, p. 291): used as food. — Indian turnip, ¢-/aw-sho-ga (22, p. 413):

used with spikenard and wild licorice. — Maize (19, p. 14): used as

food. — Melons, »s£e/oma£e (2, p. 291): used as food. — Nuts, pacami

(2, p. 291): used as food. — Onions, shekagosheke (2, p. 291): used

as food. — Peaches (40, p. 17): used as food. — Peas (19, p. 14):

several kinds used as food. — Potatoes, meashethake (2, p. 291) : used

as food. — Pumpkins, wadego (2, p. 291) : used as food. — Squashes

(40, p. 17): used as food. — Tobacco (40, p. 57): leaves smoked. —

Wheat, cawasgue (2, p. 291): used as food.

IROQUOIAN FAMILY.

Iroquois. the adaptation of the Iroquois word

hiro, used to tnde a speech, and oné, an exclamation.

(Charlevoix.)

6 THE AMERICAN NATURALIST. [VoL. XXXV.

TRIBES REPRESENTED.

Cayuga Mohawk Seneca

Cherokee Oneida Tuscarora

Huron Onondaga Wyandot

Cayuga.

Buckwheat (36, p. 58): used as food. — Corn (36, p. 57): used as

food. — Oats (36, p. 58): used as food. — Potatoes (36, p. 58): used

. as food. — Wheat (36, p. 58): used as food.

Cherokee.

Apples (12, p. 11): used as food. — Beans (28, p. 69): used as

food. — Beggar’s lice (12, p. 11): tea made of it to assist the memory

“ since it clings so well." — Blackberries (12, p. 11): used as food. —

Chestnut (28, p. 69): a tuberous root used as food. — China brier

(23, p. 34): roots used as a blood purifier. — Cone flower (20, p. 197):

used as a wash for sore eyes. — Grapes (5, pp. 228—29) : used as food.

— Hemp (5, pp. 228—29) : used for cords. — Hoary pea, Tephrosia vir-

giniana (12, p. 11): an infusion of it used as a wash to strengthen

the body. — Hops (5, pp. 228-29): used as food. — Horehound (8

p. 235): wild horehound used for snake bite. — Lobelia, Zode/ia

cardinalis, cardinal flower (8, p. 41): a decoction of the root used.

Lobelia syphilitica, great lobelia (23, p. 34): a blood purifier. —

May apple, Podophyllum peltatum (23, p. 34): used as medicine. —

Nettle, Jatropha ureus, white nettle (23, p. 34): roots used as medi-

cine. — Nordica (23, p. 34): the juice of its white root used as

medicine, and its roots smoked. — Oats (12, p. 11): cultivated as

food. — Peaches (12, p. 11): used as food. — Plantain (5, p. 235):

wild plantain used for snake bite. — Potatoes (12, p. 11): used as

food. — Pumpkins (28, p. 69): used as food. — Purslane, Portulaca

oleracea (20, p. 197): used as medicine. — Rye (12, p. 11): cultivated

as food. — parilla, Panax ginseng (23, p. 34): used as medicine.

Sassafras (23, p. 34) : roots used as a blood purifier. — Snakeroot (5,

p. 235): fern snakeroot carried in the shot pouch as a remedy for

snake bite. — Squashes (28, p. 69): used as food. — St. Andrew's

cross (5, p. 235): used for snake bite. — Strawberries (13, p. 11):

used as food.

Huron.

38, p. 13): cultivated as food. — Birch (13, p. 73): white

birch bark used to cover lodges. — Blackberries (33, p. 143) : used as

No. 409.] PLANTS USED BY AMERICAN INDIANS. 7

food. — Corn (33, p. 168): used as food. — Gooseberries (33, p. 143):

used as food. — Grape (33, p. 153): used as food. — Hemp (38, p. 13):

fibre used for twine and cordage. — Lichen (33, p. 142): “ripe de

roche, boiled and used as food. — Pumpkins, or c/fru//es (33, p. 16 4)

used as food. — Squashes (38, p. 13): cultivated as food. — Tobacco

(38, p. 13): cultivated, leaves smoked. — Wild rice (13, p. 7 aye “the

stalk is woven into mats for the walls of the lodges”; fruit used as

food.

Mohawk.

Corn (36, p. 52): cultivated as food.

Oneida.

Corn (36, p. 43): cultivated as food.

Onondaga.

Apple, szwa-hu-na (10, p. 115). — Basswood, ho-ho-so (10, p. 116):

used to make fine strings and mats. — Beans, 00-sah-ha-tah (10,

p. 119): used as food. — Birch, ga-nah-jeh-kwa (10, p. 116): used

for canoes. — Blackberry, sa-he-is (10, p. 116): used as food. —

Butternut, oo0-ha-wat-fah (10, p. 119): used as food. — Cat-tail flag,

oo-na-too-kwa (10, p. 118): used for mats. — Cherry, /a-c-goo-nah

(10, p. 114): wood used. — Chestnuts, oheh-yah-tah (10, p. 116): used

as food. — Chokecherry, ne-a-tah-tah-ne (10, p. 114): used as food. —

Currant, ska-heus-shkah-he (10, p. 119): used as food. — Flax, vv0-skah

(thread-like) (10, p. 117): used to make thread. — Ginseng, da-hien-

too-keh (10, p. 115). — Gooseberry, sha-heus-skah-he-goo-na (10, p. 1 14):

used as food. — Grape, oA-Aeun-&we-sa. (10, p. 114): wild grape, used

as food; oh-heun-kwe-so-goo-no, cultivated grape, used as food. —

Hickory, a-ze£, a bitter nut. ws-ee& (10, p. 1 20). Oo-sook-wah,

common variety of hickory. — Huckleberry, o-/eah-che (10, p. 116):

used as food. — Lettuce, 00-na-tah-kah-te (10, p. 115): eaten raw. —

Maize, v0-ne-hah (10, p. 119): used as food. — May apple, o-na-when-

stah (10, p. 115) : used as food. — Melon, wah-he-yah-yees (10, p. 114) :

muskmelon, used as food. Oo-neah-sa-kah-te (10, p. 114) : watermelon,

used as food. — Peaches, oo-goon-wAy-e (10, p. 114) : used as food. —

Pear, žoon-de-soo-kwis (10, p. 114): used as food. — Peas, o-na-kwa (10,

p. 119) : used as food. — Peppermint, kah-nah-noos-tah (10, p. 116).

— Plum, £a-£a-£a£-ne (10, p. 118): wild plum, used for food. — Potato,

o0-neh-noo-kwa (10, p. 119) : used as food. — Raspberry, o-nah-joo-kwa

(10, p. 116): used as food. Tew-tone hok-toon (10, p. 116) : black

8 THE AMERICAN NATURALIST. [Vor. XXXV.

raspberry, used as food. — Rose, ah-weh-ha-tah-ke (10, p. 118): wild

rose, used as medicine. — Sarsaparilla, ju-ke-ta-his (10, p. 116). —

Snakeroot, o-skwen-c-tah (10, p. 120). — Squash, v0-neah-sah-oon-we (10,

p. 114) : used as food. — Strawberry, noon-tak-tek-hah-kwa (10, p. 114):

used as food. — Sumac, vot-koo-tah (10, p. 119): leaves smoked. —

Thimbleberry, o-nah-jo0-kwa-goo-na (10, p. 116): used as food.—

Tobacco, o-yen-kwa-hon-we, Nicotiana rustica (10, p. 118): leaves

smoked. — Turnip, 0-7e-kwa (10, p. 115) : used as food. — Wake-robin,

o-je-gen-stah (10, p. 117) ; white wake-robin ; medicinal use not known

to the Onondagas.

Seneca.

Apples (37, p. 18): used as food. — Ash (34, p. 49): used to

make baskets. — Basswood (37, p.20): leaves used. — Beach plums

(87, p. 18): used as food. — Beans (37, p. 18): used as food. —

Black walnuts (37, p. 18): used as food. — Corn (37, p. 18): used

as food; husks used to make baskets (34, p. 49). — Groundnuts

(37, p. 18): used as food. — Hazelnuts (37, p. 18): used as food. —

Hickory (34, p. 49): snowshoes made from the wood. — High-

betony (head-betony) (37, p. 20). — Mandrakes (37, p. 18): used

as food. — Maple (34, p. 48): sugar made from the sap. — Mul-

berries (37, p. 18): used as food. — Ooklthaw (37, p. 18): a root

used to make bread. — Peaches (37, p. 18): used as food. — Potatoes

(37, p. 18): used as food. — Rushes (34, p. 49): used to make

baskets. — Squashes (37, p. 18): used as food. — Sumac (34, p. 49):

leaves mixed with tobacco. — Tobacco (34, p. 49): leaves smoked.

$ Tuscarora.

Beans (36, p. 69): used as food. — Buckwheat (36, p. 69): used as

food. — Corn (36, p. 69): used as food. — Oats (36, p. 69): used as

food. — Peas (36, p. 69): used as food. — Potatoes (36, p. 69) : used

as food. — Turnips (36, p. 69) : used as food. — Wheat (86, p. 69):

used as food.

Wyandot.

Beans, yah-re-sah (2, p. 294). — Corn, nay-hah (2, p. 294). — Grass,

e-ru-ta (2, p. 294). — Melons, o-nugh-sa (2, p. 294). — Potatoes, da-ween-

dah (2, p. 294). — Pumpkins, o-nugh-sa (2, p. 294). — Weeds, ha-en-tan

(2, p. 294).

CAMBRIDGE, Mass, August 31, 1900.

No. 409.] PLANTS USED BY AMERICAN INDIANS. 9

LIST OF THE WORKS FROM WHICH INFORMATION

WAS OBTAINED.

An account of the North-American Indians written for MM e

daus, a chief of the Ojibways. Leicester, 1848.

American Antiquarian Society, Transactions and Collections. Vol. i.

STICKNEY, G. P. Indian Use of Wild Rice. Am. Anthropologist.

U. S. Census Report, 1890. Report of Indians taxed and not taxed in

the United States. Washington, 1894.

ADAIR, JAMES. History of the American Indians. London, 1775.

ALLEN, Z. The Conditions of Life, Habits and Customs of the

Native Indians of America. Providence, 1880

BANDELIER, A. F. Archzological Institute of America. American

Series, 1-5. Boston, 1881-92.

. Barton, B. S. Collections for an Essay toward a Materia Medica of

the United States. Philadelphia, 1798.

BAXTER, The Abnakis and their Ethnic Relations. Maine

Hist. Soc. Coll. 2d series. Vol. iii. 1892.

BEAUCHAMP, W.M. Indian Names in New York. Onondaga Names

of Plants. Fayetteville, 1893.

. BRINTON, D. G. he American Race. New York, 1891

. CARRINGTON, H. B. Eastern Band of Cherokees of South Carolina.

1892.

Cotton, C. Tour of the American Lakes and among the Indians of

the Northwest Territory. Vol.i. London,

DRAKE, B. Life of Black Hawk and Behe of the Sax and Fox

Indians. Cincinnati, 1839.

GALPIN, S. A. Report upon the Condition and Management of

Certain Indian Agencies in the Indian Territo

GRINNELL, G. B. Blackfoot Lodge Tales. New York, 1892.

. Harnes, E. M. American Indian. Chicago, 1888.

Hanson, J. W. Sketch of the Abnaki Indians (in the History of

Norridgewock, 1849).

. Harvey, H. History of the Shawnee Indians from 1681-1854

inclusive. Cincinnati, 1855.

. HensHaw, H. W. Who were the Indians? Washington, 1889.

. Hunter, J. D. Manners and Customs of Several Indian Tribes West

of the Mississippi. Philadelphia, 1823.

HUNTER, J. D. Memoirs of a Captivity among the Indians of North

merica. London, 1823.

Jones, C. C. Antiquities of the Southern Indians. New York, 1873. -

Jones, PETER. History of the Ojibway Indians. London, 1861.

Lawson, Joun. History of Carolina. London, 1714.

THE AMERICAN NATURALIST.

. Lone, Jonn. Voyages and Travels of an Indian Interpreter and

Trader. London, 1791.

. MALLERY, GARRICK. Pictographs of the North-American Indians.

Report of the Bureau of Ethnology, 1882-83. Washington, 1886.

MoRnGAN, T. H. Houses and House Life of the American Aborigines.

Washington, 1881. :

MORSE, JEDEDIAH. A Report to the Secretary of War of the United

States on Indian Affairs. New Haven, 1822.

PALMER, EDWARD. Notes on Indian Manners and Customs. Am.

Vat. Vol xii. 1878.

Plants Used by the Indians of the United States. Am.

fat. Vol. xii 1878.

. POWELL, J. W. The A Indians, Report of the Bureau of

Ethnology, 1892-93. Washington.

. Rapisson, P. E. The Voyages of Peter Esprit Radisson. Boston,

188

. SANBORN, J. W. Legends, Customs, and Social Life of the Seneca

Indians. Gowanda, N. Y., 1878.

. SCHOOLCRAFT, H. R. Indian Tribes of the United States. Vol. i.

Philadelphia, 1884.

Notes on the Iroquois. New York, bab:

. SWETLAND, LUKE. Captivity among the Senecas. Waterville, N. Y.,

1875.

THRUSTON, G. P. Antiquities of Tennessee. Cincinnati, 1890.

WHEELER, R. A. The Pequod Indians. Westerly, R. I., 1877(?).

WHIPPLE, A. W. T. Ewbank and W. W. Turner. Report upon the

Indian Tribes. Reports of he and Surveys from the

Mississippi to the Pacific Ocean. Vol. iii Washington, 1856.

ON THE SYSTEMATIC POSITION OF THE SAND

GROUSE (PTEROCLES; SYRRHAPTES).

R. W. SHUFELDT.

THERE has been no question for many years past, in the

minds of avian taxonomers, as to the general affinities of these

birds. This opinion may be briefly stated by saying that the

sand grouse constitute a small assemblage of forms, related on

the one hand to the gallinaceous birds, and on the other to the

pigeons.

Some authors have relegated them to a distinct group,

placing it in their schemes of classification between the fowls

and the Columba. Huxley created the Pteroclomorphe for

them, and Sclater, regarding them as a family, Pteroclide,

placed them in the order Pterocletes, standing between the

Columbae and the Gallinze, and in this he has been followed by

Stejneger and others. Garrod, Fürbringer, and other authori-

ties, again, have arrayed them with the pigeons. Numerous

papers have been devoted to their osteology, but the best of

these is doubtless the one given us years ago by William Kitchen

Parker, in the Transactions of the Zodlogical Society of London

(V, 149), where they are treated in his memoir entitled “On

the Osteology of the Gallinaceous Birds and Tinamous." The

plates to his memoir illustrate the skeleton of Syrrhaptes para-

doxus, while in the text we have a more or less extensive com-

parison of the osseous system of this species with that of

Pterocles arenarius. Parker's figures are very helpful, and in

addition to them I have examined some bones of Syrrhaptes

loaned me by Professor Alfred Newton, F.R.S., and there are

also at hand the mounted skeleton (see plate) and disarticu-

lated one of Pterocles arenarius, belonging to the collections

of the United States National Museum, and other material.

One has but to glance at the skull of Pterocles to be satis-

fied that the bird is not a pigeon, while, on the other hand, it

I2 IHE AMERICAN NATURALIST. [VoL. XXXV.

brings to mind the skulls of some of the smaller grouse or

ptarmigans. The cervico-dorsal region of the skeleton also is

by no means truly columbine, though without difficulty we can

Were Qut .

eieton of Sand Grouse (Pter

— P ate i e I 2 Q ~ T M T

octes arenartus Pallas). No. 18,849, Coll. U. S. Nat. Museum.

Reduced about one-third.

see the pigeon in the pectoral limb, the sternum, the pelvis,

the ribs perhaps; but to a less extent in the shoulder girdle

and the bones of the pelvic extremities,

No. 409.] THE SAND GROUSE. 13

The form of the premaxillary is gallinaceous, the sutural

traces of its proximal frontal process being distinct throughout

life over the facial frontal region in the middle line, as we see

it in many fowls. The large narial openings are elliptical in out-

line, and made even more so by a curled osseous extension

upon either side of the inner nares, that is, above and in front

of the very large pars plana, below and in front of the frontal,

touching the nasal externally and the premaxillary internally,

while its upper part, with its free anterior edge, is in full view

upon superior aspect of the skull. A few small, irregular

vacuities may occur in the interorbital septum, but the good-

sized orbit has upon all sides well-defined bony walls, the

frontal roof overhead being well produced, the pars plana large

and concaved upon its posterior aspect, the anterior part of the

brain-case complete; while quadrate, pterygoid, and palatine

afford a fairly good osseous floor.

The postfrontal processes are more or less aborted, and the

squamosal ones are thin and lamelliform, as in the chickens, —

the two apophyses not coming in contact distally.

At the base of the skull the palatines are of extremely

slender construction, and widely separated from each other in

the middle line. They do not even come in contact across the

sphenoidal rostrum, which latter is much thickened and rounded,

being pointed anteriorly, where it is carried beyond the pars

plana.

Either maxillo-palatine is a mere rudimentary spine, so small

that the thread-like anterior rod of the corresponding palatine

almost conceals it from view, when the skull is looked at from

this side. A pterygoid is also very slender, and presents a

somewhat flattened sigmoid curve at its middle part. These

bones articulate with pteryapophysial processes at the cranium’s

base. No vomer seems to be present, and the nasal septum

is but very imperfectly performed in bone.

The zygomatic arches are slender and straight, while the

quadrates are well developed and present no very unusual

characters.

The mandible, of a V-pattern, resembles to no small extent

that bone in some of the smaller ptarmigans (Lagopus). A

I4 THE AMERICAN NATURALIST. [Vor. XXXV.

good-sized ramal vacuity is present in either ramus, and the

straight and blunt angular processes are considerably produced.

Kitchen Parker has said the “differences between the skull

of Pterocles arenarius and Syrrhaptes paradoxus are not great,

but are important. The head and face of the former are

altogether stronger, more gallinaceous and less pigeon-like,

than in the latter. The skull base has, in the Pterocles, that

peculiar breadth which arises from the struthiousness of its

structure. The upper frontal region is broader between the

eyes, and the ale of the ethmoid swell up to a greater extent

between the crura of the nasal The postorbital and squa-

mosal processes are much stronger, and make a thicker bridge

over the temporal fossa. The crossing of the posterior and

horizontal semicircular canals project in the same hemispherical

manner as in Syrrhaptes, and the tympanic ala of the lateral

occipital is equally arrested.”

“The molar arch is stronger, and the central interorbital

space is filled up;! so also are the orbito-frontal fontanelles ;

the common optic foramen is more closely and neatly circum-

scribed. There is still an oval slit, opening into both orbits,

between the ethmoid bar and the lower edge of the frontals at

their coalescence. Theantorbital lachrymal mass is equally large,

and the septum nasi as well developed and as completely ossified.?

‘The bones of the face generally are quite as strong as in

ordinary pigeons, and therefore a degree beyond what is seen

in Syrrhaptes. The double head of the os quadratum agrees

with the same structure in Syrrhaptes, and there is nothing

special to remark upon in the bones of the palatine region.

The lower jaw is altogether stronger and deeper, its bend is

more marked and further back, than in that of Syrrhaptes ;

the membranous space is of about the same size, as are also

.the angular processes." 3 There is an excellent account of the

skull of Syrrhaptes, including the bones of the tongue, etc., in

the work of Professor Parker just quoted. It is interesting to

! Not so in all specimens. — R. W. S.

? The septum narium does not always EROS ossify in all individuals of

this group ; it may, however, do so in the skulls o: old specimens. — R. W. S.

3 On the Osteology of Gallinaceous Birds and Garni (p. 204).

No. 409.] THE SAND GROUSE. I5

note that in Syrrhaptes there are sixteen cervical vertebrae;

fifteen in Pterocles arenaria; and but fourteen in Lctopzstes

migratorius. Again, in Syrrhaptes, three of the dorsal verte-

brze coóssify into one piece, the fourth dorsal remaining free.

This agrees with Ectopistes; while in Pterocles arenarius,

four dorsals coóssify to form a single piece, and posterior to

this another free dorsal vertebra is found, making five. This,

with many similar points in its skeleton, goes to show that

Syrrhaptes is nearer the pigeons than is Pterocles; yet neither

of these forms are truly columbaceous.

In the pelvis of either genus we find just such a bone as we

should expect to find in birds that are doubtless typical inter-

mediates, standing directly in their organization between two

well-circumscribed groups. In the sand grouse, however, the

lateral portions of the pelvic sacrum, at its widest part, fail to

ossify, and thus, in the dried skeleton, leave large vacuities in

that region not seen in tetraonine nor typical columbine birds.

There is usually one less sacral vertebra in pigeons than there is

in the sand grouse, the former having fourteen, while Syrrhaptes

and Pterocles have fifteen ; and Parker claims that the last has but

six vertebrae in its tail, and this is all I find in Ectopistes, while

Columba livia and Pterocles arenarius each possess seven.

Professor Parker is correct when he says, “I do not set

much value on the number of caudal vertebrz, as the last is

a series, and the tail is very apt to vary in the number of those

which shall be swallowed up in this terminal piece.”

The epipleural processes on the ribs are much broader and

deeper in Pterocles arenarius than they are in Ectopistes and

other pigeons.! :

In Pterocles the scapula are long, narrow, and tapering,

reaching, in fact to some extent overreaching, the ilia of the

pelvis posteriorly. In Ectopistes these bones are cimeter-

1 In the autumn of 1899 Professor C. O. Whitman requested me to write out

for him a complete account of all the species of the North American pigeons, in

so far as their osteology was concerned, as a contribution to the Journal of

Morphology. This I did, illustrating the memoir with several figures of the bones

of the birds of that group, and a number of the points referred to in the present

paper will therein be illustrated. It was accepted for publication and will in due

course appear in the aforesaid journal, probably some time in 1901.— R. W. S.

16 THE AMERICAN NATURALIST.

shaped, and both dilated as well as truncated behind, where

they do not reach the ilia by any means. Pterocles has a very

insignificant fourchette in its shoulder girdle, slender and of a

U-pattern. Its clavicular ends articulate with the scapula.

This they fail to do in all the pigeons I have examined, where

the bone has much the same form and slenderness, but reaches

a great deal farther down towards the carinal angle of the ster-

num. As already stated above, both the sternum and the

upper extremity of Pterocles are quite columbine in character,

especially the former. Its sternum has considerably more

pigeon than it has grouse in it, and as this bone is often seized

upon by some avian classifiers as 7Ze index of a bird’s system-

atic position and its affinities, it may account for the sand

grouse having been placed upon the columbine side of the

line in certain schemes of classification.

With a strong columbo-tetraonine tincture in it, the pelvic

limb of Pterocles arenarius has characters in it not commonly,

if ever, found in those allied groups. In P. arenarius (No. 18,849,

Coll. U. S. Nat. Mus.) the first metatarsal coóssifies with the

tarso-metatarsus; is high up on the shaft; and the basal hal-

lucial joint, with its unequal phalanx, is very rudimentary.

There are but three joints and a claw in either outer podal

digit; while the limb below the knee (there being no patella)

is well supplied with sesamoids. One great grooved one is

found back of the tibio-tarsal condyles, and two or three small

ones in the sole of the curious foot of this bird. Air does not

gain access into the shafts of the long bones of the pelvic limb

of Pterocles; and this also holds true for Syrrhaptes (Parker).

As to the systematic position of the sand grouse, it may be

briefly said that there is altogether too much grouse in the

skull of Pterocles to admit of its being arrayed with the

Columba; while, on the other hand, there is too much pigeon

in both Pterocles and Syrrhaptes to admit of placing either of

these genera in the tetraonine assemblage. The place they

really hold is an intermediate one, and this is best shown, I

think, and the ends of classification best served, by arraying

them in a separate group, — the suborder Pterocles, standing

between the Galli and the Columba.

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L MARK, DiRECTOR. No. 118.

CORRELATED, ABNORMALITIES IN THE SCUTES

AND BONY PLATES OF THE CARAPACE OF

THE SCULPTURED TORTOISE:

G. H. PARKER.

A TYPICAL carapace of the sculptured tortoise (Chelopus

insculptus LeC.) is composed of fifty bony plates, so united as

to form a strong dorsál shield, and of thirty-eight horny scutes

covering this shield ex-

ternally. These ele-

ments are arranged as

shown in Fig. 1, in which

the black lines represent

the limits of the bony

plates, and the lighter

ones those of the scutes.

The scutes form three

series: first, a median

set, which, beginning at

theanterior end and pro-

ceeding posteriorly, con-

sists of a narrow nuchal

scute, five large central

scutes, and a pair of

pygal scutes, one right

Fic. 1. — Dorsal view of a normal carapace of a male

and the other left 5: SEC sculptured tortoise. The faint whitish lines represent

; the edges of the scutes, which were removed in making

ondly, four pairs of large the preparation; the black lines mark the margins of

centro-lateral scutes, '"e*ony Plates. xi

which flank the median series, except at the anterior and the

posterior ends ; and, thirdly, eleven pairs of marginal scutes,

18 THE AMERICAN NATURALIST. [VoL. XXXV.

which bound the periphery of the carapace, except where the

nuchal and the pair of pygal scutes reach the edge. The bony

plates (Fig. 1, black outlines) are also arranged in three series.

The median one is composed of twelve plates, which, beginning

at the anterior end, are, first, a large nuchal plate; next, eight

neural plates; and, finally, three pygal plates, one behind the

other. Eight pairs of costal plates abut with their central ends

on the median series, and extend laterally well towards the

edge of the carapace. Excepting where the nuchal plate and

last pygal plate reach the edge, the carapace is bounded by

eleven pairs of marginal plates. Although the scutes and bony

plates are arranged upon similar plans, the two sets of elements

do not coincide either in numbers or in exact positions.

The first abnormal specimen to be described is one (Museum,

No. 1829) from the extensive series by which this species is

represented in the collec-

tions of the Museum of

Comparative Zodlogy at

Harvard College, Cam-

bridge. It isa male and

was collected at Lancas-

ter, Mass. My thanks

are due Mr. Samuel Gar-

man for having called my

attention to it, as well

as the authorities of the

museum for their liberal-

ity in allowing me to dis-

sect such parts as were

needed. The carapace,

which was about 15 cm.

long and 11.3 cm. broad,

Fic. 2. — Dorsal view of abnormal carapace No. 1 (Mus. had suffered somewhat

Ammo eee y The black lines give the out- from marginal fractures,

rue: but in no instance had a

whole scute or plate been lost through such injuries.

The arrangement of the scutes is given in Fig. 2, an exam-

ination of which shows that there are two abnormal regions :

NO. 409.] THE SCULPTURED TORTOISE. 19

first, the middle and posterior parts of the median series; and,

secondly, the anterior parts of the marginal series.

In the median series the nuchal and first central scutes are

essentially normal. The second’ central is irregular on its

posterior margin; the

third and fourth centrals

are apparently each

divided in two by nearly

parallel oblique lines;

and the fifth central is

irregular anteriorly. The

second, third, and fourth

centro-laterals of the right

side are also irregular in

form. To the right of

the fifth central is a small

supernumerary scute,

which may represent a

part of any one of the sur-

rounding elements except

the marginals. These are

apparently not involved

in the irregularity, whose Fte. 3. — Dorsal view of abnormal carapace No. 1. The

: à black en give iu ‘clad of the bony plates. x %.

center obviously lies to

the right of the median line in the region of the third and

fourth centrals.

When the bony plates underlying the region of irregularity

just described are inspected (Fig. 3) they are seen to present

no features essentially different from the normal arrangement,

and it is, therefore, clear that in this region the abnormali-

ties are limited to the scutes and are not associated with any

peculiarities of the underlying bony plates.

The second abnormal region in this carapace lies anteriorly

and is easily recognized by peculiarities in the marginal scutes

(Fig. 2). If the lines of separation between the first and

second centro-lateral scutes of both sides are traced laterally,

they will be found to be continuous with lines between mar-

ginal scutes. These intermarginal lines are undoubtedly a

20 THE AMERICAN NATURALIST. [VoL. XXXV.

natural pair, because posterior to each there are seven mar-

ginals and one pygal — a condition which is identical with that

in the normal carapace (compare Fig. 1). Anterior to these

lines, however, the right and left sides of the carapace are dif-

ferent ; on the right side are three marginals, on the left four.

As the left side has the normal number of marginal scutes, the

abnormality may be described as a deficiency on the right side,

an interpretation that is supported by the fact that the outline

of this side recedes more

than is usual in this

species (compare Figs.

i and: 2). Since the

right side is normal from

the fourth intermarginal

line posteriorly, the de-

ficiency may with fair-

ness be said to lie in the

region covered in this

specimen by the first

' three marginals.

An examination of the

bony plates in the mar-

ginal series of this cara-

pace (Fig. 3) shows a

condition substantially

like that just described.

done siam du mie TA bow plates, xx ^ The first four marginal

plates of the left side

occupy a region which corresponds to that covered by the first

three marginals of the right. The total number of marginal

plates on the left side is eleven, and this side may be described

as essentially normal, notwithstanding the fact that the first

marginal is triangular instead of quadrilateral. The total

number of marginals on the right side is ten, and the deficiency,

as can be seen by a comparison with a normal specimen

(Fig. 1), lies in the region covered by the first three plates.

As this is the region from which a scute is absent, the con-

clusion is warranted that, in this instance, the absence of a

No. 409.] THE SCULPTURED TORTOISE. 21

bony plate is accompanied by the absence of an overlying

scute. Thus the first specimen illustrates two conditions:

scute abnormalities «z7associated with bony abnormalities, as

seen in the median and right centro-lateral parts; and scute

abnormalities correlated with bony abnormalities, as seen in the

anterior portion of the right marginal series.

The second abnormal carapace was brought to my attention

by Mr. C. E. Preston, to whom the specimen had been assigned

for study, and who kindly prepared it for me. The animal

was a male and came in all probability from Maryland. Its

carapace, when prepared, measured

about 15.7 cm. in length and 12.1

cm. in breadth.

At first sight the carapace (Fig.

4) does not seem to be abnormal,

but a closer inspection shows that

there must be at least two abnor-

mal regions, one in the scutes and

the other in the bony plates; in

both instances the abnormalities

form symmetrical areas. The

scute abnormalities occur in the

posterior part of the marginal

Fic. 5. — Outlines of the scutes on the

posterior part of the carapace of (a) a

series. On: both sides marginals. normal sculptured, tortoise and of (7

he a to

1 to 8 are entirely normal In and fourth right centro-lateral scutes

8, 9, 10, eighth, ninth, and tenth right

scutes; 2, right pygal scute;

typical specimens (Fig. 5, æ) mar- i

IV, V, fourth and fifth central scutes.

ginal 8 is followed on either side

by four scutes, — marginals 9, 10, 11, and the pygal scute. In

the abnormal specimen (Fig. 5, 2), in place of these four scutes,

only three are present. As it is impossible to state which scute

of the four is absent, the condition may be described as a bilat-

eral deficiency of one scute posterior to marginal 8.

The bony plates of this specimen in the regions where scute

abnormalities occur are essentially normal (Fig. 4), and the

same may be said of those at the anterior end of the carapace.

In fact, judging from the number and shapes of the plates,

normal conditions may be said to exist between the anterior

edge of the carapace and the transverse line marked by the

22 THE AMERICAN NATURALIST. [VorL. XXXV.

posterior borders of the first costal plates, and between the

posterior edge of the carapace and the transverse line marked

by the anterior borders of the next to the last pair of costal

plates. The rest of the carapace is made up of four bony seg-

ments instead of the five which are typical of normal individ-

uals (Fig. 1). In this abnormal specimen, therefore, a whole

bony segment (a neural plate, a pair of costals, and a pair of

marginals) is absent, and the position of the deficiency is some-

where between the posterior edge of the first pair of costals

and the anterior edge of the next to the last pair of costals,

c, in the body of the carapace.

Such a suppression might be expected to be accompanied by

a shortening of the carapace, and, as a matter of fact, there is

some evidence that this is so. The length of the carapace

under consideration is 1 5.7 cm., its breadth 12.1 cm.; the

length is, therefore, 1.298— times the breadth. In ten normal

males taken at random the average length of their carapaces

was 16.65 cm., and the average breadth 12.68 cm. ; the average

length was therefore r.313 + times the average breadth. It

thus appears that the abnormal specimen is somewhat shorter

than the average normal specimen, both absolutely and rela-

tively to its length, — a condition to be expected if the suppres-

Sion of a segment is assumed.

Although the second specimen shows abnormal conditions

in both scutes and bony plates, the fact that the two abnormal

regions do not overlap even in part (for the most posterior

position assignable to the bony-plate abnormality is still ante-

rior to the most anterior position assumable for the scute

abnormality) might well lead to the inference that these two

irregularities were in no true sense correlated. Such a con-

clusion, however, is probably incorrect. The fact that neither

of the two abnormalities interferes with bilateral symmetry

at least suggests something more than an accidental relation,

though it in no wise meets the objection that the abnormalities

are not superimposed. This objection, however, is not as

serious as at first sight it seems to be. Harrison (998) some

time ago showed by a series of ingenious experiments on

young tadpoles that in the growth of the posterior parts of

No. 409.] THE SCULPTURED TORTOISE. 23

their bodies the germ layers undergo a curious change in posi-

tion. The tail of a developing tadpole is composed of an outer

covering of ectoderm — which ultimately gives rise to the outer

layers of the skin—and of a core of mesoderm. These two

masses of tissue grow in very different ways, so that as the

tail lengthens the ectodermic covering, which is most actively

produced anteriorly, slips posteriorly over the underlying meso-

derm, whose region of growth is chiefly at the posterior end.

Although this posterior migration of the ectoderm has been

actually demonstrated only in the tadpole, there is reason to

believe that it occurs in other vertebrates. Admitting its

existence in the turtle, it affords an easy means of explaining

the conditions described. The scutes of turtles are derived

from the ectoderm, the bony plates from the mesoderm. In

the anterior part of the carapace these ectodermic and meso-

dermic derivatives, according to the peculiarity of growth just

explained, would not undergo any separation but would retain

their embryonic positions. Hence, if the material from which

both scutes and plates arise were modified by any local influence,

the resultant scutes and plates would be found together, as in

the first abnormal specimen described. In the posterior part

of the carapace, on the other hand, the ectodermic migration

would be excessive and any early local disturbing influence

that affected both scute and plate-producing tissue would leave

its trace in the adult in the form of a region of modified scutes

posterior to a region of modified bony plates, — a condition

realized in the second abnormal specimen. Thus, from what is

known of the methods of growth of the integument and subjacent

parts in vertebrates, it is fair to assume that the abnormalities

of scutes and bony plates in the second specimen, though

separated in the adult, may be as truly correlated as those of

the first specimen, in which the modified areas still remain

superimposed.

The older anatomists have very generally pointed out the

superficial resemblances between the scutes and the bony

plates of the chelonian carapace, but they have as a rule

denied any close relation between these two sets of structures.

Gegenbaur ('98, pp. 132, 174), in his recently published volume

24 THE AMERICAN NATURALIST.

on the comparative anatomy of vertebrates, repeatedly empha-

sizes the idea of the independence of scutes and bony plates.

The conclusions to which the present studies lead do not favor

this view, but lend support to the opinion expressed in cautious

terms by Goette.(99, p. 430), and more radically by Gadow

(99), to the effect that in primitive turtles each bony plate was

associated with a single scute. Supposing such relations to

have existed, it is easy to conceive how the present conditions

could have been brought about ; for, if the migration of the

ectoderm were to be so retarded as to take place after its divi-

sion into scutes, the posterior scutes would be carried away

from the bony plates te which they belonged, and in con-

sequence of crowding some of them might be suppressed, with

the result that a carapace with a given number of bony plates

would be covered by a smaller number of scutes. Although

such an explanation of the present condition of the chelonian

carapace must be tested by experiment, the evidence derived

from the study of the abnormal specimens described above

shows that there is a closer relation between bony plates and

scutes than has been generally admitted heretofore.

REFERENCES.

Gapow, H. :

'99. Orthogenetic Variations in the Shells of Chelonia.. Zoölogical

Results based on Material collected by A. Willey. Pt. iii,

Pp. 207-222, Pls. XXIV-XXV. :

GEGENBAUR, C. ;

98. Vergleichende Anatomie der Wirbelthiere. Erster Band. xiv +

978 pp. Leipzig, Engelmann.

GOETTE, A.

'99. Ueber die Entwicklung des knéchernen Rückenschildes (Carapax)

der Schildkröten. Zzi;. J. "iss. Zool. Bd. Ixvi, pp. 407-434,

Taf. XXVII-XXIX. |

HARRISON, R. G.

'98. The Growth and Regeneration of the Tail of the Frog Larva.

dick. F Entwickelungsmechanik d. Organismen. Bd. vii,

PP. 431-485. i

THREE POLYMELOUS FROGS.

ROSWELL H. JOHNSON.

Havine found three undescribed polymelous frogs in differ-.

ent American museums, it occurred to me that the newly

discovered method of skiagraphy might give an opportunity

for the study of these cases without dissection, which is usually

not permissible with museum abnormalities. The accompany-

ing plates give the result of that work.

Case I (Figs. 1, 5, 6, and 7), a Rana palmipes, which had not

yet lost its tail, is from the collection of the Harvard Medical

School. This specimen bears two supernumerary limbs arising

from the right and left scapular regions. Of these the one on

the left is inferior in size to the normal leg of that side and is

entirely free from it. It is peculiarly devoid of pigment. In

the skiagraph (Fig. 5) a small rod-shaped bone, answering to a

clavicle or coracoid, is to be seen extending from the head of

the humerus of the supernumerary leg on the left to the skull.

The humerus extends past its joint with the radio-ulna in a

cartilaginous projection. The formation of the hand is

apparently normal The ulnar side is the more ventral and

anterior The supernumerary leg of the right side is repre-

sented by a mere stump, as some previous investigator has

cut off most of the limb. This arises from the scapula, which

is enlarged and misplaced on this side.

Case II (Figs. 3, 4, 8, and 9) is a young Rana halecina Dum.

and Bibr., which was collected by Mr. Carlos E. Cummings on

vacant land in the city of Buffalo. It is now loaned to the

Buffalo Society of Natural Sciences. Directly above the left

hind leg arises a supernumerary leg with nearly normal mark-

ings. In size this is but slightly less than the normal legs, the

reduction being greatest in the phalanges. The head of the

extra leg does not fit into an acetabulum, although it lies close

to the position where its acetabulum would be. The left ilium

26 THE AMERICAN NATURALIST. [VoL. XXXV.

divides about one-third of the distance from the anterior

end. The normal leg of the left side is connected with the

more ventral and mesal of the two parts, as would be natural

from its position. The dorsal part is apparently free from the

pubis. The distal extremity of the tibio-fibula appears to have

three terminal enlargements instead of the normal two. Three

bones of about the same size replace the normal astragalus and

caleaneum. Eight bony elements align themselves in the posi-

tion of the metatarsals, one group of six, with two very much

smaller ones at some distance. Three phalanges arise from the

terminal metatarsal of the more strongly developed end and

one from the next metatarsal. This structure certainly bears

out Bateson's statement : * In the enormous majority of poly-

melians the extra repetition consists of parts of a complemen-

tary pair."

Case III (Figs. 2, 10, 11, 12) is another young Rana hale-

cina Dum. and Bibr, in the anatomical collection of the

University of Chicago, which from an accompanying note

seems likely to have been found in the immediate vicinity.

The coloration of the body is somewhat abnormal, in that the

. Spots are smaller and more numerous than is usual. Directly

ventral to the right fore leg, which is slightly displaced dorsally,

and is smaller than the normal leg of the other side, is a pe-

duncle from which two legs arise. These are each about the

size of the left normal leg and distinctly larger than the normal

leg of the right side. Each of the supernumerary legs seems

normal in its parts and has the usual blotches. The ulnar aspects

face each other. The skiagraph reveals two fused bones in

the cone-shaped projection (indicated by asterisk) between the

supernumerary legs. It seems to me that we must regard each

of the halves of the outgrowth as again double, the internal

member of each pair having developed from pressure and.

Juxtaposition into these two fused bones in the cone-shaped

projection. This seems likely from the analogy in abnormali-

ties of crustacean appendages, such as that shown in Fig. 182,

IV, in Bateson's Materials Jor the Study of Variation, which is

taken from Maggi. In the cases cited of duplicity of append-

ages already of a double nature, the two median elements fuse

No. 409.] THREE POLYMELOUS FROGS. 27

and remain comparatively undeveloped. Should my interpre-

tation be correct, we have in this frog a hitherto undescribed

degree of polymely.

I wish to express my thanks to Dr. C. B. Davenport, under

whom this work was done.

HULL ZOOLOGICAL LABORATORY,

February 17, 1900.

BIBLIOGRAPHY.

The author has but one reference to add to those given in Bateson’s

Materiais for the Study of Variation, p. 555.

WasHBURN, F. L. A Peculiar Toad. Amer. Nat, Vol. xxxiii.

pp. 139-141. 1899.

28 THE AMERICAN NATURALIST. [Vor. XXXV.

PLATE I

Fic. 1. Case I. Skiagraph taken with the frog’s left side bearing the entire

supernumerary leg closer to the plate.

. Case III. Ventral side closer to the plate, and legs so arranged as

to be as much as possible in a transverse plane to the direction of the rays.

Fic. 3. Hind legs of Case II arranged to show the bony elements of the

supernumerary leg.

IG. 4. Case II. Dorsal side of pelvis closest to the plate. Anterior por-

tion of the body projected far above the plate.

Fic. 5. Case I. Back closer to plate. Plate and frog were inclined to the

course of the rays, so that image of the supernumerary leg would fall anterior

to the normal legs.

Iti. a es Pome Sete ee CR ra

No. 409.] THREE POLYMELOUS FROGS. 29

- IO.

- II.

aes

THE AMERICAN NATURALIST. [VoL. XXXV

PLATE II.

Left side view of Case I.

Right side view of Case I.

Posterior view of Case II.

Dorsal view of Case II.

Anterior view of Case III.

Side view of anterior part of Case III

Diagram of the toes of the extra legs of Case III.

ici

SOME CASES OF SALTATORY VARIATION.

CARL H. EIGENMANN anD ULYSSES O. COX.1

SOME specimens showing saltatory variation have been

collected at various times for the museum of the Indiana

University. It is the purpose of the present paper to place

these on record.

I. A remarkable case of meristic abnormality was collected

near Harrodsburg, Ind. The specimen is a male of the com-

mon frog, Rana pipiens Schreber, 54 mm. long, and shows the

following characters (Fig. 1) :

The forearm and hand of the right side are duplicated.

The extra part arises just behind the normal arm and passes

inward and forward to below the eye of the left side. It is

I9 mm. long from its origin to the base of the fourth finger

and 2 mm. in diameter. The normal right forearm is 12 mm.

long and 5 mm. thick. Its striking and unique feature is the

band of skin through which it passes, and which holds it as a

sling. This band is well represented by the photograph, and

is 4 mm. wide. The supplementary arm and wrist are so

placed that the fourth finger occupies the posterior place.

The second, third, and fourth fingers are as in the normal hand,

but more slender. Separated from these there are, in the place

of the first finger, two fingers, of which the first is the longer.

The free portions of the five fingers measure respectively 4, 2

34, 5, and 3mm. On the anterior margin of the base of the

first finger there is a broad callosity.

On the lower surface of the lower jaw, below the left eye, there

are two tubercles, the posterior one the longer, measuring 13 mm.

in height. There are no other indications of abnormalities.

The variation here recorded may be classed with pathological

abnormalities rather than with variations that lead to the

mutation of species. |

* 6 PRAN TE Sas Jfrom the Zoölogical Laboratory of the Indiana University, No. 38.

34 IHE AMERICAN NATURALIST VOL. XXXV.

` L

2. The second case is of much greater interest. It is a

—

specimen of Ameiurus natalis, 120 mm. long, from Turkey

Lake, Ind. It differs from normal specimens in the absence

of all traces of ventral fins. There are no scars to indicate

Fic. 1. — Ventral view of Rana Pipiens, with supplementary arm.

that the fins might have been lost as the result of a wound,

and we may safely assume that in this specimen we have a

case of saltatory variation which is perfectly bilateral. Such

a variant, if its characters should be slightly prepotent, would

give rise to a race of individuals without ventral fins, which, in

nature, would readily be recognized by naturalists as a distinct

jas)

No. 409.] CASES OF SALTATORY VARIATION. 35

genus. It is possible that some of the genera of fishes without

ventrals have arisen from such prepotent variants, a supposition

that is reinforced by the following case of variation.

3. The variation of greatest importance to be recorded here

was found in nine specimens of Ameiurus melas. These were

collected at random, out of a lot that must have numbered

fifty or more, in a small cave at Glasgow, Ky. The fact that

the specimens showed a remarkable series of variations was not

detected until after they had been brought to the laboratory,

so we can safely assume that they were collected at random

and not with respect to the variations to be recorded. The

B A

Fic. 2. — A meiurus melas, showing supplementary narial barbels (see directive lines).

specimens seen in the cave were all of approximately the size

of the individuals here recorded. Unless they represent a

dwarf race living in the small brook in this cave, they are prob-

ably of the same age. One of us visited this cave two years

ago, and he neither remembers to have seen any catfishes in

the cave stream at that time, nor are any specimens in the col-

lection made at that time. While the fact that he does not

remember to have seen them and did not collect any two years

ago is not conclusive evidence that there were no catfishes in

the cave at that time, the size of the specimens, taken together

with this, makes it probable that the present specimens are

under two years old and got into the cave either as a batch of

eggs or during their schooling stage. It is well known that

the young of the catfishes remain together as a school for

36 THE AMERICAN NATURALIST. (VoL. XXXV.

many daysafterhatching. It is not probable that they migrated

into the cave as separate individuals, unless they came from a

source as limited in extent as the cave itself. Specimens

migrating from a more extensive source would, under no prob-

ability, show the similarity of variation to be here recorded.

In common with all the other North American fresh-water

catfishes, Ameiurus melas is provided with a barbel in front of

each posterior nasal opening. In the specimens under consid-

eration there are usually one or more additional barbels behind

the first, frequently on the anterior lip of the nasal opening.

These barbels in detail are as follows, all measurements being

in millimeters ! :

B | LENGTH OF THE LEFT NASAL LENGTH OF THE RIGHT NASAL

ee 2 | BARBELS AND THE SUP- BARBELS AND THE SUP-

Pas s PLEMENTAL ONES. PLEMENTAL ONES.

1 Le l* €, 4. 16, 1 (Fig.2 A)

2 86. | £—t*, 18 11, 7.9 (Fig. 2 B)

3 78 11: 11,2

4 78 ILG 11,4 (Fig.2C)

5 78 11 and small lobe 6, 10, 2

5 d WE

6 76 11, Hoy 11

7 74 | minute, 11, 3 minute, 11, 14

8 74 10, minute 10, lobe on narial lip

9 78 2g x 10, :

à On the left side there are two supplemental barbels.

edge of the main barbel is 6 mm. long. One behind the outer edge is T-shaped.

segment is but 1 mm. long. The upper segment slants from behind

The

One behind the inner

upward and gig Behind the basal segment it is 2 mm. long, the portion

in front 4 m

b puden ar on d edge of the narial lip.

c The supplemental barbel is Y-shaped, with the basal m L pum. deb

the distal segments each 4 mm., and placed on the edge of the i

d Here there are two lobes on the narial margin, but no distinct development

of PCs

Y-shaped, with a basal segment 1 mm. long, and distal segments respectively

x uud 2 mm.

n the columns of lengths the figures representing the lengths of the various

d barbels ar

are placed in the same relation to the sevi of the main

barbel as the supplemental barbels are placed to the principal barbel.

No. 409.] CASES OF SALTATORY VARIATION. 37

The length of the normal barbelis indicated in the table by

heavy figures.

Summarizing these data, we get the following:

Number of cases with two barbels: left, 3; right, 2.

Number of cases with one barbel : left, 6 ; right, 7.

Number of cases with a Y or T shaped barbel: left, 3 ; right, o.

Number of cases with no supplemental barbel : left, o ; right, 1.

Number of cases with one on the left and none on the right, 1.

Number of cases with one barbel on each side, 4.

Number of cases with two barbels on each side, 1.

Number of cases with two barbels on the left and one on the

right, 2.

Number of cases with one barbel on the left and two on the

right, I.

Total length of all supplemental barbels, not counting minute

lobes, XV : left, 374 ; right, 244.

Average variation, B left, 4.166 ; right, 2.722.

While it is difficult from the variety of elements (number,

length, shape) that enter into each variation to make a direct

estimate of the degree of variability of each side and the degree

of correlation of the variation of the two sides, the number of

individuals is so small that an approximation can be arrived at

by a glance at the data.

The left side is much more variable than the right.

The presence of barbels on the two sides is an indication of

bilateral correlation. This is reinforced by the fact that in

five out of nine cases the number of barbels on the two sides

is the same. It would seem that the left side is leading

in the addition of barbels, in spite of the fact that in one

specimen no supplementary barbel is found on this side.

Admitting, in the absence of evidence to the contrary and

strong probability in its favor, that the specimens belong to

one brood, or, at the very least, that the specimens occupying

the narrow limit of environment in which they are found are

38 THE AMERICAN NATURALIST.

geneticaly related, we may deduce some interesting conclu-

sions from the preceding data:

I. The variation here recorded is saltatory.

2. It is bilateral, without reaching perfect bilateral correlation.

This makes it probable that primarily the variation was

introduced or induced by changes on one side. Judging

from its greater variability, the left side was primarily

concerned.

3. It is improbable that the variation originated independently

in each of the specimens.

4. The variation probably arose in one of the ancestors of the

specimens.

5. Admitting 4, the saltatory variation arising in an ancestor

was prepotent to a very high degree.

There are entirely too many probabilities in these conclu-

sions, in spite of the fact that the probabilities all verge nearer

certainty than uncertainty. The matter of the prepotency of

the variation can readily be determined by breeding some

specimens from the cave with normal specimens, which will

be attempted.

THE LARVAL COIL OF BACULITES.

JAMES PERRIN SMITH.

Historical. — The genus Baculites is widely distributed in

Cretaceous rotks, found in almost every region, and the straight

shafts of this form are locally among the commonest fossils.

But in nearly all these places only the straight, incomplete

specimens are found; so that until a few years ago Baculites

was supposed to be an ammonite that had reverted to the

orthoceran form. About ten years ago, however, Dr. Amos

Brown discovered in the Cretaceous beds of Dakota a number

of young specimens of Baculites compressus, with a larval coil

attached to the straight shaft; this he rightly interpreted as

indicating the descent of Baculites from a coiled ancestor.

Until recently the larval coil of Baculites had been found

only at this single locality near Deadwood, Dakota; but during

the past year the writer discovered a number of larval coils

of Baculites chicoensis Trask in the lower Chico beds, Upper

Cretaceous, on the Arroyo del Vallé, about eight miles south-

east of Livermore, Alameda County, California. Many of the

specimens are perfectly preserved, some with the shell on and

others in clear, transparent calcite casts, showing the develop-

ment and the specific characters as well as when the animal

was alive. In order that the early stages of the shell should

be preserved the animal must have died in early youth, for the

test is too thin and delicate to have remained uninjured while

attached to the larger shell, and not protected by it. All

specimens more than a few millimeters in length are found

with the small end broken off, as it could not have been of any

use to the animal. ;

A peculiarly fine sediment is necessary for the preservation

of these fragile forms, and they were found only in calcareous

nodules, where the amount of lime in the clay prevented the

dissolving of the calcite of the shells, and where rapid hardening

40 THE AMERICAN NATURALIST. [VoL. XXXV.

prevented their decaying or being ground up by the waves.

The young must have flocked together in quiet nooks where

the water was clear and where there was little grinding by wave

action on sands or pebbles. Such concurrence of circumstances

must necessarily be seldom found, and it is not surprising that

these delicate forms have been found in only two localities in

the world.

Retardation in Baculites. — This genus has always been

taken as a type of reversionary forms, since, although it

descended from coiled ancestors, at maturity it resembles

Orthoceras in its straight shell. It is, however, not really

reversionary, for the septa are not orthoceran, nor even nau-

tilian; they are ammonitic and complex, and grow more so

towards maturity, after passing through a goniatite stage in

early youth. The shell cannot, then, be said to have reverted to

the stage of Orthoceras nor even of Bactrites ; but it is clearly

a degenerate, retrogressive form, retarded in most characters,

while progressive in others. Its septa fail to reach the degree

of complexity attained by its not very remote ancestors; the

number of lobes and saddles is reduced, and the goniatite stage

is prolonged, the ammonitic stage being reached later in life

than was the case with its immediate ancestors.

But even this reduction of the elements of the septa responds

to the law of tachygenesis and is pushed back in the ontogeny,

so that in the earliest larval stages the full number of lobes

and saddles is never present. Also the early straightening

out of the spiral coil is progressive degeneration from a lytoce-

ran form. In the genus Lytoceras it has often been observed

that in old age the body chamber leaves the spiral a little way,

and Baculites is merely a case of inherited senile degeneration,

pushed back in individual ontogeny until the retrogressive

characters appear at successively earlier stages of growth,

reaching finally the larval stages. This is necessarily followed

shortly by the extinction of the race.

In all normal ammonites the siphuncle begins in the embry-

onic protoconch with a caecum or bulbous enlargement, which

never appears in later stages. Baculites shows retardation in

its development by a repetition of the siphonal czecum in several

No. 409.] THE LARVAL COIL OF BACULITES. 4I

chambers of the larval coil, indicating a persistence of embryonic

characters. This persistence of the siphonal czecum is seen in

the young of Lytoceras alamedense! from the same locality, and

it is interesting to note that this species of Lytoceras shows

degeneration also in the development of its septa; the genus

normally has its lobes trizenidian (three-pointed) in the early

adolescent stages, while at maturity they always become dicra-

nidian (divided into two sections); but Z. a/amedense never has

trizenidian lobes, they being dicranidian at the beginning of the

adolescent stage. In Lytoceras we have an early inheritance

of a mature character, and in Baculites a similar prematurity

of development, but accompanied by greater retardation. In

Lytoceras alamedense the septa become ammonitic at one

and five-twelfths coils, diameter 1.87 mm., while in Baculites

chicoensis the septa persist in the goniatite stage until the shaft

has extended two and a half millimeters from the larval coil,

corresponding to nearly two revolutions if the shell had been

coiled continuously in a spiral.

Another mark of retrogression is the contraction of the

whorl in the latter part of the larval coil; in the early stages

the whorl increases normally in size, but at about three-fourths

of a revolution begins to contract, until where the shaft leaves

the coil it is much more slender than the embryonic or earlier

larval whorl, and does not attain its former size until it has

grown some distance beyond the coil. Contraction or abnor-

mal shape of later whorls in ammonites has been shown by

J. F. Pompeckj? to be a manifestation of degeneration, and to

be accompanied by an early extinction of the race. In Baculites

we find the contraction of the chamber pushed back by tachy-

genesis into the larval stage, and a profound degeneration

otherwise shown; from the geological history of the race we

know that its life was short and that extinction speedily followed

upon this unnatural development of the shell.

Ontogeny of Baculites. — At maturity Baculites chicoensis

consists of a straight shaft, slightly tapering, with an ovoid

1 Smith, J. P. Proc. Cal. Acad. Sci., third series, Geology, vol. i, No. 4,

Pl. XVI, Fig. $

? Die Ammonoideen mit Anormaler Wohnkammer. 1894.

42 THE AMERICAN NATURALIST. [VoL. XXXV.

cross-section. The surface is corrugated with wrinkles or

curved ribs parallel with the striz of growth, forming a ven-

tral, shovel-like extension of the aperture. The septa are com-

plex, consisting of a divided ventral lobe, two pairs of laterals,

and a short dorsal lobe. These resemble the septa of Lytoce-

ras, but are simpler in digitation and number of lobes and

saddles. Specimens of the mature shell are known nearly a

foot in length, with scarcely any tapering of the form, so that

the extreme size of maturity or old age must have been

considerably greater than this.

The phylembryonic or protoconch stage is very much like

that of all the other angustisellate ammonites, except that the

spheroid tends to become more angular, and the internal sep-

tum begins to show traces of lobes and saddles. The siphonal

caecum is unusually large, and was seen to be within that part

of the protoconch cut off by the first septum. The limits of

the embryonic body chamber were plainly seen on several

specimens, marked by a constriction between the first and

second septa, but not following the outline of either; the

diameter at this stage was 0.53 mm. (Fig. 5).

The next step in growth was the formation of the czecum,

followed very soon by the development of the first septum ; this

marks the beginning of the larval stage, as shown in Figs. 3.

and 5. The body chamber of the first or ananepionic larval stage

consisted of an entire revolution; thus the metamorphosis of

the young animal must have been considerable. The surface

of the shell in the phylembryonic and ananepionic stages was

covered with pustules, giving a granulated appearance to it;

but at the end of the first revolution these pustules ceased

sharply at a constriction, and gave place to cross striz and

ribs (Figs. 12 and 18).

The second septum, which marks the beginning of the

metanepionic stage, has a divided ventral lobe, and the full

number of lobes and saddles that the animal possessed through-

out life; the later changes consisted merely in the gradually

increasing digitation of the septa, which, however, persisted in

the goniatite stage not only throughout the entire coil, but also

for two and a half millimeters of the straight shaft (Fig. 6).

No. 409.] THE LARVAL COIL OF BACULITES. 43

The metanepionic or second larval stage is characterized by

the sudden change in sculpture which takes place at the end

of the first revolution, where the pustules are replaced by fine

cross striae and ribs parallel with these (Fig. 18). This stage

may be arbitrarily considered to last as long as the coil con-

tinues, but the spiral widens and at a quarter of a revolution

beyond the constriction the shell leaves the coil and grows out

nearly straight. With this the paranepionic stage may be

considered to begin, and to continue so long as the septa are

goniatitic ; at the distance of two and a half millimeters from

the coil the septa begin to become ammonitic, and the larval

stage ends (Figs. 19 and 20). The impressed zone continues

in the shaft for nearly a millimeter from the coil, but before

the paranepionic stage has ended the cross-section of the shaft

becomes rounded instead of semilunular. The larval shell

seems to be always unsymmetric, at least in the large number

of specimens studied, and this lack of bilateral symmetry was

not due to crushing, but actually to one-sided growth. The

writer has observed that this is quite common in degenerate

forms of ammonoids, whilé progressive species with normally

healthy growth are exceedingly symmetric and constant in

development. :

The digitation of the septa, which begins with the indenta-

tion of the first lateral saddle at about two and a half milli-

meters from the coil, marks the beginning of the adolescent or

neanic stage. The complexity of the septa increases slowly at

first, but soon becomes more rapid as the whorl begins to be

compressed laterally, which takes place at about eighteen milli-

meters from the coil. This lateral compression may be regarded

as the beginning of the metaneanic or second adolescent stage,

which, however, cannot be sharply differentiated from the

others. The angle of increase of size of the whorl throughout

the later larval and earlier adolescent stages is considerable,

but at the distance of about thirty millimeters from the coil

the angle becomes smaller, indicating a distinct change in the

rate of growth. This may be called the last adolescent or

paraneanic stage, and forms a gradual transition to mature

conditions of growth. No sharp line of demarcation exists,

44 THE AMERICAN NATURALIST. [Vor. XXXV.

but for convenience the adult or ephebic stage may be said to

have begun when the compressed form, the greatest complexity

of septa, and the rough sculpture of maturity are visible. This

is the case at the distance of about seven centimeters from the

larval coil, when the animal has by no means attained to

maturesize. Further growth is then only increase in size and not

progression of development. Old age, or the gerontic stage,

shows itself in the obsolescence of sculpture and of the

increase in size. Only a few specimens showing senile degen-

eration were found, which is not surprising if we consider the

small chance any of the lower animals have of becoming old.

Phylogeny of Baculites. — This genus has usually been

classed as an aberrant form under the Lytoceratide, but E.

Haug ! says that the resemblance of the adult septa of Bacu-

lites to those of Lytoceras is accidental; that Lytoceras in

youth always has trienidian (three-pointed) septa, while Bacu-

lites is always dicranidian (two-pointed). But the writer? has

recently described the development of a somewhat degenerate

species of Lytoceras, in which the septa are two-pointed in the

earliest adolescent stage. This observation removes the objec-

tion to the commonly accepted derivation of Baculites, and is

especially interesting in view of the fact that Lytoceras in its

old age often leaves the coila little way. The young stages

of the species of Lytoceras studied by the writer are almost

exactly like those of Baculites chicoensis, the latter showing

only a greater contraction of the larval chamber, a premature

ornamentation of the embryonic and larval shell, and a reduc-

tion of the number of lobes and saddles. The only other

studies on the development of this genus have been made by

Dr. Amos P. Brown? on Baculites compressus. As compared

with that species, Baculites chicoensis shows greater degenera-

tion, for it leaves the spiral at the end of one revolution, while

1 Les Ammonites du Permien et du

vol. xxii, p. 410, 1894.

2 The velopment of Lytoceras

series, Geology, vol. i, No. 4, 1898.

* On the Young of Baculites Com

1891, pp. 159, 160; and The Develo

Baculites Compressus Say,

Trias, Bull. Soc. Géol. France, 3¢ sér.,

and Phylloceras, Proc. Cal. Acad. Sci., third

pressus Say, Proc. Phil. Acad. Nat. Sci.,

pment of the Shell in the Coiled Stage of

Proc. Phil. Acad. Nat. Sci., 1892, pp. 136-141.

No. 409.] THE LARVAL COIL OF BACULITES. 45

B. compressus has two revolutions in its coil. But the develop-

ment of the two species is essentially the same, and the genus

is monophyletic, in so far as observations on two species can

demonstrate it. Dr. Brown thought that the larval stages of

Baculites showed analogy with those of Crioceras and Ancylo-

ceras, and none with Scaphites. In all probability, however,

all three of these genera are polyphyletic, and have originated

from several stocks. Some species of Scaphites seem to come

from a Hoplites-like ancestor, but in studying the development

of some undescribed Scaphites from the upper Cretaceous of

southern Oregon the writer found their larval stages to be

very like those of Lytoceras and Baculites, and they probably

have a common origin.

Straight degenerate forms have appeared in the history of

the cephalopods from time to time, from the Trias upward, not

from any one stock in particular, and not genetically connected.

The mere fact that the form is abnormal is no indication what-

ever of kinship. In each case they spring from normal forms

and indicate their origin in their normally coiled young. Natu-

rally it is seldom that transitional forms from the progressive

to the degenerate are known, for the beginnings of these tran-

sitions are regarded as mere freaks of some normal species.

Further, degeneration or retardation is not necessarily accom-

panied by abnormality of form, as has been shown by the

writer in the case of Lytoceras alamedense and in the develop-

ment of Placenticeras,! where the genus is still progressive in

many characters. Whether normal or abnormal in shape these

degenerate forms are always short lived, for they represent the

extreme specialization of which the group is capable, while the

more primitive stocks or radicles persist through very long

periods, often little changed, but from time to time giving rise

to the abnormal forms as side branches. These side branches

coming off from the parent stock at no great distance in time

from each other may give the semblance of a genetic series,

but this is usually deceptive. It is thus supposable that some

of these forms have originated from the parent stock from

1 The Development and Phylogeny of Placenticeras, Proc. Cal. Acad. Sci,

third series, Geology, vol. i, No. 7, 1900.

46 THE AMERICAN NATURALIST. [VoL. XXXV.

different species and at different times, in which case the

genus would still be strictly monophyletic. But in the case

where the resemblance is merely that of shape and not of

development, as in the several species of Scaphites, the genus

is not monophyletic, and the forms of which the development

is different from that of the type cannot strictly be placed in

that genus. |

Baculites probably originated from Lytoceras, but it is not

at all likely that all species of Baculites came from the same

parent Lytoceras, nor, indeed, in the same region, for this

degenerate form is too widely distributed and too short lived

geologically for this to be probable. This supposition would

presuppose for Baculites means of distribution surpassing those

of the other invertebrates, which we know could not have been

the case, for they were not pelagic forms, but shore dwellers,

and individual species are no more widely distributed than the

gastropods and pelecypods that are associated with them.

STANFORD UNIVERSITY, CALIFORNIA.

No. 409.] THE LARVAL COIL OF BACULITES. 47 oe

48 THE AMERICAN NATURALIST. (VoL. XXXV.

No. 409.] THE LARVAL.COIL OF BACULITES. 49

EXPLANATION OF PLATES.

The Development of the Larval Coil of ZacuZites chicoensis Trask.

Fics. 1 and 2. Protoconch, front and top view, diameter 0.48 mm. Thirty

times enlarged.

IG. First or ananepionic septum, showing the siphonal caecum.

Fics. 4 and 5. Larval shell at one-fourth coil, diameter o. 58 mm., showing the

ananepionic and metanepionic septa, and the embryonic constriction. Thirty

times enlarged.

FIG Second or metanepionic septum, at one-fourth revolution, diameter

0.58 mm

Fics. 7 and 8. Larval shell at one-half coil, diameter 0.68 mm. Thirty times

enlarged.

Fic. 9. Sixth septum, at one-half revolution.

Fics. r0 and 11. Ananepionic shell, showing the embryonic constriction, the

ananepionic septum, the siphonal caecum, and the first larval body chambe

he yo animal died before further development took place. Thirty times

enlarge

Ananepionic shell showing ornamentation of the embryonic and

early larval shell, and the ananepionic body chamber. Thirty times enlarged.

d r4. Metanepionic shell at three-quarters of a coil, diameter 0.83

mm., showing contraction of the later chambers

FiGs. 15 and 16. Shell at end of the caress stage, diameter 1.06 mm.,

one and one-eighth coils. Thirty times enlarge

Fic. 17. Metanepionic shell, diameter 1.00 mm., showing periodic swelling of

the siphuncle, indicating a retardation of the phylembryonic siphonal caecum.

Thirty times enlarged.

Fic. 18. Larval coil attached to the straight shaft, showing all the stages from

phylembryonic through the paranepionic, and the beginning of the adolescent or

neanic stage. "Ten times enlarged.

Fics. 19 and 20. Front and side view of a metaneanic shell, showing the

unsymmetric shape of the larval coil, and the contraction of the metanepionic

body chamber. Thirty times enlar

Fic. 21. Composite specimen, Peta from several pieces, showing the aui

ment of the septa from the ananepionic into the adolescent stage. Ten tim

enlarged.

Mo.Bot. Garden,

1902.

VARIATION NOTES. — Nos. 1-3.

1. Frequency Curve of White Daisy. — Mr. F. C. Lucas, of

the Englewood High School, Chicago, who published a paper

on * Variation in Ray Flowers of the White Daisy," in the

American Naturalist for |

1898, has sent us counts made

on 444 white daisies collected

along the roadside at North-

wood, New Hampshire. The

results are given in the ac-

companying curve, where the sm- i] T

number of rays (the class) is

indicated by the figures at

the bottom, and the number

of individuals occurring in

(the frequency of) each class |__| EE ieee

is indicated by the vertical

scale. 3o ery optat

The curve is compound.

The principal mode is at 21,

where Ludwig found it for

European individuals. Sec-

ondary maxima occur at I3,

at 24 (— 3 x 8), and also at

29 and 31. What the latter

maxima mean is uncertain ;

Mr. Lucas calls attention to

the fact that they occurred

in his specimens from Mass-

achusetts also. We shall be glad to publish the results of

counting lots of 1000 to 2000 terminal flowers (only one from

each plant) from each of various localities.

Load ced

5 3o

52 THE AMERICAN NATURALIST.

2. Variation in the Branchial Filaments of Aquatic Lepidop-

tera Larve.1— The larvae of the Pyralid Paraponyx obscuralis

from the Illinois River at Havana have just [always?] 100

branched branchial filaments (gills) arising from the dorsum of

the middle segments. The number of branches to a gill is

modally different for the different gills, and for the corre-

sponding gill in successive moults. In each gill of the full-

grown larva, however, the number of branches is subject to

individual variation. The following table gives the modal

number of branches for each gill of the full-grown larva.

SEGMENTS. a.s.? | LES | a.i. pi. ped.

| EA EA E Bue

js |

2 6 | 5 3 5 6

3 6 5 3 5 6

4-7 4 6 4 5 5

8-10 4 6 4 5 4

II 4 6 4 5

12 | 3

| |

Inheritance in Tailless Cats. — A female Manx cat (with `

rudimentary tail) had six litters by normal male cats. In these

litters the number of abnormal (Maternal type) and of normal

(Paternal type) kittens was as follows:

Litter 1 LM

"t Sarre?

3M-t-2P

IMF

MERGE

3M t-2P

Aun pea N

The maternal (Manx or abnormal) quality was prepotent.

Also, there was a loss of this prepotency in the later litters.

Was this due to telegony?—R. AwrHoNv in Bull. Soc.

d Anthrop. de Paris, 4 sér., vol. X, p. 303, 1899.

On the Entomology of the Illinois River and Adjacent Waters,

te Lab. Nat. Hist., vol. iv, 149-273, 1895.

? a. anterior; ., posterior ; s., suprastigmal ; ż., infrastigmal; ped., pedal.

! Hart, C.

Bull. Illinois Sta

EDITORIAL COMMENT.

Pseudoscience. — The leading article of a recent issue of Zhe For-

ester, entitled * On the Possible Effects of the Gypsy Moth on Amer-

ican Forests,” is noteworthy as containing a number of far-fetched

conjectures that are dangerously near charlatanry. The following

quotation will more than substantiate our statement: “It is not

unlikely that some of the curious alterations in the distribution of

forest trees which geologists have recognized may have been due to

the development in former ages of the Gypsy Moth or other like

destructive species of insect. Thus in the early Miocene Tertiary

Europe was tenanted by a host of arboreal species closely akin to

those that now form our admirable American broad-leaved forests.

The Magnolias, the Gums and the Tulip trees, etc., were then as

well developed in Europe as they are in this country. Suddenly all

these species disappeared from the old world. There is no reason

to believe that the change was due to an alteration in climate.

There are many evidences indeed that such was not the case. It is

a very reasonable conjecture that that alteration was brought about

by the invasion of an insect enemy which may have been the ancestor

of the Gypsy Moth."

Against this * very reasonable conjecture " the words of Dr. Asa

Gray may be recalled: “Probably the European Miocene forest

was about as rich and various as is ours of the present day, and very

like it. The Glacial period came and passed, and these types have

not survived there, nor returned."

The statement also “that the naturalist who attained the unhappy

success" of introducing the gypsy moth into America did so for the

purpose of interbreeding the introduced insects “ with various native

species of moths, with the expectation of producing a hybrid which

would feed on the leaves of our numerous American species of oak

and produce a valuable kind of silk," shows an absolute lack of the

first essentials of successful hybridization ; moreover, it does injus-

tice to a man of pure science whose imagination never rioted with

vagaries.

* News"' in the American Naturalist. — Owing to the length

of time which must necessarily elapse between the preparation of

53.

54 THE AMERICAN NATURALIST.

the manuscript and the appearance of the printed page, the editors

have thought it best to drop the department of **News" which has

long been a feature in this journal. This step is taken the more

willingly since our contemporary, Science, covers the same field in so

able a manner, and with its weekly issue can present items of news

far more promptly than we are able to.

The notices of appointments, retirements, and deaths have been

approximately complete ever since the journal passed into its present

control. This feature will be retained, not as a matter of news but

as a matter of record, and will be presented every three months with

the endeavor to have this record as nearly complete as may be.

Another feature which will form part of the record, and which

appears for the first time in the present number, is the record

of gifts to what may be termed educational institutions, including

colleges, technical schools, libraries, and the like. In our present

record, which includes eleven months of the year 1900, we note the

gifts of over $16,000,000 to these purposes. Scattered through the -

pages of a weekly or monthly journal these items make but little

impression on the reader, but gathered at intervals of six months or

a year they are ample evidence of the generosity of our wealthy

people and of those who are classed among the “well-to-do.” In

contrast with this liberality of the moneyed classes of America stands

the parsimony of the same classes in the Old World. Gifts of a

similar character there are rare. An instance comes before us as we

write. During the past year Brown University raised over one million

dollars and in this way secured an additional gift of a quarter of a

million more. During two years past the University of Cambridge,

one of the two great universities of England, has been trying to

increase its funds, both for endowment and for new buildings, and

in spite of constant efforts it has been able to raise but a little over

three hundred thousand dollars (£62,500).

REVIEWS OF RECENT LITERATURE.

ZOOLOGY.

Hammar’s Layer. — Professor Hammar had previously? given

facts that led him to believe that blastomeres of many animals are

connected by a continuous membrane-like expanse of protoplasm

that forms the outermost part of each cell and passes over from cell

to cell. This cleavage would result in only partial subdivision and

not in complete isolation of cells. Moreover, the cleavage cavity

would be a hollowed-out interior space, not outside space extending

in between the cells. j

In a recent paper? he advances some new facts and considerations

in support of the thesis that this connection of blastomeres is a

protoplasmic part of the egg and not a dead “membrane,” and,

secondly, that it is a primary connection always there, and not a

secondary connection established from time to time as are the filose

threads previously described by others and now, apparently, seen

by Hammar in preserved eggs. HÀ A.

What holds Blastomeres together ?— Curt Herbst? adds to his

previous work upon the chemical environment of echinoderm eggs

and larva a series of observations upon eggs in an artificial sea-

water free from calcium. .

Eggs in various stages of cleavage thrown into such a solution

may continue to cleave and even to differentiate so that some cells

may show cilia; but the blastomeres do not cohere, but fall apart

and develop separately. This, however, takes place only when the

eggs have been shaken to remove the membrane, that else holds the

cells together to some extent.

‘When the isolated cells are put back into water with lime they

cleave and cohere, so that many little larvae result from one egg.

The cause of the falling apart of the blastomeres is not clear.

Herbst supposes they are normally held together by the tension of

1 See Review, American Naturalist, 1896, p. 597; and 1897, p. 454-

z Arch. f. mikr. Anat., Bd. lv. February, 1900.

3 Arch. f. Entm., Bd. ix, Feb. 20, 1900.

56 THE AMERICAN NATURALIST. [VoL. XXXV.

Hammar's layer, the ectoplasmic layer that passes from cell to cell,

and that the absence of lime so modifies this membrane as to prevent

it from holding the cells together. The actual moving apart of the

cells he thinks due to their individual movements. Yet the author

grants that some other factor remains for future research to discover,

since when cells that have fallen apart in water free from lime come

together again and flatten against one another in normal water, there

is no normal membrane present.

That one such factor is a pathological state of the ectosarc accom-

panied by abnormal *filose activities" seems indicated by the

author's figures. E A.

Vitality of Eggs and Sperms.—— James F. Gemmill! records

some interesting observations upon the life of sperm and egg.

Sperms of the sea-urchin, Achinus sphera, put into still water,

move only seven inches before dying, so that their power of dissemi-

nation must depend largely upon currents of water and not upon

their own locomotion.

In a tube of 2.4 mm. diameter they ascended 12.6 mm. in 7

minutes; 31.6 mm. in 20 minutes; and 44 mm. in 45 minutes. In

tubes of different sizes these sperms ascended a shorter distance in

the narrower tubes.

Sperms of an annelid were found to live longer if the water con-

tained douz//on; hence the author infers they are able to nourish

themselves. ;

To keep the sperm alive long the amount of sea-water added

must not exceed nine times the bulk of sperm. When the sperm

and water formed an opaque white liquid the sperms lived seventy-

two hours; but when the sperm was diluted with water till only

slightly turbid the sperms died in three to five hours.

That the sea-water stimulates the sperm is inferred from the observa-

tion that even immature sperms became active when put into water.

The eggs of this sea-urchin are in best condition for fertilization

during the first four hours after being discharged into the water, but

there is little loss of vitality up to nine hours. In seventeen hours

many develop abnormally, and many not at all. In twenty to twenty-

four hours only one to two per cent develop, and no development was

observed after twenty-eight hours. On the other hand, when the

eggs were fertilized immediately after discharge into the water, poly-

spermy and irregular development were apt to take place.

1 Journ. Anat. Phys., vol. xxxiv, January, 1900.

No. 409.] REVIEWS: OF RECENT LITERAFTURKE. 57

When an adult was kept thirteen hours in the air there were no

normal developments from the eggs as first shed, but when these

eggs were kept four hours in water seventy-five per cent developed.

The author interprets this as meaning that the eggs were in a “ state

of semi-asphyxia " and gradually recovered in the water.

The action of sperm upon moribund eggs is remarkable in that it

hastens their disintegration, causing them to form blister-like eleva-

tions, to become vesiculated, and to exhibit a sort of pseudo-cleav-

age. When dead the eggs are not changed by sperm and keep

their form for days. B AE

Animal Life. — This text-book of zodlogy is outside the usual

line, as might be expected in one of Appleton's **twentieth-century

text-books," written by the president of Leland Stanford University.

Like the Study of Animal Life of Arthur Thomson, it is not a labora-

tory manual but a book to be read for stimulus and instruction, not

for training in observation and deduction.

It is a book on animal * ecology," emphasizing life as adaptation.

In it structure and function go hand in hand, function leading the way.

The authors briefly consider the activities of the Protozoa, and

the lowest Metazoa, then take up chapters upon the sex and repro-

duction of animals, function and structure, the life cycle, the primary

conditions of animal life, the crowd of animals and the struggle for

existence, adaptations, animal communities and social life, commen-

salism and symbiosis, parasitism and degeneration, protective resem-

blances and mimicry, the special senses, instinct and reason, homes

and domestic habits, and the geographical distribution of animals.

Its simple elementary treatment, excellent illustrations, and the

piquancy of new facts gathered from the Pacific shores make it

a most attractive volume which should play a good part in awaking

interest in zodlogy. Yet used as “a first book” and, it may be, as

an only book, there is danger of catering to the desire to know what

we now think of animals rather than to know animals themselves.

E. A. A:

The World of the Great Forest.'— This latest product of Du

Chaillus busy pen is an endeavor to present to young people a

1 Jordan, D. S., and Kellogg, V. L. Animal Life. A First Book of Zoólogy.

New York, Appleton, 1900. 311 pp. 180 figs. :

? Du Chaillu, Paul. Zhe World of the Great Forest. With over fifty illustra-

1900. 322 pp.

58 THE AMERICAN NATURALIST. | [Vor. XXXV.

picture of animal life in the forests of equatorial Africa. The book is

divided into short chapters, each devoted to a pair, or a community

of animals, from the huge pachyderms to the smallest insects. The

animals are made to discourse very naively to each other on the

hardships and joys of their life, and to describe the special adapta-

tions that fit them for it. There is no attempt at a connected story,

or at any incidents which would not naturally result from the simple

motives which influence the actions of animals, their care for their

young, and their desire for food. The book is the story of the vicis-

situdes of wild life, the periods of plenty alternating with those of

want, — a life where only activity or ingenuity or patience can hope

to maintain itself. "This is a refreshingly wholesome point of view in

this age, when our views of animals are too much tinged by imagi-

native sentiment. The constant repetition of much the same story,

however, makes the three hundred and odd pages rather difficult

reading, especially as they are never lightened by a ray of humor.

Native names for the animals are early introduced and then used

exclusively, so that the memory must bear a constantly increasing

burden. When the *'nkengos" say that they are glad that they

have found no traces of *nginas, nshiegos, mbouvés, and kooloo-

kambas," we gain, despite the glossary, but a confused idea of the

cause of their joy. The book will hardly arouse an interest in ani-

mals in children who do not already possess it, but on the other

hand it will teach them nothing that is not true, unless it be an

exaggerated idea of the range of an animal's thought. The book

cannot fail to win a valuable place in a school or juvenile library, and

will teach those who have an interest in animals many details of the

life history of African animals in particular, and a very just concep-

tion of wild life as a whole. The illustrations are all good and some

are excellent. R. H

Mooswa and Others of the Boundaries.! — The author of Mooswa

and Others of the Boundaries says in his introduction: * Perhaps this

story is too simple, too light, too prolific of natural history, too some-

thing or other — I don't know; I have but tried to tell the things

that appeared very fascinating to me under the giant spruce and the

white-barked poplars, with the dark-faced Indians and the open-

handed white trappers sitting about a spirit-soothing camp-fire.”’

The suspicion here intimated that he has perhaps not succeeded in

1 Fraser, W. A. Mooswa and Others of the Boundaries.

P Ne Illustrated by Arthur

Heming. New York, Charles Scribner’s Sons, 1900. pp-

No. 409.] REVIEWS OF RECENT LITERATURE. 59

imparting the fascination to his readers seems to us justified. The

story is certainly “light,” but whether it is “too prolific of natural

history " may be questioned.

The scene of the story is laid in the forests of the Athabasca, in

which a lad of fourteen spends the winter in the charge of a half-

breed trapper. The interest of the story lies in the successful efforts

of the animals of the region to save their skins from the trapper, and

later in the endeavors of Mooswa, a moose whom the boy once

befriended, to save the boy from starvation.

If the book is an attempt to convey, in the form of a story, knowl-

edge of the ways of animals, it is a failure, for whatever truth it may

contain is obscured by a bewildering amount of romance. Thus

when the fox is caught in a trap, the beaver gnaws off his foot, and

the Canada Jay sews the skin over the stump with his beak. If the

author has tried to make a good tale, after the pattern of the Jungle

Books, he has failed through lack of the requisite literary skill. The

story is presumably intended for boys, but even boys, if fed on Kip-

ling and Seton-Thompson, would tire of the idle chatter which is put

into the mouths of the principal interlocutors. The book is by no

means bad; there are humorous situations, and even moments of

interest, approaching excitement. A decade ago it might have

proved acceptable, but the standard of excellence in such matters

has been set too high, by the creators of Mowgli and Wab, for any

but skillful artists to hope for success in the field. The illustrations

are far superior to the text. R. H.

Miller's Key to the Land Mammals of Eastern North America.' —

Probably no recent contribution to the literature of North American

mammalogy will be so gratefully and widely welcomed as Mr. Mil-

ler’s brief synopsis of the land mammals of eastern North America.

So great has been the increase in our knowledge of the subject dur-

ing the last fifteen years, so radical the changes in nomenclature, so

different the present methods of investigation, and so scattered the

literature that has been the outcome of this renaissance, that only

the few specialists engaged in the work could hope to keep in touch

with the subject. The general student hence found himself hopelessly

lost in intricate labyrinths in any attempt he might make to adn

clear conception of the results thus far reached, in even a limited

! Miller, Gerrit S. Key to the Land Mammals of Eastern North America,

Bulletin of the New York State Museum, vol. viii (October, 1900), No. 38,

PP. 59-160.

60 THE AMERICAN NATURALIST. [VorL. XXXV.

field. Mr. Miller's Key gives at once a bird'seye view of the

scene, and directs the inquirer where further information may be

obtained.

The geographical scope of the work includes *the entire mam-

malian fauna of the Atlantic slope of North America north of the

southern boundary of the upper austral zone," or the Atlantic slope

from Hudson Bay to the southern boundary of what is commonly

known as the * Carolinian Fauna." The Mississippi drainage area

is thus wholly excluded. A brief introduction defines the life zones

of the region and the plan of the Key, followed by a synopsis, giving

a classified list of the higher groups, species and subspecies. Then

comes the Key proper, supplying diagnoses of all the groups from

class to subspecies, subfamilies and subgenera excepted. Under

each species and subspecies are given “references to (1) the first

publication of the specific or subspecific name, (2) first use of the

binomial or trinomial combination, and (3) a recent monographic

paper in which the form is described in detail.. . . The type

locality is given in parenthesis after the first reference. The

accented syllable of all technical names is marked by an accent;

and the derivation of each name is placed in parenthesis at the end

of the diagnosis.” A paragraph, in larger type, follows the diag-

nosis, giving the distribution of the species or subspecies under

consideration. The Key is thus intended to give the correct nomen-

clature for all the forms treated, with a clue to their identification,

and a brief statement of their geographical ranges, generally indi-

cated by a reference to the * life zone ” they inhabit. This admirable

brochure is thus what it claims to be, a “key " to the subject, and an

aid to the acquisition of further information. That the work was

greatly needed, has been well done, and will prove a “boon” to

seekers of knowledge in this field, it is needless to further affirm.

The Key was originally planned to form part of Mr. Miller’s

recently published “ Preliminary List of the Mammals of New

York” (see American Naturalist, April, 190o,

“soon grew to the proportions of an independent paper,” and was

finally extended to include a larger area. The Key, we regret to

see, repeats the few errors of nomenclature of the list, to one of

which attention has already been called in this journal (Ze. cit.,

P. 318), namely, the highly questionable basis of the specific name

" americanus ” for the Virginia deer. Another case is the use of the

specific name * hudsonica Desmarest ” (1803) for the otter. As we

have already shown (Bull, Amer. Mus. Nat Hist, Vol. X, 1898,

pp. 316-318), but

No. 409.] REVIEWS OF RECENT LITERATURE. 61

PP- 459, 460), canadensis Schreber (1778) is perfectly tenable for this

species, for those who accept names based on plates, as does Mr.

Miller in the case of the mink, Putorius vison (Schreber), which rests

on the same basis as Zutra canadensis (Schreber) for the otter.

Moreover, the name * canadensis " had been in almost universal use

for this species till two years ago, when an unfortunate attempt was

made to replace it by “ hudsonica,” of twenty-five years later origin.

Another case that may be mentioned is the use of the generic

name ‘“‘ Rosmarus " for the walruses instead of the prior name “ Odo-

benus.” Rosmarus Scopoli dates from 1777; Odobenus Brisson

from 1756, becoming perfectly tenable from Brisson's second edition

of his Règne Animal, published in i762, or fifteen years before the

tenable date of Rosmarus. We can hardly believe Mr. Miller has

given these points due consideration.

There are no really new innovations in the technical nomenclature,

but a new subspecies of the common deer is described on p. 83,

under the name Odocoileus americanus borealis. The number of

species treated is ros, with thirty-three additional subspecies.

JA A

Pectoral Girdle of Reptiles. — Professor Max Fürbringer's' long

interrupted studies on the comparative anatomy of the pectoral

girdles and their muscles and nerves in vertebrates have been con-

tinued in a fourth part devoted to these organs in the Amphisbaenia

and the reptiles. The first 150 pages are devoted to an extended

account of the skeletal elements of the pectoral girdles, breastbone,

and humerus, including these parts in the fossil as well as in the

recent representatives of the groups under consideration. This is

followed by an account of the nerves to the shoulder muscles, after .

which a very exhaustive description of the shoulder muscles them-

selves is given. A final discussion of over 150 pages deals in a

comparative way with the facts brought forward in the descriptive

part and concludes with remarks on the phylogenetic relationships

of the groups of reptiles to one another, to the birds, and to the lower

vertebrates as shown by the structure of the parts described. The

illustrations, some seventy figures in all, are beautifully clear and

exact, and with the text constitute a work of monumental ge

1 Fürbringer, M. Zur vergleichenden Anatomie des Brustschulterapparates

und der Schulterumskiln, Jena. Zeitschrift für Naturwissenschaft, Bd. xxxiv

(September, 1900), pp. 215-718, Taf. XIII-XVII.

62 THE AMERICAN NATURALIST. [VoL. XXXV.

The Neurone in Anatomy and Physiology. — The neurone theory

of the structure of the nervous system as promulgated by Waldeyer

in 1891 has been subjected to nearly ten years of rigorous criticism,

and the outcome of this, so far as the present standing of the theory

is concerned, has been well presented by Professor Verworn,' in his

address before the Seventy-Second Meeting of the German Natural-

ists and Physicians. The fundamental postulate of the neurone

theory, namely, that nerve fibres are processes from ganglion cells

and that the so-called ganglion cell with these processes constitutes

the real cellular unit of the nervous system, is clearly stated at the

outset. The possibly closer union of these units than has heretofore

been admitted, particularly by the adherents of the contact theory, is

considered in the light of the recent work by Apathy and by Bethe

and pronounced still uncertain. The whole issue of this discussion

is rightly shown to be of secondary importance so far as the stability

of the neurone theory is concerned.

From the physiological side the author makes an excellent presen-

tation of the question as to the significance of ganglion cells. The

recent arguments of Bethe and of Steinach, to the effect that central

nervous operations are possible without ganglion cells, are shown

to be inconclusive, and many important observations made on

animals subjected to nerve poisons are adduced to show that central

nervous operations are dependent on ganglion cells for more than

a supply of nutritive material. The essay concludes with the state-

ment that the anatomical and physiological investigations of the last

ten years have left the neurone theory on a firm basis and is unques-

tionably one of the best recent estimates of the present standing

of that theory. P

Avian Helminths. — An important contribution on the frequence

and distribution of the internal parasites of birds has recently been

published by Wolffhügel? In all 630 hosts belonging to 73 species

of birds were examined. Most of them were native in the country

immediately bordering on the southern Rhine, but a few came from

the collection of the Basel Zodlogical Garden. In all 180 birds

proved to be uninfected ; the rest harbored of cestodes 35 species in

231 hosts, of trematodes 19 species in 124 hosts, of nematodes 26

species in 252 hosts, and of Acanthocephala 11 species in 41 hosts.

1 `

Verworn, M. Das Neuron in Anatomie und Physiologie. Jena, G. Fischer,

. 54 pp.

* Wolffhügel, K. Beitrag. sur Kenntnis der Vogelhelminthen. Dissertation.

Freiburg, B. 1900. 204 pp., 7 double plates.

No. 409.] REVIEWS OF RECENT LITERATURE, 63

Different kinds of birds varied very greatly in number and type of

parasites sheltered, and forms common in the one would be entirely

absent or rare in the other. Full results of the examinations are

given in a series of tables which show the number, condition, loca-

tion, and name of the parasites collected and the date, locality, col-

lector, and name of the host. The second half of the paper is

devoted to an anatomical description and discussion of some of the

less known cestodes found. The descriptions are full and contain

many new points which are well illustrated on the plates. One new

species, Hymenolepis tetraonis, was discovered in the quail, in which

it is apparently very common. H. B. W.

Revision of the Ticks. — Of this work by Neumann,’ a third part

has just appeared. It covers the tribe of the Ixodz, including the

eyeless genera Ixodes, Haemalastor, and Aponomma, and the genera

Hyalomma and Amblyomma which possess eyes. Analytical keys

for each genus, based on the characters of the male, of the female,

and of the nymph, and full bibliographic references make the work a

mine of information. Inasmuch as the ticks from the collection of

the Bureau of Animal Industry were placed in the hands of the

author for this revision, it has a peculiar value for American students;

this usefulness is greatly enhanced by the full references given under

geographical distribution to the individual states of the Union from

which the specimens have been collected. With delicate courtesy

the names of new species taken from labels written by the late George

Marx are used and the species credited to that author; many of his

drawings are also incorporated in the article, although for the text

Professor Neumann is alone responsible. The most important

change in the nomenclature of American forms is the suppression of

. Ixodes unipunctata Packard, the Lone Star Tick, as synonymous with

Amblyomma americanum Koch. The illustrations of the revision are

good, the text clear and concise, and the work is evidently carefully

done, making it altogether the most important contribution in this

group since the monograph of Koch. A fourth part to include

additions, corrections, and general considerations of a taxonomic

character to conclude the work will appear soon. H. B. W.

The Coccidze of Brazil. — As recently as 1897 Dr. H. von Ihering

catalogued the Coccidze of Brazil, but he was able to enumerate

1 Revision de la famille des ixodidés, Mém. Soc. Zool. France, tome xii (Paris,

1899), pp. 107-294, 63 figs.

64 THE AMERICAN NATURALIST. [Vor. XXXV.

only twenty-one species. Mr. Adolph Hempel has since that time

been actively engaged in their study, and as a result he has pub-

lished a work entitled * As Coccidas Brazileiras,” in which he

describes no less than 131 species as occurring in Brazil. This

work, which was received by the present writer on Sept. 26, 1900,

appears in Vol. IV of the Revista do Museu Paulista, and is, unfor-

tunately, in Portuguese. By reason of its place of publication and

the language in which it is written, it may escape the. attention of

some coccidologists, but it is in reality one of the most important

contributions to the study ever produced.

The new genera described are Cryptokermes, Stigmacoccus, Apio-

coccus, Tectococcus, Tectopulvinaria, Pseudischnaspis, and Diaspi-

distis. Stigmacoccus, though placed in the Coccinz, is doubtless a

Monophlebine, and singularly enough, it appears to be identical with

Perissopneumon, Newstead, described from India in Entomologists’

Monthly Magazine, November, 1900. The simultaneous discovery

on opposite sides of the world of this striking and distinct type is

remarkable. Pseudischnaspis is an offshoot from Chrysomphalus,

and will include, besides the Brazilian species, P. Jongissimus (Ckll.)

and P. bowreyi (CkIl.), hitherto referred to Chrysomphalus.

T. 1» AS C COCKERELL.

Notes. — The development of the common tubularian, Parypha

crocea, has been worked over by C. M. Allen (Biol. Bull., Vol. I,

p. 291). Each sporosac is an outgrowth of the body wall of the

polyp, and since it shows evidence of four radial canals, it must be

regarded as a much reduced medusoid. The genital cells, both male

and female, are derived from the ectoderm of the medusoid. The egg

grows by absorbing adjacent cells. Its nucleus is said to be absorbed

at an early stage and is later re-formed from the scattered fragments.

Segmentation is very irregular and is often outrun by the nuclear

divisions. The ectoderm and entoderm are differentiated by delam-

ination. The embryo escapes as an actinula with both basal and

buccal tentacles.

The segmentation of that portion of the neural tube which forms

the brain in teleosts has been studied by Charles Hill (Zool. Jahro.,

Bd. XIII). The region destined for the forebrain is represented by

three segments, that for the midbrain by two. These segments early

disappear and are replaced by secondary expansions which have

been mistaken for segments. The segments of these two portions

of the brain are serially homologous with those of the posterior

No. 409.] REVIEWS OF RECENT LITERATURE. 65

part which are six in number and persist to a much later stage in

ontogeny. Of these the first gives rise to the cerebellum, and the

remaining five to the medulla. The brain thus consists of eleven

neural segments.

As a suggestion of what might be done, the work of T. B. Pieri

(Arch. Zool., Exp. III, p. xxix) is interesting. Having shaken the

sperm of sea-urchins in a bottle of water for one-quarter of an

hour, he found the sperms apparently dead. When passed through

a filter the liquid, water and sperm, was added to sea-urchin eggs

and these developed into many-celled stages. The author would

fain see here the action of a soluble ferment which he would call

ovulase ; this truly would be “féconde en consequences biologiques et

philosophiques." However, he sees clearly that the experiments fail

in that the filtration did not remove the sperms and that some of the

sperms may not have been killed.

The free-swimming copepods of the Woods Holl region have

been described in the United States Fish Commission Bulletin for

1899 by Professor W. M. Wheeler. The paper contains accounts of

thirty species, several of which are new, as well as excellent tables

for the identification of these crustaceans.

The peripheral distribution of the cranial and first two spinal

nerves of the salamander, Spelerpes bilineatus, has been worked

out by Miss M. A. Bowers. The paper, which is published in

Vol. XXXVI of the Proceedings of the American Academy of Arts

and Sciences, is illustrated by four figures in which the nerve com-

ponents are brought out by appropriate colors.

Special Bulletin No. 4 from the United States National Museum

consists of a monograph of the American hydroids belonging to

the family Plumularidz by Professor C. C. Nutting. The memoir

includes a general account of the group and descriptions of some

120 species, nearly half of which are new to science. Itis illustrated

by thirty-four plates, containing over 300 figures.

The peculiar trematode, Macraspis elegans Olsson, has been

studied recently by Jágerskióld (Kg/. Vet-Akad., pp. 197-214, Stock-

holm, 1899). Good illustrations give the form, which varies greatly

as between old and young individuals. The entire structure, apr

cially that of the reproductive system, shows such similarity with

that of the digenetic trematodes that the author is not inclined to

Separate the Aspidobothrida so widely from them as has been the

custom hitherto.

66 THE AMERICAN NATURALIST. |. [Vor. XXXV.

A new genus of ectoparasitic trematode, Aporocotyle simplex, has been

discovered by Odhner (Centralb. Baki. u. Par., x. Abt., Bd. XXVII,

p. 62) on the gills of Pleuronectes. It stands in sharp contrast with

all forms of the group hitherto described, in that it lacks entirely

the suckers and all other specialized apparatus for attachment so

characteristic of the group.

The endoparasitic trematodes of Chelonia are treated by Braun

in two articles (Centralb. Bakt. u. Par., x. Abt., Bd. XXV, p. 714, and

Bd. XXVI, p. 627). A considerable number of species, both old

and new, are carefully described. Among the latter is an American

form, Jonostoma renicapitate, which has not been noted since the

original scanty description of Leidy.

The species of Filaria found in human blood are discussed by

Von Linstow (Zool. Anz., Bd. XXIII, p. 76). The characteristics

of each supposed species are given in full, with citations from some

rather inaccessible authorities. The distribution of each is also

considered.

Mr. Willis S. Blatchley's Twenty-fourth Report of the Geological

Survey of Indiana contains, besides important geological and mining

matter, a number of valuable papers on the local natural history.

E. B. Williamson contributes a descriptive catalogue of the Dragon-

flies; R. E. Call, an illustrated catalogue of the Mollusca, and

Stanley Coulter a catalogue of flowering plants and ferns. Mr.

Blatchley continues his -useful notes on the reptiles and batrachians

of Vigo County.

BOTANY.

Two Recent Mushroom Books. — The last half decade has been

` notable for the number of new mushroom books and papers, and

even more for the increase in fungus-eating in this country. Up to

the time that the late Mr. Gibson turned his happy faculty of pen

and pencil to the subject, most people had held a vague but fixed

idea that none but the expert mycologist could turn mycophagist at

large without the probability being great that his friends would ulti-

mately record in sadness the final result of some last experimental

eating of a species “ supposed to be" wholesome, — only Morels

"the" Mushroom, and a few others grossly marked being safe ot

the layman's consumption. By his clear descriptions and exquisite

No.4o09..] REVIEWS OF RECENT LITERATURE. 67

illustrations Mr. Gibson made possible the recognition of a few —

but sufficient — common edible species, while the fact that nearly all

of the fatal cases of toadstool poisoning are caused by Amanita mus-

caria, and A. phalloides and its closest relatives, led him to brand the

genus Amanita so forcibly that few of us now care to eat any volva-

bearing agaric, however wholesome and delicious experience may

have shown it to be. The only really weak point in his Our Edible

Toadstools and Mushrooms lies in the very emphasis of this most

wholesome warning against all amanitas, which causes the minor

caution against lurid Boletuses, the emetic Russulas, and other

* suspected " species to be overlooked, — a caution reiterated emphat-

ically in Professor Farlow's review of the book in the columns of

Garden and Forest.

Mycological clubs and amateur mycophagists have wonderfully

multiplied and thriven under the stimulus of this book, which, with

its selection of a few unmistakable edible agarics and its branding

as deadly of the Amanitas, provides a sufficiency of fungus food for

ordinary culinary purposes, with a good mapping out of the safest

lines of exploration for the venturesome who must go farther. So

far as I know, no fatal or extremely serious cases of toadstool poison-

ing traced to species not of Amanita have occurred in this country

in the last few years, except that a well-known phycologist, turned

mycophagist, slipped in his determination of a Boletus which he

thought he recognized from one of Mr. Palmer's plates, and, with

his family, paid a severe, if not the extreme penalty for the error ;

and that one fatal mistake and several less serious ones have been

made in considering Agaricus (Lepiota) morgani, an agaric rather

common especiall in the West, as fit for food, as its congeners

appear to be. :

The latest important contributions to mushroom literature are by

Professor Atkinson ! of Cornell University, a teacher of cryptogamic

botany, and Captain Mcllvaine,? who for the past twenty years has

been well to the front among the fungus-eaters in this country.

Both of these writers are evidently aiming at the same purpose,

! Atkinson, G. F. Studies of American Fungi. Mushrooms, Edible, Poison-

ous, etc. Ithaca, N. Y., Andrus & Church, r9oo. vi + 275 pp. with 200 photo-

graphs by the author and colored plates by F. R. Rathbun. ;

? McIlvaine, C., and Macadam, R. K. Zvadsteols, Mushrooms, Fungi, Edible

and Poisonous. One Thousand American Fungi: How to Select an Cook the

_ Edible; How to Distinguish and Avoid the Poisonous. Indianapolis, The Bow

dn Company, 1900. xxxvii + 704 pp. Pl. LXVII, and many illustrations in

ext.

68 THE AMERICAN NATURALIST. | [Vor. XXXV.

Professor Atkinson stating that he has tried to present the important

characters which it is necessary to observe, in an interesting and

intelligible way, and to illustrate these by life-size photographic

reproductions of the larger fungi, the selection of species being made

with a view to the representation of the more important genera,

chiefly those containing edible species ; while Captain Mcllvaine,

regretting the absence of any book giving the genus, names, and

descriptions of the prominent American toadstools, the edibility of

which has been tested or the poisonous qualities of which have been

discovered, has attempted to give such information for every species

known to be esculent in North America.

Both books are illustrated by colored plates and process repro-

ductions from photographs, which, particularly in Captain MclIlvaine's

book, are supplemented by diagrammatic drawings. The illustration

of pileate fungi is a subject about which opinions may and appar-

ently do differ widely. Few colored plates, not even excluding those

of the olden time, which were hand-tinted on a lithograph or engraving,

represent the colors any too naturally, and it must be said that the

illustrations in color in these two books, though often pleasing to the

eye, do not materially affect the truthfulness of this statement. On

the other hand, uncolored drawings fail to represent characters

which, though imperfectly shown in an ordinary colored plate, may

be sufficiently closely suggested in it to serve their purpose. The

two books in hand contain a wealth of photographic illustration which

in excellence is scarcely surpassed-by Mr. Lloyd's well-known photo-

gravure sheets of certain American fungi, and which may be taken

as representing nearly or quite the best that can be done by process

work; and yet, exquisite as some of this work is, and faithful as the

photographic portrait must of necessity be, it is doubtful whether the

technical characters which, no less than the gross characters, need

to be brought out in illustrating the pileate fungi, are as well shown

in the greater number of cases as they could be by an artist's skill,

guided by the unimpeachable accuracy of the camera and controlled

by the fresh dissection.

Both books are primarily intended for the fungus-eater, and yet

their scope is very different, and in both cases extends further than

that of Mr. Gibson's book already referred to. Professor Atkinson,

while covering the genera pretty fully, devotes a great deal of space

to comparatively few, but representative, species, poisonous OF

edible, while Captain McIlvaine, with nearly or quite equal fullness

of treatment, attempts to account for everything. It is easier to

No. 409.] REVIEWS OF RECENT LITERATURE. 6

9 9

prepare a monographic treatment of an entire group, provided one

have the material and the literature at hand, than to make and describe

a selection of interesting things from that group, since in the former

case, barring errors of omission, provided the work be well done,

whatever is sought is sure to be found, while in the other case inev-

itable disappointment awaits the person hoping to learn about some-

thing which the author did not consider it desirable or expedient to

include. Doubtless Professor Atkinson’s book will so disappoint

many people, and yet, for even the laboratory student of pileate

fungi, it will prove of great value. On the other hand, Captain

Mcllvaine's book, lacking the critical touch of the expert mycologist,

though it contain the names of plants sought, will probably lead to

a certain amount of error; yet it too is a book which should be

found on the departmental shelves of every American institution in

which mycology is taught. For the novice in fungus-eating, both,

though helpful, are likely to prove confusing, since the distinctions

made between species in the larger book may not prove easy to

make with the fresh plants, in many cases, while the number treated

in the smaller book, though restricted, is sufficiently great to embar-

rass ordinary people by tempting them into difficult paths; and no

book of a scope greater than that of Mr. Gibson's, in which only

thirty edible species are included, is likely to supplant it for the

amateur American mycophagist. The present books, like Gibson's,

contain numerous recipes for preparing and cooking edible species,

and, for the most part, these promise easily made and palatable

luxuries where fungi can be obtained in the fresh state.

Perhaps, in view of the uncertainties attending the use of fungi

as food, it may be as well to state that in addition to the avoidance

of amanitas, even including the wholesome ones for the sake of

greater safety, all species unpleasant to the taste or acrid, all

Boletuses, and all specimens which show the slightest trace of

discoloration or which have been allowed to become in the least

stale, should be left to the person who proposes to derive sufficient

pleasure from dangerous experimentation to justify in his own mind

the tampering with unnecessary and sometimes great risks. Pro-

fessor Atkinson, in speaking of the unwholesome species, quotes from

chemists in a way to show that in addition to muscarine, the deadly

alkaloid of Amanita muscaria, and phallin, the more deadly toxalbumin

of Amanita phalloides and A. verna, choline, an alkaloid which in

decomposition gives rise to muscarine or a related alkaloid more

deadly than itself, and helvellic acid, likewise a most energetic

70 THE AMERICAN NATURALIST. [VoL. XXXV.

poison, have been isolated from a considerable number of species

regarded ordinarily as dangerous only in a minor way or merely sus-

picious; and there seems little reason to doubt that much of the

ambiguity attending fungi of this class comes from the conversion,

in their incipient decay, of a minor and perhaps scanty poisonous

substance into a much more dangerous one, so that personal idiosyn-

crasy or differences between individuals in strength of heart action

seem capable of accounting for the divergence of opinion as to the

edibility of a number of the dangerous species, like Boletuses,

Gyromitra esculenta, certain Russulas, Lepiota morgani, and, indeed,

the Amanita muscaria itself. dw

North-American Pteridophytes. — A sixth edition of Professor

Underwood's handbook of the ferns and fern allies occurring north

of Mexico, which appears to have been carefully revised, has

recently appeared and is likely to meet with ready sale. In it are

incorporated records of the occurrence in one flora of several species

not before recorded for it, and descriptions of several species regarded

as new toscience. The author's recent comprehensive investigations

of the priority status of generic names in the ferns have been con-

sistently followed up in this book by the rehabilitation of the well-

known species of Cystopteris in the genus Filix, and of what has

been known as /echnum (or Lomaria) Spicant in the genus Struthi-

opteris, while Aspidium is now replaced by Dryopteris, Polystichum,

Phanerophlebia, and Tectaria. 1.

Notes. — An interesting note by Professor Kellerman, on an Ohio

station for Cissus ampelopsis or Ampelopsis cordata, with illustrations,

appears in the first number of a new journal, Zhe O. S. U. Naturalist,

published by the biological club of the Ohio State University, which

also contains a list of additions to the Ohio flora, notes on collecting

and preserving microscopical plants, and a paper by Kellerman on

a foliicolous form of Ustilago reiliana, which species is believed to

possess the characters of Cintractia rather than of Ustilago proper.

Viola alabamensis, a new purple-flowered acaulescent species, is

described by Pollard in a recent issue of Proceedings of the Biological

Society of Washington. |

A revision of the Cactacez of Paraguay, by Schumann, is being

published in current numbers of the Monatsschrift für Kakteenkunde.

1 Underwood, L. M. Our Native Ferns and their Allies, with Synoptical

Descriptions of the American Pteridophytes North of Mexico. New York, Henry

Holt & Co., 1900. x + 158 pp., 35 ff., and frontispiece plate. Price, $1.00.

No. 409.] REVIEWS OF RECENT LITERATURE. 71

G. P. Burns publishes an illustrated anatomical study of the

Stylidiacee in ora for October.

The purple cone-flowers, Echinacea purpurea and E. angustifolia,

have been hybridized for cultural purposes, as appears from a note,

with illustrations, by Kohler, in Die Gartenwelt for October 20.

Curtis's Botanical Magazine for November contains illustrations of

Erigeron leiomerus and Cypripedium guttatum, of the North-American

flora.

A revision of the species of Platanus by Usteri appears in No. 20

of the Mémoires de P Herbier Boissier.

A short popular article on Monstera deliciosa, with photographic

illustrations, by Theodosia B. Shepherd, is contained in The Land of

Sunshine for September—October.

A review of the Rocky Mountain Melanthacee by Rydberg

appears in the Bulletin of the Torrey Botanical Club for October.

Several species, and one genus, Stenanthella, split off from Stenan-

thium, are described as new.

K. M. Wiegand presents a revision of the tenuis group of Juncus,

in the Bulletin of the Torrey Botanical Club for October. Thirteen

species are recognized.

A well-illustrated “ Short Account of the Big Trees of California "

constitutes Buletin No. 28 of the Division of Forestry of the United

States Department of Agriculture.

Keys to species of Abies and Picea based on leaf anatomy are

given in a paper by H. B. Dorner, reprinted from the Proceedings of

the Indiana Academy of Science for 1899.

The distribution of Chilian Coniferae forms the subject of a paper

by Karl Reiche in the current volume of the Verhandlungen des

deutschen wissenschaftlichen Vereins of Santiago. Six genera, Podocar-

Pus, Dacrydium, Saxegothea, Araucaria, Fitzroya, and Libocedrus,

are considered.

The variations in Lycopodium clavatum, and their bearing on phy-

logeny, are discussed in a paper illustrated with three plates, by R.

A. Robertson, in Part IV of the current volume of Transactions and

Proceedings of the Botanical Society of Edinburgh.

The second fascicle of de Wildeman and Durand's * Contributions

à la flore du Congo," published as a part of the Annales du Musée du

. Congo, of Brussels, has recently been completed by the issuance of a

72 THE AMERICAN NATURALIST. [VoL. XXXV.

second part, and contains descriptions of a considerable number of

new species.

Professor Ascherson contributes a synopsis of the higher vegeta-

tion of Helgoland to Vol. IV, N.F., of the Wissenschaftliche Meeresun-

tersuchungen of the Commission for the Scientific Investigation of the

German Sea. |

Part I of the third volume of Boerlage's Handleiding tot de kennis

der flora van nederlandsch Indie, comprising the orders Nyctaginacez

to Casuarinacez, has recently appeared.

A series of contributions to the knowledge of the trees of Java, by

Koorders and Valeton, is appearing in current numbers of the

Mededeelingen uit 's Lands Plantentuin, of Buitenzorg.

A paper on “Some Plants of West Virginia," by E. L. Morris, is

published in the Proceedings of the Biological Society of Washington,

under date of October 31.

A number of papers on the plant geography of North America,

presented at the New York meeting of the American Association for

the Advancement of Science, are published in abstract in recent

numbers of Science.

A delightfully simple elementary statement of “ How Plants Live

Together,” by Bailey, constitutes Zzacher’s Leaflet, No. 19, of the

Cornell University Experiment Station.

An ecological comparison of the arctic and antarctic floras, by

Delpino, is reprinted from the publications of the Æ. accademia delle

Scienze dell istituto di Bologna for 1900.

The structural and superficial modifications induced in a number

of succulents, when grown with a liberal supply of moisture, are

considered by Brenner in an illustrated paper in ora for October.

Die Gartenwelt for October 27 contains an interesting statement

by M. Correvon of the successful manner in which he cultivates

Alpine plants in the crevices of a wall in the suburbs of Geneva, and

is illustrated by several hal

| f-tones, — among them a superb portrait

of Saxifraga longifolia.

Bulletin 46 of the Agricultural Experiment Station of the Univer-

sity of Nevada, which is No. 2 of the nature-study bulletins of that

institution, deals with the flowers and fruits of common trees and

shrubs. Bulletin 47 of the same series considers clover seeds and

their impurities. Both are well illustrated. |

No. 409.] REVIEWS OF RECENT LITERATURE. 73

The harvesting and preparation of balsam of Peru from Myroxylon

Fereire are described by Preuss in Der Zropenpflanser for November

with process illustrations.

Further notes on the plants known as Peyote and Ololiuhqui, by

Dr. Urbina, are contained in recent numbers of the Anales del Museo

nacional de México.

The botanical origin of coca leaves is considered quite fully by

Rusby in Zhe Druggist’s Circular for November.

Two papers on marl, of botanical interest, are published in No. 6

of the current volume of the Journal of Geology by Professor C. A.

Davis.

Dr. Kuckuck, in Bd. IV, N.F., of the Wissenschaftliche Meeresunter-

suchungen of the Commission for the scientific investigation of the

German Sea, describes his method of cultivating algz in the open

seas.

A voluminous account of the older Mesozoic flora of the United

States, by Professor Lester F. Ward, is separately published from

Vol. XX of the Annual Report of the United States Geological Survey.

An elaboration of the fossil cycads of the Yale museum, by

Professor Ward, is reprinted from the American Journal of Science.

A biographic sketch of Torrey, and an account of the work of

the Torrey Botanical Club, appear in the October Bulletin of that

organization.

A portrait of Ernest Roze is published in No. 7 of the current

volume of the Bulletin de la société botanique de France.

PALEOBOTANY.

A New Book on Fossil Plants.' — Dr. Scott's important contri-

butions to our knowledge of fossil plants are too well known to

students of palzobotany to need any introduction. The present

Work is a very satisfactory summary of much of his former work, and

the substance of it was first presented in the form of a series of

lectures delivered at University College, London, in 1896. The

lectures, however, have been entirely recast and brought up to date.

! Scott, D. H. Studies im Fossil Botany. London, Adam and Charles

Black, 1900. xiii + 533 pp., 8vo, 151 figs.

74 THE AMERICAN NATURALIST. [VoL. XXXV.

The plants dealt with are, for the most part, the vascular Palzo-

zoic plants, — Pteridophytes and Gymnosperms, — but some space

is also given to the Mesozoic types.

The first chapter is partly devoted to an exposition of the aims of

palzobotany and explains the different forms in which fossil plants

have been preserved; the latter part of the first chapter and the two

following are devoted to the Equisetales.

The earliest forms of Equisetales (Archzocalamites) occur in the

upper Devonian. These oldest types were in many respects allied

to the sole living genus, Equisetum. From this stock arose the

much more specialized Calamites of the Carboniferous, which showed

a secondary growth of the vascular bundles and more specialized

fructifications. The peculiar fossils described under the name

* Annularia " are supposed to be the smaller leafy twigs of Calamites.

The structure of the latteris often preserved most beautifully and

shows great similarity to that of Equisetum, with which the spo-

rangia also have much in common. The discovery of heterospory

in certain species is an interesting point, but it was apparently much

less marked than among the Lycopods and Ferns. Dr. Scott is very

positive in maintaining the strictly pteridophytic nature of all the

Calamariez. No Calamites are found above the Permian, the Meso-

zoic Equisetales being for the most part closely allied to Equisetum.

Chapter IV deals with the Sphenophyllales, which Dr. Scott con-

siders are entitled to rank as a fourth class of the Pteridophytes,

having certain affinities with both the Lycopods and Equisetales.

Of existing genera, Psilotum approaches Sphenophyllum in the

character of the vascular bundles, but as practically nothing is

known of the fossil Psilotacez, it is questionable how close the

relationship really is.

e very remarkable fructification known as Cheirostrobus, which

apparently combines calamarian and lycopodiaceous characters, is

considered by Dr. Scott to belong to the Sphenophyllales and to

confirm his view that “the Sphenophyllales were the highly modified

representative of an ancient stock from which both Lycopods and

Horsetails have diverged."

Chapters V-VII deal at length with the very abundant remains

of Lycopodiales. As was the case with the Equisetales, the group

culminated in the Palzozoic era, and in the later formations only

the smaller and less specialized types are encountered. Hetero-

spory, which still occurs in Selaginella, was very pronounced, and in

the case of certain forms closely resembling typical Lepidostrobus,

No.409.] REVIEWS OF RECENT LITERATURE. 75

seeds were actually developed. In spite of this, Dr. Scott is not

inclined to admit that there is any connection between the arbores-

cent Paleozoic Lycopods and the modern Conifers, although this is

suggested by the similarity in habitof thetwo classes. He concludes

that there is no satisfactory proof that the early Lycopods gave rise

to any group of the higher plants — a conclusion with which all

botanists will not agree.

The Ferns are the subject of two chapters, in which the different

types are clearly treated. Abundant remains of Ferns, often most

beautifully preserved, occur in all the formations from the Devonian

onward. Unlike the other three phyla of Pteridophytes, the Ferns

have held their own, and at present constitute a very important

element of the vegetation of many regions, especially in the mountains

of the tropics.

The fructifications are in many cases well preserved, and it is clear

that the earlier Ferns were mostly types allied to the existing Marat-

tiaceæ, which are thus shown to be a very old type, — a conclusion

reached independently by the writer some years ago, from a study

of the living forms. The fossil Marattiacez, however, showed far

greater diversity than the few existing genera. The other existing

group of Eusporangiate — the Ophioglossia,— which in certain

respects seems to present very primitive characters, is very unsatis-

factorily known in a fossil state, perhaps due to the very slight

development of firm tissues in most of them.

The occurrence of leptosporangiate Ferns in the Paleozoic rocks

is rare, and their affinities doubtful. A small number of types,

perhaps allied to modern Gleicheniacez and Osmundacez, have been

discovered, but it is not until the Mesozoic is reached that any con-

Siderable number of these are encountered. Last of all to appear

are the Polypodiacez, preéminently the modern fern type.

Among the most interesting of the Mesozoic Ferns were the

Matoninz, now represented by the single genus Matonia of the

Malayan region. This is a synthetic type, combining characters of

the Cyatheacee and Gleicheniacez.

One of the most important results of recent work with the Pala-

ozoic fossils is the discovery of a group of plants intermediate between

the true Ferns and the Cycads. These Cycadofilices have been

extensively studied by Scott and Seward in England, as well as

Y Several continental workers. Among the best known genera are

Lyginodendron, Heterangium, and Megaloxylon. Many of theseforms

which are formed from the lower Carboniferous and the Permian,

76 THE AMERICAN NATURALIST. [VoL. XXXV.

have been described as Ferns, the best known being the genera

Neuropteris and Alethopteris.

The earliest of all true seed-bearing plants were undoubtedly the

remarkable group, the Cordaitez, as to whose affinities there has

been much discussion. Their remains occur abundantly from the

Devonian through the Carboniferous. They present certain conif-

erous features, especially in the character of the secondary wood,

while, on the other hand, their structure recalls the Cycads, which they

resemble in the structure of the leaves. Unlike most fossil plants,

the flowers and fruits have been preserved with extraordinary per-

fection, even to the pollen grains which are found within the pollen

chamber, much as in the case of living Cycads. Most extraordinary

of all, so perfectly are the pollen grains and ovules preserved, that the

antheridia and archegonia are still recognizable!

Whether the Cordaitee really represent a type intermediate

between Cycads and Conifers, may perhaps be questioned, but they

certainly are one of the most interesting of all the groups of

fossil plants.

The Cycads, although occurring sparingly in the later Palaeozoic

formations, are especially characteristic of the Mesozoic, where, as is

well known, they formed one of the principal plant types. It is

evident that the Mesozoic cycadean forms were much more varied

than the existing genera, which show comparatively little variety of

structure. -

While some of the fossil forms approach closely their living

representatives, both in this character of the vegetative and repro-

ductive parts, others are extremely different, this being especially

true of the Bennettiteæ. These combined typical cycadean vegeta-

tive characters with fructifications of a very different kind, and not

readily comparable to that of the true Cycads. The seeds, which

have been very perfectly preserved, show a large dicotyledonous

embryo, nearly filling its cavity. These remarkable fossils are espe-

cially abundant in our own Potomac formation, and from Jurassic

and Cretaceous formations of the Black Hills, from which Professor

Lester Ward has described many new species.

Of the true Cycadaceæ, Cycas probably goes back at least to the

Lias.

The Coniferæ are but briefly treated. The earliest typical Conifers

seem to have been allied to the Taxodieæ, to which the fossil genus

Voltzia of the Upper Permian and Triassic seems to be allied. The

Permian genus Walchia, which has been supposed to be allied to the

No. 409.] REVIEWS OF RECENT LITERATURE. T

Araucariez, is only known from vegetative remains, the fructification

being quite unknown. Unmistakable Araucariee are not known

anterior to the Jurassic. The Abietinee are probably somewhat

more recent, but Cupressinew are found in the Jurassic. The

Taxinez (exclusive of the much more ancient Ginkgo) are first met

with in the Cretaceous.

The extraordinary genus Ginkgo, which with the Cycads represents

the oldest existing type of seed-bearing plants, and is now represented

by the solitary species, G. biloba, is recognized by Dr. Scott as the

representative of a distinct order, Gingkoacea, which is represented

by numerous forms in the later Palaozoic and earlier Mesozoic

formations. Dr. Scott agrees with Seward in assuming a somewhat

near relationship between the Gingkoacea and the Cordaitez,

The concluding chapter is occupied with a summary of the

general conclusions presented in the preceding chapters. We can

hardly agree with the conclusion that the great antiquity of the

Pteridophytes, and the absence of the remains of Bryophytes in the

Paleozoic formations, is a sound argument for the entire independ-

qu st the great divisions of Archegoniates. There certainly is

| m enience of any other forms from which they could possibly have

orici the evidence of comparative morphology is over-

,. emingly in favor of a common origin for all Archegoniates. The

. Questions of their interrel

; B um printed, and the illustrations are well

"e gh s pful in elucidating the text. We can heartily

ogni ee to all botanists interested in the fascinating

E o Pope D. H. C

RECORD OF GIFTS, APPOINTMENTS, RETIREMENTS, AND

DEATHS FOR THE YEAR 1900.

Educational Gifts for 1900. — Below we have summarized the gifts

to colleges, institutes, and libraries which have come to notice during

the eleven months of 1900, ending November 30. In its compilation

we acknowledge especial indebtedness to Science. Scattered here and

there the items have little significance, but gathered in this form they

are ample evidence of the generosity of the American people.

Amherst College, $10,000, by the will of Edward N. Gibbs.

Barnard College, New York, $100,000.

Bates College, $20,000, for a library building, from Joseph A. Coran.

Beloit College, a conditional gift of $200,000, from one of its trustees

Berea College, $50,000, from Dr. D. K. Pearson ; $150,000, by subscription.

Blackstone Library of Branford, Conn., $100,000, by the will of Timothy

T. Blackstone.

Boston University, $50,000, by the will of O. H. Durrell.

Bowdoin College, $150,000, from Gen. Thomas H. Hubbard, for a library

ildi

uilding.

Brooklyn Institute of Arts and Sciences, $15,000, by the will of Joseph C.

Hoagland.

Brown University, $1,120,000, by subscription ; $250,000, "from John D.

Rockefeller; $3500, by the will of Charles H. Smith; a conditional

gift of from $8000 to $30,000, by the will of the late A. D. McLellan ;

$25,000, by the wills of each, John Nicholas Brown and Harold Brown.

Carleton College, $50,000 from Dr. D. K. Pearson.

Carnegie SERENA Pittsburg, Pa., funds for its enlargement from Andrew

» not to exceed $3,000,000.

Catholic University of America, $20,000, by the will of Mrs. Rebecca

; $50,000, from Michael Cudahy.

Chicago xi Institute, $50,000, by the will of the late Sidney A. Kent.

Clark University, an indeterminate amount, by the will of the founder,

Jonas Clark.

College of Physicians and Surgeons of Chicago, $25,000, from Dr. W. E.

Quinn, for its library ; and $25,000, from Dr. D. A. K. Steele, for the

pathological laborato

College of Physicians of Philadelphia, $5000, by the will of Dr. J. M.

a Costa. ;

Colorado College, $50,000, from Dr. A. K. Pearson

GIFTS, APPOINTMENTS, RETIREMENTS. 79

Columbia University, $10,000, for books, from an anonymous donor ;

$100,000, by the will of the late Dorman B. Eaton, for a professorship

of municipal science and administration ; $100,000, from John D.

Rockefeller, to endow the chair of psychology.

Cooper Union, New York, residuary legatee of the late John Halstead, the

amount approximating $250,000 ; $300,000, from Andrew Carnegie ;

$200,000, from Abram S. Hewitt and Edward Cooper.

Cornell University, $80,000, anonymously, for a building for anatomy and

physiology.

Covington, Ky., $40,000, from Andrew Carnegie, for a public library.

Danielson, Conn., $15,000, for a public library, by the will of Edwin H.

James Milliken gives in two gifts $716,000, for an industrial school in

Decatur, Ill. Citizens have given $100,000 to the school, and

$100,000 more is promised.

Earlham College, $25,000, from Frances T. White.

East Orange, N. J., $50,000, from Andrew Carnegie, for a public library.

Emporia College, $50,000, from Andrew Carnegie, for a library building.

Essex Institute, $10,000, from the estate of the late Walter Dixon.

Fairmount College, $50,000, from Dr. D. K. Pearson.

Fargo College, $50,000, from Dr. D. K. Pearson.

Hackensack, N. J., public library, land and a building to cost from $30,000

to $40,000, from William M. Johnson.

Hampden Institute, $100,000, by the will of Collis P. Huntington.

Harvard University, $100,000, by the will of the late Dorman B. Eaton, for

the chair of science of government; $100,000, by the will of Mrs.

Caroline Brewer Croft, for researches into the: cause and cure of

cancer ; $2000, by the will of Barthold Schlesinger.

Haverford College, $40,000, by an alumni subscription for a gymnasium.

Kenyon College, $10,000, from Mr. Samuel Mather of Cleveland ; $15,000,

from Mr. J. P. Stevens, for a library fund ; $5000, by the will of John

Sherman.

Lafayette College, $45,000, by the will of Joseph E. Smaltz.

Lehigh University, $300,000, by the will of Frank Williams.

McGill University, $200,000, from Sir William C. MacDonald.

McKenzie College, $75,000, from Dr. D. K. Pearson. E

Marinette, Wis. $50,000, from Isaac Stephenson, for a public library

building. .

Massachusetts Institute of Technology, $2000, by the will of Barthold

Schlesinger.

Mechanics Institute of Rochester, N. Y., $200,000, from George a

Middlebury College, $50,000, from Ezra SS a science building.

Mt. Holyoke College, $200,000, from Dr. A. K. Pearson.

New York Pn S uM $20,000, and one-twelfth of the residual

estate of Judge Charles P. Daly.

80 THE AMERICAN NATURALIST. [Vor. XXXV.

New York University, $100,000, from Miss Helen Gould, for the erection

of a * Hall of Fame for Great Americans "; $2500, from Professor

and Mrs. E. R. Shaw, for a scholarship ; $20,000, by the will of the

late Robert Schell.

Oberlin College, $60,000, from Louis H. Severance, for a chemical labora-

tory; $50,000, from Dr. and Mrs. Lucius C. Warner, for a men's

gymnasium ; $75,000, by the will of Mrs. Caroline E. Haskell;

$40,000, by the will of William Osborne ; $5000, by the will of John

Sherman.

Ohio Wesleyan University, $10,000, from Mrs. Elizabeth Mebarry ; $35,000,

by the will of Mrs. Eliza Chrisman.

Onarga College, $20,000, from Dr. A. K. Pearson.

Peabody Academy of Science at Salem, $26,000, towards an addition to its

uilding.

Philadelphia, a building valued at $1,000,000, to the city, by P. A. B.

Widener, for a free library and art gallery.

Philadelphia Academy of Natural Sciences, one-sixth of the estate of the

late Charles E. Smith, estimated at $500,000.

Andrew Carnegie proposes to give buildings and an endowment of $1,000.

ooo for a polytechnic school in Pittsburg.

Princeton University, $45,000, by the will of Joseph E. Smaltz ; $40,000,

by the will of Dr. John S. Sayre, a portion to be used for fellowships in

applied electricity and applied chemistry.

Radcliffe College, $2,000, by the will of Barthold Schlesinger.

Ripon College, $15,000, from O. H. Ingham, towards the school of science

uilding.

Rush Medical College, $50,000, from Dr. Nicholas Senn.

Rutgers College, $10,000, by the will of the late Robert Schell.

Public Library, St. Albans, Vt., $10,000, by the will of Gov. J. G. Smith.

St. Lawrence University, $24,000, from a friend. .

Salem Public Library, $10,000, by the will of the late Walter Dixon.

Spellman Seminary, $180,000, from John D. Rockefeller.

Syracuse University, $25,000, by the will of the late Erastus F. Holden.

Teachers College of New York, $25,000, by the will of Miss Eliza T.

Bryson, for a scholarship.

Tufts College, $45,000, by the will of Walter Dixon, also a third of the

residuary estate.

'Tulane University, $50,000, from Mrs. Caroline Tilton, for library purposes.

Union College, $10,000, from members of the Mather family.

University of California, $10,000, for books, $100,000 additional, subject to

an annuity, and $150,000 in real estate, from Mrs. Jane K. Sather.

University of Chicago, $125,000, from A. C. Bartlett; $50,000, by will of

the late Sidney A. Kent.

University of Pennsylvania, $2 50,000, from the estate of H. H. Houston ;

$50,000, for dormitories, the residue unrestricted ; $5000, by the will

No.409.]] GIFTS, APPOINTMENTS, RETIREMENTS. 81

of Dr. J. M. Da Costa ; $250,000, from Mrs. Thomas McKean, for the

law school building.

University of the South, $50,000, from George W. Quintard, of New York.

University of Topeka (proposed), about $250,000, by the will of Mrs. Eliza

Chrisman.

Vanderbilt University, $250,000, by the will of the late Mary J. Furman.

Washington University, property yielding from $120,000 to $130,000 à

year, from Samuel Cupples and Robert S. Brookings.

Washington and Lee University, $3000, by the will of Mrs. Juliet S. Bush-

ford, for a scholarship.

Wellesley College, about $100,000, by the will of Captain George S. Towle ;

$109,000, by subscription ; $100,000, from John D. Rockefeller.

Yale University, $1000, from Professor G. J. Brush; $2500, from an

anonymous donor; $5000, from Mrs. Isaac H. Bradley, for lectures ;

$700, by the will of Dr. James Campbell, for the senior prize ; $1000,

from Mrs. H. F. English, for prize fund in art school; $1000, from

ex-President Dwight, for the art school ; $30,000, from W. E. Dodge.

It is difficult to say how much of the bicentennial fund, amounting to

over $1,000,000, is to be counted as the gifts of the year 1900.

Yankton College, $50,000, from Dr. D. K. Pearson.

York, Pa., $50,000, from Andrew Carnegie, for a public library.

APPOINTMENTS.

Dr. D. Anisito, professor of botany and zoólogy in the medical school at

Asuncion, Paraguay. — Dr. George H. Ashley, professor of natural history

in the College of Charleston, S. C. — Charles E. Banker, assistant in

normal histology, Columbia University. — W. Bergt, professor extraordinary

of geology in the Breslau Technical School. — Dr. R. H. Chittenden, pro-

fessor of physiology in the medical school of Yale University. — Dr. E. B.

Copeland, professor of botany in the University of West Virginia. — Dr.

A. S. Eakle, instructor in petrology in the University of California. — Dr.

Paul Eisler, professor extraordinary of anatomy at the University of Halle.

— Dr. E. Ficalbi of. Messina, director of the zoólogical collections of the

University of Padua. — Carlton P. Flint, assistant in anatomy in Columbia

University. — William E. Ford, instructor in mineralogy in Yale University.

— Dr. B. T. Galloway, custodian of the grounds of the United States —

ment of Agriculture. — Dr. Gasparini, professor of anatomy in the medical

school at Asuncion, Paraguay. — L. C. Glen, professor of geology at Vander-

bilt University, Nashville, Tenn.— Dr. A. W. Grabau, lecturer in geology

in Tufts College. — Dr. Lawrence E. Griffin, instructor in zoblogy. in -

Western Reserve University. — Dr. Joseph B. Grzybowski, docent for

paleontology in the University of Cracow. — Dr. G. Gürich, professor o

geology and mineralogy in the University at Breslau. — Dr. R. Hesse,

82 THE AMERICAN NATURALIST. [Vor. XXXV.

professor extraordinary of zoólogy in the University at Tübingen.— Dr. T.

C. Hopkins, professor of geology in Syracuse University. — Dr. F. H.

Howard, instructor in physiology in Williams College. — Dr. F. Insfran,

professor of histology in the medical school at Asuncion, Paraguay. — Dr.

5. Kástner, professor extraordinary of anatomy in the University at Leipzig.

— Cyrus A. King, instructor in botany in Indiana University. — Dr. L.

Kolderup-Rosenvinge, docent for botany in the Kopenhagen Technical

School. — Dr. Kolkwitz, docent for botany in the agricultural school at

Berlin. — Professor A. Kossel of Marburg, professor of physiology at

Heidelberg. — Professor K. Lampert, curator

collections at Stuttgart. —

vertebrate paleontologist of the United

- Oustalet, professor of zoólogy in charge

e d'Histoire Naturelle at Paris. — Percy

J. Parrot, entomologist to the experiment station at Geneva, N. Y.—

Dr. P. Pefia, professor of physiology in the medical school at Asuncion,

Paraguay. — Dr. N. von Raceborski, professor of botany in the agricultural

school at Dublany, Galicia, — Professor A. Richter, director of the botani-

university at Klausenburg. — Dr. F. Römer

of Breslau, curator of the Senckenberg Museum at Frankfurt a. M. — Dr.

G. E. Rogers, demonstrator in anatomy in the University of Cambridge. —

86, rector of the university at Bonn. — Dr.

Sauer, professor of mineralogy in the Stuttgart Polytechnic School. — Dr.

Alfred Schaper,

reslau. — Dr. F. Schulz, professor extraordinary of physiology in the

university at Jena. — Dr.

assistant in zoólogy in

assistant in zoólogy in the Johns

mith, professor of anatomy in the

Dr. R. Wilson Smith, professor of bot-

Toronto. — Dr. O. Zur Strassen, professor

the university

No. 409.] GIFTS, APPOINTMENTS, RETIREMENTS. 83

RETIRED.

Professor T. G. Bonney, from the chair of geology in University College,

London, after a service of thirty years. — T. Nelson Dale, instructor in

geology in Williams College. — Dr. H. von Eck, professor of geology in

the Stuttgart Technical School. — Carl Gegenbaur, professor of anatomy

in the university at Heidelberg, at the age of 74. — Dr. B. Klunzinger,

professor of zoólogy in the Stuttgart Technical School. — Dr. E. Schmidt,

professor of anthropology and ethnology in the university at Leipzig.

DEATHS.

Sir Henry Wentworth Dyke Acklund, the well-known anatomist and

physician, October 16, aged 85.— Dr. John Anderson, zoólogist and

former curator of the Indian Museum at Calcutta, at Buxton, England,

aged 66. — Paul Blanchet, explorer, of yellow fever, in Senegal. — Dr. G:

Clautrian, assistant in the botanical institute of the University of Brussels,

at Davos, Switzerland, May 23, aged 37. — Dr. A. B. Frank, professor of

botany in the Agricultural School in Berlin, September 27, aged 61.—

Dr. S. Gheorgieff, professor of botany in Sofia, Bulgaria, May 22.—

I. Ingenitzky, entomologist, at Nowovossiisk, Russia, May 20. — Abbé A. B.

Langlois, botanical „collector, at St. Martinsville, La., August 1. ios wih

Joseph Mik, student of Diptera, at Vienna, October 13, aged 62. — Victor

Lopez Sevane, ornithologist and entomologist, at Corufia, Spain, July 14. —

General Sir R. Murdoch Smith, director of the Museum of Science and Art,

at Edinborough, June 3. — Miss Margaret Stokes, an Irish archeologist. —

r. A. von Strombeck, geologist, in Braunschweig, July 25; aged pr. —

Professor G. H. F. Ulrich, geologist and director of the School of Mines at

Otago, New Zealand, in May, as the result of an accident while collecting.

CORRESPONDENCE.

To the Editor of the American Naturalist :

Sır, — Under the name of Xenichthys xenurus, and afterward that

of Kuhlia xenura, a fish in the U. S. National Museum, No. 4356,

which was found in a bottle labeled * San Salvador," was described

by Jordan and Gilbert. I have since had serious doubts whether

this specimen really came from San Salvador and have thought that

it was derived from some Asiatic source. Mr. Barton A. Bean has

compared it at my request with the description of Kuhlia malo from

the Hawaiian Islands. According to Mr. Bean it agrees perfectly

with this species, except that the depth of body is 314 times in

length, while in the adult of the other it is from 224 to 3. This

difference is doubtless due entirely to difference in age. The pec-

toral fin has the measurement recorded by Boulenger for Kuhlia

malo, and the black on the posterior margin of the caudal is very

apparent. Kuhlia xenura should therefore be stricken from the list

of American fishes. It is probable that the type came from Hono-

lulu, where Kuhlia malo is very abundant. DsI

STANFORD UNIVERSITY, CALIFORNIA,

November 15, 1900.

PUBLICATIONS RECEIVED.

BAILEY, L. H. Botany. An Elementary Text for Schools. New York,

Macmillan, 1900. xiv, p. 8vo, 500 figs. $1.10.— Du CHAILLU, PAUL.

The World of the Great Forest. How Animals, Birds, a dein talk,

think, work, and live. Illustrations p C. R. Knight and J. M Ne

ork, Scribner's, eua xiii, 323 8vo. $2.50. Ei doter pne E, and

HEIDER, K. Text of the P p Loredabdlts Translated from the

bna by imt Bernard. Revised and edited with additional notes by

Martin F. Woodward. Vol. iv, Amphineura, Lamellibranchia, Solenoconcha,

Soin Cephalopoda, Tunicata, Cephalochorda. London, Swan, Sonnen-

chein & New baie Macmillan, 1900. xi, 594 pp. 8vo, 312 figs. $4.50. —

E aan W.- Das Tierreich. ro. Lieferung, kien Oligocheta. Berlin,

Friedlander, 1900. xxix, 575 pp» iia 13 figs. - 35 m s.— MiGULA, W. A. de

s Vorlesungen über Bakterien. Dritte ens duin und theil-

weise neu bearbeitet. Leipzig, POI 1900. vi, 186 pp. 8vo, 41 figs.

3.60 marks. — MITCHELL, P. CHALMERS. Thomas Henry Huxley. A Sketch

of his Life and Labors. New VE Putnams, 1900. xvii 297 pp» 8vo. Illus-

trated. — PEPOON, H. S., MITCHELL, W. R., and MAXWELL, F. B. Studies of

Plant Life. A Series of Exercises for the Study of Plants. Boston, D. C. Heath

& Co. 1900. xii, 95 pp., 8vo. $0.50.— Scorr, D. H. Studies in Fossil Botany.

London, Adam and Charles Black, 1900. xiii, 533 pp» 8vo, 151 figs. $2.75.—

TiLLMAN, S. E. A Text-Book of Important Minerals and Rocks with Tables

for the Determination of Minerals. New York, John Wiley & Sons, 1900. viii,

176 pp., 8vo, 38 figs. $2.00. — UNITED STATES WAR DEPARTMENT. Report on

the Census of Cuba, 1899. Washington, Government Printing Office, 1900. 786

Pp., plates, maps, etc. — VERWORN, M. Das Neuron in Anatomie und Physio-

logie. Jena, Fischer, 1900. 54 pp. 8vo, 22 figs. 1.50 marks. — WALLACE, A. R.

Studies, Scientific and Social. Two volumes. London, Macmillan, 1900. 1%,

532 pp. 88 figs., and viii, 535 pp. 8vo, 25 figs. (?). $5.00.— WALTER, H. E,

tory Exercises for the Use of High Schools. Boston, D. C. Heath & Co., 1900.

vi 106 pp., 8vo. $0.50. Teacher's Book of Suggestions to accompany Studies

of Animal Life. pp. xxxi.

` ASHE, W. W. New North-American Plants — Some New Species of Cratæ-

gus. Notes on Some Dichotomous Panicums. N. C. Agr. Exp. Sta. ull.

No. 175. August, 1900, pp. 109-116. — KiNcCAID, T. Papers from the Harriman

Alaska Expedition. VII, Entomological Results ; (1) the Tenthredinoidea. Proc.

Wash. Acad. Sci. Vol. ii, pp. 341-365. — KiNcaip, T. Papers rs from the Harri-

man Alaska Expedition. VIII, Entomological Results; (2) the Metamorphoses

of Alaskan Coleoptera. Proc. Wash. Acad. Sci. Vol. ii, pp. 367-388, Pls. XXII-

XXVI. — Knicut, W. C. Preliminary Report on the Artesian Basins een Wyo-

ming. Wyo. Exp. Sta., Bull. No. 45. June, 1900, pp. 107-251, 14 plates

LAMBE, L. M.- Sponges from the Coasts of Northeastern Canada and Greenland.

86 THE AMERICAN NATURALIST.

Trans. Roy. Soc. Can. [2]. Vol. vi, sect. iv, pp. 19-48, 6 plates. — LUCAS,

F. A. A New Fossil n Leuciscus turneri, from the Miocene of Nevada.

Proc. U. S. Nat. Mus. Nol. i, pp. 333, 334, Pl. VIII. — Lucas, F. A. The

Pelvic Girdle of Zeuglodon. aatis Atoides (Owen), with Notes on Other

Portions of the Skeleton. Proc. U.S. Nat. Mus.

V-VII. — MILLER, G. S., Jk. A Second Collection of Bats from the Island of

Curacao. Proc. Biol. Soc. Wash. Vol. xiii, pp. 159-162. — MILLER, G. S., JR.

A New Gerbille from fatua Turkestan. oan Biol. Soc. Wash. Vol. xiii,

pp. 163, 164. — Morris, E. L. Some Plants of West Virginia. Proc. Biol. Soc.

Wash. Vol. xiii, pp. 171-182. —OsBorn, H. The Genus Scaphoideus. Jour.

Cincinnati Soc. Nat. Hist. Vol. xix, No. 6, pp. 187-209, Pls. IX, X. — PERKINS,

G. H eport of the State Geologist on the Mineral Industries of Vermont,

1899-1900. Burlington, 1900, 83 pp. 29 figs. — WHITEAVES, J. F. On Some

Additional or Imperfectly Understood Fossils from the Cretaceous Rocks of the

Queen Charlotte Islands, with a Revised List of the Species from these Rocks.

Geol. Surv. Canada, Mesozoic Fossils. Vol. i, Pt. iv, pp. 263-307, Pls. XXXIII-

, XXXIX. — WrrHERS, W. A. Another Warning in Regard to Compost Pedlars.

N.C. Agr. Exp. Sta., Bull. No. 173. a ue pp. 83-116.

Brooklyn Medical Journal. Vol. November. — /usect World.

Vol. iv, No. 10. October. sur nier pun Vol. ii, e 5,6. Novem-

ber. — Journal of Zoóphily. Vol. ix, No. 11. November S. U. Naturalist.

Vol. oe No.1. November. — Popular Astronomy. Vol. viii, No. 9. November.

9s N.S, vol vii, No. 78. November. — Stelunca. Bull.

Spee "oos vi, Nos. 21, 22.— Springfield Ebo ubi Vol xx,

o.8. December.

Vol. xxiii, pp. 327-331, Pls.

(No. 408 was mailed December 28.)

TO COLLECTORS

I have a few fine, perfect Specimens of P ADEE

Argo (paper Nautilus), about 2% inches, at $1.00 each.

Also a large number of rare, scarce shells.

List submitted on application.

J. F. POWELL, Waukegan, Ill.

MARINE BIOLOGICAL SUPPLY DEPARTMENT

Preserved material of all types of ae, = Class

work or for the museum. For sg

and all information, addres

_ GEO. M. GRAY, CURATOR - - WOOBS HOLL, Mass.

DISSECTI NG MICROSCOPES

of every size, style and price, suited

for all kinds of work.

A new series of lenses

for dissecting work have

recently been perfected by

us which have unusually

long working distance and

large flat field. These

lenses are offered at

prices less than any lenses of equal quality heretofore

in use. The new Dissecting Stands are all nickeled

metal with glass stage. ~ Catalog free.

Sample copy JOURNAL OF APPLIED MICROSCOPY on request.

BAUSCH & LOMB OPTICAL co.

ee Btreata,

25th Street m ioci E ROCHESTER, N. Y. — Piae d ;

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Published by GINN & COMPANY

= _ JOURNAL OF MORPHOLOGY

sa A Journal of Animal Mo rpholo ogy. Devoted principally to embryological,

-. anatomical, and histological subjects. gg by C. O. Whitman, Head Professor

; ukee, Wis.; F. R. , Univ of Chica ; Ho Ayers, University

of Cincinnati; T. H. Morgan, bim Max Colleg c G. Conklin, University of

ennsyl ; E. B. Wilson, Columbia University. Crown 8vo. Three numbers

` per volume of 100 = pages each, with fro e to den double plates. Sub-

volume ; single n ES $3.50. Agents: for Great

Scription price, T

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: All manuscripts, books for review, RE etc., il 3 sent to the

American Naturalist,” Cambridge, Mass. Annual subscripti: net, in

advance. sd copies, 35 c pes |o subscription; Ee All business

— should be be sent direct to Ginn & Company; Boston.

AMERICAN JOURNAL OF PHYSIOLOGY

See vete the American P Physiological Society by H. P.

R. H. C E eres New Haven;

; to teac M

P e

NO. 410

THE

AMER

NATURALIST

A MONTHLY JOURNAL |

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

AE CONTENTS :

= À Scharff’s History of the European Fauna z 2$ QUU T ROM

2 EE On th: Question 2

and the Int 1

The American Naturalist.

ASSOCIATE EDITORS:

J. A. ALLEN, Pu.D., American Museum of Natural a New York.

E. A. ANDREWS, PE. D., Johns Hopkins University, Sse

WILLIAM S. BAYLEY, Pu.D., Colby University, Waterv:

CHARLES E. BEECHER, Pu. D. Yale University, New eee

DOUGLAS H. CAMPBELL, PH.D, Stanford ae

JH. COMSTOCK, S.B., Corneil University, Jt

WILLIAM M. DAVIS, M. E., Harvard Univ nce Cantik

ALES HRDLICKA, M.D., Mew York 2

D. S. JORDAN, LL.D., Stanford Univer.

CHARLES A. KOFOID, Px.D., Dach d Jllinois, Urbana.

J. G. NEEDHAM, PH.D., Lake Forest Univer sity.

ARNOLD E. DEW, deem Princeton Univer

FRANK RUSSELL, Pu.D., Harvard University, Cambridge.

s ISRAEL C. RUSSELL, LL.D., University of Michigan, Ann Arbor.

WIN F. SMITH, S.D., U. S. Department of Agriculture, Washington.

area STEJNEGER, Smithsonian Institution, Washin ngton.

W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, Pu. D., University of Nebraska, ina.

WILLIAM M. WHEELER, PH.D., University of Texas, Austin.

‘Tue AMERICAN NATURALIST is an illustrated monthly magazine

of Natural History, and will aim to present to its readers the leading

: . facts and discoveries in Anthropology, General Biology, Zoolo.

. Botany, Paleontology, Geology and Physical Geography, and Miner.

alogy and Petrogr phy. The contents each month will consist of

leading. original articles containing accounts and discussions of new

erates reports of scientific expeditions, biographical notices of

T reviews of recent erus; and a final department for

ic ; news and personal noti

dist: who have anything interesting to say are invited

ntributions, bu the editors will endeavor to select

oni that which is is of truly scientific value and at the

en so as to be intelligible, i instructive, and interesting

ntific reader.

d n cripts, books for | review, Sees = should be

iness be sent | direct to the

r pt a s $4.00, avi, in advance, Sago copies, 35 cents.

oF m TOR

THE

AMERICAN NATURALIST

VoL. XXXV. February, 190r. No. 410.

SCHARFF'S HISTORY OF THE EUROPEAN

FAUNA.

LEONHARD STEJNEGER.

IN a recent volume of the Contemporary Science Series, Dr.

Scharff! has given us, under the above title, an elaborate

attempt to trace the origin of the present fauna of Europe and

to some extent also its flora. He has followed up the various

migrations which have taken place in that continent chiefly

Since Pliocene times, and indicated the routes which in his

opinion they must have followed, as well as the obstacles which

barred their way in other directions. This leads to a discus-

sion of the conditions of climate and distribution of land and

water during the glacial epoch. He has thus brought face to

face in one volume some of the most important problems in

zoology, botany, and geology, and whether we agree in his con-

clusions or not, we owe him a debt of gratitude for the admi-

rable manner in which he has laid the whole question before us

and for the mental stimulus which his presentation of it is sure

to infuse into the discussion.

‘Scharff, R. F. The History of the European Fauna. London, Walter

Scott, 1899. viii + 364 pp. 8vo, 21 figs. and maps in text.

88 THE AMERICAN NATURALIST. [Vor. XXXV.

In the introductory chapter Dr. Scharff, starting out from the

present fauna of Great Britain and Ireland, demonstrates its

heterogeneous composition, embracing, as it does, animals, the

affinities or geological history of which point to their southern,

northern, or eastern origin, and shows that in all probability the

southern forms must have extended northward from the conti-

nent long ages ago, while the others arrived comparatively

recently. He then discusses the means of dispersal possessed

by the various animals, especially *the occasional means," as

Darwin called them, and ventures to think that both Darwin

and Wallace have somewhat overestimated its significance, and

he reaches the conclusion that five per cent would be a high

estimate for the animals which have reached British soil by

accidental means. This proportion he regards as insignificant,

in fact as utterly negligible.

In the second chapter, which is headed * Preliminary Con-

siderations," Dr. Scharff endeavors to show how to determine

approximately the original home of an animal so as to be able

to study the component elements of the European fauna. As

one of the results he announces that what was formerly be-

lieved to have been one great northern invasion now resolves

itself into two distinct ones — the Siberian and the Arctic.

An examination of the present distribution of mammals, snails,

and earthworms shows that the British Islands have been

connected with one another and with the continent; Spain

with Morocco across the Straits of Gibraltar; Greece with

Asia Minor, and so forth. The British fauna forms the key

to the solution of the wider problem of that of Europe, five

elements being recognizable, of which the Lusitanian is the

oldest and the Siberian the most recent. A discussion of the

climate of Europe during the glacial period follows, in which

Dr. Scharff tries to maintain that so far from being of an Arctic

nature the climate was mild, possibly even milder than at pres-

ent, to which result he is mainly led by a contemplation of the

striking and most remarkable mingling of a northern and a

southern fauna during the Ice Age. An extensive glaciation,

as advocated by modern geologists, is consequently denied and

evidence brought forward to demonstrate the marine origin

No. 410.] THE EUROPEAN FAUNA. 89

of the boulder clay as opposed to its character as bottom

moraine. Under such climatic conditions portions of the fauna

and flora! were enabled to continue their existence in localities

reached by them in preglacial times.

The British fauna is taken as a convenient starting point

and is treated of in the third chapter. Examples are given of

the more noteworthy forms belonging to the three foreign ele-

ments of which it is composed, viz., the northern, eastern, and

southern, as well as a small endemic one, and adhesion given

to the almost unanimous opinion of biologists that the bulk of

the British fauna and flora (biota? !) is attributable to invasions

by land from the continent. As for the relative age of these

invasions and the geological periods in which they entered the

British Islands, Dr. Scharff feels convinced that the south-

western or Lusitanian fauna, and also the flora, must have

arrived before the glacial period and survived the latter. The

Alpine and Oriental invasions arrived next. After these

came the Arctic, and finally the eastern, or Siberian. The

geological age of the latter is most easily traced because of

the more complete fossil evidence at hand. As the Siberian

invasion arrived in Germany after the deposition of the lower

boulder clay, consequently after the first portion of the glacial

period had passed, it would seem to follow that the Forest

Bed in England, which geologists hold to be preglacial, must

be interglacial, corresponding to the Loess formation of central

Europe. The chapter concludes with statements of facts

Showing a continuous coast line to have existed between

France and Ireland.

With the fourth chapter Dr. Scharff takes up in detail the

discussion of what he calls the * Arctic fauna." The basic

! The author, like many other writers on similar subjects, has felt the need of

* comprehensive term to include both fauna and flora which will not only designate

the total of animal and plant life of a given region or period, but also any treatise

“pon the animals and plants of any geographical area or geological period. As

Such a term I would suggest Biota, not only because its original significance cor-

ers the above definition, but also because of its brevity and obvious relationship

A the term « Biology " as embracing Zoólogy and Botany. Biotic would theo

MY “pertaining to or treating of a biota,” as, — a biotic publication, a biotic

egion.

gO THE AMERICAN NATURALIST. [VoL. XXXV.

supposition upon which rests the whole theory of the Arctic

immigration, in so far as such terrestrial animals are concerned,

which require a continuous land connection, consists in the

assumption that. America and Europe were connected far north

between Scandinavia, Spitsbergen, and Greenland until toward

the end of the glacial period, while Norway again was continu-

ous with Scotland across the North Sea, and England (and

Ireland) with France. The earliest traces of such an immigra-

tion he finds in certain so-called American plants and fresh-

water sponges which are found on the northern and western

coasts of Ireland, in the Hebrides, in Scotland, and in North

America. “The geographical distribution of some of the

Arctic species is referred to in greater detail," to use Dr.

Scharff's own words in his summary of this chapter, “to show

how the relative age of their entry into Europe can be deter-

mined. Two forms of reindeer, resembling the Barren-Ground

and Woodland varieties, have been met with in European

deposits, but only the former occurs in Ireland and the south

of France, whilst eastward the other becomes more common,

and finally is the only one found. It is believed that the

Barren-Ground is the older form as far as Europe is concerned,

and that it came to us with the Arctic migration, and that the

other reindeer reached Europe much later from Siberia, when

Ireland had already become detached from England. The

range of the Arctic hare is equally instructive. It must have

been a native of Europe since early glacial or preglacial times

— before the common English hare had made its appearance

in central Europe. Along with other Arctic forms it entered

northern Europe directly from the Arctic regions by means of

the former land connection which joined, as I remarked, Lap-

land with Spitsbergen, Greenland, and North America.” The

stoat, or ermine, is another distinctly northern animal which

Dr. Scharff regards as having reached Ireland and England at

two different times; the one from the Arctic regions as a

northern migrant, the other with the Siberian fauna from the

east, and its range is made to include

Greenland and Spits-

bergen.

rg He also thinks that there can be no doubt that the

British grouse belongs to the northern migration, and the

No. 410.] THE EUROPEAN FAUNA. QI

ptarmigan and snow bunting are also reckoned as invaders

from the north. Several other animals are mentioned as

Arctic; thus the various lemmings, though it is not quite clear

whether he regards these as belonging to the Arctic immigra-

tion, since they are also included in the animals constituting

the Siberian contingent. The character of the flora is also

alluded to as strengthening the theory of an Arctic (7.e., North-

America- Greenland-Spitsbergen- S linavia-Scotland) route,

though a postglacial connection between Europe and Green-

land is not insisted on, as the present flora of that country

may have survived the glacial period in the Arctic regions.

Professor Forbes's opinion, that the occurrence of certain

shore Mollusca, both on the coast of Finmark and Greenland,

shows that these two countries were not long ago joined, is

alluded to, and the view that the continental boulder clay is a

marine deposit is again brought forward as fitting so much

better with the known facts of distribution.

The Siberian invasion forms the subject of Chapter V, in

Which a number of British animals, living and extinct, are

traced to a Siberian origin, chiefly mammals, and some birds.

Among many of the lower vertebrates and invertebrates there

are but few species which in Dr. Scharff's opinion have reached

England from Siberia. They may have had their original

homes in the Alps, in eastern Europe, or in central and

southern Asia, and have joined in their westward course the

later, more quickly traveling mammals. No less than twenty-

Six species of the Siberian mammals, according to Scharff,

penetrated as far west as the British Islands, and nine of these

still inhabit Great Britain. Some of the remaining seventeen

species probably lived only for a very short time in England,

and the rest gradually became extinct one by one. Our author

enumerates them as follows!: Canis lagopus, Gulo luscus,

* Mustela erminea, * M. putorius, * M. vulgaris, * Sorex vulgaris,

Lagomys pusillus, * Castor fiber, Spermophilus eversmannt, $.

erythrogenoides, Cricetus songarus, Myodes lemmus, Cuniculus

torquatus, * Mus minutus, * Arvicola agrestis, * A. amphibius, A.

or did so within

! Those marked with an asterisk still inhabit Great Britain,

historic times.

92 THE AMERICAN NATURALIST. [VoL. XXXV.

arvalis, * A. glareolus, A. gregalis, A. ratticeps, Equus caballus,

Saiga tartarica, Ovibos moschatus, Alces latifrons, A. machiis,

Rangifer tarandus. It will be noticed that this list contains

species which have also been quoted as Arctic immigrants,

in which case, however, Dr. Scharff regards them as having

arrived at different times both from the north and from the

east. It will also be observed that several species which from

their present distribution are regarded as Arctic, — as, for

instance, Canis lagopus and Ovibos moschatus, the Arctic fox

and the musk ox, — are given as Siberian immigrants, the latter

obviously because it has not been found in Norway, and conse-

quently cannot have come by way of the latter country from

Greenland. In discussing the route by which these animals

arrived and their origin, Dr. Scharff returns to his preterea

censeo, that a mild climate prevailed then in central Europe;

and in reply to Nehring’s “steppe theory ” he states that we

have really no idea under what precise climatic conditions the

Siberian mammals lived in their original home, and offers evi-

dence from other synchronous animals, such as the mollusks,

to show that they afford no proof of a steppe character of the

country at the time when they were alive. Additional support

is derived from the evidence of a connection between the Cas-

pian and the White Sea which would have prevented the

Siberian fauna from spreading westward in Pliocene and early

glacial times. But on disappearance of the marine connection

a way would have been opened into central Europe. The

marine character of the boulder clay is again reverted to, and

the age of the English Forest Bed determined as interglacial,

contemporaneous with the German interglacial beds also con-

taining the bones of these Siberian migrants.

While the so-called Siberian invaders came from the east,

they entered Europe by a more northern route than a large

number of animals which also came from Asia,

but by way

of Asia Minor.

They compose what Dr. Scharff calls the

Oriental migration, which is treated of in detail in Chapter VI,

and of which he himself has given the following summary :

. They originated in central, southern, and western Asia.

It is not easy to discriminate in all cases between the Oriental

No. 410.] THE EUROPEAN FAUNA. 93

migration and the Siberian. To a certain extent, even an

entry of northern Asiatic species has taken place by the

southern route, and vice versa. On the other hand, southern

species might have come to Europe by the southern route —

that is to say, to the south of the Caspian —and also by the

northern, which lay to the north of that great inland sea.

The red deer [Cervus elaphus] is a good example. It arrived

on our continent by both routes. However, there is a racial

difference in the members of the two migrations. The small

race now found in Corsica, Sardinia, northwest Africa, and

western Europe is probably the older of the two, while the

larger one — resembling the American wapiti deer — arrived

very much later from Siberia.!

“The mammoth, wild boar, badger, the dippers, and pheas-

ants are all Oriental species which have come to us from the

southeast ; but there are also reptiles and amphibians, and

a host of invertebrates. Not all the animals, for instance,

which have reached us in England from the southeast are of

Asiatic origin. There is an active center of distribution in

southeastern Europe itself, from which species radiate out

in all directions. This fact is well illustrated by the genus

‘Clausilia. Species from this center, and also from the Alps,

joined the Oriental stream in their northward course.

“In reviewing a number of instances of Oriental species in

Europe one is struck by the peculiarity of their having appar-

ently followed two distinct routes. All entered from Asia

Minor, which is proved to have been connected with Greece

until recent geological times. From here some seem to have

proceeded straight west, others northward. Further study

reveals the fact that the first route was followed by a much

older set of migrants at a time when the Mediterranean area

Was greatly different from what it is at the present day.

Greece was then joined to southern Italy, Sicily, and Tunis.

The latter was also connected with Sardinia and Corsica, and

the Straits of Gibraltar did not exist. Under such geographical

ger race of deer)

10 a d i that it (że. the lar

n page 250 it is stated that it appears that it ( Scharff, how-

also reached England. Cervus elaphus is not enumerated by Dr.

ever, as one of the Siberian migrants on page 202. — L. S.

94 THE AMERICAN NATURALIST. [Vor. XXXV.

conditions a direct migration on land from southern Greece to

Spain was not only possible, but was actually undertaken by

a very large number of Oriental species."

Chapter VII treats of the Lusitanian fauna in detail. As

already mentioned, Dr. Scharff regards this element as the

oldest in the British Islands, its antiquity being indicated by

the discontinuous distribution of so many of its species. As

Lusitanian, he regards species having their origin in the

southwestern portions of Europe, “or on the mysterious

lands which lay beyond it." Not all the species which have

entered Great Britain from that direction, however, are true

Lusitanian, inasmuch as many of the Oriental invaders are

supposed to have traveled as far as Spain by way of the

Mediterranean route, and then, proceeding northward, to have

reached France, Ireland, and England. Only one mammal

is treated of as undoubted Lusitanian, viz., Oryctolagus cuni-

culus, the rabbit ; but several birds are mentioned as having

a southwestern origin, prominent among which are the Dartford

warbler (Melizophilus undulatus), the pied wagtail (Motacilla

lugubris) and the genus Fringilla. Among the lower verte-

brates there are more species of this kind. Thus the ances-

tors of the amphisbzenian lizard (Blanus cinereus) are supposed

to have entered Europe by way of the sunken * Atlantis” ;

other probable Lusitanians are the Discoglossoid toads. and

the salamandine genus Chioglossa. Various butterflies, beetles,

and spiders may be similarly traced to a southwestern origin,

but especially a large number of land mollusks, notable among

which is the spotted slug (Geoma/acus maculosus), which is known

only from Portugal and few localities in southwestern Ireland.

The Alpine fauna forms the subject of the last chapter, the

eighth. The history of this part of the fauna begins with the

rise of the Alpine island in the Miocene sea. This island,

being first connected with the mainland to the east, naturally

received its first inhabitants from that direction ; and Dr.

Scharff regards them as having formed part of the older

Oriental invasion, many species of which, by long isolation

and the elevation of the Alpine country, became modified into

very distinct species, thus forming as it were a new fauna.

No. 410.] THE EUROPEAN FAUNA. 95

At least this is the way I understand Dr. Scharff when he

speaks of indigenous Alpine species. Following these came

the newer Oriental invasion. During Pliocene times dry

land gradually supplanted the sea to the north of the Alps,

and their Biota (fauna and flora) poured into the plain. At

that period Arctic species from the north (via Scandinavia,

Great Britain, and France) and Lusitanian forms from the

west found their way to the Alps. The true Siberian types

came much later, vzz., in the Middle Pleistocene, making

their appearance at the foot of the Alps, though it is doubtful

whether many of them ever reached the mountains. Thus

he accounts for the presence of the so-called Scandinavian

species in the Alps, and vice versa, in two ways — first, an

early northern invasion directly from Scandinavia to the Alps

via England, and, second, by both Scandinavia and the Alps

receiving a share of the Siberian colonists, parts of which

went north, while others went south. Dr. Scharff is thus

led to disagree with Forbes’s theory, once quite generally

accepted, that this similarity between the mountain faunas of

northern and southern Europe was due to a gradual forcing

south of the northern species and north of the southern ones

by the expanding glaciers on both sides, by their mingling in

the intermediate territory and subsequent retreat to their old

homes when the glaciers. receded, the northern forms mixed

with some Alpine species and vice versa. Nehring’s Tundra

theory is also dissented from in so far as it regards the Siberian

animals as forming the nucleus of the faunas of these two areas.

Dr. Scharff ends the summary of this chapter, and thus the

book, with the following remarks: “ One of the most impor-

tant conclusions obtained by this study of the flora in con-

junction with the fauna, is that I have emphasised in most of

the preceding chapters — viz., that the glacial period in Europe

was not a time of extreme cold, and that its destructive

effect on the animals and plants was by no means such as Is

currently believed." :

This last sentence is in a measure a clue to Dr. Scharff's

Whole book, which must be read in the light of it. It must

be admitted that he has made out a good case, from the

96 THE AMERICAN NATURALIST. [VOL. XXXV.

standpoint of the biologists, against the view that would ascribe

to the glacial period so severe a cold and so enormously and

universally developed an ice sheet, or ice cap, that all life

became extinct and every inch of ground hidden by a thick

covering of perpetual ice. I doubt, however, that this is

currently believed by the biologists of to-day. I do not think

I am much out of the way when I say that most of us regard

the conditions and climate of the Ice Age in Europe to have

been on the whole much like the conditions and climate of

Greenland and the island world north of Hudson Bay at

the present time. It would therefore be possible for us to

subscribe to Dr. Scharff's conclusion as above worded were

it not that in the book itself he goes much farther, requiring,

as he does, a climate in central Europe at least as mild as

that of the present day or even milder. The present reviewer

at least cannot admit that the known facts relative to the dis-

tribution of animals and plants during Pliocene and Pleisto-

cene times require such a hypothesis for their satisfactory

explanation.

It may be true *that with a comparatively slight change of

the atmospheric conditions in the British Islands, we might

have glaciers back again on all our highest ranges in England,

Scotland, and Ireland" (p. 69), and it may also be true that

Falsan, whom Dr. Scharff quotes with evident approval, is

right when he says “that the mean annual temperature of

France during the glacial period was approximately from

6-9" C. perhaps more. This," Dr. Scharff continues, “is

the actual mean annual temperature of the southwest of

Sweden and Norway, or the north of Scotland." Of course

this statement is correct enough, but the whole question

assumes a somewhat different aspect when we consider that

nu M - Falsan means a lowering of the

cR Nn c. riim s * This would mean for Berlin

Vienna, a January isotherm lik genae gemis uan

Gobat qid dt dust " et vi of the south end of

between Sitka and the Ali aa we Pipetite ae

laska peninsula. Under the same

conditions Edinburgh would have a summer like the extreme

No. 410.] THE EUROPEAN FAUNA. 97

northern Iceland and Bering Straits and a January temper-

ature like that of Reikiavik.

On page 183 Dr. Scharff makes the following statement:

* Everybody knows that northern and Arctic species can live

perfectly well in a temperate climate, but that it is almost

impossible to acclimatize southern animals in an Arctic or

even temperate one. We have in this circumstance almost a

proof, therefore, that the climate cannot have been very cold.

Though a cold sea bathed the shores of eastern England, and

even eventually invaded a portion of northern England, the

warm ocean on the west must have effectually prevented any

great lowering of temperature." Of course a good deal

depends on what we understand by “great.” At the period

of which Dr. Scharff treats he admits that a cold Arctic cur-

rent came down along the eastern base of Scandinavia, wash-

ing the east coast of England on the west and the northern

shore of central Europe on the south. Now, England has even

to-day a warm ocean on the west coast; does he imagine that

if a cold current — like the Labrador coast current — were to

strike the east coast of England the temperature of the British .

Islands would not be “greatly " lowered? On the two sides

of the Atlantic the warm and the cold currents make a differ-

ence in the annual temperature of the countries of the same ,

latitude of about 10? C. Such a lowering of the annual tem-

perature in Europe east of Ireland would bring the climate of

South Greenland, Labrador, and the Hudson Bay countries

to middle England, Holland, central Europe, down to Vienna,

the northern:edge of the Black Sea, and the northern forth

of the Caspian Sea ; it would also render Lapland and ‘northern

Finland equal to Baffin Land, while the climate of Spitsbergen

would be as severe as that of Grinnell Land or both sides of

Smith Sound. Under similar conditions France and northern

Italy would have the same yearly temperature as Newfound-

land, Winnipeg, and the Aliaska peninsula. The conditions

Which must have prevailed in Germany when land and water

Were distributed approximately as indicated by Dr. Scharff on

is map on page 170 remind one strikingly of a leen

of Hudson Bay and Baffin Bay, a combination certainly no

98 THE AMERICAN NATURALIST. [Vor. XXXV.

liable to ameliorate the climate which in the adjoining coun-

tries cannot have been much different from that of Baffin

Land. This conception is of course widely different from an

absolutely unbroken ice sheet with a temperature destructive

of all higher life, but it is also greatly different from the idea

of Dr. Scharff. Moreover, it is not inconsistent with a possi-

bility of even a considerable number of southern forms having

survived the glacial period in various sheltered nooks and

corners. The humming bird occurs even as far as Sitka,

almost in sight of the gigantic glaciers, and forests are known

to grow above a substratum of ice. The Lusitanian species

in Ireland may well have survived the glacial period, even

including the spotted slug (Geomalacus maculosus). The fact

that it occurs in Cork and Kerry Counties of Ireland to-day

does not prove that it lived there throughout the glacial period,

and that consequently the glacial climate was mild or milder

than now, for on page 156 Dr. Scharff has a chart showing

Ireland to have extended probably a couple of hundred miles

farther south, this extension forming a large peninsula which

must have been washed on both sides by comparatively warm

currents. It is quite possible that the more tender species of

the Lusitanian fauna enjoyed here a congenial climate during

the greatest glaciation, retreating to their present stations in

Ireland as the sea rose and the glaciers receded. It is therefore

scarcely necessary to postulate a temperate climate for Europe

during glacial times. Not only did considerable changes of

the climate take place during that period over large areas, but

there must of necessity also have been a great variation inside

this vast area according to local conditions, while in the adja-

cent countries not directly subjected to the glaciation these

local variations must have been vastly greater. We see even

to-day isolated spots having a southern temperature within the

limits of countries with a northern climate, and on the other

hand similar northern oases in regions bounded by isotherms

indicating a southern climate.

One of the principal reasons which has led Dr. Scharff to

assume such a mild climate in Europe at that time is the

mixture of southern and northern forms in several deposits.

NO. 410.] THE EUROPEAN FAUNA. 99

The occurrence of the bones of the hyena, the reindeer, and

the hippopotamus in the same caverns at Kirkdale, of the lion

and northern voles near Paris, of “the mammoth, the woolly

rhinoceros, horse, ox, reindeer, Arctic fox, lemming, and Pica "

in the same deposit at Thiede, seems to him ample proof of

a southern climate, as he is of the opinion that a northern

species can much easier exist in a southern climate than a

southern species in a northern one. Dr. Scharff continues

(p. 74): “If, in a central European deposit, occurs a mixture

of northern and southern forms of animals, the presence of

the latter is more remarkable than that of the former. Logi-

cally, we should look upon the occurrence of southern species

in the north, therefore, as supporting the view that a mild

climate had induced them to travel northward." In order to

show, however, that this “extraordinary mixture of northern

and southern types of animals" is no indication of a warm

climate, I will call Dr. Scharff's attention to the fact that we

have a striking example even at the present day. It may seem

a paradox to many, yet it is a fact that there is a district in

eastern Asia where the tiger’ (which we are used to regard

as much a tropical animal as the lion and the rhinoceros)

hunts the wild reindeer. That part of Manchuria where this

“extraordinary mixture’’ occurs has an annual temperature

of between o and — 2° C. ; in other words, like south Greenland,

south Labrador, and central Kamchatka. Surely, this woolly

tiger might easily have lived at Thiede during the glacial

period at the time of the deposition of the bones of the mam-

moth, the woolly rhinoceros, etc., and it is just as sure that

the presence of its tropical bones in the German deposit

would not prove that central Europe then had a subtropical

climate,

It seems to me that Dr. Scharff’s whole train of reasoning

in this matter rests on a misconception. I have above quoted

the following axiom of his: «Everybody knows that northern

and Arctic species can live perfectly well in a temperate

climate, but that it is almost impossible to acclimatize southern

animals in an Arctic or even temperate one.” On page 74 he

1 Felis tigris mongolica (Lesson).

IOO THE AMERICAN NATURALIST. [Vor. XXXV.

says: ‘Breeders of animals and those acquainted with zoó-

logical gardens know perfectly well that it is much easier to

keep a northern species in a southern climate than a southern

species in a northern one." Similarly he returns time and

again to the curious delicacy of Arctic plants in botanical

gardens as a proof that their presence in central Europe dur-

ing glacial times is no evidence of an Arctic climate. He even

adduces the fact that Dryas octopetala, one of the most typi-

cally Arctic plants, grows wild in profusion on the coast of

Galway, in Ireland, at sea level, as strengthening “the view,

not only that the Alpine flora is of preglacial origin, but that

the climate of Europe during the glacial period was mild."

That the Arctic and Alpine floras survived the glacial

period there is but little reason now to doubt, but that con-

clusion is not inconsistent with an Arctic climate in which all

these plants even now flourish. But from this admission to

the assumption that the climate was mild is a long cry and the

two conclusions are not logically connected. Much less is the

inference justified, which Dr. Scharff evidently draws, that

the animals and plants which we now find characteristic of the

Arctic regions originated during the glacial period during the

mild climatic conditions and afterwards were driven off into

the Arctic or the Alps by more vigorous invaders from Siberia

accommodating themselves to the much severer climate. With

regard to the plants, here are Dr. Scharff's own words (p. 239) :

“This fact [7.¢., the difficulty of wintering these plants in the

Botanical Gardens] suggests that the Alpine and Arctic plants

really did not originate in countries with cold temperatures.

They probably made their first appearance long before the

glacial period — perhaps in early Tertiary times — chiefly in

the Arctic regions, which at that time had a mild climate.

They have since become adapted to live in cold countries

where they flourish, provided they receive sufficient moisture

in the summer, and are protected from severe frost in the

winter by a covering of snow." Apart from the fact that this

theory is diametrically opposed to his own idea of the difficulty

of southern species to accommodate themselves to a northern

climate, as expressed above, both views, and in fact the whole

No. 410.] THE EUROPEAN FAUNA. IOI

argument, rest upon the fundamentally wrong idea that whole

assemblages of species — or single species for that matter —

can accommodate themselves to so different conditions without

changing their specific identity. Surely the examples which

Dr. Scharff mentions, while referring to the difficulty of keep-

ing southern animals and plants alive in northern climates,

and, vice versa, the greater ease with which northern animals

survive in temperate countries, refer to individuals only, though

he speaks of species. It is a pretty well established fact by

this time that the distribution of an animal or plant (species)

is limited within a certain life zone beyond which it cannot

proceed with impunity, and that this life zone is bounded by

certain isotherms of the propagating season, boundaries which

may differ with each species but which are fairly inflexible

within the species. This Zzzv, which Dr. Merriam has so suc-

cessfully defined and elaborated, renders it certain beyond a

doubt that the presence in central Europe of a breeding and

propagating assemblage of animals and plants practically identi-

cal with that of the Arctic and sub-Arctic regions of to-day

indicates a corresponding climate during the propagating sea-

son; in other words, the life zone of these Arctic and sub-

Arctic species was at some period during the glacial epoch

located in central Europe, plants, animals, temperature, and all.

Another reason for Dr. Scharff's adoption of the theory of

a mild glacial climate is the alleged marine origin of the

boulder clay. "Various deposits of marine invertebrate fossils

in stratified beds are cited as proof that the boulder clay is not

of the nature of a ground moraine. While this question prob-

ably is one in which the geologists are more directly concerned,

I may say that, in my humble opinion, the most reasonable

. explanation is that the boulder clay is of a dual origin, — that

part of it is deposited on land by the ice sheet, while other

Portions were formed at the bottom of the sea, dropped by

floating ice and bergs. There can be but little doubt that the

€normous weight of the Scandinavian ice cap depressed the

land to the south of it, so as to bring it under the level of

the sea. There was probably always a more or less extensive

Sea skirting it to the south and east, preceding the glaciers as

102 THE AMERICAN NATURALIST. [VoL. XXXV.

they extended southward, following them in their retreat to the

north, receiving and distributing the ice floes and icebergs with

their deposits in various places according to the varying con-

ditions, such as changes in currents and winds and the oscil-

lations of the rise and fall of the earth’s crust. A sea like that,

shut out from connection with the Atlantic Ocean to the west,

only connecting with the Arctic Sea to the north, and fed by

the melting ice and snow of the surrounding countries, would

present features something between the Baltic and Hudson

Bay. Its waters would naturally be brackish, and conse-

quently deficient in marine life, except in a few favorable

localities. The supposition of such a sea would meet all the

requirements the biologists can put to it; it would explain the

varied conditions of the boulder clay and the presence as well

as the scarcity of the distinctly marine deposits. It would

also meet their demand for an effective barrier north of the

Caspian Sea to the invasion of Siberian forms during the

earlier part of the glacial period, as a combination of glacier

and sea is as effective for this purpose as either of these

agents alone. On the other hand, it certainly cannot be taken

as an indication of a mild climate, possibly milder than our

present one.

While I have thus been unable to accept one of Dr. Scharff's

more general and fundamental propositions, viz., the one which

relates to the glacial climate, there is another of his more special

conclusions from which I must also dissent, vzz., the northern

origin of the invasion, which he styles the Arctic migration.

As already explained above, Dr. Scharff accounts for the

presence of certain Arctic animals in Ireland, Scotland, and

other parts of western Europe, including the Alps and the

Pyrenean peninsula, by a supposed immigration from America,

via à continuous land connection between Greenland, Spits-

bergen, Norway, Great Britain and Ireland, and France, this

immigration being only subsequent in age to the Lusitanian

fauna, and distinctly older than the Siberian immigration, which

came to Europe much later from the east, though also consist-

Ing, to a great extent, of northern types closely allied to those

composing the Arctic invasion.

No. 410.] THE EUROPEAN FAUNA. 103

There is at once a fatal objection to this theory, vis., that

there can have been no such land connection during Pliocene

or Pleistocene times between Greenland and Spitsbergen. It

was formerly generally believed that the Arctic Sea was a fairly

uniformly shallow basin, but as one of the most important

results of Nansen's Fram expedition we know now that the

sea north of Spitsbergen and Francis Joseph Land is very

deep, certainly more than 1600 fathoms, while to the west of

Spitsbergen, between it and Greenland, we have soundings as

deep as 2650 fathoms. A wide channel between these coun-

tries, certainly not less than 1500 fathoms in depth, connects

the North Atlantic deep with the polar basin. It can be said

with the utmost certainty that an elevation sufficient to bridge

this and thus connect Greenland with Spitsbergen has not

existed during the geological periods mentioned.! It is highly

probable that the extreme elevation in that part of the world

at that time did not exceed the present 300-meter line. At all

events, there must have been a gap between land and land of

at least 150 miles, a distance quite sufficient to bar all migra-

tion of the mammalia which Dr. Scharff includes in his Arctic

fauna.

But apart from this insuperable barrier, there is a good

reason why Spitsbergen cannot have been in the route of these

animals, v/z, that with one exception they do not occur in

Spitsbergen, nor were they ever known to occur there. The

Arctic hare is certainly absent, and the records of a lemming

and the ermine are highly dubious. Even if it should be true

that a lemming occurs there, it is pretty safe to say that it 1s

Tt will scarcely do to regard the dead shells of shoal water forms, such as

Yoldia arctica, which the Danish “ Ingolf ” expedition in 1896 dredged between

Iceland and Jan Mayen Island in depths between 500 and 1300 fathoms, as p m

ofa Corresponding depression since glacial times. It is incredible that these dead

Shells which are scattered all over the surface of the bottom of the North Atlantic

Ocean should: have been lying there loose all these thousands or ten —

years without being covered up or destroyed. I certainly agree with those w 5

hold that these shells have been dropped there by shore ice floated out to sea.

will be noted, moreover, that even the supposition of an extreme rise of I j

fathoms does not affect our argument as to the continuity of Greenland nae

Spitsbergen, since they would still be separated by a deep channel at least 15

miles wide,

104 THE AMERICAN NATURALIST. | [Vor. XXXV.

Cuniculus torquatus, the presence of which would rather work

against Dr. Scharff's theory, since it does not occur in Scandi-

navia at all. The Spitsbergen reindeer, on the other hand, is

so different from the one of Norway that it does not seem

probable that one is the lineal descendant of the other. Add

to this that the sea between Norway and Spitsbergen is more

than 300 meters deep, and an exchange of terrestrial mammals

between these two countries becomes highly improbable.

Before proceeding any farther I wish to state that while

disagreeing with Dr. Scharff in his supposition of a continuous

connection between Greenland and Spitsbergen and between

the latter and Norway, over which his Arctic invaders might

have traveled, I am in full accord with him in regard to the

land bridge between Norway and Great Britain across the North

Sea, and also in his conclusion that most of the immigrants

which he designates as Arctic belong to an invasion different

from and considerably older than the great Siberian immigra-

tion. That I also accept the continuity of Great Britain with

Ireland and France, the breaking down of this connection

between the two former prior to that between England and

France, goes without saying ; but I do not exactly agree with

him in his views as to all the higher vertebrates which he

attributes to this invasion.

To any one who is familiar with the present and past dis-

tribution of the Norwegian lemming (Myodes lemmus}y it

must be somewhat surprising to find it included by Dr. Scharff

among his Siberian colonists, rather than among the Arctic

invaders. The reasons are obvious, vzz., first, because this

rodent does not occur in America and Greenland, whence the

Arctic invasion is supposed to have come; second, because it

has not as yet been found fossil in Ireland, where, according to

the theory, it should have occurred if it arrived in Great Britain

before the land connection with Ireland broke down, an event

later than the arrival of the Arctic invaders according to Dr.

Scharff's chronology. This negative evidence, however, would

have been counterbalanced, I imagine, by the fact that remains

1 i i :

I adhere in this review to the nomenclature used by Dr. Scharff without

regard to my own preference in the matter, in order to avoid confusion.

No. 410.] THE EUROPEAN FAUNA. 105

belonging to the Norwegian lemming type have lately been

discovered in Portugal, since it is one of Dr. Scharff’s criteria

of a Siberian migrant that it does not occur south of the Alps

or Pyrenees, were it not for the impossibility of ascribing an

American origin to this species. As I have shown that none

of the other Arctics can have come that way, and as the lem-

ming otherwise agrees so well with them in their present and

past distribution, I think there can be no difficulty about refer-

ring them to the same category.

Nor can I endorse Dr. Scharff's treatment of the various

ptarmigans (Lagopus) On page 336 he sums up his conclu-

sions to the effect that ** Z. albus and L. mutus appear in our

continent chiefly! as Arctic migrants." His reason for so

regarding the former is given on page 334, where it is stated

that “no doubt the British grouse is a descendant of the Scan-

dinavian willow grouse. The latter is known also to inhabit

Greenland and Arctic North America, and is even found beyond

Bering Straits in northern Siberia." In the first place, as the

British grouse (Lagopus scoticus), with its brown wing-feathers,

is almost certainly nearer the original common ancestor of the

willow grouse,? it would be the Scandinavian willow grouse

(L. albus) which must be regarded as the descendant, if there

really be a direct line of descent between the two, and not vice

versa. In the second place, it is easy to show that the geographi-

cal distribution of Z. a/bus is not that of an Arctic species in

Dr. Scharff's sense. It will be noted that he gives Greenland

among the localities where this species is found. But it is one

of the peculiarities of the distribution of this species that it

does not occur in any part of Greenland. It is also absent in

Spitsbergen and Iceland, and a route of immigration from

America by any one of these countries is consequently out of

! On page 334 he regards the latter both as Arctic and Siberian; see also page 142.

* The Probability that the dark wing color of Z. scoticus might be the result of

reversion is very remote. In a species so variable in its coloring we should in

such à case expect a very frequent cropping out of so universal a character imme-

diately and comparatively recently preceding the last change: It must be m

bered that the white wing-feathers are characteristic of all the other species and

Orms of the genus, and that they are not subject to seasonal color changes like

the rest of the plumage.

106 THE AMERICAN NATURALIST. [VoL. XXXV.

the question. Altogether this species is not found in most of

the truly Arctic regions, as it is absent, not only in the above-

mentioned islands, but also in Novaya Zemlya, Francis Joseph

Land, Taimyr Land north of 72°, New Siberian Islands, Wran-

gel Land, and the Aleutian Islands. Its northern limit conse-

quently agrees almost exactly with that of the wolverine (Gulo)

which Dr. Scharff counts among the Siberian invaders.

With regard to the Tundra ptarmigans (Lagopus rupestris and

mutus) our knowledge is unfortunately not quite so complete,

owing to the difficulty of distinguishing correctly between these

two forms; but, as Palmén has already indicated, it is highly

probable that the latter does not occur east of the White Sea,

that in fact it is confined (with its local races) to the Scandi-

navian mountains with their Lapland spurs, Scotland, the Alps

and the Pyrenees, while Z. rupestris (with its various forms and

subspecies) extends over the entire Siberian and North Ameri-

can Tundra and Barren Ground, as well as Spitsbergen, Green-

land, and Iceland. The former, therefore, is nearly uniform in

its distribution with Myodes lemmus, while the latter corre-

sponds fairly well to that of A. odensis (and its local forms).

The significant fact in this connection, in so far as the ptarmigans

are concerned, is that both the Spitsbergen, the Greenland,

and the Iceland forms belong to the Siberian and American

L. rupestris, while the Scandinavian and Scotch (also probably

subfossil English and Irish) ptarmigans, with those of Switzer-

land and the Pyrenees, form the Z. mutus group. These,

therefore, cannot have come from America via Greenland and

Spitsbergen (or Iceland).

But while thus this group of animals, which Dr. Scharff has

called Arctic immigrants, by both physical and distributional

reasons is barred from the route America-Greenland-Spitsl

Norway-Scotland-western Europe, it may be partinently asked,

By what road did they reach western Europe, Scotland, and

Norway ?

Let us first determine where they did not come from.

Having eliminated Greenland and Spitsbergen, there are to

the northward only two countries which need be further

investigated, vis., Norway and Iceland.

No. 410.] THE EUROPEAN FAUNA. 107

The latter might have been dismissed in a few words, were

it not that Dr. Scharff, in his history of the European fauna,

practically has no reference to the fauna of Iceland, the origin

of which certainly is as European as that of Great Britain itself.

Almost the only allusion to the famous island is a brief para-

graph to the effect that if a land connection existed between

Greenland and Scotland in that direction “it must have been

in very early Tertiary times." Granting that there was no

such continuous land bridge any more at the beginning of

Pleistocene times or even during the Pliocene, by which the

larger herbivorous animals could have migrated from Greenland

to Iceland and Scotland, it is evident that the conditions must

have been much different from what they are indicated to be

on Dr. Scharff's diagrammatic maps on pages 156 and 170, in

which the present sea level is maintained at Iceland and eastern

Greenland, while the continental platform is raised about 200

fathoms at the western coasts of Ireland, Scotland, Norway,

and Spitsbergen. Under such a distribution of land and water

Iceland, though still an island, must have been much larger,

while the Faróe Islands, forming a large island of nearly the

present size of Ireland, were separated from Scotland by a

comparatively narrow channel, and numerous islets on the high

ridge between the Faróe Islands and Iceland constituted a

series of stepping-stones to the latter. Such a state of affairs

would of course effectually block the way of the mammals,

without being a bar to many other animals, as the birds, for

instance. To any one familiar with the land birds of Iceland

it is perfectly plain that it would require at least that much of

an interrupted land connection to make it possible for them to

have developed a highly frequented migration route across that

now nearly trackless ocean. I will mention only one example,

viz., the large-winged race of the common wheatear (Saxicola

ænanthe leucorhoa). This race, characterized by a length of wing

Greenland and adjacent

ca and to migrate in

he Shetlands, Great

tern edge of France

rning in spring the

of over 100 mm., is known to breed in

portions of northeastern Arctic Ameri

winter over Iceland, the Faróe Islands, t

Britain, and probably thence along the wes

and the Pyrenean peninsula to Africa, retu

108 THE AMERICAN NATURALIST. | [Vor. XXXV.

same way to its Arctic home. For this bird to have found a way

to America there must at the time have existed a route fairly

well.outlined by islands more extensive and more numerous

than now, and the very fact that this route of the extension of its

distribution (for it belongs to an exclusively Old-World genus)

became an annual migration-route points plainly to the exist-

ence of such an interrupted land bridge some time in the

glacial period. However, the total absence of the reindeer,

the hare, the lemming, the ermine, and the musk ox in Iceland

is incontrovertible proof that the Arctic mammalian invasion

into western Europe did not come by way of Iceland.

There remains then only Norway as the last possible home

of these animals if they arrived from the north at all. The

question, however, at once presents itself: If they arrived in

Scotland from Norway by way of the North Sea bridge, by what

route did they then come into Norway? It is of course out of

the question to suppose that they originated in that country

situated at the extreme northwest periphery of the Old World,

as they are all closely allied to species of arctogcean derivation.

On the other hand, there seems to have been no other land con-

nection at that period between Scandinavia and the rest of the

Eurasian continent than that with Scotland. It is pretty gen-

erally agreed that the sea then covered the lowlands of northern

Russia to the east, thus effectually cutting off any communi-

cation between Siberia and Lapland. Apart from the consid-

erations which have influenced that conclusion it would be

very difficult to explain the absence from the Scandinavian

peninsula of a number of both Arctic and Siberian animals

had there been a land connection in that region during the

earlier and middle stages of the glacial period. The total

absence of Cuniculus torguatus and the musk ox is particularly

significant.

It would then appear that we are compelled to conclude that

the mammals and birds in question did not come to Great

Britain and Ireland from the north at all. They certainly did

not come from the west, and it is equally certain that they

formed no part of the Lusitanian fauna. There seems then to

be no other way by which they could have reached England

No. 410.] THE EUROPEAN FAUNA. IO9

and Ireland except over the land connection with France.

But how did they get into France? Where did they originally

come from? Various reasons preclude any hypothesis of these

animals being part of the Oriental invasions, ànd the offshoot of

the later, the Alpine fauna. Their relationships are decidedly

Siberian, not central or south Asiatic, and the lemmings, as

well as the willow ptarmigan, are quite foreign to the Alps.

I have already expressed my agreement with Dr. Scharff that

they do not form part of the great later Siberian invasion ; they

were present in northern Europe long before that event, as

shown by their history in the British Islands.

It goes without saying that the relationships of these Arctic

animals are decidedly northern and, as already remarked,

equally closely Siberian, and I have no doubt that northern

and western Siberia was their home, before they invaded

western Europe. I am therefore compelled to recognize two

distinct Siberian invasions (or rather three, since a third one

is in progress to-day) widely separated in time.

The first Siberian invasion (Scharff's Arctic) took place

early, probably before the first great glaciation had reached its

maximum. Neither ice nor water had yet shut off the passage

north of the Caspian Sea and along the northern edge of the

central European mountain ranges. Thus the Tundra reindeer,

the variable hare, the ermine, the Norwegian lemming, the

ptarmigan, the willow grouse, and others penetrated westward

to France (and the Pyrenean peninsula) and over the then con-

tinuous England, Ireland, and Scotland to western Norway

without leaving any traces in the continental boulder clay

which was deposited only after they had passed. The maxi-

mum of the first glaciation then barred the further ingress of

any more Siberians; the boulder clay was deposited next, and

on the retreat of the glaciers or rise of the land, — or rather

both, — the second Siberian invasion, among which were several

of the same species as the first, took place over the boulder clay.

It is my impression that the musk ox and the banded lemming

(Cuniculus torquatus) formed part of the first invasion, but for

some reason or other failed to reach sufficiently far west and

north early enough to pass into Ireland or Norway. It may

IIO THE AMERICAN NATURALIST. [VoL. XXXV.

also be questioned whether the willow grouse reached Norway

at that time, as such an assumption would involve the dilemma

of either regarding Lagopus scoticus as a reversion since glacial

times or else of supposing the Scandinavian and the present

Siberian Lagopus albus to have originated independently and

yet specifically identically from the brown-winged ancestor,

none of which propositions I am at present prepared to accept.

To me it appears most probable that Lagopus scoticus belongs

to the first Siberian invasion reaching Great Britain and Ireland

only, but not penetrating to Norway, and that Lagopus albus,

the white-winged willow grouse, belongs to the second invasion

entering Scandinavia from the south. I do not know that

anybody has been able to distinguish the bones of these two

forms, and it appears most likely to me that the fossil remains

which Milne-Edwards records from France and Italy as L. albus

really belong to LZ. scoticus. The occurrence of the latter south

of the Alps is a fairly good indication that it belonged to the

first invasion. However that may be, the others must have

reached Norway, where they still survive, by the above route,

and while the willow grouse, the Norwegian lemming, and the

Tundra reindeer died out in central Europe, the remaining

species, except the musk ox and Cuniculus, saved themselves

in the southern mountain ranges and in congenial parts of

Ireland.

It is quite probable that several other species, members of

the older Oriental invasion, joined the preglacial Siberian immi-

gration in France. As one of these I regard the red deer of

Scotland and Norway, which doubtless forms a small-antlered

Face, or subspecies, of Cervus elaphus. Its distribution in

Norway is highly interesting and suggestive, occurring, as it

does, only along the western coast from Stavanger up to about .

Namsos (65° north latitude). It is here confined to the outer

coast lint, chiefly to the larger islands, while it is entirely

absent in the interior or eastern Norwa

consequently,

the west and

: y. Ibe deer there,

is restricted to that part of the country lying to

north of the backbone of the great ice cap which

extended from southwest to northeast across the Scandinavian

peninsula during the glacial period. To the south of this ice

No. 410.] THE EUROPEAN FAUNA. III

divide red deer occur at present only in southern Sweden,

but these belong to the great central European race, and it is

highly interesting to note that a subfossil antler of this large

form also has been found in southeastern Norway, in Ringerike,!

consequently on the south side of the ice divide. This large-

antlered deer, therefore, probably entered the Scandinavian

peninsula from the south at a much later period, while the

smaller Scotch form came early across the North Sea bridge

and settled on the islands off the west coast of Norway. It

wil be seen that this hypothesis agrees pretty well with Dr.

Scharff's views, expressed on page 250, as follows: “ There

were probably two distinct migrations of the red deer into

Europe, an older one coming from Asia Minor into Greece,

which stocked Sardinia, Corsica, Malta, and North Africa in

the first place, when these were still connected with one

another. This same migration likewise affected western con-

tinental Europe, the Irish red deer being probably the descend-

ant of this very ancient stock. The latter entered the island

when it was still part of the continent. The later migration

of a larger form came from Siberia and spread mainly over

eastern and central Europe, but it appears that it also reached

England, although there is no evidence of any of these Sibe-

rian deer having ever inhabited Ireland." It will be seen that

the case is exactly parallel to that of the deer in Norway. Dr.

Scharff also calls attention in this connection to the double

invasion of the reindeer, but under my supposition that both

of the latter came from the east the parallelism is still more

striking. n

One isolated fact appears at first sight to be antagonistic to

my theory of western Norway having been invaded from Scot-

land and not vice versa, as Dr. Scharff thinks, vés., the occur.

rence of a reindeer in Spitsbergen. I have above expressed

the opinion that there was not à direct land connection

between Spitsbergen and northern Norway at the time when

Dr. Scharff's Arctic migration must be supposed to have taken

place. Whence then did the Spitsbergen reindeer come from

if not from Norway ? Unfortunately, the interrelationship of

1 Collett. Nyt Mag. Naturv., Bd. xxxvi (1898), p. 360.

II2 THE AMERICAN NATURALIST. [Vor. XXXV.

the various forms, or subspecies, of reindeer is as yet too

imperfectly known! to permit us to answer this question with

any degree of certainty, but I wish to call attention to the fact

that the Spitsbergen reindeer (Rangifer spitzbergensis) is too

different from the typical wild Scandinavian stock to make it

probable that the former is a direct descendant from the latter

or vice versa. On the other hand, the rise of the land in that

region 300 meters would still see Norway separated by the

sea from Spitsbergen, at the same time leaving the latter

connected by dry land with Novaya Zemlya, and there is at

present nothing known which would prevent us from assum-

ing that the reindeer originally came to Spitsbergen from

Novaya Zemlya and Taimyr Land in Siberia.”

In defense and elaboration of his theory of an Arctic migra-

tion by way of Arctic America, Greenland, Spitsbergen, Lap-

land, Dr. Scharff also discusses the travels of the insects and

plants and comes to the conclusion that they argue for a land

connection along this route. I think it altogether likely that

such a dispersal took place, and would even call attention

to the fact that the plants along the north coast of Spits-

bergen show a greater percentage of Arctic-American plants

than those of southern Spitsbergen as corroborative of this

theory, but I must insist that an unbroken land connection is

not at all necessary for the dispersal of plants and insects

along that route, if currents of air and sea were favorable.

Dr. Scharff in his preliminary chapter would reduce this kind

1 If these exceedingly important questions are ever to be solved, it is neces-

sary that they be investigated at once by some competent authority. Not only are

the wild reindeer becoming scarce everywhere from excessive hunting, but they

are also being mixed up in the various localities to such an extent that extreme

care will have to be exercised in using whatever material can now be brought

together. Thus in Norway large herds of the Lapland tame reindeer have been

located on the southwestern fiells, where they mingle with the wild ones of a

possibly different stock; in Spitsbergen also tame Lapland reindeer have become

feral, as some of the draft animals which Professor Nordenskiold brought with

him from Finmarken ran away; tame Asiatic reindeer have been brought by the

thousands across Bering Straits to America, and a herd of Lapland reindeer have

also been introduced into Alaska to add to the confusion.

2 Dr. Scharff in his discussion of the migrations, etc., of the reindeer (pp. 149-

158) seems to argue out of the premises that the Barren-Ground reindeer do

not occur in Siberia, but this is a mistake.

No. 410.] THE EUROPEAN FAUNA. II3

of dispersal to a minimum because the many happy coinci-

dences necessary for a species to establish itself on a distant

shore must be exceedingly rare. But granting this, we must

not forget that while the recorded actual observations of

direct accidental transplantation and subsequent establishment

are few and far between, our entire experience covers scarcely

a couple of centuries, while the periods of which we here treat

are reckoned in thousands, possibly tens of thousands of years.

And how many lucky chances may not have presented them-

selves in ten thousand years! Moreover, the distances with

which we have to do in the present case only look formidable

in the Mercator projection, while in reality they probably did

‘not exceed 150 miles. I believe it to be a distinct mistake to

suppose that all classes and kinds of animals and plants have

followed the same lines of distribution in time and space. As

a matter of fact, it may be even said that there are scarcely

two species which have exactly the same history. Each one

must be worked out separately, and too sweeping generaliza-

tions are dangerous at the present state of our knowledge.

It is one of the distinct merits of a book like Dr. Scharff's

that it makes plain these defects in our knowledge. It is a

kind of stock-taking by which we find out just how our busi-

ness stands. It must then be admitted with regret that the

status is not as satisfactory as one might have reason to expect. —

There is yet a great uncertainty as to the exact and detailed

distribution of many of the larger and more important animals

in the Arctic regions and in Europe. The grosser facts are

known of course in a general way, but they are not sufficient

for the purpose. The finer details are still unknown, or if

known in some isolated cases are unavailing because they are

as yet only isolated. American mammalogists, for instance,

have studied some of the species here treated of in so far as

they relate to American forms, but this knowledge is at pres-

ent a dead one, because the corresponding Asiatic and Euro-

pean forms are still in chaos, or vice versa. Dr. Scharff's book

shows that there is still much necessary work to be done by

the “splitter” of species and subspecies, but it also shows

that this work must be done with some purpose in view and

I I4 THE AMERICAN NATURALIST. [Vor. XXXV.

not in the ordinary perfunctory manner for the mere naming

and labeling of museum specimens. The time has come when

the interrelationship and the sequence in time and space of

the various subspecies must be studied. A perusal of Tke

History of the European Fauna should make clear to the

average describer and namer of animals that his work is only

the means to an end. It brings before him the very

problems for the solution of which his work is necessary,

and the more conscious he becomes of the uses to which it

is to be put, the better and more reliable it is apt to be

performed.

I do not know whether Dr. Scharff is to be commended or not

for having withstood the temptation to correlate the immigra-

tion of man into Europe with that of the other post-Pliocene

mammals. The wanderings of the plants are alluded to in

order to strengthen the views expressed relating to the origin

of the fauna. The travels of primitive man must to a great

extent have followed much the same lines as the other mam-

mals, the same natural barriers being nearly as effective in his

case as in theirs. It might be said that it would require

volumes to exhaust this subject alone, but that is almost

equally true of all the various groups of animals. Then again

it might be objected that our knowledge of the wanderings of

prehistoric man in Europe is very limited and uncertain. I

think, however, it can be safely asserted that it is not any

more so than our knowledge of the migrations of the animals

and plants in the same period, and I venture to suggest

that there is a remarkable similarity between the migra-

tions we have discussed above and those of the various

European subspecies of man.

The first point to be observed is that the dolichocephalic

brunet Mediterranean, or Atlanto-Mediterranean, race in its

distribution both in time and space clearly corresponds to Dr.

Scharff’s “ Lusitanian fauna," The agreement is not only a

general one, but in some details almost startling, as seen if one

compares, for instance, Scharff’s map on page 7 with any map

showing the distribution of the brunet type in the British

Islands.

No. 410.] THE EUROPEAN FAUNA. 115

The next point is the almost self-evident absence of an

* Arctic immigration ” by way of Greenland-Spitsbergen-Nor-

way-British Islands. On the other hand, the “ Alpine fauna,"

the offshoot of the Oriental immigration, is plainly represented

by the brachycephalic Alpine race of unmistakable Asiatic

relationship. These “round-barrow men" in their westward

push reached the British Islands, though they have left but

few traces, except their bones, behind them there. Ireland

may have been separated at the time of their invasion, since

they do not seem to have reached that island, but they pene-

trated to the extreme of Scotland, the Shetlands, the Faroe

Islands, and, as I believe, by that route — the North Sea bridge,

either yet intact or only broken to the extent of furnishing

stepping-stones — to western Norway, where to the present day

this Alpine population holds the extreme west coast to almost

the identical extent as the red deer, a most suggestive distri-

bution when we compare it with what has been said above

about the probable route of immigration of that animal in

Norway. It is true that there are traces of an Alpine popula-

tion in other parts of southern Norway, especially in Smaale-

nene, and that a migration along the Danish peninsula has

been suggested. It is even likely that part of the broad-

headed dark Norwegians have come by the latter route

as well, but this view is not necessarily antagonistic to the

above theory.

Finally the long-headed blond Nordic, or Teutonic, race,

the last to arrive, corresponds substantially to the “ Siberian

invasion.” 1! Whether any part of this can be paralleled with

the preglacial, or first, Siberian stream which I have indicated

above, I am not now prepared to say, but the question should

be carefully looked into.

1 This view does not antagonize the theory of the African origin of the doli-

that the portion of the latter which

chocephalic race. On the contrary, I hold

ultimately developed into the blond branch originally expanded from eastern

Africa to western and central Asia. Here, in the high altitudes, I tke it, the

bleaching began, which after the race joined the second Siberian weary puo

more and more pronounced as it progressed westward and ssa bead , ub n

reached its extreme development of blondness at the extreme nort western p

of its range, viz., in Sweden.

116 THE AMERICAN NATURALIST.

-It will be seen that Dr. Scharff's book has tempted me even

beyond its own limits. This suggestiveness must serve not

only as an excuse for this transgression, but also for the length

of this review. It is one of the greatest merits of the book.

UNITED STATES NATIONAL MUSEUM.

ON THE QUESTION OF AN ARBOREAL ANCESTRY

OF THE MARSUPIALIA, AND THE INTER-

RELATIONSHIPS OF THE MAM-

MALIAN SUBCLASSES.

A REVIEW OF CERTAIN PHASES OF PRESENT AND PAST OPINION.

B. ARTHUR BENSLEY.

RECENT researches have seemed to indicate that the theory,

proposed by Huxley, of a genetic succession of the former repre-

sentatives of the Monotremata, Marsupialia, and Placentalia,

is untenable, and have thus reopened the most fundamental

question of mammalian zoólogy, as to the interrelationships

of the primary subclasses and the possibility of a di- or

polyphyletic origin.

The recent publication by Dollo (99) of the results of his

studies on the foot structure of the Marsupialia, which he

interprets as confirming the growing idea of a placental origin

of the group, renders this an opportune time to review and

comment upon certain features of the evidence which has

lately been produced in opposition to Huxley's theory.

I. Tue EVIDENCE OF AN ARBOREAL ANCESTRY OF THE MAR-

SUPIALIA AND ITS BEARING AS INTERPRETED BY DOLLO.

The question of arboreal ancestry is of considerable impor-

tance in determining the relationships of the Marsupialia, and

Dollo’s results may accordingly be reviewed in some detail.

On page 191 of his paper we find the problem involved set

forth as follows:

“Car si cette étude démontre que les ancétres immédiats des

Marsupiaux étaient arboricoles, il sera prouvé, du méme coup,

que ces Mammiféres ne peuvent représenter la souche des

Placentaires.

117

118 THE AMERICAN NATURALIST. [VoL. XXXV.

* En effet, personne ne soutiendra que zous ceux-ci ont passé

par une phase arboricole. Et quant à ceux qui en sont là

aujourd'hui, ce sont des types sporadiques, qui ont pris nais-

sance indépendamment : la chose est évidente.

* Si, donc, la nature du pied des Marsupiaux nous amène à ce

résultat, que leurs ancétres immédiats étaient arboricoles,

cette conclusion contribuera à appuyer ce que nous ont appris

le placenta et la dentition du lait, notamment :

«A savoir, que les Marsupiaux constituent un rameau latéral

trés specialisé, et non le groupe générateur des Euthériens

actuels."

The essential element of Dollo’s theme is therefore an

attempt to prove, by means of the structure of the foot, that

the Marsupialia are primarily arboreal animals and that in

respect to the possession of an arboreally modified foot they

are too specialized to have constituted the source of the

Placentalia.

Dollo recognizes in the marsupial hind foot certain charac-

ters of arboreal adaptation. These are (a) opposability of the

hallux, (6) predominance of the fourth digit, with reduction

and syndactylism of the second and third, and (c) recession of

the claws. These characters represent successive stages in

the development and perfection of a prehensile foot. Their

purpose is essentially that of placing the great toe in opposi-

tion to the outer digits, notably the fourth, in order to make

the foot an efficient organ for grasping and holding fast to the

limbs of trees.

For convenience Dollo considers the marsupials in two series,

— those which have the great toe well developed and func-

tional, and those which have it reduced, or atrophied. This

division is naturally one of habit ; the former are animals which

have retained the arboreal habit, while the latter are animals

which have abandoned it. In these the great toe, being no

longer useful, has become reduced;

To the first series belong the American opossums (Didel-

phyida) and the Australian phalangers (Phalangerida). In

the Didelphyide (Fig. 1, a), with the exception of the sub-

genera Micoureus and Philander, opposability of the hallux is

No. 410.] THE MAMMALIAN SUBCLASSES.

VOMIT E Uy

Fic. I, a-i. Qah .

Mars runt as red by Dollo

a Didelphys pay porri &, Fistawpor celebensis; c. Tarsiges r rostratus ; d, S bani a

j'aime Antech Er D votes di ur sinus 2

moschatus, (After Dollo) ) For uus see Tie

Peragal, anon ; i, Hypsiprymnodon

I20 THE AMERICAN NATURALIST. [VorL. XXXV.

the only arboreal character presented, the other digits remain-

ing normal. But in Micoureus and Philander there is, in addi-

tion to the opposability of the hallux, an enlargement of the

fourth digit. In the Phalangeridz (Fig. 1, 4) there is super-

added to both of these a reduction and syndactylism of the

second and third digits.

One member of the Phalangeridz, Tarsipes (Fig. 1, c), which

is unique in many other respects, is the only marsupial pre-

senting the latest stage of arboreal modification. Tarsipes

shows, in addition to all of the arboreal characters of the

typical phalangers, a recession of the claws of the fourth and

fifth digits. :

To the second series belong the remaining members of the

Australian fauna, the dasyures and their relatives (Dasyuridz),

the wombats (Phascolomyidz), the bandicoots (Peramelidz),

and the kangaroos and wallabies (Macropodida). These ani-

mals are mainly terrestrial, but they all show arboreal modifica-

tions of the foot.

The dasyures have a functionally tetradactyl foot, derived

from a pentadactyl one by reduction of the hallux. Various

stages of this reduction are illustrated in the family ; Smin-

thopsis murina (Fig. 1, d), for example, shows a well-formed

hallux, while Antechinomys laniger (Fig. 1, e) has none at all.

Wherever the hallux is indicated, however, it is of the opposa-

ble type. As in the Didelphyidz, this is the only arboreal

character represented in the family.

In the wombats (Fig. 1, f) arboreal characters are more

numerous. The hallux is here reduced to a tubercle, but its

transverse position proves it.to have been formerly opposable.

In addition, there is a slight syndactylism and reduction of the |

second and third digits.

The bandicoots have a functionally monodactyl foot, derived

from a pentadactyl one by an enlargement of the fourth digit

and a corresponding reduction of the others, notably the first,

second, and third. It shows all of the arboreal characters of

that of the typical phalangers. The least departure from the

phalangerine condition is seen in Perameles doreyana (Fig. 1, g)

in which the only pronounced modification is a reduction of the

No. 410] ` THE MAMMALIAN SUBCLASSES 121

hallux. The greatest departure from this condition is seen in

Peragale leucura (Fig. 1, 4) and in Cheropus castanotis, in both

of which the hallux is entirely obliterated, the fourth digit

greatly enlarged, and the remaining digits much reduced.

The exclusive predominance of the fourth digit in Choeropus

Dollo regards as significant evidence of former arboreal habit,

since in other cases in which there is a tendency towards mono-

dactylism, as, for example, in the horse, it is not the fourth

digit which predominates, but the third. The fourth digit must

have been enlarged, and the third reduced, in the Peramelidze,

in connection with arboreal habit, before the tendency towards

monodactylism appeared in connection with terrestrial, curso-

rial habit ; so that the already enlarged fourth digit became still

more predominant, while the third became still more reduced.

The kangaroos and wallabies also have a functionally mono-

dactyl foot, resembling closely that of the bandicoots, and

showing a similar range of modification. The musk-kangaroo

(Hypsiprymnodon moschatus) (Fig. 1, 2), for example, which is

the most generalized type of the family, has a foot which, like

that of Perameles doreyana, is but little modified away from

the phalangerine condition. The true kangaroos, on the other

hand, have an extreme monodactyl foot, like that of the higher

bandicoots.

Dollo considers separately the South American form Coeno-

lestes (Epanorthidae) and the Australian burrowing form Noto-

ryctes (Notoryctida). With regard to the former he quotes

the description of the foot given by Thomas, as “ hallux short,

clawless, and not properly opposable ; other digits subequal,

the fourth slightly the longest.” He recognizes this case as

doubtful, but thinks it entirely probable that Coenolestes also

is of arboreal derivation.

In Notoryctes, finally, he finds indicated predominance of

the fourth digit, and reduction and syndactylism of the second

and third. These characters, he states, have not been adap-

tively developed in connection with a burrowing habit because

they are not found in other burrowing animals, such as the

moles (Talpa and Chrysochloris). . They point rather to à

former arboreal habit.

122 THE AMERICAN NATURALIST. [Vor. XXXV.

Dollo accordingly finds in all of the Marsupialia examined a

more or less extensive arboreal modification of the foot, and

from this he draws the following conclusions :

All of the Marsupialia were at one time arboreal.

In the possession of a modified foot they are too specialized

to have constituted the source of the Placentalia.

They are, on the other hand, Aighly modified Placentalia, in

which the placenta has been lost.

They doubtless arose from very primitive Placentalia, in which

an imperfect placentation resulted in frequent accidents, such as

premature birth, and it was probably to avoid these accidents

that the placenta disappeared.

According to this conception there would be four types of

development in the Mammalia, as follows: (1) An oviparous

condition in the Monotremata, and formerly in the Prototheria.

(2) A viviparous condition, zz which the placenta has not yet

appeared; found in none of the existing forms, but formerly

present in the Metatheria. (3) A viviparous condition, with

placenta, in the Placentalia, and formerly in the Eutheria.

(4) A viviparous condition, zs which the placenta has disap-

peared, in the Marsupialia.

II. Huxiey’s HYPOTHESIS IN THE LiGHT OF RECENT

RESEARCHES.

In order to define clearly the position of Dollo and other

investigators on the question of the relationships of the Mam-

malia, it is necessary to notice in some detail the view of

Huxley as expressed in his famous paper of 1880, with which

modern speculation on this subject may be said to have begun.

Huxley recognized three primary and, in a sense, hypo-

thetical divisions of the Mammalia, which he designated as

Prototheria, Metatheria, and Eutheria, these groups being

represented at the present day by the Monotremata, Marsu-

pialia, and Placentalia, respectively. The existing Monotre-

mata and Marsupialia he regarded as highly modified away

from their original types, but the most generalized of the

existing Placentalia (Insectivora) as only slightly so. He

No. 410.] THE MAMMALIAN SUBCLASSES. 123

recognized a genetic succession of Prototheria, Metatheria,

and Eutheria, the Metatheria being intermediate with re-

' spect to the possession of certain prototherian and certain

eutherian characters, among the former notably a non-placental

allantois. To the Marsupialia, in passing from the purely

metatherian condition, he assigned the adoption of the following

special characters: a prehensile foot, associated with arboreal

habit; a marsupium ; a reduced milk dentition; a foreshortened

period of uterogestation.

Huxley therefore implied the following plan of succession :

the Prototheria giving rise to the Metatheria, and becoming

specialized as Monotremata; the Metatheria, in turn, giving

rise to the Eutheria, and becoming specialized as Marsupialia ;

the Eutheria becoming specialized as Placentalia.

In opposition to this Dollo and others regard the Metatheria

as giving rise to the Eutheria or primitive Placentalia, and the

latter as giving rise, on the one hand, by retention of the pla-

centa, to the modern Placentalia, and, on the other, by loss of

the placenta, to the Marsupialia. Other investigators, notably

Osborn, Hubrecht, and Seeley, recognize a separate origin of

the main mammalian stems.

With this introduction we may examine certain features of

the evidence on which these views are based.

I. The Question of a Placental Origin of the Marsupialia.

In the first place, what is the bearing of arboreal ancestry ?

If we assign to arboreal habit any value as a differential

character of the Mammalia, we may recognize two possible

arrangements, based on the views of Huxley and Dollo, in

which this factor might appear. For the sake of TEM

these may be indicated in tabular form as on following page.

According to the former plan arboreal habit would be asso-

ciated with the differentiation of the Marsupialia and Placen-

talia. According to the latter it would also be an index —

following Dollo of a placental (eutherian) origin of ‘the

Marsupialia. This will make it clear that we have two distinct

Problems to decide: one as to the value of arboreal ancestry

I24 THE AMERICAN NATURALIST. [VoL. XXXV.

HUXLEY. DOLLo.

Placentalia Marsupialia Placentalia Marsupialia

Terrestrial Arboreal Terrestrial Arboreal

Metatheria Metatheria Eutheria Eutheria

| / oS

a Eutheria

j (primitive Placentalia)

/ |

Metatheria Metatheria

as an index of a placental origin of marsupials, the other as to its

value as a factor of differentiation of marsupials and placentals.

For the solution of the former we have not farto seek. We

may at once accept Dollo’s statement that the ancestors of the

Marsupialia were, in the possession of a modified foot, too

specialized to have given rise to the Placentalia, because it is

impossible to. make any other assumption. In doing so, how-

ever, we do not in any way compromise Huxley’s view; for it

must be remembered that Huxley derived the Placentalia, not

from Marsupialia, but. from Metatheria. The prehensile foot

he regarded as a special marsupial character. On the other

hand, we cannot accept the evidence of arboreal ancestry as

indicating a placental origin of marsupials. It scarcely needs

to be observed that the assumption by non-placental Meta-

theria of an arboreal habit would account for, and inas satisfac-

tory a way, the arboreal modifications of the foot which Dollo

has demonstrated throughout the existing forms.

. The latter problem yields much more satisfactory results.

It is on this account, in fact, that Dollo's results have been

considered in detail If the former existence of an arboreal

habit throughout the Marsupialia were susceptible of decisive

proof,’ we would have a fairly reliable index of the cause of the

2 ee Oe p evidence of arboreal ancestry as suggestive rather

- Dollo has demonstrated conclusively that the Australian

Marsupialia are of arboreal derivation, but a study of their characters, notably

th *

s Pp des y teeth and feet, convinces me that they have arisen by a rapid adap-

iation from opossum-like ancestors. If such is the case, they are not at

No. 410.] THE MAMMALIAN SUBCLASSES. 125

original separation of marsupials and placentals, because on a

basis of arboreal habit we can explain the more important of

the special characters which now distinguish the former. It is

of interest to notice in this connection the opinion of Huxley :

“I think it probable, from the character of the pes, that the

primitive forms, whence the existing Marsupialia have been

derived, were arboreal animals; and it is not difficult, I con-

ceive, to see that, with such habits, it may have been highly

advantageous to an animal to get rid of its young from the

interior of its body at as early a period of development as

possible, and to supply it with nourishment during the later

periods through the lacteal glands, rather than through an

imperfect form of placenta."! In other words, Huxley regarded

arboreal habit as explaining two important characters of marsu-

pials, namely the prehensile foot and premature birth. Now

to these we may safely add the reduction of one of the denti-

tions, because, as Leche ? has indicated, this is probably attrib-

utable to the peculiar suckling conditions which are brought

about by premature birth. We may also add the marsupium;

Klaatsch (92) has shown that the perfect marsupium is a special

character of the Marsupialia, and its *perfect development in

this group may best be explained on a basis of arboreal

habit. Naturally these suggestions cannot be regarded as

at all final, but they point strongly to an association of the

special characters of marsupials with arboreal habit.

Passing by the results of Dollo, we find the idea of a pla-

cental origin of marsupials also expressed by Wilson and

Hill (97). These writers, in fact, originated this conception

and produced what is perhaps the most significant evidence in

its favor.

The valuable joint contribution of Wilson and Hill (97) on

the tooth development of Perameles, published in 1897, tended

to confirm the idea, conceived by Huxley (80), but elaborated

all representative of the Marsupialia in general In the Jurassic fauna of me

northern hemisphere, and the Miocene fauna of diprotodonts of South America,

we have two important groups for which an arboreal habit cannot as yet be

proven. i

1 Huxley ('80, p. 656).

? Leche. Morph. Jahrb., Bd. xix, p. 525.

126 THE AMERICAN NATURALIST. [Vor. XXXV.

notably by Röse and Kükenthal,! of the former existence of a.

complete double dentition in the Marsupialia, and also indi-

cated their tooth series as directly comparable to those of the

Placentalia. Prior to this the permanent antemolar series of

marsupials had been regarded as homologous not with the per-

manent series of placentals but with their milk series? The

results of Wilson and Hill seemed accordingly to suggest a

fundamental relationship between the two groups in respect

to dentition. .

Similarly, the brilliant discovery by Hill (97) of an allantoic

placenta in Perameles, which was announced at about the same

time, appeared to indicate a further relationship in respect to

placentation, the Marsupialia having been previously regarded

as entirely non-placental.

These writers in discussing the bearing of their results,

although maintaining a moderate position throughout, recog-

nized these resemblances as primary, and indicated their pref-

erence for a derivation of the Marsupialia, by retrogression

both of milk dentition and placenta, from primitive Placentalia.

Now, although it must be admitted that decisive proof of a

former diphyodont condition in the Marsupialia would be of

considerable confirmatory value in the presence of other evi-

dence, it would not of itself suggest placental origin. The

possibility of such a condition was clearly perceived by Huxley

in advocating the reverse. On page 655 of his essay he says:

* I think that there can be no reasonable doubt that the existing

marsupials have undergone a like suppression of the deciduous

teeth, in the course of their derivation from ancestors which

possessed a full set." Huxley, in fact, astutely realized that,

in order to successfully derive the Marsupialia and Placentalia,

it was necessary to assume the existence of a diphyodont

condition in their metatherian ancestors.

We may accordingly pass over the evidence derivable from

the tooth development of Perameles and consider in detail only

that presented by Hill alone from a study of the placenta of

! See Wilson and Hill (97, pp. 433-438).

? Róse, Kükenthal, Leche ; see Wilson and Hill (97, p. 582). This view has

been recently supported by Dependorf (98).

No. 410.] | THE MAMMALIAN SUBCLASSES. 127

this animal and the modifications of the allantois in other

marsupials.

Prior to Hill’s discovery of the placental allantois of Pera-

meles, other investigators! had examined the modifications of

the allantois in certain other forms. Owen, Selenka, and

Semon had described for the phalangers (Phalangeridz) and

kangaroos (Macropodidz) (Fig. 2, c) one condition in which

the allantois was reduced and enclosed by the yolk-sac. Selenka

had found a similar condition in the opossum (Didelphyidz).

Caldwell and Semon, on the other hand, had found in the

koala (Phascolarctos) (Fig. 2, 4) another condition in which

Fic. 2, a~c.—The main modifications of the marsupial allantois arranged in the order of their

primitiveness as interpreted by Hill. (č may represent the type from which æ and c have

been derived.) ; :

4, placental allantois of Perameles (from Parker and Haswell, after Hill); 4, “respiratory

allantois of Phascolarctos cinereus; c, the reduced allantois of Æpyprymnus rufescens

(Macropodidz), (4 and c after Semon); y.s., yolk-sac; aZ., allantois.

the allantois was well developed and freely exposed to contact

with the embryonic envelope, much as in the Reptilia.

In his concluding remarks Hill considers these types and

endeavors to decide as to which one of them is the most primi-

tive; in other words, to determine whether a placental allantois

is being developed secondarily in the Marsupialia, or whether

the placental condition was formerly present throughout the

group and has been subsequently lost in all forms (so far as

known) but one. |

He argues thus: that the reduced condition of the allantois

is necessarily secondary; that although it is possible to derive

1 See Hill ('97, p. 438)-

128 THE AMERICAN NATURALIST. [VoL. XXXV.

the placental allantois of Perameles from one of non-placental

type, like that of Phascolarctos, the reverse procedure is the

more natural one, because Perameles is of more archaic aspect

than Phascolarctos, and the latter is much more retrogressive

in dentition ; that it is entirely improbable that such a struc-

ture as an allantoic placenta should have developed twice-

independently in the Mammalia (Ze. independently in Pera-

meles) On these grounds he would regard the placental con-

dition of the allantois as the primitive one for the Marsupialia

and the non-placental allantois of most existing forms as

secondarily derived by retrogression.

That the typical condition of the allantois has arisen by

reduction, and is therefore secondary, naturally admits of no

reasonable doubt, for no one would suppose that the allantois

would be incipient in animals which have passed through an

oviparous phase. That the evidence derivable from the organi-

zation of the Marsupialia may be interpreted as proving a

reduction of the allantois from a placental to a non-placental

condition is, however, open. to objection, both on general and

on special grounds. In opposition we may maintain that there

is no possible, and at the same time legitimate, way of assuming

retrogression of the allantois on a basis of retrogression in

other structures, especially those of adaptive character.

On general grounds, changes in the embryonic membranes

may not be closely correlated with adaptive changes (progres-

sive or otherwise) in the general organization. For example,

the modifications of the placenta in placental mammals are not

directly related to the adaptive (ordinal) characters of the

animals in which they occur; several modifications may occur

in the same group (e.g. Edentata), a condition which may

possibly be due, as Hubrecht ! has suggested, to the greater

youth of the placenta as compared with other structures of the

animal organization, but which is more probably due to the

fact that the allantoic placenta is not directly affected by

the various evolutionary factors which determine the appear-

ance of adaptive characters in the animal.

But even admitting such a correlation, retrogression of

! Hubrecht (89, p. 588).

No. 410.] THE MAMMALIAN SUBCLASSES. ‘129

placenta could not be assumed on a basis of retrogression in

single structures. For example, if we assume retrogression of

the allantois from a placental to a non-placental condition on a

basis of the more retrogressive character of the teeth of Phas-

colarctos as compared with those of Perameles, as suggested

by Hill, we may also assume the reverse on the evidence of

the feet, for, as Dollo has shown, the foot type of Perameles

is a direct natural advance on that of Phascolarctos.

The greatest objection, however, might be taken on the

special grounds that the reduction and modification of the

teeth which has taken place in Phascolarctos is not retrogres-

sive (* decadent," Hill, p. 435) at all, but is, on the other hand,

progressive and very comparable to that. which has taken place

in ungulate placentals; and, further, that as far as the greater

primitiveness of Perameles or Phascolarctos is concerned there

is little to choose, for although the former is decidedly more

archaic in general dentition, its individual upper molar teeth

are well evolved (quadritubercular), and its foot is much more

specialized.

In all, it may be said that efforts to explain the reduction of

the allantois, whether from a placental or non-placental condi-

tion, by reference to retrogression of the general organization

are foredoomed to failure; for whatever may be true of the

Marsupialia in general, the Australian members of the group,

far from being degenerate, show a progressive adaptive radia-

tion (doubtless made possible through freedom from competi-

tion) which parallels, and is in general very comparable to

that which has taken place during the Tertiary period in

placental mammals.

And in view of this a parallelism of placenta, though not

genetically related to them, need not be more surprising than

the extensive parallelisms in otherstructures. In other words,

there is no adequate reason why the placental allantois of

Perameles should not be regarded as having arisen by progres-

sive modification from one of non-placental (sauropsidan) type.

These suggestions naturally do not imply that the placenta

of Perameles is not primitive, or that in general the reduced

allantois of the Marsupialia has not degraded from one of

I 30 THE AMERICAN NATURALIST. [VOL. XXXV.

placental type, but that, as far as present evidence goes, the

alternative view is just as plausible

The question now to be decided is whether or not the latter

explains the facts of the case equally well.

Admitting the reduced allantois to be secondary, and recog-

nizing the importance of the yolk-sac stage in marsupials, is it

not just as probable that the reduced allantois has proceeded

from one of sauropsidan type as that it has proceeded from

one of placental type? A failure, due to whatever cause, of

the allantois to form a placental connection furnishes a more

plausible explanation of the premature birth of marsupials than

that of Dollo (99, p. 203), which attributes it to the loss of

placenta in the violent gymnastics of an arboreal life. And it

thus affords a simple explanation of the reduction of the allan-

tois, inasmuch as premature birth, instituted through the failure

of the allantois to form a placental connection, provides not only

for the nutrition of the embryo but also for its respiration.

This condition being perfected, the allantois, which is a provision

for respiration during the later embryonic stages, having its

function usurped, has naturally become reduced.

Why the allantois should have failed to form a placental

connection cannot at present be explained. The question

naturally suggests itself, Is not this also attributable to arbo-

real habit? It is not improbable that there may have been

something in the assumption by metatherian animals of an

arboreal habit which, for mechanical reasons, may have made

a placental formation impossible.

While still concerned with the former relationships of mar-

supials, it is of interest to notice certain facts concerning their

ancestry which have been derived from paleontology.

In a recent paper on the origin of mammals, Osborn (9)

has dealt with this subject and has represented in the form of

a chart the probable geological and phylogenetic relationships

of the mammalian subclasses. In this the Prototheria and -

Eutheria! are depicted as primary groups in the Triassic, the

! Following Gill's division of 1872 (cf. Gill,'72). The Eutheria as defined by

Gill are not assigned a placental or non-placental character. Hence they may be

; interpreted as equivalent either to the Metatheria of Huxley or to his Eutheria

as interpreted by Wilson, Hill, and Dollo.

No. 410.] THE MAMMALIAN SUBCLASSES. I 31

former appearing at a slightly later period than thelatter. The

Eutheria are depicted as giving rise, somewhere in the upper

part of the Triassic, to the Marsupialia and Placentalia, which

are represented in the Jurassic by the Triconodonta and Insec-

tivora primitiva, respectively. The significant feature of this

is the time relation which is implied by the recognition of

marsupial and placental types among the Jurassic Mammalia!

If we may argue from the probable sequence of events, it

would appear that we have an important clue to the characters

of the Jurassic forms, namely the single tooth change of Tri-

conodon, providing that the latter is comparable to that of

existing forms. For if the reduction of one dentition is, as

Leche suggests, attributable to premature birth, and the latter

is, as Huxley and Dollo suggest, in turn dependent upon arbo-

real habit, then already in the Jurassic period the marsupials

must have had their special characters well under way ; so that

the common ancestors of marsupials and placentals must be

placed, as Osborn has indicated, at some time prior to the

Jurassic period. Naturally this gives us no clue to a placental

or non-placental origin of marsupials, but the distinction, if

valid, is an important one, because it avoids the confusion which

would result from an attempt to identify and characterize the

common ancestors among the more complex faunas of later

periods. But more especially it minimizes the importance

of the arboreal characters of the marsupials of later periods

and renders necessary a knowledge of the foot structure of

Jurassic forms before decisive proof of arboreal ancestry may

be obtained.

2. The Question of a Polyphyletic Origin of the Mammalia.

As indicated above, the view of Dollo, Wilson, and Hill

opposes that of Huxley merely in reversing the relationships

of marsupials and placentals while retaining the idea of a gen-

eral continuity of the Mammalia. The views of other writers,

on the other hand, substitute for continuity an independent

origin and development of the main mammalian stems.

'91, p. 115).

! Proposed in 1888 ; cf opposite opinion of Flower and Lydekker (91, p. 115)

132 THE AMERICAN NATURALIST. [Vor. XXXV.

Among the latter there is notably that of Hubrecht (95),

which recognizes a polyphyletic origin of the Mammalia from

viviparous Amphibia.

Whether or not the embryological data on which Hubrecht

bases his conclusions justify this interpretation is a question

on which the writer does not feel competent to ‘express an

opinion. It may be noticed, however, that Hubrecht's results

are differently interpreted by Assheton (98), and that his

hypothesis has not been generally accepted.

Hubrecht cites in support of his hypothesis the results of.

Gegenbaur (86), Mivart (88), Osborn (93), and Klaatsch (92).

These may accordingly be examined at the same time in rela-

tion to the view of Hubrecht and that of Huxley.

In the first place, Hubrecht regards the recognition by

Gegenbaur and Mivart of a fundamental difference between

the mammary glands of monotremes and those of the higher

mammals as implying an independent origin of the two

groups. In some respects this is open to a different inter-

pretation. Gegenbaur, while emphasizing the sudoriparous

nature of the mammary glands of monotremes as opposed to

the sebaceous nature of those of the higher mammals, was fully

aware of the limitations of this distinction, since he found

on the mammary areas of monotremes some glands of the

latter kind. In discussing the bearing of this he says (p. 35):

* Aus solchen Verhältnissen des Drüsenfeldes kann man die

Entstehung der Mammardrüsen bei Monotremen und den

übrigen Sáugern sich vorstellen. Bei den Einen ist die

eine Drüsenart, bei den Anderen die andere zur. Ausbildung

. gelangt." In other words, he perceived that from a diffuse

condition, in which both kinds of glands were present, we

may derive the monotreme condition by assuming an exclusive

` development of one set of glands, and the higher mammalian

condition by an exclusive development of the other. The

significance of this was later fully appreciated by Klaatsch

(92), on the discovery by him of both kinds of glands in the

peculiar mammary pouches of artiodactyl ungulates. On

! A more moderate position on this subject has recently been taken by

Jenkinson ('00).

No. 410.] THE MAMMALIAN SUBCLASSES. 133

page 365 of his paper he speaks of a fundamental correspond-

ence between the artiodactyl ungulates and the monotremes in

this respect.

Here we may notice the opinion of Wilson and Hill (97, p.

579): “ We do not believe that in the long run it will be found

possible to maintain the essential dissimilarity of the mam-

mary glandular organ in monotremes." This appears to be

the more appropriate attitude.

With regard to the results of Osborn, Hubrecht quotes,

among other statements, the following one, which he inter-

prets as indicating polyphyletic origin: ** All stem mammals

were related in their double succession (of teeth), in their

dental formula, and in their primitive molar form. These

features point not to a succession, but to a unity of ancestry

of the monotremes, marsupials, and placentals." !

This involves the broader question as to how convergence

of type ought to be interpreted in the Mammalia; for, in

addition to the results of Osborn, those of Wilson and Hill

(97) confirm convergence in dentition, those of Gegenbaur

(86) and Klaatsch (92), convergence in the mammary glands,

those of Klaatsch ('92, 95), finally, convergence in the acces-

sory mammary apparatus. All of these suggest unity of

ancestry.

If there were no convergence of type in the structures

under consideration, Huxley's hypothesis would immediately

become untenable, because it would then be necessary to

assume a polyphyletic origin. But if we admit convergence,

the chances are equally divided between genetic succession and

polyphyletic origin. In order to prove the latter, however, 1t

would be necessary to trace the converging structures back

to a pre-mammalian stage. In other words, it would be neces-

sary to prove that in progressive development certain struc-

tures diverged in character before the mammalian condition was

instituted ; because if any structure diverged from a type which

was still mammalian, the latter would be regarded as also pro-

totherian. None of the above-named structures have as yet

been traced to a non-mammalian type.

1 Osborn ('93, p. 20).

I34 THE AMERICAN NATURALIST. [Vor. XXXV.

. Finally, Hubrecht mentions the results of Klaatsch in con-

nection with polyphyletic origin. He quotes from Klaatsch

as follows: * Dadurch wird man zu der Annahme gedrängt,

dass die Hufthiere sich ganz direkt an jene oben dargestellte

Urform der Saugethiere anschliessen, deren hypothetische

Vertreter man als Taschenthiere oder Bursalia von den Beu-

telthieren oder Marsupialia unterscheiden kann. Es wird

denkbar, dass die Hufthiere niemals ein Marsupial-

stadium durchliefen."!

If we follow Klaatsch throughout the course of his paper,

we find that he recognizes a primary condition of the mammary

apparatus consisting of paired mammary pouches lodging the

mammary glands. Out of these are formed independently the

incipient marsupium of monotremes and the perfect marsupium

of marsupials. In the artiodactyl ungulates these pouches

show no signs of marsupium formation ; they are converted

into the tubular teat cavities. He accordingly concludes that

the artiodactyl ungulates have not passed through a marsupial

stage (Marsupialstadium).

Although this is doubtless a true interpretation, it can-

not be regarded, as Hubrecht suggests, as pointing to poly-

phyletic origin; for it is not our object in contemplating

polyphyletic origin to prove that, for example, the artiodactyl

ungulates have not passed through a marsupial stage, but

that they have not passed through a metatherian stage. The

results of Klaatsch confirm that which Huxley, notwithstand-

ing the meagerness of his evidence, perceived, namely that

the marsupium is not a transitional character but a special

character of marsupials.?

We may again refer to the opinion of Wilson and Hill ('97,

P. 579). Concerning the mammary pouches of Klaatsch they

speak as follows: «Such a rudimentary or primitive condition

must have been characteristic of the common stock of both

Metatheria and Eutheria, and was only a marsupial one in the

sense that Echidna is now marsupial.” From this it is clear

that they do not regard Klaatsch's results as at all indicative

of polyphyletic origin.

! Klaatsch ('92 p. 369). 2 Huxley (’80, p. 656).

No. 410.] THE MAMMALIAN SUBCLASSES. 135

There are still to be noticed certain phases of opinion

which chiefly concern the relationships of the Monotremata.

In a recent paper by Osborn (99) to which reference has

already been made, we find the Monotremes separated off from

the marsupials and placentals, and assigned an independent

origin on a basis of their distinctness of organization. A

similar view has also been expressed by Seeley ('96, '99).

Under the circumstances this view has been a most natural

one. The tendency of recent research has been essentially

towards the revelation of placental characters in the Marsu-

pialia (Wilson and Hill (97), Hill (97)), on the one hand, and

the revelation of sauropsidan (theriodont, Seeley) characters

in the Monotremata (Semon (94), Seeley (96,'99), van Bem-

melen ('98,'99,'00), Smith (99), Sixta (00), Hochstetter (96),

Ziehen (97). Thus the two groups have been naturally

regarded as genetically distinct. It is probable, however, that

in assigning a’ separate origin to them we overestimate their

distance apart: in the first place, on account of the deceptive

mask of placental characters which the Marsupialia, especially

those of Australia, have assumed ; and in the second place,

because, while diligent search has been made for sauropsidan

characters in the Monotremata, little has been done towards a

recognition of the prototherian characters of marsupials. That

the latter field is not unproductive may be seen from the dis

covery by Broom (97), in a foetal marsupial, of a distinct

coracoid comparable to that of the Monotremata, and the

recent results of McClure, showing that the numerous varia-

tions of the postcaval and related veins in the opossum

(Didelphys virginiana) are all modifications of a ground type

which Hochstetter has shown to be characteristic of the

embryonic Monotremata! Moreover, we should not lose sight

of the familiar osteological characters which marsupials and

monotremes have in common.

Reviewing the evidence as a whole, there seems

adequate reason for abandoning Huxley's hy pothesis.

of the facts which have recently come to light are

confirmatory than otherwise, and many others, which have been

Professor McClure.

ms to be no

Many

rather

1 These results were kindly communicated to me by

I 36 THE AMERICAN NATURALIST. [VoL. XXXV.

regarded as antagonistic, are open to a different interpretation.

Since it has not been definitely disproven, and especially since

no other view has been advanced which explains all of the

facts of the case as well, we may justly regard Huxley’s theory

as entitled to first leration, even at the present day.

COLUMBIA UNIVERSITY, NEW YorK.

BIBLIOGRAPHY.

'98 ASSHETON, R. The Segmentation of the Ovum of the Sheep, with

Observations on the Hypothesis of a Hypoblastic Origin for the

Trophoblast. Quart. Journ. Micr. Sci. Vol. xli, pp. 205-262.

'98 BEMMELEN, J. F.van. On Reptilian Affinities in the Temporal Region

of the Monotreme Skull. Proc. Internat. Congr. of Zool, Cam-

bridge.

'99 The Results of a Comparative Investigation concerning the

Palatine, Orbital, and Temporal Regions of the Monotreme Skull.

Verh. d. Kon. Akad. van Wetenschappen te Amsterdam. pp-

81-84. :

'00a Uber den Schädel der Monotremen. Zool. Anzeiger. Bd.

xxiii, Lief. 622, S. 449-461.

'O0b Further Results of an Investigation of the Monotreme Skull.

Verh. d. Kon. Akad. van Wetenschappen te Amsterdam. S. 930-

932.

'97 Broom, R. On the Existence of a Sterno-Coracoidal Articulation in

a Foetal Marsupial. Journ. Anat. Phys. Vol. xi, N.S., pp. 513-

515.

'98 DEPENDORF, T. Zur Entwickelungsgeschichte des Zahnsystems der

Marsupialier. Zoologische Forschungsreisen in Australien und

dem Malayischen Archipel. Bd. iii, Lief. 2, S. 245—402. Jena.

'99 DoLLo,L. Les ancêtres des Marsupiaux etaient-ils arboricoles ? Mis-

cellanées biologiques. pp. 188-203. Paris.

'91 FLOWER, W. H., AND LYDEKKER, R. Mammals Living and Extinct.

ndon.

'86 GEGENBAUR, C. Zur Kenntniss der Mammarorgane der Monotremen.

Leipzig.

74 GILL T. Arrangement of the Families of Mammals. Washington.

'97 HiLL, J. P. The Placentation of Perameles. Quart. Journ. Micr. Sci.

Vol. xl, pp. 385-442.

No. 410.] THE MAMMALIAN SUBCLASSES 137

'96 HocHSTETTER, F. Beiträge zur Anatomie und Entwickelungsgeschichte

des Blutgefásssystems der Monotremem. Zoologische Forschungs-

reisen in Australien und dem Malayischen Archipel. Bad. ii, Lief. 3,

S. 191-300. Jena.

'89 Huprecut, A. A. W. Studies in Mammalian Embryology. I. The

Placentation of Erinaceus europzus, with Remarks on the Phylogeny

of the Placenta. Quart. Journ. Micr. Sci. Vol. xxx, pp. 283-404.

Die Phylogenese des Amnions und die Bedeutung des Tropho-

blastes. Verh. d. Kon. Akad. van Wetenschappen te Amsterdam.

'80 HuxLEv, T. H. On the Application of the Laws of Evolution to the

Arrangement of the Vertebrata, and more particularly of the Mam-

malia. Proc. Zool. Soc. London. pp. 649-662.

'00 JENKINSON, J. W. A Re-investigation of the Early Stages of the Develop-

ment of the Mouse. Quart. Journ. Micr. Sci. Vol. xliii, pp. 61-81.

'92 KLAATsCH, H. Uber Mammartaschen bei erwachsenen Hufthieren.

Morph. Jahrb. Bd. xviii, S. 349-372.

Studien zur Geschichte der Mammarorgane. 1. Theil: Die

Taschen- und Beutelbildungen am Driisenfeld der Monotremen.

Zoologische Forschungsreisen in Australien und dem Malayischen

Archipel. Bad. ii, Lief. 2, S. 77-188. Jena.

00 McCLumr, C. F. W. The Variations of the Venous System in Didel-

phys virginiana. Anat. Anzeiger. (In press.)

'88 Mivart, St. G. On the possibly Dual Origin of the Mammalia,

Proc. Roy. Soc. London. Vol. xliii, pp. 372-379-

'88 OSBORN, H. F. The Structure and Classification of the Mesozoic

Mammalia. Journ. Acad. Nat. Sci. Philadelphia. Vol. ix, No. 2,

pp. 186-285.

'93 —____ The Rise of the Mammalia in North America. Stud. Biol.

Lab. Columbia College. Zodlogy. Vol. i, No. 2, pp. 1-45-

The Origin of Mammals. Amer. Journ. of Sci. Vol. vii, pp.

92—96. :

'96 SEELEY, H. G. On the Complete Skeleton of an Anomodont Reptile

(Aristodesmus riitemeyeri Wiedersheim) from the Bunter Sandstone

of Reihen, near Basel, giving New Evidence of the Relation of the

Anomodontia to the Monotremata. Proc. Roy. Soc. London. 1896.

pp. 167-169. :

The Origin of Mammals. Jnternat. Congr. of Zoöl, Cam-

bridge. (Printed abstract. iR

'94 Semon, R. Die Embryohiillen der Monotremen und Marsupialier.

Zoologische Forschungsreisen in Australien und dem Malayischen

Archipel. Ba. ii, Lief. 1, S. 19-58. Jena.

'00 SIXTA, V. Vergleichend-osteologische Untersuchung über den Bau

des Schädels von Monotremen und Reptilien. Zool. Anzeiger.

Bd. xxiii, Lief. 613, S. 213-229.

138 THE AMERICAN NATURALIST.

'99 SMITH, G. E. Further Observations on the Anatomy of the Brain in

the Monotremata. Journ. Anat. Phys. Vol. xiii, N.S., pp. 309-

342.

'97 WILSON, J. T., AND HILL, J. P. Observations on the Development

and Succession of the Teeth in Perameles ; together with a Contri-

bution to the Discussion of the Homologies of the Teeth in Marsu-

pial Animals. Quart. Journ. Micr. Sci. Vol. xxxix, pp. 427-588.

'97 ZIEHEN, T. Das Centralnervensystem der Monotremen und Marsu-

pialier. 1. Theil: Makroscopische Anatomie. Zoologische For-

schungsreisen in Australien und dem Malayischen Archipel. Bd.

iii, Lief. 1, S. 1-187. . Jena.

For references to papers of Róse, Leche, Kükenthal, and others, on the

dentition of the Marsupialia, see Wilson and Hill ('97, p. 582).

For references to papers of Owen, Caldwell, Selenka, and Semon, on the

marsupial allantois, see Hill ('97, p. 438).

THE THEORIES OF THE ORIGIN OF THE ANT-

ARCTIC FAUNAS AND FLORAS.

ARNOLD E. ORTMANN.

WHILE preparing a report on the fossil Tertiary invertebrates

of Patagonia collected by the Princeton expedition, the writer

was led to collect the literature on the subject of the resem-

blance of the southern faunas and floras (of South America,

South Africa, Australia, and the Antarctic islands), and was

very much surprised by the vague and sometimes incorrect

rep tations of the existing theories relating to this fascinat-

ing zoógeographical question. Many authors do not quote their

predecessors at all, while others refer to them only in a very

general way, occasionally misstating their views or giving

incorrect or defective quotations. I have, therefore, tried to

collect everything that has been written on this topic, and

think it will be worth while to give here a condensed report

on the subject.

One of the first to call attention to the resemblance of

southern life, in this case to that of the flora, and certainly the

first to advance a theory, was Hooker! He is of the opinion

that we could possibly explain the fact of the existence of

identical plants in southern lands widely distant from each

other by the assumption that there was once a connection of

these parts by land. This theory, first expressed very cau-

tiously, and at that time much disputed, was again more ener-

getically propounded by Hooker,? and, in the last paper, with

reference to the Darwinian view of the origin of species.

In the course of time this theory was almost forgotten, at

1 Hooker, J. D. The Botany of the Antarctic Voyage of H. M. Discovery Ships

Erebus and Terror (Flora Antarctica), Pt. ii (1847), p- 211- |

? Hooker, J. D. Introductory Essay to the Flora of New Zealand (1853),

Pp. xxiii ff. ; and On the Flora of Australia; its Origin, Affinities, and Distribu-

tion, Botany of the Antarctic Expedition, Pt. iii, vol. i (1859), pp. xvii and civ.

139

140 THE AMERICAN NATURALIST. (VoL. XXXV.

least it is not referred to by the next writers to be mentioned,

whom we should regard as the founders of the theory of the

Antarctica. .The first of them is Ruetimeyer.! He states dis-

tinctly that we should take a part of the present faunas of.

South America, South Africa, and Australia for remnants of

an old fauna that spread over a larger extent of the Antarctic

continent, and that this Antarctic continent was the center of

origin of a peculiar Antarctic fauna.

Practically the same idea — but without reference to Rueti-

meyer — was set forth a little later by Hutton?; that is to say,

he also assumed the former larger extension of the Antarctic

continent, and its connection with the southern ends of the

present continents.

In 1875 Gill? relying on his studies on fishes, constructed

his Eogaea, which was apparently conceived as a large conti-

nental mass, embracing Africa, South America, and Australia ;

but no mention is made of the Antarctica entering it.

In opposition to all these theories, which construct land

bridges, where there is now deep water, Wallace,* consistent

with his views on the permanence of land and oceans, entirely

repudiates these opinions, and believes that the faunal elements

common to the southern continents are remnants of a formerly

more extensive distribution, and have been pushed into the

southern ends of the land by the competition with other

animals.

In favor of this view, Hutton? abandons his first theory of

a connection by an Antarctic continent. But he still main-

tains that there must have been a connection between Aus-

tralia and South America, and he constructs a bridge across

the mid-Pacific, assuming a large Pacific land mass, which was

+ i L. eus die Herkunft unserer Thierwelt. Basel, 1867.

2 Hutton, F. W. the Geographical Relations of the New Zealand Fauna.

Trans. jet ew Zidane x. vol. v (1873) ; reprint in dan. Mag. Nat. Hist., Ser. 4,

vol. xiii (1874).

Gill, T. On the Geographical Distribution of Fishes, Ann. Mag. Nat.

Hist., Ser. 4, vol. xv (1875)

* Wallac mo A. R. The Geographical Distribution of Animals, vol. i (1876).

5 Hutton, F. W. On the Origin of the Fauna and Flora of New Zealand, Wew

Zealand partis Sci, January, 1884; reprint in Ann. Mag. Nat. Hist., Ser. 5, vol.

xiii, June, 1884.

No.410] ANTARCTIC FAUNAS AND FLORAS. I4I

joined, on the one side, to Australia and New Zealand, and,

on the other, to Chili. |

Von Ihering ! takes up again the Antarctic communication,

but at the same time he accepts, at least in part, Hutton's

Pacific continent. According to him, a large land mass

extended from South America across the Antarctic regions

to Australia, from whence it continued into the Pacific conti-

nent, and even communicated with Asia (Archinotis). The

Pacific continent was not directly connected with Chili.

While all these writers expressed their opinions on the

Antarctica only in a very general way, Forbes? was the first to

give it a more definite shape by drawing a map of it. He

constructs his Antarctica by raising the land to about the

present two thousand fathoms line, which results in an enor-

mous extension of the Antarctic land masses.

In opposition to this, Hedley? restricts the Antarctica con-

siderably, and admits only narrow connections of it with the

other continents. He does not think that New Zealand was

ever joined to the Antarctica by land, and does not believe in

Hutton's Pacific continent, although a part of the Pacific islands

were once connected with Australia and New Zealand. Of

the latter relations he gives a map in his second paper.

Finally, Osborn* gives a map of the possible extent of the

Antarctica by raising the land to the 3040 meters line. This

attempt resembles somewhat that of Forbes, but shows a

1 Ihering, H. von. On the Ancient Relations between New Zealand and South

America (Trans. New Zealand Inst., vol. xxiv, 1891), and Die Ameisen von Rio

Grande do Sul (Berlin. entomol. Zeitschr., Bd. xxxix, 1894). See also Science, v,

December 7, 1900.

Von Ihering has referred to this subject in numerous other papers, and it is

extremely difficult to collect all of them, since a large number have been pub-

lished in out-of-the-way places. Lists of them have been Ae by himself in

Engler’s Botanische E Bd. xvii (1893), p. 9, and Science,

Fo rbes, e Chatham nt g eir Relation to a der Southern

); abstract: Antarctica, a

C Con r

Ancient Anas Life (Proc. Roy. Soc. N. S. Wales, 1895), and A Zoógeo-

graphic Scheme for the Mid-Pacific (Proc. Linn. Soc. N. S. Wales, 1899)-

4 Osborn, H. F. The Geological and Faunal Relations of Eur

America during the Tertiary Period, etc., Science, April 13, 1900.

ope and

I42 THE AMERICAN NATURALIST.

tendency to restrict the boundaries of this continent to reason-

able dimensions, thus approaching Hedley's ideas.

Other writers have only added new facts to the material

already at hand that favors the assumption of a former conti-

nental connection of the southern regions, and they have all

accepted the general idea of Hooker and Ruetimeyer, without

making any material change in it; therefore I do not consider

it necessary to mention them here.

We may sum up all theories advanced, and put them into a

table in the following way :

I. Theories assuming a land connection between the respective parts.

This general idea was first expressed by Hooker (1847). It has

been accepted by all subsequent writers except Wallace.

1. The land bridge is placed across the present Antarctic continent,

first by Ruetimeyer (1867) and by Hutton (1873). It was

accepted by Von Ihering, Forbes, Hedley, Osborn.

(a) Forbes constructs his immense Antarctica (1893).

(^) Hedley restricts it to reasonable limits (1895).

2. Gill constructs his Eogzea, a continent uniting Africa, South America,

and Australia, but leaving out the Antarctica (1875).

3. Hutton connects Australia and South America by his mid-Pacific

continent, but denies the existence of an Antarctic connection

(1884).

II. Theory of Wallace (1876) rejecting any land connections whatever

between the respective parts.

I do not want here to go into any further detail, since my

only purpose is to give an account of the existing theories with

proper references, and to classify them according to their

contents. But I may state here that in the forthcoming

report on the Patagonian fossils mentioned above, I shall accept

Hooker's general idea, as well as Ruetimeyer's Antarctica

theory, with the restrictions put upon it by Hedley, and thus

we may call it 7e Hooker-Ruetimeyer-Hedley theory.

PRINCETON UNIVERSITY.

THE GENERIC NAMES MYRMECOPHAGA AND

DIDELPHIS.

OLDFIELD THOMAS.

THERE have recently appeared in America two papers on

these generic names, the one in this journal, by Mr. J. A. G.

Rehn,! and the other, criticising the first, by Dr. J. A. Allen?

The points raised in both appear to me to need further dis-

cussion, especially with regard to the somewhat ready manner

in which the species usually quoted as the type of each name

is thrown aside as unrecognizable.

I. MYRMECOPHAGA.

With regard to Myrmecophaga tridactyla, which, if determi-

nable, would admittedly be the type of the generic name, Mr.

Rehn, while discussing Linnzeus’s references to Seba and Ray,

altogether ignores the very first quotation of all, that to Marc-

grave (* Tamandua-guacu, Marcgr. bras. 225"). On examina-

tion Marcgrave's animal proves to be beyond all question the

great ant-eater of Brazil, and on the general principle of taking

the first important reference as the basis for Linnaeus's names

we must clearly accept the great ant-eater as the type of the

genus Myrmecophaga.

Linnaus's words, “ macula nigra a pectore versus latus ducta

.and “cauda lata,” are also diagnostic of the great ant-eater,

mistaken as are his statements about the digits and mammary

formula.

It is unfortunate that the specific name, dating from the

tenth instead of the twelfth edition of the Systema Nature,

must be zridacty/a instead of the familiar jubata; but even this

1 Am. Nat., vol. xxxiv, p. 575, July, 1900.

2 Bull. Am. Mus. N. H., vol. xiii, p. 185. October, 1900.

143 "

I44 THE AMERICAN NATURALIST. [Vor. XXXV.

is a lesser evil than the transference of Myrmecophaga to the

Tamanduas and the dubbing of the great ant-eater with a new

generic name.

Dr. Allen says that while differing about Didelphis, he

agrees with Mr. Rehn in regard to Myrmecophaga, but as he

seems also not prejudiced (as I am) against dismissing old

names as unrecognizable, and has probably not looked up the

vital reference to Marcgrave, I do not feel justified in accept-

ing Mr. Rehn's conclusions, even when backed by so great an

authority as he is.

As a result I claim that Myrmecophaga tridactyla Linn.

should be the name for the great ant-eater, Uroleptes and

Cyclopes remaining as before for the other genera of the

family.

II. DipErPnurs.

In this case Dr. Allen most rightly refutes the necessity

asserted by Mr. Rehn for calling all the large opossums

Sarigua instead of Didelphis, but I fail to be convinced as to

his application of the specific name marsupialis to the Virginian

opossum. This name marsupialis hangs primarily on Seba's

figure, and both authors claim that the latter is unrecognizable,

Dr. Allen going on to make the remarkable statement that

it is *not an American animal, but a species of Phalanger

from Amboyna.” It is true that Seba said his animal was

from Amboyna, but even at that date mistakes could be made

as to locality, and no one familiar with Phalanger, its general

appearance, dentition, or mammary formula, could for one

moment suppose that Seba’s figure and description were based

on a member of that genus. Among the whole of the Mam-

malia the peculiar arrangement of the mammze is alone abso-

lutely diagnostic of a Didelphis, nor have I any doubt what

opossum it is meant to represent.

Seba's South American animals nearly all came, as was

natural, from Guiana, and the figure will perfectly suit the

ordinary dark opossum of northeast South America, for which

Dr. Allen has used the name D. karkinophaga.

No. 410.] MYRMECOPHAGA AND DIDELPHIS. 145

This conclusion, if accepted, will have the double advantage

of retaining the familiar term virginiana for the Virginian opos-

sum, and abolishing so jaw-breaking a name as harkinophaga.

In the necessity of renaming Philander I am compelled to

acquiesce, though I may express a regret that in giving the

name Caluromys Dr. Allen has departed from the modern

practice of restricting the ending -mys to members of the

Rodentia.

But of his transference of the species cinereus and alstoni

from Marmosa to Caluromys I find it more difficult to approve,

for the points that he mentions as allying them to the latter are

all found in different degrees in one or other of the larger

species of Marmosa, including M. murina, while neither cinereus

nor alstoni present those which are more truly characteristic of

Caluromys. The general shape of the skull of the latter, the

small rounded molars, the great sabre-like canines, the curi-

ously shaped lower jaw, and many other characters make up

an ensemble to which, as it appears to me, neither of the

species referred to shows any real approximation. I should

therefore consider them both as members of the genus

Marmosa. i

BRITISH MUSEUM, NATURAL HISTORY.

THE APPLICATION OF DIDELPHIS MARSUPIALIS

LINNAEUS.

JAMES A. G. REHN.

REcENTLy Dr. J. A. Allen published a paper, ** Note on the

Generic Names Didelphis and Philander” (Bull. Amer. Mus.

Nat. Hist., XIII, pp. 185—190), in which he takes exception to

several statements recently published in this journal (XX XIV,

pp. 575—577) by the present writer. A very hasty examination

of Dr. Allen's paper convinced me that some of his conclusions

must have been hastily formed, as they show a lack of detail.

A more painstaking scrutiny has disclosed several interesting

points in regard to the references cited by Linnaeus under his

Didelphis marsupialis; as, like Dr. Allen, I have changed my

views regarding the recognition of this name.

To quote from Dr. Allen : **It is a recognized rule of nomen-

clature that a name applied to a composite group, whether

Specific or generic, must be conserved for some one of its

components when the group is later subdivided.” With this I

agree perfectly, and, as Dr. Allen argues, think we should give

each reference cited under the name a close examination, the

description being too indefinite to furnish anything. This Dr.

Allen has done, but not as thoroughly as it should be.

To take the references mentioned above, they are as follows :

Philander Seb. Mus., I, p. 64, t. 39.

Opossum Tyson, act., 290, p. 1565.

Carigue Laét, Amer., 551.

Carigueija brasiliensibus Marcgr. bras., 222; Pis. bras., 323;

Jonst. quadr., 136, t. 36.

Maritacaca Pis. bras., 323; Ray quadr., 182.

Tlacuatzin Hernand, mex., 330.

Concerning the first, the Philander of Seba, we find some-

thing of interest even if it does not affect the whole. An

147

148 THE AMERICAN NATURALIST. [Vor. XXXV.

examination of this work shows that if Dr. Allen had exam-

ined it closer he would have modified his statement that **the

Philander of Seba . . . isnotan American animal, but a species

of Phalanger from Amboyna." The first part of the descrip-

tion of this animal is quoted from Valentino (Amdboinensis

Animal., p. 273), and while it very probably relates to a species

of Phalanger as Dr. Allen states, the latter part of the text

(which is that of Seba) undoubtedly refers to an opossum.

Such statements as **Caput, canino simile, longum protendit

rostrum, acutisque dentibus . . . dorsum longis pilis, setaceis,

atro spadiceis, hispidum est . . . cauda, longa, squamis rhom-

boideis tota tegitur," surely do not refer to a Phalanger. Seba

makes the statement that Valentino had made a great error in

describing the animal as he did; accordingly he gives his idea

of the same, which, with the plate (undoubtedly a Didelphis),

shows that Seba had nothing but an opossum in mind.

The second, Tyson's work, has been already closely examined

by Dr. Allen, and nothing further can be said regarding it.

The third reference, the Carigue of De Laét, I cannot examine,

and therefore will pass over, as very likely the animal is a South

American opossum which would not be identifiable.

The Carigueija of Marcgrave, Piso, and Jonston is an opos-

sum of South American origin, but both the description and

the locality are too indefinite to be considered.

The fifth, the Maritacaca of Pisonis, I cannot examine, but

Ray has quoted it in its entirety, from which the same con-

clusions as the foregoing would be entertained.

The last, the Tlacuatzin of Hernandez (Anim. Mex., p. 330,

1651), is an opossum which represents the form inhabiting the

southern end of the tableland of Mexico, the Cordilleras, and

Vera Cruz, as Mr. Nelson (Science, N.s., VIII, p. 897) has

pointed out regarding Spermophilus variegatus Erxleben, which

was based on an animal described by Hernandez. To quote

from Mr. Nelson: * At the time when Fernandez made his

observations the main area of Spanish occupation in Mexico

was the southern end of the Mexican tableland, about the valley

of Mexico, and thence eastward across the plains of Puebla,

throughout the Cordillera to the hot lowlands of Vera Cruz.”

No.410] DIDELPHIS MARSUPIALIS LINNAEUS. 149

According to the evidence set forth, the point is raised, —

should Tyson’s reference be given precedence over that of

Hernandez?

Disregarding the dates of the references (that of Hernandez

being forty-seven years older), we will proceed with the work

by elimination.

The opossum “from Virginia" was separated by Kerr in

1792, while the southern Mexican animal, which, if it has

been separated, can only be either californicus or breviceps

Bennet, which were not described until 1833. The only other

species of the genus which is found anywhere near the locality

in question is D. aurita, which has not (to my knowledge) been

recorded north of Guatemala. The elimination of Tyson’s

reference by the D. virginiana Kerr leaves us only the

reference of Hernandez as the base of Didelphis marsupialis

Linnzeus.

ACADEMY NATURAL SCIENCES, PHILADELPHIA.

THE SNAKES OF NEW YORK STATE: AN

ANNOTATED CHECK LIST.

EDWIN C. ECKEL.

No paper on the ophidian fauna of this state has appeared

since Baird’s “Serpents of New York,” published in the Seventh

Annual Report on the Condition of the State Cabinet, Albany,

1854. That which is now presented is merely a list of those

species which have been mentioned more or less definitely as

occurring within our limits.

The notes which I have added are of two classes. In the

case of species whose presence within the state is doubtful,

I have given the authority for their inclusion. In this con-

nection I have endeavored to give credit, for the introduction

of new species into our faunal lists, to those to whom it is due.

My own observations on the snakes of this state have been

confined largely to that portion of the state which I traverse

during my geological field work. With this area and with its

fauna I am fairly well acquainted, having spent my summers

for the past twelve years within it. I have, therefore, added

a few notes upon the occurrence and abundance of the various

species in southeastern New York. This area, as the term is

used here, has geological rather than geographical boundaries.

Roughly defined, it includes the counties of Orange and Rock-

land, on the west of the Hudson, and those of Westchester,

Putnam, and the southern half of Dutchess, on the east of the

river. It is an area of crystalline rocks, bordered on the north

by a usually well marked depression based on limestones and

shales. In one case at least: (e herpetology mcus to be

affected by the geology. The rattlesnake is rarely seen out-

side of the Highlands, while the copperhead is abundant in

the great Cambro-Silurian lowland which adjoins the High-

lands on the north. The ophidian fauna of the region 1s rich

ISI

152 THE AMERICAN NATURALIST. (VOL. XXXV.

in both species and individuals, while in the Catskills, not far

away, there is a notable scarcity of snakes.

I have adopted the terminology and classification of Cope,

as set forth in his Crocodilians, Lizards, and Snakes of North

America, though a note under the heading of Eutenza sirtalis

expresses my dissent from his method of handling the sub-

species of that group.

—

Carphophiops amenus (Say).

Diadophis punctatus (Linn.).

So far as my observation goes, very scarce in the southeastern

counties.

Heterodon platyrhinus Latreille.

Very common in sandy regions in Orange nx and southern

Westchester County. I have never encountered the melanistic form

which has been variously described as Æ. niger and H. p. niger, but

Cope notes a specimen from Scarboro, Westchester County, and there `

is now in the New York Zoólogical Park a living specimen taken in

Sullivan County.

Liopeltis vernalis (De Kay).

More common, | believe, in Orange County than east of the Hudson.

Zamenis constrictor (Linn.).

A very common species, both relatively and absolutely, in south-

eastern New Yor

. Coluber vulpinus (B. & G.).

Jordan (Manual Vertebrates Northern United States, eighth edition,

P. 195) gives the range of this species as “ Massachusetts to Kansas,

and north." Cope, however (Crocodilians, Lizards, and Snakes of

North America, p. 831), states that it is not known from east of

Illinois, and I have been unable to find any record of its occurrence

in or near New York.

bas

Coluber obsoletus obsoletus Say.

Occurs in Orange County (and probably east of the Hudson), though

much scarcer than Zamenis constrictor.

Pityophis melanoleucus (Daudin)

Included as a possible straggler over our southern border.

Osceola doliata triangula (Boie).

A rather common snake in Orange County ; less abundant, I believe,

east of the Hudson.

Osceola doliata clerica (B. & G.).

Cope quotes a specimen (U. S. Nat. Mus., No. 1407) whose locality

is given as “(?) New York.” As this state is far beyond the most

No. 410.] THE SNAKES OF NEW YORK STATE. 153

northern range of the subspecies, so far as any other records go, the

label may be considered as in error.

lan]

. Ophibolus getulus getulus (Linn.).

Baird (* Serpents of New York," 1854) states that “ the chain snake

is quite maritime in its northern distribution, being rarely found in the

Northern States, except near the coast.” De Kay had mentioned its

occurrence on Long Island. In a * Catalogue of Reptiles and Fishes,

from St. Lawrence County, procured for the State Cabinet of Natural

History by Franklin B. Hough” (Fifth Annual Report on Condition

of State Cabinet, Albany, 1852), I find that Dr. Hough collected

specimens of this species from Rossie, St. Lawrence County, and that

he quotes it as “of common occurrence in this portion of the state.”

These specimens cannot now be found, but I am inclined to accept

the record, as John Gebhard, then Curator of the State Cabinet, had

too thorough a knowledge of the ophidian fauna of the state to have

allowed an obvious error to stand in the catalogue.

. Natrix fasciata sipedon (Linn.).

Abundant in all the counties of southeastern New York. The normal

ground color here is dark grayish to black. I have never seen indi-

viduals of the brownish or yellow-brown color described by some

authors, and shown by casts at several prominent museums.

WwW

. Natrix fasciata erythrogaster (Shaw).

Cope quotes a specimen (U. S. Nat. Mus., No. 9984) as from West-

field Falls, Connecticut. This is far to the north and east of the

usual range of the subspecies, and may be an error in the label.

. Natrix leberis.

I have never seen individuals of this species in southeastern New

York, though much of my field work has been in localities which would

seem to furnish a highly favorable environment.

I5. Storeria dekayi (Holbrook).

This and the next species are abundant in southeastern New York,

though both are so inconspicuous and retiring as readily to escape

observation. As late as 1898 I have found a specimen of S. occi-

pitomaculata on New York Island.

- Storeria occipitomaculata (Storer). i ;

Described by De Kay as extra-limital, this species was included in

the fauna of the state by John Gebhard, Jr. (Fourth Annual Report

on Condition of State Cabinet, Albany, p. 23, 1851), as specimens e

been obtained by Dr. Fitch in Washington County. Tue jm td

than this publication B aird, in his “ Serpents of New York’ (Seventh

Annual Report on Condition of State Cabinet, Albany, p. 124, 1554),

“ formally introduced” the species as “an inhabitant of New York.

I54 THE AMERICAN NATURALIST. [Vor. XXXV.

17;

[^]

-—

2I.

Eutenia saurita (Linn.).

In Westchester and Putnam counties this species appears to be even

more abundant than Zw£ezia sirtalis. It is rarely found out of the

immediate vicinity of water, though it is not so truly aquatic in its

habits as is /Vazrzx fasciata sipedon. On being alarmed this latter

serpent will usually go below the surface, for a time at least. Eutenia

saurita, on the contrary, when flushed from its place of concealment

in the reeds or grass along the shore, will remain on the surface, not

going far from the bank of the stream or pond, and watching for a

favorable opportunity to regain the shore.

. Eutenia brachystoma (Cope).

The single specimen on which this species is founded was taken

at Franklin, Venango County, Pennsylvania. The species may, there-

fore, be found within our limits.

. Eutenia sirtalis graminea (Cope).

No specimens are recorded from this state, and I have never seen

an individual of E. sirtalis even approaching the coloration of this

subspecies as described by Cope, but as specimens are described from

both Ohio and Massachusetts, the subspecies is here included.

. Eutenia sirtalis ordinata (Linn.).

Eutenia sirtalis sirtalis (Linn.).

Both this subspecies, and that immediately preceding, are common

in southeastern New York. My own observations upon the various

forms of Ewtenia sirtalis occurring in New York have led me to

feel that the subspecific grouping proposed by Cope is not entirely

satisfactory.

In addition to those here listed we have in this state, I believe, at

least two forms of equal rank with Æ. s. ordinata and E. s. sirtalis.

Whether any of these forms of E. sirfa/is occurring in New York are

sufficiently distinct to be given subspecific rank is still, I think, an

open question.

had hoped to be in a position, before the publication of this

paper, to discuss this subject in a more definite manner, but I have

not been able to accumulate sufficient material to do so. This being

the case, I have used Cope's grouping and terminology in this list.

Eutenia sirtalis obscura.

The type specimen of this subspecies was obtained at Westport,

New York

Ancistrodon contortrix (Linn.).

Occurs in swamps and low grounds in Orange and Dutchess Counties,

but scarcer in the oo Most of the many snakes annually killed

as “ Copperheads " a: Is of the species Heterodon platyrhinus,

Ophibolus da ianen, or even Natrix fasciata sipedon.

No.410] THE SNAKES OF NEW YORK STATE. 155

. Sistrurus catenatus catenatus (Raf.).

Described by De Kay as extra-limital, but added by Gebhard (Sixth

Annual Report on Condition of State Cabinet, Albany, p. 22, 1853),

as specimens had been sent in from Genesee County. There appears-

to be no recent definite record of their occurrence in that region.

. Crotalus horridus (Linn.).

Still occurs in Orange and Rockland Counties, but very rare and

possibly extinct east of the Hudson in this state. Cope notes a speci-

men collected in 1878 at Katonah, Westchester County ; and I have

been informed that one was killed in 1887 near White Plains.

NEW York STATE MUSEUM,

December 3, 1900.

(No. 409 was mailed January 31, 1901.)

TO COLLECTORS

I have a few fine, perfect specimens of Argonauta

Argo (paper Nautilus), about 2% inches, at $1.00 each.

Also a large number of rare, scarce shells.

List submitted on application.

J. F. POWELL, Waukegan, Ill.

MARINE BIOLOGICAL SUPPLY DEPARTMENT

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VOL. XXXV, No. 411 ^ ds MARCH, Igor

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

I. The Males of Some Texan Ecitons ue

Professor W. M. WHEELER and W, H. LONG 1 ——

II. Ses Morphology of the Hinge Teeth of Bivalves . . Dr. W. H. DALL

The Pharyngo-Œs eal Lung of Desmognathus

2 ees z Professor H. H. WILDER 183

HARRIS

IV. Notes on the Habits of Cambarus immunis sacs J. ARTHUR

V. Artificial Incubation of Alligator Eggs . Dr. A. M. REESE

VI. The Colors of Northern Apetalous Flowers .

VII. Prehistoric Vorkshops at Mt. Kineo, Maine 13

VIII, m = Recent Literature: Zod/ogy, Two I Papers on North- 221

n Mammals, Eighteenth Antal Report of P the Fishing Board for

hic Representation of Rock Com

and Parsons's Ele-

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Rod =i Ae x * 1 aphs of Sedimentary Rocks

Ix. Pablicasions NR

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AMERICAN NATURALISI

Vor. XXXV. March, rgor. No, 411.

THE MALES OF SOME TEXAN ECITONS."

WILLIAM MORTON WHEELER AND WILLIAM HENRY LONG.

SiNcE the observations on Eciton published in the June

number of the American Naturalist for 1900 were undertaken,

the recognition of: the sexual forms of the Doryline ants has

made some progress. European myrmecologists, notably Pro-

fessor Forel and Professor Emery, have thrown considerable

light on several of the species belonging both to the Old

World (Dorylii) and American branches (Ecitonii) of the

subfamily. i

The doubts entertained in the senior author's former paper

concerning the identity of André's Pseudodichthadia incerta

have been dissipated. It now appears that this insect is really

the female of Eciton cecum Latr. Ina note at the very end

of his splendid work on the ants of Central America and

Mexico? Forel quotes the following admission from a letter

from M. André: “ Je tiens à vous dire que je suis depuis long-

temps convaincu que ma Pseudodichthadia incerta, est. bien

1 Contributions from the Zological Laboratory of the University of Texas,

o. II

2 Biol. Cent. Am. Hymenoptera Formicida, vol. iii (1 899-1900), P- 160.

158 THE AMERICAN NATURALIST. | [Vor. XXXV.

la femelle de l’Eczton cecum Latr. J'ai été trop prudent en

n'affirmant pas tout de suite cette assimilation, mais la présence

d'une ğ dans le tube contenant la 9 et la connaissance plus

étendue qu'on a aujourd'hui des Dorylides ne laissent plus

aucun doute à cet égard.”

Professor Emery has just published a revision of the genus

Eciton. For some time past he has recognized in the insect

known as Labidus latreillei Jurine, and a number of other

forms which seem to have merely varietal or synonymic value

(Labidus sayi Hald.; L. atriceps F. Smith; L. jurined Shuck;

L. Servillei Westwood; L. smita D. T.; L. pilosus F. M.;

and L. fulvescens Blanch), the male of Eciton cecum. We are,

therefore, acquainted with the three phases of the commonest

and most widely distributed of the ecitons, a species which

ranges from Utah and Texas to southern Brazil and has been

very generally known in the worker phase to entomologists for

more than a century.?

Eciron (EciroN) cccuM LATREILLE.

[sj ? Formica omnivora Oliv. Encycl. Method. Ins., vol. v (1791), No. 6, p. 496.

9 Formica ceca Latr. Hist. Nat. des Fourmis, tome ix (1802), p. 270, Fig. 56.

9 Eciton vastator Smith. Journ. Ent., vol. i (1860), p. 71.

Q Eciton erratica Smith. Loc.cit,p.71. Bates. Natur. Amazons, vol. ii (1863),

gj Wycteresia ceca Roger. Berl. Ent. Zeitschr. (1861), p. 22.

[s] miii rubra Buckley. Proc. Ent. Soc. Phil. (1866), p. 335.

$E ton omnivorum Emery. Bull. Soc. Ital, vol xxiii (1891), p. 163; ibid.,

E xxvi i (1894), p. 179, T. II, Figs. 9 a-d; Zool. Jahrb., Abth. f. Syst.

Bd. viii (1894), p. 258.

Q Eciton cecum Mayr. Wien. Ent. Zeitschr. (1886), p. 119; Verh. zool. bot. Ges.

Wien., Bd. xxxvi (1886), p

9 Peudodihthad incerta kee grues aux Fourmis (1885), p. 8, Fig. 1-5;

en. Europ. II (1886), p. 840, fig.

$ m ales Jurine (1807); Emery, Nuovi Studi sul Genere Eciton, Mem. -

Accad. Sci. st. Bologna (1900), p. 9.

1 Nuovi Studi sul Genere Eciton, Mem. letta alla R. Accad. delle Sci. dell’

Jstituto di Bologna, 25 marzo, 1900, 18 pp., 1 tav.

? The remarkable sexual trimorphism of this i insect, together with its variability,

atleastin the worker and male phases, is largely responsible for the following

interesting synonymy compiled from the contributions of Forel and Emery. For

her mention of the literature the reader is referred to Dalla Torre's Catalogus

| Hymenopterorum, vol. vii, Formicide, 1893, pp. 1-7.

No. 411.] MALES OF SOME TEXAN ECITONS. 159

$ Labidus. sayi Hald. Stansbury’s EZxplor. of Utah (1852), p. 367, Pl. IX,

‘igs. 1-3. Emery, Zec. cif. (1900), p. 9.

P. Labidus servillei Nestw. Arcana. Ent., vol. i (1842), p. 75 tome xx, Figs. 2,

2a; Emery, Bull. Soc. Ent. Ital., vol. xxii (1890), p. 39; Emery, Nuovi

Studi, etc., Joc. cît. (1900), p. 9.

& Labidus jurinei Shuck. Emery, doc. cit. (1900), p. 9.

$ Labidus smithi Dalla Torre. Emery, loc. cit. (1900), p. 10.

4 Labidus pilosus F. Sm. Emery, oc. cit. (1900), p. 10.

$ Labidus fulvescens Blanch. Emery, loc. cit. (1900), p. I0.

4 Labidus atriceps F. Smith. Emery, loc. cit. (1900), p- 16.

Both before and since learning of André’s conclusion we

have sought diligently but in vain for the female of Eciton

cæcum in-the vicinity of Austin, notwithstanding the species

is so common that we rarely spend a few hours collecting

and observing ants without happening on two or three of its

colonies, These are found, as a rule, under clusters of stones

which lie with their edges in mutual contact. But as the

species is entirely subterranean it is not an easy matter to

find the breeding chambers. The narrow burrows run along

under the stones for some distance and then dip down into the

soil. Frequently the galleries are found under large stones

that have fallen from stone walls, and in these cases the bur-

rows almost invariably extend under the wall where they are

inaccessible to the observer. For several days after a rain

E. cecum may often be found under the stones in open fields,

but during dry weather it seems to prefer the more sheltered

and therefore moister localities. The males of E. caecum,

which may be readily identified by the aid of the table in.

Emery's latest paper, are common about the electric lights

at certain times of the year. Messrs. Melander and Brues

took them in considerable numbers on March 27, less fre-

quently on April ro and 20 of the past year. All of these

specimens are referable to Labidus sayt, although somewhat

exceeding the dimensions of the specimens described by

Haldemann. :

Eciton cecum, which may be regarded as the typical species

of the genus sensu stricto (workers with denticulate claws), 1s,

as we have said, the only Eciton of which the male, female,

and worker phases are all described. In the present paper

160 THE AMERICAN NATURALIST. [Vor. XXXV.

we wish to call attention to the discovery of the males of

E. opacithorax* Emery and of E. schmitti Emery? The male

of the latter species, together with the female described in the

senior author's former paper, completes the series of sexual

phases of a species belonging to another natural group of

ecitons (subgenus Acamatus Emery), the workers of which

have smooth claws.

A male Eciton found dead in a spider's web, Oct. 3, 1899,

by Mr. C. T. Brues was referred by the senior author to

E. schmitti, mainly on account of its coloration, but it was not

till October 13 of this year that it was possible to obtain posi-

tive proof of the truth of this conjecture. Late in the after-

noon of the latter date Mr. A. L. Melander, Mr. C. T. Brues,

and the senior author were collecting insects near Mt. Bonnel,

a few miles from Austin, when Mr. Brues saw a winged ant

perched on a large flat stone beneath a shrub (Zzsem/Zardtia

amorphoides). lt was at once recognized as the Eciton of the

spider's web. On drawing near, two or three other specimens

were seen moving about among a lot of workers of E. schmitti

which were issuing from a hole in the soil about the roots of

the Eisenhardtia and appeared to be on the point of starting

on one of the sorties so characteristic of these ants and their

congeners. We raised the stone and found beneath it a flour-

ishing colony of Æ. schmitti, comprising no less than a hundred

of the huge (Ze, when compared with the workers) winged

males. Many of these were literally covered with workers and

1 Emery (Nuovi Studi, etc., /oc. cit., p. 15) is now inclined to regard the former

species, menm described by him as a subspecies of Æ. californicum Mayr, as a

distinct species.

- acing of the species designated as E. sumichrasti in the senior author's

former paper have been examined by Professor Emery, who pronounces the spe-

cles to be the rien allied Æ. schmitti Emery. The senior author has since

taken the true Æ. sumichrasti in Mexico Dirt Morelos) and is satisfied

that Fuse Emery's identification is corre On seeing the males of Æ

schmitti, Professor Emery was inclined to d them as belonging to Labida:

nigrescens Cresson. If this is true Emery’s name of the species must be relegated

e the synonomy. But some of these males were sent to Mr. Fox for comparison

with Cresson's types and were pronounced to be different, being *too hairy for

seins: which is entirely brownish. The first segment of wigrescens is shorter

the wings light fuscous.” This has induced us to retain Emery's name of

e species for the present.

No.411.] MALES OF SOME TEXAN ECITONS. 161

were being hurried by them into some deep water-worn cavi-

ties on the under surface of the stone and into the galleries

which the ants had excavated in the hard soil. We succeeded,

nevertheless, in capturing a large portion of the colony and in

transferring it, together with some of the soil of the nest, to

a strong canvas bag. The colony, first established in a large

glass jar, was later placed in a Lubbock nest. The material

of the original nest was searched on the spot for the pupz of

the males and for the female, but without success. The fol-

lowing is a description of the male of E. schmitti drawn from

fresh specimens.

Eciton schmitti Emery (Figs. 1 and 2, b), male. Length of

body, 11-13 mm.; length of fore wing, 10-11 mm. Head,

thorax, petiole, extreme base of first abdominal segment, ven-

ter, antennz and legs, except the tarsi, black ; abdomen, tarsi,

hypopygium, tips of mandibles, and, in some specimens, the

flagellum of the antennz, the knees and the tips of the tibiae

fulvous red. Wings blackened, with black veins and stigma, the

costa and some of the veins yellowish red at the extreme base;

maxillae and labium yellow.

Head shining, clothed with long fulvous bairs arising from

coarse punctures. These hairs are longest on the vertex,

posterior orbit, mandibles, first antennal joint and clypeus.

Mandibles rather long, curved at the base only, slightly broad-

ened in the middle, with convex inner edge ending in short,

rather blunt points. Antenne longer than the head and

thorax; tip of the somewhat incrassated scape scarcely reach-

ing the lateral ocellus; second joint small, remainder of the

cylindrical flagellum opaque, its basal somewhat thicker than

1 The nest contained two species of ecitophiles, viz., some small trichopterygid

beetles and more than a dozen specimens of a small active staphylinid with pale pro-

notum and elytra (Zcitopora tenella Wasm. n. sp. in litt.). Some of the latter

lived for nearly a month in the artificial nest. They sought the dry portions of

the nest and seemed to elude the ants by their rapid mon kie verd

lurked near the entrance in cracks in the soil. When, during the morning hours,

the ants were dormant in a compact cluster in the center of the nest, uu pt

crept from their hiding places and moved about the galleries with less trepi —

When the ants left the nest towards evening to move about in files on ud plat-

form, the staphylinids sometimes accompanied them and then often ran off into

the moat and were drowned.

162 THE AMERICAN NATURALIST. [Vor. XXXV.

its apical half; joints of approximately equal length, except at

the tip, where they are somewhat longer but not compressed.

Clypeus without teeth below, frontal carinze not very promi-

nent, separated by a distinct facial furrow. Eyes and ocelli

small and not protruding. The distance between the two

lateral ocelli is about the same as the distance between the

lateral ocellus and the eye. Thorax gibbous in front, flattened

on the dorsal surface,

which is subopaque;

pleurz and pectus shin-

ing. Dorsum with fine

and rather dense pleure,

with scattered and lar-

ger piligerous punc-

tures. Whole thorax

covered with long ful-

vous pile somewhat

shorter and more ap-

pressed on the mesono-

tum and scutellum ; long

and erect on the pleurz

and pectus. Anterior

half of mesonotum trav-

ersed by a longitudinal

, smooth line which seems

3 to lie in a shallow groove

l at its anterior end. A

somewhat similar line

is found on either side,

extending over the pos-

terior two-thirds of the

mesonotum, as far back as the scutellum, and another indis-

tinct line traverses the posterior third of the scutellum, which

is flat on its dorsal and convex on its posterior surface.

Metanotum declivous and slightly concave on its posterior

surface. Petiole subopaque above, shining below, nearly twice

as broad as long, flattened transversely across the middle of

its dorsal surface, but distinctly concave in front of and behind

Fic. 1. — Eciton schmitti Emery. Male.

No.411.] MALES OF SOME TEXAN ECITONS. 163

this region ; posterior angles only moderately projecting ; ante-

rior portion with sparse pile, posterior edges, and especially the

posterior angles, with rather long, dense, and appressed ful-

vous pile; ventrally the pile is thin and more erect. Abdo-

men cylindrical, subopaque above (except at the subconstricted

posterior edges of the segments which, like the venter, are

glabrous), covered with minute piligerous punctures ; pile on the

dorsal surface of the segments short and appressed, especially

towards the base of the abdomen, posteriorly and on the ven-

ter, especially in the region of the hypopygium, it is longer and

more erect. Hypopygial plate shining with two short, slightly

recurved teeth at its tip. Legs rather small and feeble, pos-

terior pair not reaching to the end of the abdomen. Middle

metatarsus a little more than half as long as the middle tibia.

Coxze, femora, and tibize shining, with rather erect fulvous pile,

pile on the tarsi short and appressed. Tibial spurs red, claws

with extremely rudimental denticles. Wings clothed with

minute black pile, anterior pair reaching to the tip of the

abdomen. For venation see Fig. I.

On October 27 the junior author was so fortunate as to dis-

cover a flourishing colony of E. ofacithorax Emery under a

large stone in a dilapidated wall at the edge of some woods, on

the sloping banks of Shoal Creek at Austin. This colony also

contained a great number of males but was ransacked in vain

for a queen. The workers had stored their nest with a

considerable number of small carabid beetles that had evi-

dently been captured on one of their marauding expeditions.

E. opacithorax is considerably smaller and appears to be some-

what rarer in this locality than E. schmitti. We insert a

description of the male, which, like the male of schmitti,

as a species of Labidus.

appears to have escaped description

i : Length of body,

Eciton opacithorax Emery (Fig. 2, €) male.

10-11 mm. ; length of fore-wing, 8-9 mm. "oh

Body glabrous throughout ; head, thorax, petiole, first abdom-

and tibiae, black; lateral and

inal segment, venter, femora, Sons

dorsal regions of abdomen, including the posterior edge ot the

inner edges of mandi-

first abdominal segment, hypopygium, |

bles, knees, and tarsi, dull red. Antenna usually black, but

164 THE AMERICAN NATURALIST. [Vor. XXXV.

the flagellum in some specimens more or less reddish. Wings

slightly blackish, covered with minute black pile; stigma red-

dish, veins dirty yellow. Pile covering the body grayish ful-

vous, long and erect on the mandibles, antennal scape, head,

thorax; pleurze, pectus, scutellum, base of petiole, and femora,

elsewhere very short and appressed.

Mandibles short, covered with small piligerous punctures,

curved only at the base, thence becoming broad and flat, with

distinctly convex inner margin and terminating in abrupt rather

blunt tips. Head covered with punctures like those on the

mandibles. Eyes and ocelli very small, not projecting ; cheeks

and posterior orbital region broad. Lateral ocelli separated

by a space about equal to that which separates each lateral

ocellus from the eye. Frontal caring prominent, arcuate

below the lateral ocelli, and separated by a rather broad

longitudinal furrow. Anterior half of mesonotum with a dis-

tinct smooth longitudinal line and distinct lateral furrows

extending to the insertion of the scutellum. Scutellum with-

out a furrow on its posterior surface. The piligerous punc-

tures covering the mesonotum and scutellum are small and

evenly distributed, those on the pleurz a little larger and more

scattered. Metanotum declivous, slightly concave, finely punc-

tate, and bearing little pile. Petiole twice as broad as long,

convex above and behind, more concave anteriorly ; posterior

angles not projecting and covered with very short appressed

pile, contrasting with the long and often dense erect pile on the

basal ventral and basal dorsal surfaces of the same segment.

Abdomen short, cylindrical, and laterally compressed, its surface

minutely punctate; pile everywhere, even at the tip, and on the

hypopygium, very short and appressed. Hypopygium armed

with the usual short, slightly recurved teeth. In some speci-

mens the red of the abdomen even on the dorsal surface is more

or less suffused with black, especially near the middle of the

different segments. Fore-wings reaching to the tip of the abdo-

men or even a little beyond. Venation very similar to that of

E. schmitti (vide Fig. 1). Legs short, posterior pair not reaching

the tip of the abdomen. Middle metatarsus a little more than

half the length of the middle tibia. Denticles of claws vestigial.

No.411.] MALES OF SOME TEXAN ECITONS. 165

At first sight the male of Æ. opactthorax is very similar to the

male of E. schmitti, but comparison discloses a number of con-

stant differences. Apart from its distinctly smaller size, the

thorax and scutellum of the former species are more glabrous and

therefore appear to be of a deeper black color. The pile is shorter

on the abdomen and posterior portion of the petiole, the wings

are distinctly smaller, less blackened, and have paler nervures.

The junior author has also taken the males of two other

forms of Eciton at Austin, but hitherto only about the lights in

the evening. One of these, taken October 29, agrees very

closely with the description of Æ. (Labidus) harrisi Hald.

The other, which was flying in considerable numbers August 1,

is smaller and more reddish, with very pale wings. Professor

Forel pronounces it to be Æ. Za7ris?, but as the color is not

that of the type, it must be regarded as a variety. The heads

of both forms, drawn to the same scale, are represented in

Fig.2, 2 and Z. Besides the various species already mentioned

we have recently taken at Austin the workers of E. californi-

cum Mayr. and E. pilosum F. Sm. The latter species has not

before been recorded from the United States. With the single

exception of Æ. carolinense, all the species of Eciton known to

occur north of Mexico have been found in Texas, and it is very

probable that Æ. carolinense will be found in the eastern por-

tion of the state. The species of the genus occurring north

of Mexico may be tabulated as follows, according as they are

known from the male or worker phase alone or from both :

WORKERS. MALES.

1. Eciton cecum Latr. Labidus sayi Hald.

2. E. sumichrasti Norton. Unknown. :

3. E.schmitti Emery. E. schmitti Em.

4. E. californicum Mayr. Unknown.

5. E.opacithorax Emery E. opacithorax Em.

6. E.carolinense Emery. Unknown

7. Unknown L. harrisi Hald

8. Unknown L. mexicanum F. Sm

9. Unknown L. nigrescens Cress.

to. Unknown. L. minus Cress.

E. spoliator Forel.

L. melshemeri Hald.

Unknown.

. Unknown.

. E. pilosum F. Sm.

—

i-r

—

Qə

166 THE AMERICAN NATURALIST. [Vor. XXXV.

The males of Æ. schmitti, after living in good condition in the

artificial nest for a few days, began to die off, at first a few each

day, then more, till by the evening of October 22, nine days after

their capture, they were all dead. During this period, how-

ever, it was possible to make a few observations on their habits.

ns. &, E. harrisi; b, E. schmitti; c, E. cocum ;

Fic. 2. — Heads of male ecito

d, E. harrisi, var.; e, E. opacithorax.

In his former paper the senior author recorded the fact that

the odor of the workers of E. schmitti, like that of other

Species of the genus, is rank and disagreeable, whereas the

odor of the queens is mild and pleasant.

1 " ^

" The peculiar odor of Eciton workers was noticed by Sumichrast (Notes on

e Habits of Certain Species of Mexican Hymenoptera, etc. Note 1, On the

No. 411.] MALES OF SOME TEXAN ECITONS. 167

It was interesting, therefore, to find that the males have the

same pleasant odor as the females. We believe that this

peculiar property acts as a powerful attraction to the workers,

causing them to cling in great numbers to the bodies of the

fertile sexes. Certainly no other ants with which we are

familiar appear to be so fond of their queens and males. The

workers seem never to tire of fondling and licking the latter.

They lick even their large mandibles and the broad membranes

of their wings, and when the males move about in the nest

the workers ride on their backs and sides. Sometimes the

males are loaded down so heavily with workers that they can

neither walk nor fly. On such occasions they are often seen

to relieve themselves of their too affectionate attendants by

suddenly flirting their wings and tossing the workers to a

distance. Even dead males are often fondled for hours.

Although the males have beautifully developed strigils on their

fore-tibiae, they do not appear to use them while in the nest,

probably because the continual grooming which they receive

from the workers makes attention to their own persons unneces-

sary. In this respect the behavior of these male ants is in

marked contrast with the behavior of the male Ponerinz

(Pachycondyla harpax, for example).

So far as could be observed the males made no use of their

huge mandibles, structures which cannot fail to excite interest,

because the mandibles of male ants are usually so small or

even rudimental as compared with the mandibles of the workers

and queens. The male ecitons were never seen to fight

with one another or with the workers, nor did they take food

of any form during their captivity. Since the female Eciton

is wingless and sluggish, the mandibles of the males can hardly

function as clasping organs. We are inclined, therefore, to

regard them as secondary sexual characters belonging to the

same category as the much larger mandibles of the male stag-

beetles (Lucanidz) or the cephalic and pronotal horns of many

male chafers (Scarabzide). It is not improbable, however,

i . Ent. Soc.

Habits of the Mexican Species of the Genus Eciton Latr., Trans. Am En

i buted to the ants’ living in rotten

(June, 1868), pp. 39-44). but erroneously attri

wood.

168 THE AMERICAN NATURALIST. (VoL. XXXV.

that when the mystery which still envelops the origin of the

female Eciton and the time and place of her impregnation has

been dispelled, the mandibles of the male may be found to

have some as yet unsuspected function.

During the morning hours the ecitons remained very quiet,

hanging in clusters in an angle formed by the soil and the

glass of the jar, or between the glass and the earth of the

Lubbock nest after being removed to the latter. The males,

concealed beneath the mass of workers, were usually huddled

together, with their heads hidden in holes in the earth and

their abdomens and the tips of their wings projecting. They

appeared to be sleeping. Between one and two o'clock in

the afternoon the whole colony awoke. The males would

begin to climb the glass sides of the jar in perfectly perpen-

dicular paths, falling to the bottom from time to time but

again persistently ascending. The workers would often file

up the: sides of the jar with the males walking at the same

rate of speed in their ranks. The activity of the colony

appeared to reach its height between five and seven o'clock,

and then to subside as the night came on, when the colony

again clustered with the males in its midst. Even a lamp

kept in the room with the nest, if not too near it, failed to

interrupt their slumbers. This striking rhythm of alternating

activity and rest was observed on each of the nine days during

which the males lived, and was continued by the workers

alone for some weeks afterwards.

The numerous males of FE. schmitti and E. opactthorax taken

in the two nests above described showed relatively little varia-

tion in size and coloration. They were all equally fresh and

active and had evidently all hatched but a short time before the

nests were discovered. This induces us to add some remarks

concerning the supposed dimorphism of the males of Eciton, if

only for the purpose of corroborating Emery's statements on

this subject. For the sake of bringing the matter clearly

before the reader, we translate most of Emery's remarks.

“In my memoir on the larvz of ants?] omitted a discussion

! Nuovi Studi sul Genere Eciton, Zoc. cit., pp. 4, 5.

? Intorno alle larve di alcune F. ormiche.

No. 411.] MALES OF SOME TEXAN ECITONS. 169

of the singular observations published by W. Miiller,* which,

as recorded, tend to show that the larvae of Eciton are of

two forms, one like those subsequently described by myself,

the other furnished with setigerous tubercles, like the larvae of

the Ponerinæ. When, during the past autumn at the Munich

meeting of German naturalists, I presented the results of my

observations, Forel took exception to them, on the ground that

the dimorphism observed by W. Müller in Eciton went to

prove that the cutaneous tubercles could not be peculiar to the

larvae of the Ponerinz. I was thus compelled to examine

Miiller’s observations, for the purpose of ascertaining whether

or not Forel’s conclusion and those of the author himself were

legitimate.

« Among many thousand typical larvee of Eciton, Müller

observed a very few much smaller larvæ, covered with tuber-

cles and some dark-colored cocoons containing similar larvae

and pupz and differing from those of the ecitons. The tuber-

culate larvae were of different sizes and forms, and most of the

dark-colored cocoons woven by these larvee measured 8 mm.,

but some of them only 6 mm., in length. As it was not known

at that time that the larvae of the Ponerine are of this struc-

ture, he could not suppose that he was dealing with larve

stolen from the nest of other ants. Müller and Forel, who

examined the material collected, thought there was an evident

connection between the two observations to be discussed, a

fallacious connection which, nevertheless, as an element of

suggestion, has dominated the whole thought of these authors.

«In the midst of the mass of worker ecitons Müller found

a deálated and damaged specimen of the insect then known as

Labidus burchelli, the male of the Eciton which was the sub-

ject of his observations. On examining a pupa taken from one

of the dark-colored cocoons and hence derived from a tuber-

culate larva, Forel observed that, whereas the form of the head

and thorax was that of a worker ant, the tip of the abdomen

presented appendages, which, though poorly preserved in the

specimen studied, recalled by their position and configuration

1 Beobachtungen an Wanderameisen (Zciton hamatum) in Kosmos, 10. Jahrg.,

Bd. xviii (1886), pp. 81—93-

170 THE AMERICAN NATURALIST. [VorL. XXXV.

the copulatory appendages of the male sex. This pupa must,

therefore, be a male, though very different from the Labidus,

as shown in the figures which I repróduce in the plate

accompanying this memoir (Fig. 30, a, 2, c). Believing the

determination of the sex to be correct and desiring an explana-

tion of the facts, the specimen was supposed to be a hetero-

morphic male of Eciton, and the tuberculate larva were

therefore regarded as male larve, the smooth ones as belong-

ing to ants of the femalesex. Thus arose the strange doctrine

of the dimorphism of the males in the genus Eciton. The

new heteromorphic male was, from analogy with the termites,

designated as a supplementary male.” :

From an examination of W. Miiller’s figures, Emery con-

cludes that the pupa which gave rise to the above view was

a normal worker pupa of some ponerine ant, presumably a

species of Pachycondyla which had been appropriated by the

ecitons. That Emery has drawn the correct inference from

Miiller’s data is very clear from the following observations

made on Æ. schmitti during the past spring. These observa-

tions are transcribed from the notebook of the senior author.

May 25 we came upon a colony of Æ. schmitti under a large

flat stone. The ants had dug their galleries to a depth of

several inches in the moist black soil and had literally packed

them with larvae and pupe. The latter were more abundant -

and were at once seen to be the slender, naked worker pupa

of the Eciton. The relatively small number of larva were

nearly mature and closely resembled the figures of Eciton

larvee published by Emery. There were, however, many

naked pupa and larve of very different shapes and sizes.

Some of the former had wing-cases, and for a few days the

senior author lived in the pleasant anticipation of being able

to hatch the male ecitons, since much of the nest had been

captured and placed in a large glass jar. The ecitons at once

set to work and collected the slender larvae and pupz and then

turned their attention to the others, which were finally brought

together in the same place. Some of the winged pupa and

i Intorno alle Larve di Alcune Formiche, Mem. della Accad. delle Scienze dell’

Istituto di Bologna, 7 maggio, 1899, 2 tav.

No.411.] MALES OF SOME TEXAN ECITONS. I7E

the obese larvae from which they were evidently derived were

examined more carefully, and it was found that many of them

were queen larvae and pupz of some one of three species of

Pheidole whose nests had been noticed a short distance from

the Eciton nest. Other larva and pupz appeared to belong

to another common ant, Solenopsis geminata. There could be

no doubt that the ecitons had stolen all these myrmicine

progeny, which they proceeded to devour a few days later

till all had disappeared.

That this habit of pillaging the nests of other ants is shared

also by the tropical ecitons is shown by the following obser-

vation recorded by Sumichrast!: “It is probable that the

Eciton attacks the larva and pupe of other ants to make

them serve as food for the nourishment of their own larvae or

for sustaining themselves. I surprised one day, in the first

hours of a somber and rainy morning, a considerable assem-

blage of zepeguas (No. 36), fastened one upon another like a

swarm of bees, and entirely still. Having dispersed them, I

perceived in the place which they covered with their bodies a

quantity of little white larvae, brought away, doubtless, from

‘the nests of some Myrmicide. At another time I witnessed

the pillage of a nursery of other ants by a quite numerous band

of workers minores of No. 68; alarmed by the reprisals which

I made on their account, they took to flight, some of them

carrying between their mandibles as many as three larvae at

once." ? ;

The habit of seizing the larvae and pupæ of other ants and

of storing them in their nests for several days till required for

food is only a special phase of a more general habit of the

ecitons and probably also of some other ants, for the ecitons

do the same with the other portable insect booty which they

1 Loe. cits p.42. Au f

2 Recently at Querétaro in Mexico the senior author surprised a troop o

Eciton crassicorne Sm. pillaging a large nest of the agricultural ant (Pogono-

this powerful Eciton was found

myrmex barbatus). The temporary nest of

under a large stone only a few yards away. PYR SN

agriculturals. At Cuernavaca the same species of Eciton was seen pillaging

some small Pheidole nests and dragging away carabid beetles from under the

stones. i

172 THE AMERICAN NATURALLIST. [VoL. XXXV.

capture on their marauding expeditions. When they migrate

toa new nest they carry this booty with them, as shown by

the following observation made in the city of Galveston during

the past June. A large colony of Æ. schmitti, which had

been living under the front doorstep of the house in which the

senior author was stopping, decided to change their quarters.

In the early twilight the entire colony moved out in regular

file through the garden and entered a new nest which was

being dug only a few yards from the old one about the roots of

some violets. While carefully scrutinizing the file of ants for

the purpose of detecting any ecitophiles which might be march-

ing with them, the ecitons were seen to carry considerable

numbers of dead carabid beetles (small species of Harpalus and

Pterostichus). These evidently represented their store of food

for the time being. Since this observation was made, the

senior author has had frequent opportunity to feed the ecitons

in artificial nests with termites and the larve of ants (Campo-

notus and Pachycondyla!). In all cases many of these larva

and termites were carried about or stored in one corner of the

nest for several days before they were eaten.

Postscript.

The first female Eciton known to have been taken in the

United States was discovered by Rev. P. Jerome Schmitt in

April, 1894, on the grounds of St. Mary’s College, Belmont,

Gaston County, N.C. The insect was found under the bark of

a tree and concealed under a large mass of workers half rigid

with the cold. Eighty to one hundred specimens of an ecito-

philous beetle (Ecitonusa schmitti Wasm.) were found scattered

through the cluster of workers. The latter were identified by

Wasmann and Emery as belonging to Æ. opacithorax Emery.

The description of the female has been delayed till the present

time, as Rev. Mr. Schmitt happened to be more interested in

other fields of entomology. At our request he has most gener-

ously sent us the insect for examination and description, and has

also permitted us to use the accompanying cut, prepared for him

some years ago under the direction of Messrs. Schwarz and

No. 411.] MALES OF SOME TEXAN ECITONS. 173

Pergande. Through this kindness Rev. Mr. Schmitt has enabled

us to complete our description of the sexual forms of Æ. opaci-

thorax, the male of which is described above. Comparison of

the female of this species with the female of Æ. schmitti

shows such striking similarity that unless the specimens had

been taken with workers of two very different species, one

would scarcely regard them as specifically distinct. The fol-

lowing slight differences can be noticed: The female Æ. ofaci-

thorax has the head somewhat more shining, with smaller

punctures, which are not confluent as in Æ. schmitti; the

eyes are smaller, the hairs on the dorso-lateral portions of the

head are shorter, the occiput is distinctly less depressed in

the middle line, and

the pronotum is some- : fs:

what rounder and more gs >W

N » »

«33 s>

convex above. The

thorax is less conflu-

ently punctate, more

evenly hairy, and the

petiole is less concave

in the middle. The

size and color of the

two species are very

nearly identical; even

the large dark blotches

on either side of the

metanotum are present

in Æ. opactthorax.

If we may be per-

mitted to generalize from the study of the two species of

Eciton, of which we have seen all three phases, it may be

said that the fertile forms of the different species of this genus

are even more difficult to distinguish from one another than

the workers. For this reason, and also on account of the much

greater rarity of the males and females, myrmecologists will

probably continue to regard the worker ecitons as of greater

taxonomic importance.

UNIVERSITY OF TEXAS, AUSTIN, TEXAS, Nov. 15, 1900.

Fic. 3:— Eciton opacithorax Emery.

THE MORPHOLOGY OF THE HINGE TEETH OF

BIVALVES.

WILLIAM H. DALL.

Tug most notable step in advance for the study of bivalve

mollusks which has been made for many years is due to the

researches of the late F. Bernard, which are included in half a.

dozen papers, of which the earliest appeared in 1895.

Ever since the time of Schumacher and Lamarck, in the

early years of the century, the interlocking projections of the

dorsal margins of the valves of pelecypods have been recog-

nized, under the name of hinge teeth, as affording characters

of the highest value in systematic classification of the subdi-

visions of this group. All works treating of these animals

have utilized the more or less obvious features of the adult

hinge teeth in framing diagnoses of families and genera.

Systematists contented themselves, however, with noting the

number, position, and division of the teeth into so-called “ car-

dinals” and “laterals ” and ignored the fact that large groups

have teeth of special types, which can hardly be closely homol-

ogized with the teeth characteristic of several other groups.

This method of treating the teeth was unquestioned for more

than, half a century and became so fixed in the minds of sys-

tematists that modern efforts to suggest a new point of view

have up to the present time been almost futile. Steinmann, in

his Lehrbuch der Paleontologie, Fischer, in his Manual (1887),

and especially Neumayr, in his Morphologie Bivalven Schlosses

(1883), emphasized the necessity of distinguishing between

certain fundamental classes or types of hinge teeth, and to

Steinmann we owe the suggestion of a formula for indicating

in brief form and almost graphic manner the number and rela-

tions of the teeth on any given hinge. The writer, in 1889, and

Neumayr, in a posthumous paper issued in 1891, applied these

principles to the general classification of bivalves, necessarily

175

176 THE AMERICAN NATURALIST. [VOL XXXV.

in a tentative manner, but further advance, or even the

confirmation of steps already taken, could be made only on

the basis of observed facts of development in concrete cases.

Munier-Chalmas (1895) propounded certain theories of proto-

typic hinges, but in the absence of published observations

these attracted slight attention. In the same year the writer

elaborated his previous thesis, insisting on the modifications

due to dynamic causes acting directly on the hinge, on the

origin of hinge teeth from initiative due to external sculpture

in the dorsal region, and on the duplex nature of the so-called

ligament, which many years before had been recognized by

Récluz, whose observations fell on sterile soil. The existence

in many nepionic bivalves of a primitive hinge, since named

the provinculum, was also pointed out. Almost simultane-

ously the first of Bernard's papers appeared, for the first time

furnishing facts in relation to the development of the hinge

teeth in a comparative series of different genera ; and thence-

forward, until the time of his premature decease, he continued

to bring together invaluable data in the same line, covering a

large number of genera and opening the way to a field of

research of extraordinary promise, throwing an entirely new

light on the development of hinge structure and making a

permanent breach in the barriers of a conservatism which had

so long retarded the appreciation of efforts for a more scientific

consideration of the facts of hinge structure. As in all first

attempts, the shackles of outgrown methods were not wholly

discarded, and much more study is required before a final

system can be hoped for, but so much has been done that it is

no longer possible for the former inertia to prevail. Several

papers bearing on this subject have recently appeared. One

of them, by W. von Vest,! can be briefly dismissed, as it con-

tributes nothing new in the way of facts and is chiefly devoted

to the introduction of a host of new technical German terms,

which in turn are reduced to symbols, which when formulated

; Ueber die Bildung und Entwicklung des Bivalven-Schlosses, followed by

Entwurf einer Einteilung der lebender Bivalven nach dem Schlossbau, Verh. des

Siebenburgischen Vereins für Naturw. zu Hermannstadt, Bd. xlviii (1899), pp-

25-150, Pls. I-II.

No. 411.] HINGE TEETH OF BIVALVES. 177

comprise five or six lines of difficultly intelligible letters and

figures, requiring three or four times as much more space

devoted to * Erklärung der Formel," and, when all is said and

done, hinder rather than help. How long will it be before it

is universally understood that symbolization which has not the

merit of brevity is not only a foolish waste of ingenuity and

time but is absolutely without any chance of acceptation ?

In this connection it may be noted that Munier-Chalmas

proposed a system of notation for the hinge of bivalves which

erred not only by involving much purely hypothetical assump-

tion but also by inordinate length. A better was proposed by

Bernard, of which some modification is suggested by Dr. Fritz

Noetling! in an interesting and suggestive paper which has

lately reached us. Bernard began work on the teleodont

forms (Heterodonta of Neumayr) and under the influence of

Munier-Chalmas. As many of the Teleodesmacea have com-

parable hinges, it was natural that Bernard should suppose that

all hinges were comparable and that the formula he proposed

should be based on the assumption that from the mere position

on the hinge plate the homologies of the individual teeth could

be taken for granted, which we believe to be a very mistaken

idea. Bernard in his later work realized that error, but unfortu-

nately was not spared to consider the revision of his symbols.

As a matter of fact even in the Teleodesmacea Bernard showed

before his death that several distinct systems of tooth develop-

ment can be recognized which cannot be interchangeably homol-

ogized. Much more is it impracticable to homologize teeth of

such groups as the Prionodesmacea and Teleodesmacea. In

groups where it is practicable, — to be determined by observa-

tion and not by theory, — Bernard's system will doubtless, with

some modification, continue to be used. It is, however, very

liable to lead the student unacquainted with its pitfalls into an

unprofitable wilderness of theory, when, for the present, facts,

and again facts, are imperatively demanded. If we eliminate

theory, it still remains important that there should be some

symbolical way of presenting the facts to the eye without

1Notes on the Morphology of the Pelecypoda, Paleontologia Indica, N-S.

vol. i (1899), pp- 1-43, No. 2. Calcutta.

I 78 THE AMERICAN NATURALIST. [VorL. XXXV.

extended verbal description, and for this we must fall back on

some modification of Steinmann's system, which for clearness

and brevity can hardly be excelled. It is essential also that

the student should not forget that the teeth are not, so to

speak, individualities or specific organs, no matter how true

they, in the main, are bred from generation to generation.

Apart from their genetic origin as the selected results of

the dynamic influence of external sculpture on limited areas

of the shell (this sculpture itself being a selected conse-

quence of the dynamic influence of the irregularities of the

edge of the mantle upon the shell it secretes), the teeth are

essentially prominences of a structure of homogeneous material

modified by strains and impacts, and an excess or deficiency in

the dynamics of the hinge margin in any individual will directly

affect the hinge. The proof of this is most obvious in the

differences of the lateral lamella which will be found in exam-

ining a large series from different localities of such a type as

Tellina, for instance.

One important result of Bernard's researches was to show

that cardinal and lateral are the disunited segments of an

originally continuous lamina, and hence of genetically identical

origin in certain teleodonts. In others the cardinals and

laterals probably are distinct in origin (Tellina), though these

developments so converge that in the adults they would be

homologized by any student. The dynamic influence of the

ligament when internal is very great and differs at different

stages of growth in the same individual. In general it pre-

vents the formation of the ** hook," from which cardinal teeth:

are developed, on the laminz at the posterior end of the shell,

but in some Leptonacea and Crassinella, where the internal

resilium is nearly central prominences which are genetically

identical with the so-called cardinals are formed also behind the

beaks. The inclination of the teeth to a perpendicular from

the beaks is a function of the torsion of the umbonal extremity

of the valve and may differ strongly in otherwise closely related

species. The cyclodont type of hinge largely results from such

torsion. This inclination therefore should not be discussed as

if it was an essential character of the teeth in themselves.

No. 411.] HINGE TEETH OF BIVALVES. 179

For those who may be interested to look into this subject it

will be useful to explain the formulz proposed by Bernard and

Steinmann, with their subsequent modifications.

In Bernard's scheme the teeth are numerals, — those of the

right valve with odd, and of the left valve with even numerals ;

the cardinals have Arabic, the laterals Roman numerals. The

character of the teeth is further indicated by capital letters, and

their origin by lower case italic letters corresponding to the

primitive lamina from which they are supposed to have been

detached in the course of development. Bernard begins with

the median cardinal and numbers both ways. In most pele-

cypods the cardinals are all anterior, though the laterals are

either anterior or posterior. He begins with the ventral lat-

erals and numbers upwards. One of the interesting points

made by Noetling is that the ventral lateral laminae are some-

times the younger, and therefore the homologies which may be

inferred from the numeration beginning below are liable to be

disturbed, whereas if the numeration of the laterals began from

. above, the dorsal laminz being, as he contends, the older, this

source of error would be eliminated.

In our opinion liability to error is inherent in the system

which attempts to. combine theoretical assumptions as to age

and origin with a numerical indication of facts. Laterals may

undoubtedly be added above as well as below in the course of

growth, for the dorsal margin of the hinge plate is added to as

well as the ventral; the hinge, when the ligament is opistho-

detic, or posterior to the beaks of the valves, increases at both

edges as long as the shell continues to.grow.

Bernard indicates the sockets by a colon and uses the letter

L between vertical bars to indicate the place of the ligament.

The portion of the ligament which is internal is (so far as

known) always the resilium, so it would be better to use R as

a symbol, the letter L being already appropriated to symbolize

the lateral lamina. Vertical bars are also used to separate the

anterior laterals from the cardinal series, though, as the index

letter L or C is already in use as an indicator, the bars do not

seem to be needed. If we take as an example the case of

a bivalve having three cardinal teeth and two anterior and

180 THE AMERICAN NATURALIST. [VoL. XXXV.

posterior laterals in each valve, Bernard’s formula would be as

follows, — the anterior end of the hinge being at the left end

of the formula.

Right valve LA I : HI :| Ca3a:1:3p: RILPELHI:

wert ave LÆT- IV | Cas ba pep Ap | RÍLP:H:IV

The Steinmann formula for the same shell is:

Rlolo:1orororlolo

Lolol-orototrrolol.

Ordinarily this formula is written in the opposite direction,

bringing the left valve above the line, but I have reversed it

here to make the formule more comparable. In the Steinmann

formula we put l for a lateral and 1 for a cardinal ; o for a socket,

and a period to set off the anterior laterals from the cardinals.

In this formula nothing is left to theory: the figures symbolize

exactly the number and relative position of the parts of the

hinge and a diagram could hardly do it better; of course it is

capable of enlargement : for instance, bifid teeth may be sym-

bolized by a fraction %, instead of a single digit ; the amor-

phous irregularities which occur on such a hinge as that of

Venus mercenaria may be symbolized by an X, and the solder-

ing of adjacent teeth by the insertion between their symbols

of a+sign. All the facts of the hinge of an adult shell can

be indicated by this formula. It is only when the facts of the

development of any particular hinge have been ascertained

that we can use the Bernard formula with equal confidence

and accuracy.

The initiation of denticulations on the hinge plate by

external sculpture, as pointed out by Neumayr, is illustrated

in many of the curious Paleozoic bivalves which he called

Paleoconchs. These forms, though not in themselves the

oldest known pelecypods, or prototypes, doubtless illustrate

the method by which in prototypic forms the organs in ques-

tion were evolved. Moreover, in the living Mytilidze, especially

the more minute and deep-water forms, the same process in

various stages can be directly observed. In some of them, as,

for instance, Dacrydium, the development of hinge teeth stops

No. 411.] HINGE TEETH OF BIVALVES. ISI

with the completion of the crenulations which in many other

pelecypods form only the nepionic hinge or provinculum. The

provinculum is common to all the Prionodesmacea, bivalves

which include Neumayr’s Taxodonta (Arca, Nucula, etc.) and

Dysodonta (Mytilus), and other ancient types (Unio, Perna,

etc.). A few of the Teleodesmacea (Heterodonta and Des-

modonta of Neumayr) also retain traces of it. The marvelous

thing in the development of the hinge is that in the vast

majority of hinges, including such primitive types as Nucula,

the hinge does not go on developing from the provinculum, but

the latter stops short and, as it were, after an interval, a wholly

new set of teeth, taxodont or other, begins to develop. In the

taxodonts there is a multiplication of similar teeth, often more

or less A-shaped, separated by a gap from the most adjacent

of the provincular teeth, even when closely similar to them.

The explanation of this discontinuity in development is the

most interesting question to be solved in connection with the

pelecypod hinge. In the Teleodesmacea instead of a multi-

plication of similar teeth we have a few, usually with one limb,

of the A disproportionately elongated. The apex of the A being

adjacent to the umbo, the proximal portions develop into the

so-called cardinals on the anterior side of the ligament. Only

im forms (Crassinella) having a very symmetrical development

is there any indication of a cardinal behind the ligament. The

distal end of the long limb of the A, usually the upper or outer

limb, develops into a so-called lateral tooth. This process is

characteristic of the forms (Venus, Cyrena, Mactra, etc.)

included by Bernard in his cyrenoid type of heterodonts.

In others the connection between the processes called cardi-

nals and laterals is practically wanting; the laterals arise

separately from the first, as far as observation can determine

at present. These are types having the hinge much elongated

and the laterals distant (Tellina), or absent (Macoma), or in

which the teeth are what I have called cyclodont (Cardium,

Isocardia), arising from below the hinge margin and practi-

cally without a hinge plate. Most of these were included by

Bernard in his lucinoid type of heterodonts. There is nothing

to deter us from believing that all these types are fundamentally

182 THE AMERICAN NATURALIST.

of common origin ; the few-toothed ledas approach the hetero-

dont type, sometimes very closely; but the divergencies must

have begun very early in the history of pelecypods and the

differentiation is now very marked. It is earnestly to be

hoped that such researches as those of Bernard will be con-

tinued until light is thrown on all the various problems for

which we now know no suggestion of a solution.

UNITED STATES NATIONAL MUSEUM.

THE PHARYNGO-CESOPHAGEAL LUNG OF

DESMOGNATHUS.

HARRIS H. WILDER.

SINCE the discovery of lungless salamanders in 1894, numer-

ous investigators have endeavored to ascertain the method of

respiration in these forms. Camerano (1894), after a series of

physiological experiments upon Sfelerpes fuscus, came to the

conclusion that integumental respiration was no better devel-

oped than in other Amphibia, and that the respiration must

be mainly bucco-pharyngeal. Directly along this line Maurer

(1897) showed that in lungless forms the pharyngeal capillaries

pass beyond the corium and actually invade the epidermis, a

condition unique among vertebrates. Hopkins (1896) supplied

an interesting point of negative evidence, that in lungless sala-

manders the pulmonary vein is entirely wanting, and that cor-

respondingly the left atrium is reduced to a rudiment. This

view was slightly modified by Bruner (1900), who showed that

what Hopkins had taken for the septum atriorum was really a

valve and that a proper septum did not exist, hence the single

atrium of lungless forms is probably due to a confluence of the

original two.

A careful study of the blood vessels of both lunged and

lungless salamanders was conducted by Bethge (1897), with

the result that he located in the latter (Spelerpes fuscus) a

pharyngeal plexus of capillaries, the vessels of which seemed

swollen at irregular intervals. Miss Woldt (1897) found a

pulmonary artery in Plethodon, supplying both cesophagus

and skin.

Thus far, however, the investigation seems to have centered

about the organs of circulation, and, consequently, a curious

set of facts along a different line has been entirely overlooked.

Although these have been met with and described by various

investigators, no conclusions have been drawn from them.

183 .

184 THE AMERICAN NATURALIST. (VoL. XXXV.

The first of these is the obvious fact, to which my attention

was first directed by Dr. H. S. Pratt, that lungless forms

breathe in the same way as do other salamanders ; that is,

they indulge in rhythmic respiratory movements during which

the throat fluctuates in and out with considerable rapidity.

A second closely related fact is that recorded by Bruner

(1896), that, in common with salamanders with lungs, the sev-

eral species of lungless forms investigated possess a muscular

apparatus for opening and closing the anterior nares. Although

in lunged forms this apparatus is directly connected with the

respiratory act, Bruner, influenced by the general opinion that

lungless forms could not really breathe, explained the presence

of this apparatus in such genera as Desmognathus and

Spelerpes as solely a protection against water. The third fact,

which has remained entirely unexplained until now, is the

presence in lungless salamanders of well-developed respiratory

muscles identical with those possessed by other Urodeles and

inserted into the walls of the pharynx and cesophagus (Wilder,

1896). - >

The meaning of these incongruous and apparently unrelated

facts has at last been made clear through the careful investi-

gations of one of my assistants, Miss Anne Ide Barrows, whose

preliminary report on the subject has just appeared in the

Anatomischer Anzeiger (Bd. XVIII, Nos. 18, 19) Miss

Barrows has just concluded a long and exhaustive study of the

entire circulatory system of Desmognathus fusca and has suc-

ceeded in demonstrating that not only the pharynx, as shown by

Bethge in Spelerpes, du¢ also the entire esophagus, is supplied

with a dense capillary plexus, the vascular area of which is of

Sufficient extent to transform the entire pharyngo-esophageal

region into a functional lung of at least as great respiratory

power as that of the paired lungs of normal salamanders. The

plexus is formed mainly by four arteries, two external maxil-

laries on the dorsal, and two pharyngeal on the ventral side.

Posteriorly a few branches of the gastric artery contribute to

the formation of the plexus.

It is worthy of note that a large part of the plexus is formed

by arteries from the fourth or respiratory arch, and that much

No. 411.] LUNG OF DESMOGNATHUS. 18

of the aérated blood is not returned directly to the heart but

enters ,the hepatic portal system through a pair of cesophageal

veins.

The demonstration of this respiratory organ puts the nec-

essary meaning into the three facts enumerated above, for

since there is a functional lung there must be respiratory

movements performed by respiratory muscles, and thus the

true function, both of the small muscles about the anterior

uu 4 d

| E (5

nil speed de Ned) One, S ! pua; / - Art. Pharyngea.

muscle gS d

— € nL Portion of

Pulmonary arch.

Esophageal ets m oou 4 trt. Gastrica anas-

muscle. N, H tomoses here with

the pulmonary

ch.

are

bo-~4rt. Gastrica.

| ~F. esophagea.

bier Ben of pure and — of ere ae Aveo ied the plexus of blood

. The b

the author, based upon one published

in set The blood vessels were added by Miss Nano.

nares and the large internal set, becomes evident. Concern-

ing these latter, the respiratory movements induced by them are

of two kinds: an act of inspiration which expands the pharyngo-

cesophageal region and an act of expiration which contracts

it again. The muscles employed in inspiration are evidently

those which arise from various dorsal and lateral points and

become inserted along the sides of the pharynx and cesopha-

gus, called by me the dorso-laryngeus, scapulo-pharyngeus, and

186 THE AMERICAN NATURALIST.

cesophageal muscle. These by their contraction draw apart the

walls of the alimentary canal in this region, and the external

air rushes in through the nostrils to fill the vacuum thus formed.

In my previous paper, not having the present point of view, I

said little about the muscles used in expiration, but it is now

obvious that the well-developed muscular layer which surrounds

the pharynx in some of the species, and which I have figured

in Fig. 2 of the article referred to under the name of pharyn-

geal muscular sheet, must have that function. It is also prob-

able that in some species the digastricus pharyngis becomes a

powerful expiratory muscle.

Summarizing the results thus far obtained, we may state

the following: Desmognathus fusca breathes mainly by means

of a definitely localized portion of the anterior part of the ali-

mentary canal, which may be known as the pharyngo-wsopha-

geal lung. The walls of this organ are richly supplied with

blood from a capillary plexus which is irregularly reticular in

its pharyngeal portion, and in its esophageal portion consists

mainly of very numerous longitudinal vessels which run paral-

lel with the mucous folds. Rhythmic inspiratory and expiratory

movements of this organ are caused by two sets of muscles, the

one dilating and the other contracting the lumen.

These motions are accompanied and assisted by respiratory

movements of the floor of the mouth, as in other salamanders.

The mouth is normally closed during respiration, and the nostrils

are used for the passage of air. The anterior nares are equipped

with a regulating apparatus, as in other salamanders, and for the

same purpose.

Although the above summary rests upon recent investigation

upon a single species, it is probable that a similar organ exists,

with some slight modifications, in the other lungless sala-

manders. The bibliography referred to here merely by the

author's name is given in full in Bruner's latest article on the

subject in the Journal of Morphologv, February, 1900, to which

the reader is referred.

SMITH COLLEGE, Nov. 16, 1900.

NOTES ON THE HABITS OF CAMBARUS

IMMUNIS HAGEN.

J. ARTHUR HARRIS.

In the present notes I intend to give only a few observa-

tions made on the habits of one of our common crawfishes,

Cambarus immunis Hagen. The material which I have exam-

ined seems quite variable, and a part of it would probably fall

under Faxon's variety, spinirostris. It does not embrace all

the data I have at hand, and as more is being collected as

rapidly as opportunities for field work are presented, I hope in

the near future to give a much more complete account of the

habits of this and other forms. These few points are pre-

sented here simply because they seem to throw some light

upon one or two questions concerning the habits of the craw-

fish, a field which I feel sure is deserving of careful study.

C. immunis is decidedly a mud-loving species. Faxon! says :

«Mr. H. G. Hubbard has found it in muddy pools and ditches

connected with the Detroit River, Michigan. According to Mr.

Hubbard, it does not form burrows but conceals itself among

weeds." Herrick,? in his description of C. signifer [= C. immu-

nis], says: “ Found by hundreds in a shallow pool known as

Grass Lake in Richfield, Hen. County." Forbes? says: ** This

is the commonest species of central Illinois. It is especially

frequent in the muddy ponds of the prairies, whence it may be

drawn by the hundred with a small seine." Hay * says: “ This

species is a mud-lover, being found in great numbers in muddy

1 Faxon, Walter. A Revision of the Asad Mem. Mus. Comp. Zool. Har-

ame MdL watt x (1885), No. 4.

Papers on the Crustacea of the Fresh Waters of Minnesota,

die eu on d Geol. of Minn., 1882.

3 Forbes, S. A. List of Illinois o with Descriptions of New Species,

Bull. IH. Ww. Lab. of Nat. Hist., vol. i,

W. P. The Crawfishes of the Suis of a Twentieth Ann. Rept.

Ind. pud Survey, 1896.

187

188 THE AMERICAN NATURALIST. [Vor. XXXV.

ponds in the early spring. I have always found them in the

greatest abundance in ponds which became perfectly dry dur-

ing the summer months, but where the crawfish go during this

time I have never been able to ascertain. Doubtless great

numbers of them are eaten by birds and other animals, and

great numbers of them perish; yet by the next spring they

are as abundant as ever and of about the same size."

C. immunis is our most common species in Douglas County,

Kansas, and is found in almost precisely the same habitat

as reported above for Illinois, Indiana, Michigan, and Min-

nesota. Roadside ditches and ponds in pastures are favo-

rite places. In this county we have ponds in an old river

bed which contain immense numbers of this species. These

ponds are six to eighteen inches deep, depending upon the dry-

ness of the season. They are very muddy and have usually in

the shallower parts, which are apt to become dry during the

summer, a rank growth of Polygonum and plants of like habit.

From such a pond I took, one day in October, 1898, in com-

pany with Mr. C. D. Bunker and another collector, about fifteen

hundred specimens, of which only a very few were less than

two and one-half inches in length. There were not more

than a half dozen small specimens in the whole lot; in other

ponds, however, I have made collections which showed almost

all stages of development. In many of the ponds C. immunis

seems to be the only species, but in others we find also

C. gracilis Bundy and C. virilis Hagen (?), but these do not

occur in nearly so great numbers as does C. immunis.

According to my observations, C. immunis is a burrowing

and, at least to a certain extent, a chimney-building species.

Burrowing, however, appears to be only resorted to when there

is danger of the drying up of the ponds or on the approach of

winter. Lack of opportunity for examining the ponds in all

different conditions without doubt accounts for the failure of

Hubbard’ or Hay? to observe the burrowing habit of this spe-

cies. Hay's suggestion of crawfish perishing upon the drying

up of the ponds seems to me to be largely ungrounded, since

I have never found any considerable numbers of dead ones in

! Faxon, W. Loc. cit. 2 Hay, W. P. Loc. cit.

No. 411. CAMBARUS IMMUNIS HAGEN. 18

many places which I have visited in all stages of drying up.

Possibly birds may eat a few, but probably these are mostly

water birds which could easily get the crawfish in the shallow

water at any time.

In early September, 1900, I visited a pond, which earlier in

the season had many crawfish in it, and found there were many

burrows around the edge and but few animals in the pond.

Upon digging out one of these burrows, which had a chimney

about four inches high and five inches in diameter at the base,

I found a large first-form male and a somewhat smaller female.

The burrow was about fifteen inches deep and about one and

one-half inches in diameter; at the bottom it was expanded

into a cistern-shaped chamber about three and one-half or four

inches in diameter. It appeared that the burrow had been

carefully sealed by the animals. Some time later I examined

the pond when it was almost dry and found no crawfish, but

plenty of chimneys in various stages of disintegration. At

about the same time in another pond, which contained more

water, the crawfish were abundant. About the middle of

October, 1900, I examined the burrows of C. immunis around

the first pond mentioned above. Some of the burrows had

chimneys, others were simply open at the top; probably those

which had no towers were old burrows whose chimneys had

been disintegrated by the weather. The chimneys were of

different forms, some being almost regular, while it was evi-

dent that others were formed simply by the mud being thrown

out of the burrow. Sometimes the mud appears to be just

thrown out on one side, then, after a considerable amount of

material has accumulated, the remainder is thrown out wherever

convenient, forming an irregularly circular mound one to three

inches high and six to nine inches across, or sometimes a long

ridge with the shaft in the middle, the material here being

thrown out in both directions. Sometimes the work is stopped

before enough mud has been removed to make it inconvenient

to throw it all to one side; thus forming a crescent-shaped

pile. The highest chimneys noticed were about five inches

high and three inches in diameter at the top. These were

sometimes much inclined from the perpendicular. On the

I9O THE AMERICAN NATURALLIST. [Vor. XXXV.

whole, I think that, for this species at least, the theory of

Tarr! and Shufeldt,? that the chimney is simply the deposi-

tion of the material brought up from the burrow, is correct.

The mud is brought up in very soft condition, and I think that

even if the burrow were made in the sloping side of a ditch

there would be little danger of the pellets rolling down, as

suggested by Abbott ;? in fact I have examined several burrows

on the sides of ditches, presumably in the same kind of posi-

tion as those examined by Mr. Abbott, and I have not been

able to convince myself that anything except the easiest

method of disposing of material from the burrows could have

prompted the animal to build chimney-like structures. The

quantity of material brought up is very considerable, amounting

in some cases to nearly two hundred cubic inches.

As suggested above, the accidental sealing of the burrows

would seem to be impossible. I believe that I found the

“ dumps " in enough different stages of construction to war-

rant the conclusion that the opening through the tower is left

open until the last, when it is closed by stopping with mud

brought from below. The opening is thus filled solidly from

the top, or near the top, to almost the level of the ground,

where the shaft has a concave hemispherical end, which is

not so smooth as Shufeldt's* description of the burrow of

C. diogenes would seem to indicate. And I cannot agree with

him that the animal might have used the lateral tail fins in

finishing this off, since I have never seen impressions in the

mud which would indicate such a method, while there are

many marks which would indicate that the animal had run the

ends of the chelz into the soft mud. In at least one case

which I examined, the burrows had been filled to somewhat

below the surface of the ground with clay so tightly packed

that it would seem impossible that it might have fallen in by

accident. The shaft usually goes straight down, but in some

! Tarr, Ralph. Habits of Burrowing Crayfish in the United States, ature,

vol. xxx (1884), p. 127.

? Shufeldt, R. W.

Chapters on the Natural History of the United States.

New York, 1897.

3 Abbott, C.C. Are the Chimneys of Burrowing Crayfish Designed? Amer.

Nat., vol. xviii (1884), p. 11 Lvl t Shufeldt, R. W. Loc. cit.

No. 411.] CAMBARUS IMMUNIS HAGEN. IQI

cases may be somewhat sinuous. They are quite deep, one

being followed down for four feet without reaching the end.

I am not able to convince myself that there was a ledge

upon which the animal could rest, as stated by Hubbard? for

C. argillicola, although in some cases the burrow seemed to be

expanded in a way which might serve for this purpose. In

one case examined quite early in the fall, the main shaft went

straight down ; at a distance of about six inches from the sur-

face a branch went off at almost right angles and ended

somewhat higher than it originated about eighteen inches from

its connection with the main shaft. It might seem that the

animals do not like to stay down in the bottom of the burrow

very long, at least early in the fall, since when a small exca-

vation extending somewhat below the water mark was made in

one of the burrows the animal came to the top and darted

back two or three times when I attempted to catch him, which

I finally succeeded in doing. The burrow was left for a few

minutes and when again examined a medium-sized female had

crawled out of the water and was standing on the almost

perpendicular side of the excavation.

Hay's? theory, that the burrows of C. diogenes are made to

escape the dry months of summer, seems undoubtedly the

explanation for the summer burrowing of this species, while

of course the purpose of burrows made or enlarged upon the

approach of winter is evident. It is certain that the burrows

are not adopted as retreats, while the eggs are being hatched,

for I have taken the females in the open ponds in the fall,

apparently soon after the eggs were laid, and they come out

early in the spring — about March 20 —to complete the

process of hatching the eggs.

UNIVERSITY OF KANSAS,

Nov. 10, 1900.

1 Faxon, Walter. je aep of New —— of Cambarus, Proc. Am. Acad.

Arts and Sci., vol. xx (1884), p. ? Hay, W. P. cit.

ARTIFICIAL INCUBATION OF ALLIGATOR EGGS.

ALBERT M. REESE.

THE writer received, about the first of August, 1900, a lot of

alligator eggs from southern Georgia, and in order to get

a series of embryos at different stages of development, it

was necessary to find some means of incubating the eggs

artificially.

As is well known, the Florida alligator lays her eggs, about

thirty in number, in a so-called nest, which she constructs of

sticks, leaves, earth, etc., on the banks of the pond or stream

in which she lives. The eggs are laid in the cavity of this

nest, and are carefully covered and allowed to incubate by the

heat of the sun. When the young alligators are about ready

to hatch, they make a curious squeaking noise, which attracts

the mother's attention, and she uncovers the eggs so that the

young alligators may not be smothered in the nest after they

escape from the eggs.

The lot of eggs above mentioned was sent by express from

Georgia to Baltimore, and arrived in good condition, being

packed in damp leaves and Zu us, probably from the native

nest. Several of the eggs were at once opened, and were

found to contain living embryos in an advanced state of

development.

As the incubator which is ordinarily used for chick eggs

was not in operation, an attempt was made to imitate nature,

and the eggs were put into a large bucket of the loosely packed

humus in which they had been received. Two thermometers

were thrust into the umus, one deep down to the bottom of

the bucket, and the other near the surface. The whole artifi-

cial nest was covered with thick cloths and. put in the sun to

warm. Before sunset the nest was brought into a small room

which could be tightly closed to prevent any very decided fall

in temperature. The weather was intensely hot at this time,

193

I94 THE AMERICAN NATURALIST. [Vor. XXXV.

so that no difficulty was experienced in keeping the eggs

at a sufficiently high temperature. By very careful watching,

the temperature of the eggs was kept fairly constant, the

extremes being 32? C. and 40? C. In spite of this care, the

embryos within the eggs were kept alive only a little over one

week, and whether they were killed by too great heat, too little

air, or too great variation in temperature, it was not easy

to decide. It would seem that in nature the eggs must be

subjected to a much greater range of temperature than they

were in this case, so it seemed probable that it was lack of air

that caused the death of the embryos.

Towards the end of August a second lot of eggs was

received, but the contained embryos were in such an advanced

state of development that most of the eggs were opened and

the young alligators preserved for anatomical study. "When

the shells were opened the young alligators would wriggle out,

snapping their little toothless jaws and blinking their eyes,

although there was still a mass of unabsorbed yolk protruding

through the abdominal wall as large as a pigeon's egg.

A few of these eggs were packed in a small box of damp

humus, to prevent drying, and were kept in an incubator at a

temperature of 37? C. On opening the incubator a couple of

weeks later, curious squeaking sounds were heard coming from

the inside of the eggs, the sounds which, in nature, tell the

mother that her young are about ready to hatch and should be

helped out of the mass of earth and leaves in which they are

buried. These sounds are audible at a distance of fifteen

yards or more, so that even when the eggs are buried in the nest

the parent is probably able to hear the call of her young. The

next day after the first sound was heard one of the alligators

broke out of its shell, and a couple of days later two more

hatched; the rest of the eggs proved to be infertile.

During the act of hatching, the young alligators would snap

at the fingers, or any small object, quite savagely; but after

finally escaping from the egg they could not be induced to do

so. The umbilical scar persisted for some time after hatching,

but gradually disappeared. The alligator is about 20 cm. in

length when hatched, and that an animal of its bulk should

No. 411.] INCUBATION OF ALLIGATOR EGGS. 195

have been contained in so small an egg is quite astonishing.

These three artificially hatched alligators are now living, in

apparent good health, in a glass-covered box in the laboratory.

They are fed, about once a week, on small bits of raw meat

which are thrown into the shallow pan of water in their box.

It is probable that had the first lot of eggs been treated as was

the second lot, the desired stages of development might have

been obtained.

JOHNS HOPKINS UNIVERSITY.

THE COLORS OF NORTHERN APETALOUS

FLOWERS.

JOHN I TOVELL.

THE dicotyledons, which include about two-thirds of flower-

ing plants, were divided by Jussieu and Enlicher into three

divisions — the Polypetalz, the Gamopetalz, and the Apetalz

—an arrangement familiar in most of our floras. The highest

type of a flower, according to De Candolle, was one that had

all the organs present, but inserted separately upon the recep-

tacle; for it may be argued, as A. Gray remarks, that fusion is

an arrest of development, and therefore an indication of low

rank, or less perfection, than the contrary. Accordingly the

Polypetala were placed at the head of the vegetable kingdom.

The morphological doctrine that the flower is a metamorphosed

bud or branch, and that the union of its parts marked an upward

progress, was not made the guiding principle in the arrange-

ment of plant families in a lineal series until a comparatively

recent date. The miscellaneous group of dicotyledonous fami-

lies, known as the Apetalz, were believed to be retrogressive

or degraded forms derived from both the Polypetala and Gamo-

petala, which had once possessed petals and conspicuous

flowers. This division, as originally constituted, was made

the receptacle for families the affinities of which were obscure ;

and, in the absence of knowledge as to its true position, was

placed as a sort of appendix after the Gamopetale. It required

the united labors of Braun, Hanstein, ‘and Engler to determine

and place in their proper collocation those families which are

clearly reductions, and to point out that those remaining

are not derived from the higher orders, but are primitive in

character. They are naked blooming, according to Eichler,

rather than abortive.

In many of the true apetalous families the perianth has

remained rudimentary, and in those of lowest rank has never

197

198 THE AMERICAN NATURALIST. (Vor. XXXV.

been differentiated into calyx and corolla. It is either want-

ing, or is represented by a number of small scale-like bodies,

indefinite in number, sometimes present in the staminate and

replaced by bracts in the pistillate flowers; or it may be so

ambiguous as to leave its morphological significance in doubt.

Gradually in the rise of the flower from its primordial stage

the perianth becomes more and more important, until in the

pink family, which occupies an intermediate position, the

corolla is large and conspicuous. The flowers are very gen-

erally wind-fertilized, and, as may be observed in the grasses

and sedges, this is not favorable to the high development of

the floral envelopes. Mechanical difficulties are also presented

by the aggregation of the flowers in a dense inflorescence.

For instance, in the staminate ament of the alder, where the

calyx is present, I found by actual count in an ament two and

a half inches long seventy-seven flowers; while in the stami-

nate ament of the willow (S. discolor), of about half this length,

there were two hundred an seventy flowers, or seven times as

many flowers to an inch in length. There is no room for

a perianth, and the office of protection has been assumed by

scales and woolly hairs.

Though the larger part of the families are anemophilous or

self-fertilized, their coloration is highly interesting as showing

what colors the bracts and perianth would develop naturally as

the result of chemical and physical influences, and in a limited

number of cases the effect upon such flowers of insect visitors

at a later period.

Lowest in rank of the choripetalous series stands the order.

of the Piperales, with perfect flowers in slender spikes destitute

of a perianth, and divided, according as the carpels are sepa-

rate or distinct, into the Piperaceze and Saururacex. The

former is a tropical family, but the latter is represented in

eastern America by Saururus cernuus, lizard's tail, a name sug-

gested by the slender spike with drooping apex. The flowers

are fragrant, with white stamens, and probably attract insects.

A large number of shrubs and trees have the flowers in

aments and are frequently referred to as the Amentacez.

They are widely distributed throughout the temperate and

No. 411.] NORTHERN APETALOUS FLOWERS. 199

northern regions, and have been developed to endure severe

climatic conditions. They are or were wind-fertilized, the

moncecious or dicecious flowers appearing in early spring, when

there are no leaves to intercept the pollen.

There are about thirty-five species of the Juglandacez, which

include the walnut and hickory, large trees valuable for timber

and fuel. The flowers are green; the staminate form a long

drooping ament, while the pistillate are solitary, or few in a

cluster. The perianth is present in both forms. In Juglans

there are four narrow petals in the sinuses of the calyx; but

in Hicoria they are wanting. An abortive ovary rarely occurs

in the staminate flowers.

In the Myricacez, or bayberry family, neither the male nor

female flowers possess a perianth, but its place is taken by

several bracts near the flower.

The Salicacez form a large family confined almost entirely

to the north temperate and arctic zones. It includes the pop-

lar and the willow. The poplar has no perianth, but the recep-

tacle is extended to form an oblique, cup-shaped disk. In early

spring the pollen is expelled forcibly a short distance by the

elastic purple anthers. In the genus Salix so variable are the

species and so freely do they hybridize that any entirely satis-

factory treatment from a systematic standpoint is impossible.

The Swedish botanist Anderson, whose standard monograph,

published in the Prodromus of De Candolle, was the work of

nearly twenty-five years, declared that he never saw two speci-

mens of Salix nigricans, which has one hundred and twenty

synonyms, that were exactly alike. In Great Britain the num-

ber of species have been placed all the way from twelve to

eighty. This genus is exceedingly interesting to the evolu-

tionary botanist, since it so fully refutes the ancient dogma of

the constancy of species. Though the flowers were formerly

anemophilous, they are now fertilized by insects. There is an

abundance of honey, a sweet perfume, and the bright yellow

stamens render the blossoms very conspicuous. As an evi

dence of their attractiveness, it may be mentioned that they

are collected in England for decorative purposes on Palm

Sunday, and are offered for sale in New England cities by

200 THE AMERICAN NATURALIST. [Vor. XXXV.

street flower-venders. In S. candida and. S. purpurea the

anthers are red, and in the former the style is also dark red;

but usually the anthers are yellow. The scales display con-

siderable range in coloration; in S. purpurea and several other

species they are purple; in S. nigra, yellowish ; in S. myrtz/-

loides, greenish-yellow, capsules reddish-green ; in S. uva-ursi

the scales are rose red at the tip; in S. humilis and SS. tristis,

dark red or brownish. The twigs also vary much in color; as

green, white-woolly, yellow, brown, red, crimson, and purple.

The willows are very attractive to insects and on a warm day

they may be observed hovering in clouds about the bright yel-

low sprays of bloom. The pistillate aments are not so con-

spicuous as the staminate, and in the case of S. discolor attract

a smaller number of visitors. On the flowers of this plant I

have collected five bees, nine flies, and two beetles. Species

of Andrena, seeking food for their young, and flies are very

common.

The inflorescence of the Betulaceze, birch family, is ane-

mophilous, and usually moncecious There is no corolla. In

Carpinus, Ostrya, and Corylus the calyx is present in the pis-

tillate flowers but wanting in the staminate; conversely, in

Betula and Alnus the staminate flowers have a calyx and the

pistillate have not. Originally the calyx was doubtless present

in both forms. Its presence or absence in the one sex or the

other of the different genera has been largely influenced by

mechanical conditions. In the fertile flowers of the hornbeam,

hop-hornbeam, and the hazel there are but few blossoms in the

capitate inflorescence, while the sterile flowers are more densely

aggregated. An opposite condition prevails in the birch and

alder, where, though both kinds of aments contain numerous

flowers, they are more densely aggregated in the fertile than in

the sterile. Where declinic flowers are solitary, or few in a

cluster, as in Fagus, Castanea, Asparagus, and Ribes nigrum,

both forms possess a perianth. The hazel (Corylus) derives its

English name from the color of the nuts, as “in hue as hazel-

nuts," Shakespeare, 7; aming of the Shrew, II, 1.1 The stami-

nate aments are yellow, and in autumn the leaves also become

1 Enc. Brit., vol. xi, P- 548.

No. 411.] NORTHERN APETALOUS FLOWERS. 201

a handsome yellow. The thread-like stigmas of the pistillate

are bright scarlet. In C. purpurea the leaves, husk, and pellicle

of the kernel are purple. The hornbeam has the anthers pale

yellow and the inner bark yields a yellow dye. In the genus

Betula the flowers are greenish-yellow and the leaves are a pale

yellow in spring and a bright yellow in autumn. The aments

of Alnus are a reddish-brown, and the leaves turn to a dull dark

brown. The styles, as a rule, in early flowering spring plants

are crimson, a coloring which, by converting light rays into heat,

favors the growth of the pollen tubes. In several instances I

have seen the male flowers of Alnus incana visited by the honey-

bee for pollen. Müller states that he has seen numerous honey-

bees collecting pollen on the male flowers of the European

species of hazel (Corylus avellana).

In the Fagaceze, or beech family, there are no petals, but the

calyx is present in both kinds of flowers. In the beech, one

of our handsomest trees, the greenish-yellow calyx is bell-

shaped, 5—8-cleft in the staminate, but 6-lobed in the pistillate

flowers. The inflorescence is in small clusters. In the chest-

nut (Castanea dentata) the flowers are exceedingly abundant

and give a yellowish tinge to the whole tree. The staminate

flowers may contain an abortive ovary and the pistillate five to

twelve abortive stamens; the former are in aments, the latter

clustered several in an involucre. The flowers of the oak are

greenish, sometimes reddish, as well as the scales of the invo-

lucre and the leaves in autumn. The species of Quercus are

exceedingly variable. Of Q. robur there are twenty-eight varie-

ties, while several other species have from eight to ten. No

sharp line of demarcation is possible, as they grade into cach

other by many intermediate forms. At the time of his revi-

sion of the family De Candolle wrote, ** It is difficult to believe

that above one-third of the actual species in botanical works

will remain unchanged." The wood of the Fagacec is com-

monly brown or reddish-brown, the inner bark of Quercus velu-

tina (tinctoria) is orange and yields a yellow dye.

It has been shown that the scales and perianth of the Amen-

taceze present a wide range of coloring, including nearly every

hue save blue. If the flowers were once entomophilous, as

202 THE AMERICAN NATURALIST. [Vor. XXXV.

has been maintained, and are the result of extensive degenera-

tion, then these colors may be the relics of an earlier higher

stage. But if the perianth has always remained rudimentary,

and the form of the inflorescence has been developed in con-

nection with wind-fertilization, then the coloring is due largely

to chemical and physical conditions. It is desirable to con-

sider briefly the origin of the ament, for which the Fagaceze

present special advantages. The ament, though frequently

referred to as a spike, is in reality a contracted panicle. It is

composed of clusters of flowers with a common involucre

arranged around a central axis or rachis, and is, consequently,

a branch system with the lateral axes of the first and second

order, which would bear solitary flowers, aborted or eliminated.

In the oak the female flower still remains solitary, and with

the involucre of many bracts represents a non-developed

branch; further steps are presented by the beech with two

flowers, and the chestnut with several in an involucre. The

association of these clusters along a common rachis would pro-

duce an ament, the production of which is the result of con-

traction and concentration, of elimination of axes, and arrested

development. There is no evidence that the perianth was ever

large and well developed. The primitive flowers were probably

perfect and possessed a simple and undifferentiated perianth,

which in certain genera has been wholly or in part replaced by

bracts orscales. The Piperales are regularly perfect, and rudi-

mentary ovaries and stamens are of frequent occurrence in the

Amentacez, especially in the Fagacec. The causes which

have led to the separation of the sexes are still involved in

much obscurity, though it is well known that nutrition and

climate influence differently the stamens and pistils. It is

evident, however, as Darwin has remarked, that cross-fertiliza-

tion must have been assured before the flowers became declinic,

since otherwise the species would have perished. When the

antiquity of these families, their wide geographical distribution,

the vast number of individuals,— in the case of the birch form-

ing vast forests in Russia, — as well as their floral structure, are

considered, there seems no reason to suppose that the flowers

were ever entomophilous and conspicuous.

No. dtr.) NORTHERN APETALOUS FLOWERS. 203

The order of Urticales does not show any advance in the

structure of its flowers over the Fagales. The order is a

large one, comprising some fifteen hundred species, distributed

chiefly through tropical regions. The flowers are small, in

inconspicuous spicate or axillary clusters, greenish, and ane-

mophilous. In Ulmus (elm), of the Ulmaceze, the flowers are a

reddish-purple, and the wood of U. fulva is reddish also. The

Moracee, mulberry family, is composed of trees and shrubs

with milky juice. The flowers are greenish, but in lieri

the calyx or receptacle becomes fleshy and bright-colored ;

Morus, red-purple and white; in Toxylon, ps elidel

and in Broussonetia the drupes are red. Of the tropical genus

Ficus there are six hundred species, three of which occur

in the Southern States. Its manner of fertilization has been

the subject of much discussion. The hollow, pear-shaped

receptacle is lined with male and female flowers, and pollina-

tion is effected by small wasps, which force their way into the

cavity forthe purpose of depositing their eggs. In fruit the

enlarged receptacle becomes deep purple, purplish-red, orange,

yellow, and whitish, and is eagerly devoured by birds, espe-

cially parrots. The greenish fruiting bracts and achenes of

— lupulus (hop) bear numerous yellow glands from z§5

to 4l, inch in diameter. When fresh they are filled with a

yellow liquid containing wax and resins and a bitter- tonic

medicinal principle called /upu/im. About one ounce may be

obtained from one pound of hops. These glands doubtless

protect the flowers from attacks of aphides. Glandular-leaved

peach and nectarine trees are less subject to curl, to mildew,

and to the attacks of aphides than the non-glandular (Darwin,

Animals and Plants under Domestication, Vol. I, p. 364). The

hop in some years is attacked by vast numbers of Aphis humuli,

and is also subject to blight from a parasitic fungus. The

flowers of the nettle family are green, wind-fertilized, and the

pollen is scattered by the explosion of the anthers.

The order Santalales includes about seven hundred and fifty

species, which are most abundant in tropical regions, only six

species being found in the Northern States. The two northern

species of the Loranthaceæ, mistletoe family, are parasitic

204 THE AMERICAN NATURALIST. [VOL XXXV.

plants which contain chlorophyll, and are yellowish or brown-

ish green. Tropical species of Loranthus produce magnificent

flowers 10-20 cm. in diameter and display most gorgeous

orange and purple colors. The flowers of the Santalacez,

sandalwood family, are perfect, the calyx is greenish-white

or purplish, and at least one species in Europe has been seen

to be visited by the honey-bee.

The colors of the Balanophoracee, a tropical family parasitic

on the roots of forest trees, which belongs also to this order,

are of much interest. The plants, of which there are about

forty species, resemble fungi, such as toadstools, producing

flowers. They were, according to Kerner, made the subject

of many fanciful speculations by the nature-philosophers, by

whom they were considered as “in the position of a hiero-

glyphic key between two worlds." The entire plant of the

American genus Langsdorffia is pale yellowish or, in the case

of the scales, waxen yellow, orange, or red. The genus Bala-

nophora occurs in the eastern hemisphere and is vividly

colored a deep yellow, red, or purple. In Helosis the floral

spadix is purple or blood-red ; in Corynea turdici, which lives

on the roots of Peruvian-bark trees, the purple spadix is sup-

ported by a white shaft. The coloring of the inflorescence of

Lophophytum leandri * cannot be exceeded," says Kerner, “ in

variety, its rachis being pale reddish-violet, the bract scales

gamboge, the ovary yellowish, the styles red, and the ovaries

white." The entire plant of Sarcophyte sanguinea from the

Cape of Good Hope presents a most striking appearance,

owing to the blood-red coloring of all its parts.

'The flowers of the Aristolochiaceze, or birthroot family, are

adapted to Diptera, especially to small gnats. The calyx is

highly specialized and in Aristolochia, familiar in A. sipho,

Dutchman's pipe of cultivation, is prolonged into a tube with

a contracted throat, either straight or shaped like the letter .5,

which is set on the inside with reflexed hairs. Flies can creep

inside easily, but when they attempt to escape they are pre-

vented by the hairs, which form an impassable grating. As soon

as the anthers have dehisced, the hairs wither, the calyx shrivels,

and the imprisoned insects are set free. The mechanism

No. 411.] NORTHERN APETALOUS FLOWERS. 205

of Aristolochia, which was first studied by Sprengel, ‘was

long," says Müller, “ the only example known of a temporary

prison for insects." Progressive steps toward this structure

are presented by Asarum and Heterotropa. The flowers of

Aristolochia are lurid purple, with a yellowish-green tube; the

tube of Asarum is greenish or brown-purple, puberulent with

purple hairs; or in A. macranthum the tube is mottled with

violet within. According to Kerner, South American species

of Aristolochia are of immense size and are used as caps by

children at play; in color they are a combination of cream and

deep maroon purple. Purplish coloring of bracts and sepals is

of so common occurrence, even extending to the whole plant,

that in this family it has doubtless been developed directly

from the primitive green without passing through any inter-

mediate stage. The flowers should be compared with the

pitcher-like leaves of Sarracenia and the spathes of Arum, as

all three serve as traps for small flies and are lurid purple, a

color probably attractive to these insects. Rudimentary petals -

occur in Asarum canadense.

The flowers of the Polygonacez, or buckwheat family, are

devoid of petals and are wind-fertilized, as in Rumex, or au-

togamous or entomophilous, as in Polygonum. In the genus

Eriogonum, growing west of the Mississippi, the campanulate

calyx is usually yellowish or white, sometimes changing to

pink. The calyx of Rumex (sorrel) is small and commonly

green, but in R. venosus it is red, and the fertile panicles of

R. acetosella turn reddish; but the achenes, or fruit, are usu-

ally red, and frequently the stems and leaves, as in A. san-

guinea. Red butterflies, which are attracted by red coloration,

were often seen by Miiller in the Alps seated upon the plants

when in seed, and a species of Halictus was observed fre-

quently collecting pollen. The plants of the Polygonacec

tend to develop red rather than yellow coloration; but in

Rumex persicarioides (golden dock) the fruiting calyx becomes

orange-colored, and the roots of several species of dock are

yellow.

Of the thirty-eight species of the genus Polygonum (knot-

weed) ten are white, ten red, three purple, and fifteen green.

206 THE AMERICAN NATURALIST. [Vor. XXXV.

Conspicuousness is gained by the union of the flowers in axil-

lary or terminal clusters. The green flowers are very small,

odorless and honeyless, and self-fertilized. The white and red

are more conspicuous, may contain honey, and are visited by

few or numerous insects; for instance, P. persicaria has a

white or red calyx, secretes honey sparingly, and is visited by

many flies and small bees. The species of this genus most

excellently illustrate the successive steps by which a green

perianth may become conspicuous. In the common door-weed

(P. aviculare) the margins of the sepals are white, turning

pink, while the centers remain green. The flowers are self-

fertilized, but are occasionally visited by flies. In P. convol-

vulus the two inner divisions of the perianth are entirely white,

but the outer are keeled with green; the calyx of P. virgini-

anum is usually green, tinged or tipped with white; P. persz-.

carta has in the same spike green, white, and red flowers ;

while P. orientale, cultivated from India, has large, bright rose-

colored flowers. In the case of the familiar buckwheat (Fago-

yrum fagopyrum) the waving fields of white bloom are very

conspicuous. The dimorphic flowers possess perfume, and

insects manifest special preference for the honey; in Germany

Müller has enumerated forty-one visitors, of which twelve are

bees. Cross-fertilization is insured and self-fertilization ren-

dered difficult. The marked tendency of both the vegetative

and floral organs in this family to develop reddish coloration is

evidently due to bright sunlight and the chemical constitution

of the sap, for the Alpine bistort (P. viviparum) of the White

Mountains often has little red bulblets in place of the flesh-

colored flowers. In P. scandans and P. dumetorum the calyx |

is yellowish-green.

The Chenopodiacez is a large family consisting chiefly of

herbs of a homely aspect. To it belong the garden beet and

the pot-herb spinach. The green flowers are very small, usu-

ally clustered, and in many species unisexual. The calyx is

usually present, but is wanting in the pistillate forms of some

genera. As in the Polygonacez, certain species show a tend-

ency to develop red coloration; Chenopodium rubrum has a

red calyx, and the inflorescence of C. album often turns reddish

NO. 4H. T NORTHERN APETALOUS FLOWERS. 207

in autumn, as well as the stem. In ZZwm capitatum (straw-

berry blite) the reddish calyx becomes bright red and juicy in

fruit, the globular, axillary heads resembling a strawberry ; in

Salicornia herbacea the whole plant turns bright red in autumn,

* forming vividly covered areas in the salt marshes, hence

called Marsh samphire.” The large converging lobes of

Salsola kali become rose-colored, and the leaves and outer

branches of S. zragus also turn bright red at maturity. Many

of the species are halophytes, most abundant by the sea and in

the salt marshes of Central Asia and in the basin of the Great

Salt Lake of Utah. The pollen is dust-like and the flowers

are anemophilous, or autogamous, though rarely visited by

pollen-eating flies. Dondia americana has purple-green sepals.

The red coloration noticed in the two preceding families is

also highly developed in the Amaranthacee. This is not so

much observable in our native species as in cultivated forms.

The foliage of the ornamental Amaranthus is richly variegated

with deep red, yellow, and green, “and the flowers are dark red.

In Amaranthus hypochondriacus from Mexico the entire plant

is tinged with red; while the whole plant of A. melancholicus

from eastern Asia is purplish. The crests of the flowers of

Celosia cristata (cockscomb) from India are rose, crimson, yel-

low, and white; in Gomphrena globosa (globe amaranth) the

dense round heads are crimson, orange, purple, and white.

In this family the sepals are — or united at base, or in

Froelichia form a tube.

In the Nyctaginacez (four-o’clock family) the calyx is cam-

panulaté or salver-form, corolla-like, with a deciduous limb.

The involucre resembles a calyx. The flowers are entomoph-

ilous and mostly pink or red in northern species; but in

Abronia fragrans the slender flowers are white, fragrant, open-

ening at night and adapted to nocturnal Lepidoptera. Florists

offer yellow, whité, and red varieties of Mirabilis jalapa. The

. sweet-scented M. longiflora is white, with a tube 15 cm. in

length and adapted to night-flying Lepidoptera. The limb

stands edgewise and is designed to render the flower more

conspicuous in the evening and not as a landing place for

insects.

208 THE AMERICAN NATURALIST. [Vor. XXXV.

The Portulacee and Caryophyllacez usually possess a

corolla, though it is sometimes wanting. The colors of the

flowers of the Portulacez are white, yellow, and red. Portu-

lacea oleracea, which, Warner fitly remarks in his charming

essays on gardening, ** grows with all the confidence of youth

and skill of old age," has yellow flowers and red stems;

P. pilosa has red flowers, and the intermediate stages between

“red and yellow are shown by the cultivated P. grandiflora. In

this latter species the white flowers have green stems, and the

yellow and red flowers red stems.

In the Caryophyllacez, a large family of some fifteen hun-

dred species, there are in the Northern States fifty-six white,

twenty-two red, two purple, and eight green flowers. The

green flowers are apetalous. Both the green and smaller

white-flowered species are low, tufted, weak herbs of a spread-

ing or ascending habit, represented by the chickweeds and

sandworts. The flowers are solitary or, at least, not densely

clustered, and usually white, or in Spergularia reddish. The

honey is freely exposed and the pollinators are chiefly flies,

beetles, and the smaller bees, such as Andrena and Halictus.

Certain species are visited also by butterflies and moths and

by the cosmopolitan honey-bee. I have never observed and

have been unable to find any record in the works of Müller

and Knuth of the visits of bumblebees. Many of the species

are dichogamous, but self-fertilization is always possible. The

. chickweed (A/szme (Stellaria) media), so widely distributed as

a garden weed, may be taken as a representative species.

The individual flowers are quite inconspicuous, but they are

numerous and, in contrast with the green foliage leaves, can be

seen at a considerable distance. I found on trial that a flower

could be distinctly seen at a distance of twenty-five feet ; but

after removing the petals it was visible only about four feet.

The plants blossom throughout the entire year, except when

prevented by severe weather, and in early spring and late

fall, when there are few other flowers in bloom, are very fre-

quently visited by flies. In April, Müller collected in Germany

six of the less specialized bees and four Diptera, and in the

middle of October I collected in Maine five species of Diptera

No. 411.] NORTHERN APETALOUS FLOWERS. 209

and Andrena. The honey is abundant. In winter the flowers

fertilize themselves. The white petals in these genera may in

part be due to the non-formation of chlorophyll, and in part to

the selective influence of the visitors in choosing the more con-

spicuous flowers. Insects have also probably aided in the preser-

vation of the petals, for in certain species of Alsine and Sagina

they are sometimes present and in other instances are wanting.

The pinks proper, or Silenez, exhibit a wonderful variety of

red shades, varying from white, through rose, pink, and deep

red, to scarlet and crimson. The petals may be dotted or

marbled with white, with a white center, surrounded with a

purple ring, as in Dianthus deltoides. The corolla is often

notched or ‘fringed and surmounted by a corona of scales.

The perfume is aromatic, and the honey is deeply concealed.

The red flowers are very attractive to butterflies, which are the

chief visitors, while the white species are adapted to night-

flying Lepidoptera. The following table, prepared from Miiller’s

Alpenblumen and Knuth's Handbuch der Blütenbiologie, shows

the importance of butterflies as fertilizers of red flowers.

VISITORS TO RED-FLOWERED SPECIES.

By | |

E Xon |

2 : i Eo

P E á B | er

S8 = E d | f

3 D " 5 A l F

BS E E: A E | g 3

i E a z

a a S15

SE ea z m a O =

Mee aedulis o. e 10 19 4 4 3 a

Lychnis fos Jovis . . . 10 3 I 4

OUR HNNE. 1 o 10-13 II 1 I 13

Saponaria ocymoides . . | 10-12 | 28 $ 3 ai 38

Dianthus sylvestris. . . | 18-25 I I

pric

= 1

_D. atrorubens ; 13-15 4 5

D. deltoides i. 12-14 9 2 II

D. carthusianorum . . . 12 9 6 3

The carmine flowers of Silene acaulis, which grows in

the higher Alps, are so frequently visited by butterflies that

the power of self-fertilization has been nearly lost. Both the

species of Lychnis have bright red, beautiful flowers and

210 THE AMERICAN NATURALIST. [Vor. XXXV.

attract many butterflies. The handsome red flowers of Sapo-

naria ocymoides are very abundant on sunny slopes in sub-

alpine regions and are sought by twenty-eight species of

butterflies. The species of Dianthus have the calyx tube so

long and narrow that the honey can be reached only by

Lepidoptera. The flowers are rose or dark red, elegantly

marked, of large size and great beauty. The association of

bright red coloration with a slender calycine tube and fertiliza-

tion by butterflies is not a coincidence, for throughout the

Caryophyllacez in proportion as the flowers increase in

conspicuousness the power of self-fertilization is lost.

* As the honey gets more deeply concealed and access more

directly limited to butterflies, we find," says Müller, “pari

passu among the Caryophyllacez increasing development of

sweet scents, bright red colors, fine markings round the

entrance of the flower, and indentations at the circumference.

All these characters which are so attractive to us seem to have

been produced by the similar tastes of butterflies.” This view

is much strengthened when it is considered that the nocturnal

flowers of the genera are white, and without variegation.

Saponaria officinalis is pale pink, or, on expanding, white, with

the perfume strongest in the evening; Sz/ene nutans, S. nocti-

Jora, and Lychnis alba all have white or pale pink flowers and

are visited by night-flying Lepidoptera. Since red is invisible

at night, while white is conspicuous, it is evident that the

former color would be disadvantageous to nocturnal flowers.

Originally the flowers of these genera were probably pale pink

or whitish, as in Gypsophila, where they are small, reddish, the

tube short, the honey fully exposed, and the visitors a miscella-

neous company of flies, bees, and Lepidoptera. A part of

the species became adapted to butterflies and a part to moths ;

no new colors were developed, but the red and white were dif-

ferentiated. The sexual markings of butterflies show that they

are in a very high degree color-loving insects, and while they

visit flowers of all colors they certainly prefer bright hues to

dull, and as with rare exceptions butterfly flowers are red, it is

probable that they find this color most attractive.

Under cultivation the pinks have proved susceptible of great

No. 411.] NORTHERN APETALOUS FLOWERS. 211

improvement, and numberless splendid varieties have been pro-

duced of every shade of red, dotted and striped, or marbled

and angled with white. In Dianthus barbatus (the sweet-

william) white and two different shades of red florets may

occur in the same fascicle, the white turning pink with age.

Florists also offer pure white and yellow forms, and white and

yellow varieties bordered with red or purple. Yellow, which

is well shown in Dianthus, is a comparatively rare color in the

Caryophyllacez. `

THE COLORS or NORTHERN APETALOUS FLOWERS.

$8| o | x "

ORDERS. FAMILIES. £a E E ; = i 4

CIN OE MEUM

Soa; E l= | me) a | a] oe

. Piperales . . .| /Saururacee. . . I 1

Juglandales . Juglandacez 13 13

Myricales iu ue | M yee 4 4

= Leitneriacez I I

Salicales. . . . Salicacex 9 D 4t 43

Fagales j Betulaceae 7 II 18

vu | Fagacez 25 25

Ulmacez 3 4| 7

Urticales . $ oraceæ 6 | 6

n Urticaceze 8 8

Santalales f Loranthaceæ 2 | 2

x Santalacew . . .| 2| 2 1 | 5

_Aristolochiales. . | Aristolochiacex . IO | 10

olygonales . . Polygonacez 33 | 22 § | 11 11 74

7 ( Chenopodiaceæ 3 I 39

Àmaranthaceze m4 i 17

Phytolaccacez 1 I

Chenopodiales . . |< Nyctaginacez . 1 41.3 8

am "Aizoacee . . 1 1 2

Portulacacee . . $1 3] 3 12

Caryophyllaceer .| 8 | 56 22| 2 88

Tas . . . .l|msi9o| 51145144 384

SUMMARY.

1. The apetalous Choripetala (Saururacea-Aizoacea) are

of primitive character; and are, or were, autogamous or ane-

mophilous. In the formation of the dense inflorescence

212 THE AMERICAN NATURALIST.

characteristic of many genera there has been much contraction

attended by the elimination of axes, leaves, bracts, and even

of the perianth, but the flowers are not degraded entomoph-

ilous forms. Consequently, when the flowers possess bright

colors they are not the relics of a higher stage of coloration

developed by the selective tastes of insects, but are due to the

chemical constitution of the nutritive fluids and the action of

light and heat.

2. The absence of blue is noteworthy. Yellow is not com-

mon, but is well shown in the scales and calyx of Betula. The

inner side of the calyx of Mollugo verticillata is whitish, and

in Polygonum the margins or entire calyx is white. Red is

very common and occurs in many genera. Purplish flowers

also occur, as Sesuvium maritimum. There is evidence derived

from this group of families that if anthophilous insects were

devoid of color sense, they still would have developed white,

yellow, red, and purple flowers, though they would be less

frequent and of duller shades.

3. A number of genera have become entomophilous, and

this change in the manner of fertilization has been attended

by an increased conspicuousness of the flowers. The anthers

of Salix, formerly a wind-fertilized genus, have become a

brighter yellow; in Aristolochia and Asarum the calyx is a

lurid purple attractive to small Diptera; and in Polygonum

and several other genera clear white or deep red. Insects

have not in these instances produced new colors, but have

intensified those already partially developed.

4. Petals are usually present in the Portulaceze and Caryo-

phyllacez, and are white, yellow, red, or purple. The smaller

white flowers of the Caryophyllacez are visited by flies, beetles,

and the short-tongued bees, which may have aided in the pres-

ervation of the petals and in rendering them a clearer white.

The larger red and white flowers are correlated with the visits

of Lepidoptera; the red species with butterflies, the white

nocturnal forms with moths.

WALDOBORO, ME.

PREHISTORIC WORKSHOPS AT MT. KINEO,

MAINE.

C. C. WILLOUGHBY.

THE porphyritic felsite of Mt. Kineo, Moosehead Lake, was

one of the chief minerals used for the manufacture of chipped

implements by the tribes of central and southern Maine. Chips

and broken implements of this stone were found in nearly all

of the camp sites and shell heaps which I have examined in

that state. The oldest New England people of whom we

have knowledge, and whose art remains were taken from the

very ancient graves explored by me in Hancock County, Maine,

in 1892-94, used knives of this

mineral.

Although erratic bowlders of

this stone furnished a limited sup-

ply of material, the chief source

was the great cliff of Kineo.

The southern side of this moun- PRAE Rr

tain is a mile or more in length

and rises nearly perpendicularly to a height of several hun-

dred feet. Its opposite side slopes gradually to the wooded

plain forming the northern portion of the peninsula.

In connection with other archaeological work in Maine car-

ried on under the auspices of the Peabody Museum of Harvard

University, the writer made two visits to Mt. Kineo, for the

purpose of locating Indian workshops and learning the manner

in which the rock was quarried or otherwise obtained.

The talus slope at the foot of the great cliff of Kineo

(Fig. 1, 2) is from two hundred to three hundred feet in width and

extends the entire length of the mountain. Patches of ever-

greens interspersed with deciduous trees are growing near its

base, but its surface is practically free from soil. The slope

of the talus is composed of comparatively small fragments

213

214 THE AMERICAN NATURALIST. [VoL. XXXV.

intermixed with larger masses of the rock. This talus is

constantly forming, and the colored patches along the face

of the cliff mark the places from which masses of felsite have

recently fallen, which are shattered as they strike the rocks

below, and the larger pieces rolling down the slope are chipped

and broken into innumerable forms. The recently fractured

pieces are easily distinguished from those which have been

long exposed to the action of the atmosphere. The fresh frac-

ture presents a green surface sprinkled with small dots and

squares of gray feldspar crystals. Upon long exposure the

surface becomes a uni-

form dirty gray. Upon

this slope one can gather

bushels of chips, flakes,

and pseudo-implements

wholly the work of

nature, which, if placed

unlabeled on the shelves

of a museum, would be

accepted without ques-

tion as the work of man.

For comparison a series

of these natural forms is

shown with an equal

number from Indian

J workshops at a distance

Fic. 2. —* map of Mt. Kineo and ad showing from the mountain. In

ocation of Indian workshop

selecting the natural

forms care was taken to include only those lying on the sur-

face near the top of the talus and having the green color

indicating freshly fractured stone. Only a few of the many

examples collected are shown. The natural forms are illus-

trated on Pl. I, z to 7. The artificial forms from the workshops

are shown upon the same plate, a to f. Typical artificial chips

showing the “ bulb of percussion” are seen in e and f, while 4

and / show natural chips having corresponding bulbs. These

* bulbs of percussion" are generally accepted as conclusive

evidence of the artificial origin of such flakes.

AS Ei HH

j ; ; N

f E: EAR AT ge 2 D <

(s e, D t1 Ax vs 2 i

CoO to Z

CP mes rs -—

€ \ At x

No. 411.] PREHISTORIC WORKSHOPS. 215

At points at the lower edge of the talus slope and in several

places on the low peninsula south of the mountain were indica-

tions of former occupancy by the Indians. Chips and rejectage

occurred in many places, but the principal workshops were

located at the points indicated upon the sketch map (Fig. 2).

Workshops 1 and 2, near the eastern and western ends of the

cliff, were evidently the principal blocking-out shops. These,

being near the water, had unfortunately been disturbed and

partially destroyed by the waters raised by damming. A large

amount of chips and general shop refuse was found at the

eastern shop (1). Large discarded worked nodules lay in beds

of chips. Ashes and charcoal occurred at intervals. A few

hammer stones were found, all of felsite. The rejectage of

this shop indicated principally the production of large imple-

ments. Very few nearly completed implements were found. |

Workshops 3 and 4 differed principally from those at the

foot of the talus in the size of the rejectage. Both had been

somewhat disturbed by the damming of the outlet of the lake

and the consequent washing away of portions of the shore.

Workshop 4 had been nearly obliterated, but the abundance

of refuse along the beach showed the types of implements

manufactured there. In both these workshops medium and

small *turtlebacks " predominated.

The ruder forms of rejects collected from the different

workshops are illustrated upon Pl. II, æ to e. These occurred

in great abundance, the larger examples being nearly all from

the shops at the foot of the talus slope. Types of the second-

ary forms are shown on Pl. III, e to 4. These were much less

abundant than the ruder forms. The largest specimen, e,

measuring eighteen inches in length, was found by Mr. L. L.

Hubbard in workshop 3, and presented by him to the Peabody

Museum.

Comparatively few implements broken in the last stages of

formation were found. The lengths of the perfect examples

of which these were a part would range from about three inches

to ten inches. Unbroken finished or nearly finished imple-

ments were very rare. They were of the types shown upon

Pl. III, a to d.

216 THE AMERICAN NATURALIST.

I could find no evidence that the rock used was detached

from the main mass by the Indians. The material was evi-

dently taken from the talus, fractured pieces being selected of

the size and form most readily chipped into the implement

desired.

That most of the products of the Kineo workshops were

intended for transportation and to be finished at a distance is

evident not only from the workshop refuse itself but from the

chips and more highly specialized forms of this material, both

broken and perfect, which are found in nearly all the burial

places, village and camp sites which I have examined in central

and southern Maine. Small chips of Kineo felsite are very

abundant in nearly all the village sites in the valleys of the

Kennebec and Penobscot rivers and their tributaries, and also

in the camp sites and shell heaps of the inlets, smaller rivers,

and islands along the coast between these rivers, and for some

distance east of the Penobscot and west of the Kennebec.

The broken or discarded implements found in company with

the chips in these places are more commonly small knives and

projectile points of various forms, together with scrapers and

perforators, types common in most prehistoric Indian village

sites, but absent or only occasionally found at the Kineo

workshops.

PEABODY MUSEUM OF AMERICAN

ARCHJEOLOGY AND ETHNOLOGY,

Cambridge, Mass.

—————— 4

PLATE I.— a-f, typical rejects and chips from Indian workshops, Mt. Kineo, Maine.

£-4, natural forms from the talus

[VorL. XXXV.

THE AMERICAN NATURALIST.

j iy M

219

PREHISTORIC WORKSHOPS.

No. ATI]

"eure *oourw IA ‘sdoysy10m uvrpu] woaz sjuouropdurr peusrag pue (sjoo[o1) sur10j Á1epuooss — *TII SLLV'Id

suf pousiung

AN LS

REVIEWS OF RECENT LITERATURE.

ZOÓLOGY.

Two Important Papers on North-American Mammals. — The

literature relating to recent work on North-American mammals is so

scattered, and the results have been the outcome of investigations

by such a number of different workers, and based on such varying

amounts of material, that it is a great gain when a competent author-

ity on any given group can go over it and coórdinate the efforts of

his predecessors in the light of, practically, all of their material,

combined with a vast amount in addition. In other words, the

monographic revision of any of the larger genera of North-American

mammals by an expert is a distinct advance, for which all mammalo-

gists may well feel grateful. It is with pleasure, therefore, that we

call attention to two recent contributions of this character — Mr.

Vernon Bailey's * Revision of American Voles of the Genus Micro-

tus," and Mr. W. H. Osgood's “ Revision of the Pocket Mice of the

Genus Perognathus."

Mr. Bailey's revision! of the American voles, or meadow mice, is

“ based on a study of between five thousand and six thousand speci-

mens from more than eight hundred localities, including types or

topotypes of every recognized species with a known type locality,

and also types or topotypes of most of the species placed in syn-

onymy." With such material at command, and with a wide experi-

ence with the animals in life, and personal knowledge of the actual

conditions of environment over a large part of the range of the group,

Mr. Bailey has had peculiar advantages for his work, and his results

are subject to revision only at points where material is still deficient,

or from some other point of view. This revision, while obviously

not final, presents a new starting point for future workers, and is

likely to be a standard for many long years to come

The little animals here treated are the short-tailed field mice,

! Revision of American Voles of the Genus Microtus. By Vernon Bailey,

Chief Field Naturalist, Division of Biological Survey, U. S. Department of Agri-

culture. Prepared under the direction of Dr. C. Hart Merriam, Chief of the

Division. North American Fauna, No. 17, pp. 1-88, with 5 plates and 17 text-

figures. Issued June 6, 1900.

221

THE AMERICAN NATURALIST. [Vor. XXXV.

familiarly typified by our common “meadow mice” of the Eastern

States. The group is divisible into several well-marked subgenera,

formerly generally known under the generic term “ Arvicola,” which

has had to give way to the less known but older term “ Microtus.”

The group is especially distinctive of the northern hemisphere north

of the tropics, and is found throughout North America from the

mountains of Guatemala and southern Mexico northward, increasing

numerically, both in species and individuals, from the south north-

ward till it reaches its greatest abundance in the middle and colder

temperate zones, again declining thence northward to the Arctic

coast. They are vegetable feeders, and often do considerable dam-

age to trees and crops ; they are active in the winter, forming long

burrows or tunnels under the snow ; they are also very prolific, breed-

ing several times a year, young being found throughout the warmer

months.

The seventy species and subspecies recognized by Mr. Bailey are

arranged in nine subgenera ; between the extreme forms the differ-

ences are strongly marked, but the intermediate forms present grad-

ual stages of intergradation. The subgenus Neofiber, of Florida,

embracing the round-tailed muskrat, and the subgenus Lagurus, of

the semi-arid districts of the northwestern United States, present the

most striking contrast, not only in size but in many other features.

The former is perhaps the largest known vole, while the latter group

includes the smallest.

Mr. Bailey’s paper, being a synopsis rather than a monograph,

leaves much to be desired in point of detail, but is admirable in its

way, and covers the ground with as much fullness as his prescribed

limits would permit. Of the twenty-six synonyms cited, it is notice-

able that thirteen relate to our common eastern meadow mouse, and

date from the early authors, while two other eastern species furnish

three others, also of early date. Only six of the remaining ten are

of recent date, showing that of some fifty-five forms described within .

the last ten years, by nine different authors, forty-eight meet with

Mr. Bailey’s approval. Four of the remaining seven are identified

with earlier names which for many years have been considered

indeterminable, but which Mr. Bailey claims to have established on

the basis of topotypes.

While he may be correct in these determinations, it would have

been of interest to his fellow-specialists if he had stated the basis of

his determination of certain type localities, notably those of Richard-

son's species, described as from the * Rocky Mountains," or similarly

No.411.] REVIEWS OF RECENT LITERATURE. 223

vague localities. If he has some *' inside history " to fall back upon,

it is only fair that the secret should be made public.

It may be said further, in the way of gentle criticism, that it is

hardly fair wholly to ignore such knotty points as the allocation of a

few names which he omits, since they form part of the literature

of the subject, as, for example, ypudeus ochrogaster Wagner, Arvi-

cola noveboracensis Richardson, and some of Rafinesque’s names.

Mr. Bailey describes as new two species and one subspecies.

Mr. Osgood's * Revision of the Pocket Mice”? is an equally wel-

come contribution, and has been prepared upon much the same lines,

with equal advantages in the way of material and field experience.

The pocket mice of the genus Perognathus are confined to a limited

portion of North America, being found only west of the Mississippi,

and ranging from the southern border of British Columbia south to

the valley of Mexico. They are strictly nocturnal and live in bur-

rows, are partial to arid regions and seem to thrive even in the most

barren deserts. Their habits are hence not well known, as they are

very shy and even difficult to trap. They are mouse-like in form, but

only distantly related to the true rats and mice. Their most obvious

character is the possession of cheek pouches which open externally.

The pocket mice vary greatly in size, form, and in the nature of

their pelage, which may be either soft or hispid ; but between the

wide extremes there are so many closely connecting links that it is

difficult to find any sharp lines of division, although two subgenera

are fairly recognizable. The whole number of forms here recognized

is 52 — 31 species and 21 additional subspecies, about equally divided

between the subgenera Perognathus and Chztodipus. Of these,

thirteen are here for the first time described. Out of a total of 61

specific and subspecific names applied to forms of this group, 9

are relegated to synonymy. Of these 61 names, it is interesting to

note that 52 date from 1889 or later, and that of these, eight prove

to be synonyms, three of them having become so through the identi-

fication of older names thought ten years ago to be indeterminable,

but since reéstablished on the basis of topotypes.

A previous revision of this group was made in 1889 by Dr. C.

Hart Merriam, on the basis of less than two hundred specimens —

1 Revision of the Pocket Mice of the Genus Perognathus. By Wilfred H.

Osgood, Assistant Biologist, Biological Survey, U. S. Department of Agriculture.

Prepared under the direction of Dr. C. Hart Merriam, Chief of Division of Bio-

logical Survey. North American Fauna, No. 18, pp. 1-72, Pls. I-IV, and 15 text-

cuts. Issued Sept. 20, 1900.

224 THE AMERICAN NATURALIST. [VoL. XXXV.

all of the material then available — when the number of currently

recognized forms was raised from six to twenty-one. Dr. Merriam’s

work, however, cleared the way for a better conception of the group,

rectifying important errors of nomenclature and making known many

new forms. Mr. Osgood, with fifteen times this amount of material,

seems to have settled all of the remaining doubts regarding the appli-

cation of certain early names, and, besides coórdinating the work

of his predecessors, has immensely extended our knowledge of the

group. The paper is admirable from every point of view and does

great credit to its author. TAA

The Eighteenth Annual Report of the Fishing Board for Scot-

land. — In this report Thomas Scott gives an interesting local list

of the fishes of the Firth of Clyde. The determination of species

seems to be accurate, and the nomenclature is more modern than

usual in British lists.

Mr. H. C. Williamson attempts by means of very many measure-

ments to ascertain whether a racial difference exists between the

mackerel of the east and west coasts of Scotland. He uses the

means employed for the distinction of races among men and lately

used by Heincke for the definition of races of herring. The Mean,

the Probable Error of the Mean, and the Standard Deviation are

derived from the formula given in Davenport’s Statistical Methods.

By these mathematical means the alleged variation in the mack-

erels of Scotland is elaborately investigated, with negative results,

the races not being sufficiently marked to require recognition.

Mr. H. M. Kyle, of St. Andrews, has a suggestive and valuable

discussion of the origin and mutual relations of the different groups

of flounders and soles. The arrangement adopted agrees in general

with that of Jordan and Evermann, which is based largely on earlier

researches of Dr. Gill. He would differ from Jordan and Evermann

in reducing somewhat the number of genera, and in separating the

Paralichthys type as a subfamily distinct from Hippoglossine. To

this subfamily, which he calls Hippoglosso-rhombinz, he would add

the allies of Syacium and Citharichthys. The soles constitute in his

view three additional subfamilies, Achirinz, Soleinz, and Cyno-

glossinzm. The affinities of these groups are obscured by making the

soles a distinct family, the three subfamilies being separately reduced

or degenerated groups of flounders. To all this there is no serious

objection, though Citharichthys and its allies seem to us rather closer

to the Psettine (or Rhombinz, as Mr. Kyle prefers to call them,

No. 411.) REVIEWS OF RECENT LITERATURE. 225

though the name “ Rhombus " is properly used only for another type

of fish). The allies of Xystreurys are, moreover, really intermediate

between Paralichthys and the Hippoglossine.

Mr. Kyle has added considerably to our knowledge of the olfactory

structures of the different groups and to our knowledge of the

shoulder girdle. His discussion of the origin of the different groups

is pertinent and sagacious. It is to be hoped that Mr. Kyle will

continue this line of work, and that he may secure specimens and

skeletons of the numerous genera which he has not yet examined.

We may note in passing that the genus Mancopsetta is of Gill.

DSL

Gill and Smith on American Moringuoid Eels.— A singular

group of eels of low structure, and distinguished among other things

by the extreme shortness of the tail and the backward location of

the heart, is the family of Moringuidæ. It has been supposed to be

exclusively East Indian, one species ranging northward as far as the

Liu-Kiu Islands of Japan.

Dr. Gill and Dr. H. M. Smith record in Science (June 22, p. 973)

the discovery of a species of Aphthalmichthys, a genus of this group,

from a coral reef near San Juan, Puerto Rico. Further study of

this type shows that the very slender whip-like eels of the West

Indies, constituting the subfamily Stilbiscinæ, are in fact genuine

Moringuidæ. Stilbiscus proves to be identical with Moringua. Gor-

dichthys must belong to the same group and probably Neoconger

also, thus giving four genera in America, as compared with the three

(Moringua, Raitaboura, and Aphthalmichthys) found in the East

Indies. The family is thus almost as well represented in the West

Indies as in the East. All the American species are very rare.

The species from Puerto Rico is to be described as Aphthalmichthy

caribbeus Gill and Smith. D. 5.1].

Transplanting of California Trout. — Students of trout in Cali-

fornia have noticed a number of anomalies in the distribution of the

different forms. The writer has been interested in following these

out, and now wishes to place on record for the reference of future

naturalists the facts in regard to them. If the investigation had

been delayed a few years until the clues were lost, these cases would

be altogether inexplicable.

In the tributaries of Feather River, around Prattsville in Plumas

County, is found the Lake Tahoe trout, Sa/mo henshawi. I learn

226 THE AMERICAN NATURALIST. [Vor. XXXV.

that these trout were placed in Feather River by Mr. Pratt, for

whom the town of Prattsville is named.

In the Blue Lakes of Amador County are also found trout trans-

ported across the Sierras from tributaries of Lake Tahoe.

In the streams running down the east slope of Mt. Whitney about

Lone Pine are found the golden trout of Mt. Whitney, Salmo agua-

bonita, These were transported by local anglers from Volcano

Creek, the isolated mountain stream above Agua-bonita Falls, in

which the peculiar form or subspecies has been developed.

This summer Rev. Edwin Sidney Williams, of Saratoga, Cal.,

transferred twenty young trout, the species not indicated, and a

dozen chubs from Pelican Bay on Klamath Lake into the famous

Crater Lake of Oregon, an extraordinary body of water without

inlet or outlet and, I believe, hitherto without fish life. WEN

An Error Corrected. — In Jordan and Evermann's Fishes of North

America the generic diagnoses of Collettia and Aéthoprora have been

by some unaccountable accident interchanged. It is Aéthoprora

which has a luminous gland on the front of the head “like the

headlight of an engine.”

In the same family of Myctophide, Neoscopelus macrolepidotus

Johnson, dredged by the A/ake in the West Indies, was omitted by

oversight. This genus, with Scopelengys, should apparently form

a distinct family, Neoscopelidz, distinguished by the broad maxillary

with supplemental bone. DSI

Notes on Recent Fish Literature. — In the Proceedings of the

Academy of Natural Sciences of Philadelphia, Mr. Henry W. Fowler

gives an account of the fishes from the Caroline Islands presented

to the Academy by Professor Cope. Forty-five species are enumer-

ated, the following new: Cypsilurus quindecimradiatus, Thalassoma

immanis, Scarus pronus, Scarus lupus. These are illustrated by

accurate but rather coarsely engraved plates.

Mr. Fowler gives an account of the typical specimens of Ameiurus

prosthistius described by Professor Cope from Batsto River, New

Jersey. This is regarded by Jordan and Evermann as a synonym of

the Florida species of catfish, Ameiurus erebennus. But Mr. Fowler’s

account leaves little doubt of its specific distinctness.

I venture to say that other species in this group will prove to be

valid. Especially is it likely that the short-bodied type, called Amei-

urus natalis, will prove distinct from the common form which has

been called Ameiurus lividus.

No.411.] REVIEWS OF RECENT LITERATURE. 227

Mr. Fowler redescribes the great catfish of the Florida Everglades,

which has been named Jctalurus okeechobeensis by Heilprin. He

regards it as a subspecies of Ameiurus lacustris, but until these

great catfishes have been fully studied, it seems as well to regard

this, with Jordan and Evermann, as a distinct species.

Dr. Einar Lonnberg, in the Annuaire of the Zoólogical Museum of

St. Petersburg, gives an account of the discovery by Dr. G. Adlerz

of the Opah, Zampris luna, on the coast of Murman in Russia. This

great pelagic fish is occasionally taken on almost every coast in the

world, especially in the northern hemisphere (Nova Scotia, Maine,

California, Japan, Madeira, etc.). Dr. Lónnberg adopts the earlier

name, Lampris pelagicus (Scomber pelagicus Gunner, 1768), instead of

Z. luna (1788). But there was already, in 1766, a Scomber pelagicus

of Linnzus, supposed to be the same as Coryphena hippurus, and

the name given by Gunner was preoccupied.

In the Zransactions of the Connecticut Academy, Vol. X, 1900, Mr.

Garman describes three fishes from Bailey Bay, Bermuda Islands,

collected by the Yale expedition of 1898. One of these, Brosmo-

Żhycis verrilli, is described as new. This belongs to the section or

genus Ogilbia of Jordan and Evermann. Godius stigmaturus is also

recorded from Bailey Bay. The original type was from unknown

locality, but Jordan and Evermann record the species from Key

West

Dr, Einar Lonnberg writes in the acts of the Swedish Academy of

the Saibling of “Baren” Island, as collected by J. G. Andersson.

To this form he gives the name of Sa/mo umbla var. salvelino-insularis.

In the Bulletin of the U. S. Fish Commission Professor J. P. Gor-

ham describes the *gas-bubble disease" of fishes in aquaria. He

finds it due to the expansion of gases from the reduction of pressure

in removing fishes from deeper waters.

Under the title of * Les Péches du Hokkaido," the Japanese Fish-

eries Bureau gives an interesting statistical account of the great

salmon and herring fishes of the island of Hokkaido (called Yeso

on our maps, but no longer bearing that name in Japan).

Dr. Seth E. Meek has published in the records of the Field

Columbian Museum an account of the species of Eupomotis,

the group which includes the common brook sunfish. Æupomotis

longimanus is recognized as probably a valid species.

228 THE AMERICAN NATURALIST. [ VOL. XXXV.

In a well-printed and finely illustrated volume published by

Appleton, Eugene McCarthy tells of the familiar fishes of the rivers

of the United States, their habits and the way to catch them. Mr.

McCarthy writes best of the phases and places of angling most

familiar to him, his first interest being in the Ouananiche or land-

locked salmon of Lake St. John. To this useful book a preface has

been written by Dr. Jordan. D SJ

North-American Reptiles. — The annual report of the Smith-

sonian Institution for the year ending June 30, 1898, contains, in

addition to a report on the present condition of the United States

National Museum, a monograph on the crocodilians, lizards, and

snakes of North America, by the late Professor Cope. . This

noteworthy contribution covers some 1120 pages of text, and is

illustrated by 347 groups of text-figures and by 36 plates. It is pro-

vided with a separate index. After a brief introduction the groups

and subgroups of reptiles are defined and their phylogenetic rela-

tions discussed. This is followed by a series of excellent descrip- .

tions of the species of crocodilians, lizards, and snakes found in

North America. The account is accompanied by keys for the

determination of species and by tables illustrating geographical dis-

tribution. Considerable attention is devoted to the comparative

anatomy of parts important from a systematic standpoint, and these

are well illustrated by clear but simple figures which fill most of the

plates. The account is concluded by a discussion of the geograph-

ical distribution of reptiles, particularly in their relation to the North-

American fauna.

This work, in connection with the forthcoming volume by the late

Dr. Baur on turtles, and Cope’s former monograph on the Batrachia

of North America, will place North-American herpetology next to

our ornithology in compactness and completeness of its systematic

treatment. p.

Porcupine Quills. — The arrangement of the quills and woolly

hairs on the eastern porcupine (Erethizon dorsatus) has been care-

fully described by Loweg.! In an embryo 18 cm. long the integu-

ment of the dorsal and lateral aspects of the body was covered

with short transverse rows of developing quills. Each row was com-

posed of some nine quills, the middle ones being longer than those

!Loweg, T. Studien über das Integument des Erethizon dorsatus, Jena.

Zeitschr. f. Naturwiss., Bd. xxxiv (1900), PP. 833-866, Taf. XXVII, XXVIII.

No. 411.] REVIEWS OF RECENT LITERATURE. 229

near the ends of the row. The rows were so placed on the surface

of the body as to form bands transverse to the animal’s chief axis.

In any given band the rows constituting it alternate with those of

the two adjacent bands. The rows of quills break the integu-

ment up into plate-like areas, which the author interprets, in accord-

ance with the conclusions of Weber and of others, as the remains of

scutes with which the ancestors of mammals are supposed to have

been covered. Judging from the condition of the integument in

the porcupine, these scutes were large and well developed dorsally

and small and poorly formed ventrally. In the porcupine the woolly

hair makes its appearance much later than the quills, and may be

formed on the scute areas. Phylogenetically the quills represent

the primitive hairs, and these are distributed in conformity to the

primitive scute covering. The woolly hairs, on the other hand,

are a much later acquisition, and are distributed without respect to

the places once occupied by scutes. As the skin of the porcupine

contains no sweat glands, the animal will probably be found to have

a summer and a winter pelage as an adaptation to temperature

changes. P.

Human Physiology. — The last addition to the series of Temple

Primers is entitled Zhe Human Frame and the Laws of Health, and

is a translation by F. W. Keeble from the German of Rebmann and

Seiler. The first ninety-five pages are devoted to the more salient

facts of human anatomy and physiology, and the remaining fifty to

hygiene. The presentation is remarkable for its clearness and its

general freedom from misstatements such as so frequently mar texts

intended to be popular. Here and there slight slips are to be

noticed: thus, on page 24 we are told that without the influence

of the nerve the muscle cannot contract, and on page 30, in the

description of the brain, we are informed that the third ventricle

gives off three clefts, lateral ventricles, on either side. Further, on

page 139, the distinction between smell and taste is inadequately

made out, and the subject is left in the confused state in which it

exists in the popular mind. Occasionally inapt expressions are met

with, as when (p. 31) the cerebral hemispheres are said to be marked

out into two unegual halves and (p. 99) ozone is described as a con-

densed form of oxygen. Even such small defects as these, however,

l Rebmann and Seiler. Zhe Human Frame and the Laws of Health. Trans-

lated from the German by F. W. Keeble. 148 pp. The Temple Primers.

London, Dent & Co.

230 THE AMERICAN NATURALIST. [VOL. XXXV.

are comparatively rare, for the book as a whole is a remarkably

trustworthy condensation of the chief facts of human physiology and

hygiene.

As to ** Social Ascidians.’’ — In a recent brief paper M. Maurice

Caullery ! adds one more to the numerous instances brought to light

in recent years tending to obliterate the distinction between social

and compound ascidians. ‘The case now reported is likely to be

noticed more than the others have been, in that it relates to the

genus Clavelina, which is one of the ascidians most familiar to the

general zoólogist, and is usually given in Merit um as a type of

the social ascidians.

In this genus the ascidiozooids, it will be recalled, bud from a

stolon, but remain entirely distinct from one another, excepting for ;

their connection with the common stolon. In the species here

described (there are two of them) the ascidiozooids differ struc-

turally in no way from a typical Clavelina ; but, instead of being

connected to the common stolon only, they are fully imbedded in a

common testicular mass also.

It is obviously necessary, Caullery says, to establish a new genus

for these species ; and the name proposed is Synclavella.

Had the author’s acquaintance with the literature of this subject

included the case of Perophora annectens described by me seven

years ago, he would not, perhaps, have been so sure about the

necessity of a new genus for his species. In this one species I

showed (Proc. Calif. Acad. Sei, Series 2, Vol. IV, p. 37) that “in

very many, though not all, of the colonies the ascidiozooids are as

completely imbedded in a common test as they are in Botryllus or

Goodsiria."

This species is exceedingly abundant at various points on the

California coast, and one may frequently observe transitional states

between social and compound on the same rock, and apparently in

the same colony. I may now add that, after having studied them

for a number of years, I have about reached the conclusion that the

social condition is the usual one; and that the compound con-

dition occurs only occasionally, even in the same locality. At

Pacific Grove, for example, where the most perfect instances of the

compound phase have been found, I have, on several visits, failed

to find any at all of this kind. It is an interesting fact, also, that

lSur des Clavelines nouvelles (Synclavella 2.g.), constituant des cormus

d'Ascidies composées, Comptes Rendus, No. 21, May 21, 1900, p. 1418

No.411.] REVIEWS OF RECENT LITERATURE. 231

sometimes, at any rate, the ascidiozooids of these fully compound

colonies are noticeably smaller than those of the social forms.

The specimens studied by Caullery were collected by Lesson in

1825, locality not given; and by Quoy et Gaimard in 1829 on the

coast of Australia. Wa. E. RITTER.

Parthenogenesis of the Honeybee. — Weismann publishes an

interesting preliminary account of studies on the parthenogenesis of

bees, which have been carried on in his laboratory for the past three

years. The conclusions of Dzierzon, confirmed by von Siebold and

Leuckart, that fertilized eggs always produce workers (or queens),

and unfertilized, drones (or males), having been in recent years called

in question by practical bee culturists, Weismann deemed it impera-

tive to have the question reinvestigated, especially in view of its

great theoretical importance. He accordingly induced one of his

students, Dr. Paulcke, to undertake the problem, and the studies thus

begun are now being completed by Dr. Petrunkewitsch, another of

Weismann's students.

One of the most energetic of the recent opponents of Dzierzon's

conclusions has been the editor of the Nördlinger Bienenzeitung, F.

Dickel of Darmstadt. His experiments seemed to prove that nor-

mally a// eggs are fertilized. Eggs which had been laid in drone

cells were transferred to worker cells, with the result that they devel-

oped into workers, and vice versa. Dickel argued, further, that the

evidence of von Siebold and Leuckart, based on the microscopic

examination of the eggs, could not be regarded as definitive, now

that we know more precisely the phenomena accompanying fertiliza-

tion. These observers had not examined eggs immediately after they

were laid, but only after the lapse of from one to twelve hours. But

it is now known that spermatozoa lose their thread-like form within

a few hours after the eggs are laid, and are succeeded by the sperm

nucleus and sperm aster ; in fact, it has been recently asserted that

this metamorphosis takes place in bees' eggs within fifteen or twenty

minutes after the entrance of the spermatozoóün into the egg.

Nevertheless, there is no ground to doubt, says Weismann, that

von Siebold saw seminal filaments (even two to four in a single egg),

for Blochmann, making use of the sectioning method, has seen the

same, and Petrunkewitsch now confirms the observation ; but these

could have been seen only in von Siebold's freshest eggs, all of which

! Weismann, A. Ueber die Parthenogenese der Bienen, Anat. Anzeiger, Bd.

xviii, Heft 20-21, PpP- 492-499, December 5, 1900.

232 THE AMERICAN NATURALIST. [VoL. XXXV.

were, however, from worker cells. The twenty-seven drone eggs which

he examined were all *about twelve hours old," so that his failure

to find in them sperm filaments is in no way proof that fertilization

had not taken place.

The most of the material for the investigations in Weismann’s

laboratory was furnished by F. Dickel, who took the eggs from the

hives, put them at once into the preservative fluid and sent them to

Weismann. The main results are as follows: Whether the egg is

fertilized or not can be determined with certainty only when it is

killed in the stage of the second maturation spindle. Before that, in

the stage of the //rs? maturation spindle, either the sperm nucleus

is without radiations, or the sperm filament has not yet been meta-

morphosed into a sperm nucleus. In either case it is a matter of

chance, depending on location and physical condition whether the

sperm cell can be recognized with certainty. In the second-spindle

stage, on the contrary, the formation of a sperm aster is completed

and the structure can neither be overlooked nor misinterpreted.

Petrunkewitsch sectioned 123 eggs that were in the first-spindle

stage. Twenty-nine of these were from worker cells; in twenty-three

cases (79%) the sperm nucleus with radiations was present. On

the other hand, ot a single sperm aster was found in any of the

ninety-four eggs from drone cells. ‘The condition of eggs in the second-

spindle stage was still more striking. very one of the sixty-two

eggs from worker cells showed the sperm aster ; whereas of the 272

eggs from drone cells ovZy ove contained a sperm aster. It is explained

by Weismann that in this one case the queen probably made a mis-

take and deposited a fertilized egg in the wrong (drone) cell, a phe-

nomenon which bee-keepers have long recognized as occasionally

taking place.

Weismann believes, therefore, that Dzierzon’s views are fully con-

firmed, — that normally eggs laid in drone cells are not fertilized and

that those laid in worker cells are always fertilized.

Dickel’s observation, that as soon as the queen has laid an egg,

workers enter the cell and busy themselves with the egg, probably

licking it and coating it with saliva, does not warrant his conclusion

that the sex is determined thereby. What the significance of that

act may be is not known. That it is of importance seems to follow

from the results of some of Dickel’s experiments. He isolated a

comb containing freshly deposited eggs by removing the workers

from it and then enclosing it in fine gauze, without, however, remov-.

ing the comb from the hive. All such eggs perished sooner or later,

No.411.] REVIEWS OF RECENT LITERATURE. 233

frequently in late embryonic stages. It may be that here, and in

other insects whose eggs have a thin chorion, there is need that the

egg be coated with saliva to prevent its drying up. Whatever the

influence of the secretions of the three pairs of salivary glands, sex

in bees is determined by the existence or absence of fertilization.

Other conditions within the sex — whether, for example, worker or

queen shall result — may be determined (as they apparently also are

in termites, according to Grassi's observations) by the quantity and

quality of the food, including the salivary secretions. How it is that

fertilization determines sex is not known, but that it does can no

longer be denied.

Dickel has insisted that there is a difference between drones pro-

duced by fertile workers and those produced by queens, and he

believes that the former are infertile. Weismann says he knows of

no proof of this, but admits that the studies of Petrunkewitsch show

that there is a slight, though constant, difference in the early phe-

nomena of development between unfertilized eggs from queens and

those from workers, and that this may possibly be of importance.

Further study on this point is required. A difference between queen-

drones and worker-drones is a priori in no way improbable; indeed,

there is a wasp in which two kinds of drones are known to exist,

though it is not known whether these have the same or a different

parentage. m.

Artificial Parthenogenesis. — Following the lead of Morgan and

Loeb, who have shown that certain salts can induce the development

more or less complete of unfertilized eggs of the sea-urchin, Pieri *

and Winkler? have, independently of each other, tried to induce

development of the unfertilized echinoderm egg by means of sperm

extract. Their efforts have been at least partially successful, though

the observations made are less extensive than might be desired.

Pieri took fresh, sound sperm of Strongylocentrotus lividus, or of

Echinus esculentus, and shook up the same either in sea water or in

distilled water. The fluid was then filtered and examined under the

microscope. It was found to contain still a certain number of sperma-

tozoa, but these were motionless, rounded, tailless, and, so far as

direct observation could show, dead. Unfertilized eggs of the species

from which the sperm was obtained were then placed in some of this

! Pieri, J. B. (99), Exp. (3), 7, Notes et revues, Arch. de Zool., p. xxix. =

2 Winkler, H. (1900). Nachrichten v. d. k. Gesellsch. d. Wiss. Gottingen,

Math -phys. Klasse, p. 187.

234 THE AMERICAN NATURALIST. [Vor. XXXV.

fluid on a glass slide, and watched under the microscope. A certain

number of the eggs developed (without fertilization) upto the morula

stage. The fluid was found to be effective in causing the develop-

ment of unfertilized eggs, even when ten hours old. Control lots

of eggs placed in sea water or in distilled water did not develop at

all. The sperm extract made in sea water worked better than that

made in distilled water, which is not at all surprising considering the

known injurious effects of fresh water on marine organisms and their

sexual products.

Pieri admits that his experiments would be more convincing if a

centrifugal machine and a porcelain filter had been employed in mak-

ing the extract. He believes, however, the fact to be established that

an extract of spermatozoa contains a chemical substance capable of

causing the development of unfertilized eggs of the species producing

the sperm. ‘This substance he believes to be a soluble ferment,

ovulase (Dubois, 1900, Mémoires Soc. Biol. Paris, 52, p. 197). No

evidence is offered in support of this view.

Winkler’s experiments and results are similar, though slightly dif-

ferent methods were employed by him to obtain the extract, and

greater precautions were taken to exclude from it living spermatozoa.

Sperm of Spherechinus granularis or Arbacia pustulata was put into

distilled water, shaken frequently during half an hour, and the fluid

then filtered five or six times through three thicknesses of filter paper.

The fluid was then brought up to the density of sea water by adding

to it salt obtained by evaporation of sea water.

Sperm extract was also prepared by putting sperm into concen-

trated sea water (400 c.c. evaporated to roo c.c.). In this sperma-

tozoa shrunk at once, but were allowed to stand for half an hour,

being frequently shaken. The extract was filtered as already

described and then diluted to the concentration of normal sea

water.

In the extract obtained by either method a certain number of

unfertilized eggs underwent cleavage, though the process was not of

the normal sort, and it did not progress beyond the 4-cell stage.

Stained eggs showed that genuine mitoses occurred. Control lots

which were put into sea water, instead of sperm extracts of the same

density, did not develop in any instance..

Only negative results were obtained by heating sperm to 50°-60° C.

in sea water and placing eggs in the cooled fluid. Winkler expresses

no opinion as to the chemical nature of the extract substance which

causes the development of unfertilized eggs, but thinks the idea of

No.41] REVIEWS OF RECENT LITERATURE. 235

Pieri and Dubois that it belongs to the enzymes lacking in support

as yet.

The observations which have just been briefly summarized add to

the accumulating evidence that fertilization may be essentially a

process of stimulation of the egg. This, of course, is true only if one

leaves out of account the contribution of hereditary tendencies made

by the spermatozoón, which add to the variability of the offspring.

The egg, however, apparently contains everything necessary for the

production of a new and complete organism like the mother, and

needs only appropriate stimulation to start it on its course of develop-

ment (see Koulagine, 1898, Zool. Anz., 21, p. 653 ; also Delage, 1900,

Archives de Zool., Exp. (3), 7, p 525). In this direction point also

the observations of Tichomirov, who showed that the unfertilized

eggs of Bombyx mori can be made to develop, at least to an advanced

embryonic stage, either by dipping them in sulphuric acid or by brush-

ing them; likewise thé observations of Dewitz, who obtained cleavage

of unfertilized frog's eggs by treating them with corrosive sublimate ;

likewise the observations of Hertwig, who observed a similar result

following the treatment of sea-urchin eggs with strychnine ; further,

those of Koulagine (already cited), who has induced cleavage of

certain fish and amphibian eggs by treating them with the antitoxin

of diphtheria; and those of Klebs, who found that parthenospores

are formed in conjugating filaments of Spirogyra, in the presence of

salt or sugar solutions of appropriate density. Still more emphatically

is this indicated by the recent work, already mentioned, of Morgan,

and especially of Loeb. W. E. C.

PETROGRAPHY.

Graphic Representation of Rock Composition and Schemes of

Rock Classification. — The recent great increase in our knowledge

of rocks and the complexity of their chemical relationships is responsi-

ble for the attempts now being made to indicate by graphic methods

these relations and incidentally to classify rock magmas on some

chemical basis. The differential hypothesis was intended to explain

the cause of the chemical relations existing between rock masses.

It cannot serve, or at any rate it has not served, as a basis for rock

classification. Rocks as objects of study are but portions of differ-

entiated masses. A rock classification must deal with these portions,

and it must necessarily deal with well-characterized portions or

236 THE AMERICAN NATURALIST. (Vor. XXXV.

types. The difficulty in all classification schemes has been to select

such types as will express the essential differences between nearly

related rock masses and at the same time to show the relationships

existing between them. The graphic method of representing rock

analyses by diagrams serves to emphasize the characteristic chemical

features of the different specimens analyzed, and to enable the student

to recognize readily their differences and similarities. From these

diagrams a composite diagram may be constructed, and this repre-

sents a chemical type. The chemical relationship of various types

is easily read from their diagrams; and classification of the dia-

grams is a classification of rocks according to chemical composition.

The first complete diagrammatic representation of rock composition

was suggested by Brógger.! His diagrams are polygons drawn from

the ends of radius vectors whose lengths correspond to the propor-

tions of the constituents in the rock represented in the diagram.

The shapes of the polygons show at a glance the relative importance

of the principal components. They are extremely characteristic of

the different rock types, and thus may be used to exhibit chemical

relationships.

Hobbs? makes a few unessential modifications in the Brogger

diagrams and combines them to form composite diagrams or dia-

grams of rock types. This he does, however, not by actually com-

bining the individual polygons, but by averaging the analyses of the

rocks which are assumed to belong with the type in question. So far

as rock classification is concerned this method is not of great value,

since it does not exhibit the differences in composition between the

constituent members of the composite and the type produced by them.

It affords a very convenient method of exhibiting the peculiar chemical

characters of rock families and a concise method of showing their

chemical relationships with one another, but the personal element

enters so largely into the selection of the rocks that comprise the

groups represented in the diagrams that these are not of great use

in other respects. If the analyses represented in the diagrams could

be averaged in such a way as would take into account the abundance

of the various rocks, the composites would possess a high scientific

interest. The idea proposed by the author is, however, an excellent

one, especially for comparative purposes. It is bound to be received

with favor.

Harker? suggests a method of using diagrams of rock series

! Das Gangafolgs des Laurdalits, p. 255. Kristiania, 1898.

? Journ. of Geol., vol. viii (1900), p. 1. 3 Journ. of Geol., vol. viii (1900), p. 389.

No. 411.] REVIEWS OF RECENT LITERATURE. 237

with a view to determining the mode of origin of certain rock masses.

He lays off on a system of rectangular coordinates the proportions of

the chemical constituents present in the known members of a series

of rocks derived by the differentiation of the same magma, and draws

lines through the corresponding points in the figure. Then by inter-

polation the composition of intermediate members of the series may

be determined by inspection. When the lines passing through the

points representing the proportions of the constituents present in

the various rocks are straight lines, the series concerned is a “ linear

series," that is, the rates of change in the constituents are constant

and their proportion is determined by the percentage of SiO, present.

This is a “rock series" in Brógger's sense. Such a rock series,

however, Harker thinks nonexistent. The normal diagram of a

*rock series" consists of curved lines, some of which are concave

and others convex. Sometimes when a rock’s analysis is plotted the

discovery is made that it does not fit into the diagram of the series.

Such abnormal rocks may have been formed by the admixture of two

magmas, or by the solution of foreign rock fragments in a normal

magma. In either case the resulting mixture possesses a peculiar

composition, the plotting of which does not fit into the diagram.

For instance, the lavas of the Lassens Peak district, comprising

basalts, andesites, dacites, and rhyolites, form a rock series the com-

position of which may be represented by a well-defined diagram. The

quartz-basalts, however, refuse to adapt themselves to the scheme.

Their abnormal composition is clearly brought out by the plotting.

In their content of lime and potash they do not differ notably from

normal rocks of the same silica percentage, but they show a

marked deficiency in alumina and ferric oxide and to a less degree

in soda, . Magnesia and ferrous oxide are in excess.

The plotting of analyses in the manner indicated by the author,

and the interpretation of the diagrams thus made in the way out-

lined above, form a ready means of detecting rocks of abnormal

composition in a set of normal rocks from the same petrographic

province. The diagrams thus may be made a means of aiding

rock classification, since they enable one to exclude from dis-

cussion those rock species which can have no part in any series of

related rocks, and which if discussed together with normal rocks

would give rise to difficulties hard to overcome in any scheme of

classification. The author declares that a natural classification

“must be based, confessedly or implicitly, upon fundamental genetic

considerations, and primarily upon the mode of operation of the

238 THE AMERICAN NATURALIST. [Vor. XXXV.

processes of differentiation in rock magmas. Rocks resulting from

admixture must therefore be excluded from the main scheme and

be relegated to an appendix. Any discussion which tends to the

recognition of this principle and to the establishment of some crite-

rion of distinction will forward the object by disembarrassing the

problem of a disturbing element."

The most recent and most complete discussion of the chemical

elements of rocks is by Osann,' who proposes a formula for each rock

which shall represent approximately its chemical composition and at

the same time be capable of easy plotting. He also proposes a

method of plotting by which the composition of a rock is indicated

by the position of a dot in a triangular diagram. The dot in its

relation to the bounding lines of the triangle exhibits at once the

relative proportion of the important constituents in the rock mass.

A single dot thus represents an entire rock. By plotting a large

number of rocks on the same diagram their chemicalrelationships are

easily and conveniently studied. The plotting at once exhibits a

grouping of closely related rocks and differentiates those whose mineral

composition may be similar but whose chemical composition is

different. The personal element has no place in the system of

plotting. The types are determined solely by the grouping of the

dots which express by their position the composition of individ-

ual specimens. In the constructure of his rock formulas Osann

transforms the percentage analyses into molecular proportions and

recalculates to the sum of 100. The mean of two analyses of the

norite ? of Montrose Point on the Hudson River thus treated becomes

SiOz Al203 FeO MnO MgO CaO NasO K0

59.50 10.35 7.38 36 8.14 8.65 4.23 1.39

The percentage of SiO, (including ZrO, and TiO,) in the recalcu-

lated analyses is represented by S with the corresponding exponent.

By A is represented the alkalies in the molecular group (NaK)s

AlO, In the above instance 4 = 5.62. The lime is considered

as in the group CaALO, and is represented by C. But CaO may

be present in another molecular group, whereas the Al,O, occurs only

in groups Aand C. Hence C= 10.35 (molecular percentage of Al,Q;

in analyses) minus 5.62 (the proportion of Al,Q, in group 4) = 4.73-

The remaining oxides are regarded as being present in the group

(Fe, Mn, Mg, Sr, Ba, Ca)O, and this is represented by Æ, which

1 Tscher. Min. u. Petrog. Mitth., Bd. xix, p. 351-

2 Amer. Jour. Sci, vol. xxxiii (1887), p. 193.

No. 411.] REVIEWS OF RECENT LITERATURE. 239

measures the dark components of the rock mass. In the norite

F= 19.78, vig, FeO = 7.38 + MnO —.36 + MgO —8.15, and the

remainder of CaO left after subtracting 4.73 CaO (the amount

necessary to combine with the 4.73% of Al,O, in group C) from

8.65 CaO, the amount shown in the analysis. The relation of the

alkalies to one another is indicated by the letter 7, which is given a

value corresponding to the proportion the NaO bears in the analysis

to the total alkalies calculated to the unit ro. In the above analysis

the Na,O is to K,O as 4.23 is to 1.39, or as 7.5 is to 2.5. 7 thus

is 7.5. The formula of the rock is

S 2s BO. 5O, A -—ts2, Cup F — 19.78, n=

In the plotting the absolute values of 4, C, and ¥ are not used,

but instead their proportions are calculated to a total of 20 units,

and these ratios are made use of. The values 5.62, 4.74, and 19.78

are as 4: 3: 13 in the scale of 20, and the simplified formula is

559-5 @4 Ca fis, 27-5 This formula not only expresses the approximate

chemical composition of the rock for which it is calculated, but it

expresses also roughly the proportions of alkaline feldspars, lime

feldspars, and dark components present in it, and from the formula

may be calculated the analysis. The ratios a: c: fare represented

in a triangular projection by a dot, the position of which discloses at

a glance the chemical character of the rock it represents. „Rocks

of similar composition are represented by groups of dots in certain

portions of the triangle, and these groups are observed to fall natu-

rally into subgroups. The discussion of these features is reserved by

the author until after he has plotted the effusive rocks. The present

paper deals exclusively with rocks possessing the granitic texture. Of

these the author has investigated over 200. He has recalculated the

proportions of each of the analyses and has plotted them in a series

of projections. A final projection contains all the types and thus

serves as a summary of the work. Among the rock analyses studied,

40 are of granites, 36 of syenites, 37 of diorites, 28 of eleolite-sye-

nite, 27 of gabbros, 23 of essexites and theralites, 4 of iolites and

other rare basic rocks, and 11 of peridotites.

Moses and Parsons, Elements of Mineralogy, Crystallography,

and Blowpipe Analysis, etc. — The new edition of Moses and

Parsons’s Elements of Mineralogy covers concisely the greater part

of the field of mineralogy. As its title indicates, it not only treats

Moses, A. J., and Parsons, C. L. Elements of Mineralogy, So

and Blowpipe Analysis, from a practical standpoint, including a description

240 THE AMERICAN NATURALIST. [Vor. XXXV.

of the descriptive portion of the science, but it is also a very brief

but quite satisfactory treatise on blowpipe analysis. It contains

also a discussion of crystallography and a summary of the principles

of physical mineralogy. That the authors have attempted to cover

too much ground in the volume might be judged from this summary

of its contents. Everything discussed, however, is so concisely

put that the parts of the book are fairly well proportioned. The

chapters on the optical, thermal, and electrical properties of minerals

are so brief that they possess little value. But since these proper-

ties are those of least importance to a class of students beginning

the study of the science, perhaps this fault is not of practical

moment.

In the opinion of the writer the discussions are throughout the

book so compactly condensed that it cannot be used successfully as

a text-book. "There are too many points in it that need amplifica-

tion. As an accompaniment to a course of lectures on mineralogy,

however, it seems to be very well suited. Indeed, it is an excellent

book for use in this way. The volume is not as large as Dana’s

Text-Book of Mineralogy, and therefore is better suited to courses

extending through half a year; and yet, at the same time, it is not

as small in size nor as elementary in the treatment of its subject-

matter as are most of the Elementary Mineralogies on the market,

most of which are entirely. without value for class-room purposes.

The changes noted in the new edition as compared with the old

one are: the treatment of crystallography according to the newly

accepted classification of crystal forms, the addition of about one

hundred figures illustrating the combination of forms observed on

definite mineral species, the revision of the chapters relating to blow-

pipe analysis, the addition of a description of the spectroscope and

its use, the complete revision of the paragraphs dealing with the

economic uses of the different minerals and the simplification of the

part devoted to determinative mineralogy.

The book is well printed on good paper. The figures illustrating

crystals are for the most part beautifully clear, but most of the wood-

cuts supposed to represent the appearance of minerals as they actu-

ally occur in nature are poor. They might well be omitted without

affecting the value of the book a mite. W. S. B.

common or useful minerals, the tests necessary for their identification, the recog-

nition and measurement of their crystals, and a concise statement of their uses in

the arts. New enlarged edition. New York, D. van Nostrand Company, 1900.

413 pp. 664 figs.

No.411.] REVIEWS OF RECENT LITERATURE. 241

Kemp’s Handbook of Rocks, for use without the Microscope,

Second Edition.! — The publication of a new edition of the Hand-

book of Rocks so soon after the appearance of the first edition is a

guarantee that the author has furnished a book that has given satis-

faction to its users. It is the only treatise in English that aims to

give the student an accurate and scientific knowledge of rock-masses

without requiring him to master the technique of microscopical

analysis. The new edition differs from the earlier edition principally

in the glossary, which has been extended to include the new terms

proposed by petrographers in the discussions of the past four years.

The volume is of handy size and is beautifully printed on excellent

paper.

Hovelacque’s Photographs of Sedimentary Rocks.? — This album

consists of sixty-nine plates, presenting microphotographs of thin

sections of alpine limestones. These photographs were made in the

course of a full investigation of the so-called “Calcaires du Brian-

connais," in the French Alps, by Messrs. Killian and Hovelacque,

who hoped by means of microscopic study to be able to determine

more accurately the various horizons of the series in the absence of

available fossils. The work was interrupted by the untimely death

of Mr. Hovelacque, and this album is a preliminary report present-

ing his uncompleted work.

The plates represent a great variety of limestone structures, as

well as many types of the minute organisms which compose, in large

part, many of the limestones. Full explanations make each plate

intelligible, and together they make a valuable addition to our knowl-

edge of the microscopic petrography of the sedimentary rocks whose

value will, however, be much enhanced when the fuller report, which

is promised by M. Killian, makes its appearance.

1 Kemp, J. F. A Handbook of Rocks, for use without the Microscope. With a

— af the names of rocks and of other lithological terms. Second edition,

revised. New York, D. van Nostrand Company, 1900. 185 pp.

* Killian, W. Album de Microphotographies de Roches Sédimentaires faites par

Maurice Hovelacque. Paris, Gauthier-Villars, 1900. 14 pp., quarto, 69 plates.

PUBLICATIONS RECEIVED.

(Regular exchanges are not included.)

DENNERT, E. Plant Life and Structure. Translated from the German by

Clara L. Skeat. The Temple Primer dein ON J. M. Dent & Co, igoo.

viii, 115 pp., 12mo, 56 figs. 40 cents.— MIA and HAMMOND, A. R. The

Structure and Life History of the Hlegihi: Fi odii Oxford, Claren-

don Press, 1900. 204 pp. ré 36 figs. $1.90. — SCHENCK, F., and GÜR yak

Outlines of Human Physiology. Authorized [translation from the Mis pu

man Edition by Wm. D. Zoethout, with a Preface by Jacques Loeb. New York,

‘Henry Holt & Co., 1900. viii, 339 pp. 8vo, 49 figs. $1.75. — SCHNEIDER, A.

The Limitations a Yeahuüg mi Se EEN Papers. Chicago, Medical Book

a 1900. IOO pp.,

NKS, N. Papers from the Harriman Alaska pressing XI, Entomo-

oil Results ; ( di Arachnida. Proc. Wash. Acad. Sci. Vol. . 477-486,

XIX.— BANKS, N. Papers from the Harriman Alaska Egpadinan. X,

E Radia. (4) Nemopteroid Insects. Proc. Wash. Acad. Sci. ol.

ii, pp. 465-476, Pls. XXVII, XXVIII. E enses A. A. Special Instruction

in Poultry Culture. “R. I. Agr. Exp. Sta, Bull. No. 72. 36 pp. pls. — CAD-

Papers from the arean Alaska Expedition. XV, Entomo-

logical Results; (9) irs era. Proc. Wash. Acad. Sci. Vol. ii, pp. 511, 512.

— COCKERELL, T. D. A. Observations on Insects. New Mex. Agr. Exp. Sta.,

Bull. No. 35. 2 ., IO figs. ee W. P. House Flies. sig Mex.

Normal Univ. Nature Study, Bul. No. r. 2 pp. — Co ETT, D. W. Papers

from the Harriman Alaska kiaia IX, Mücalogióut Reus € Diptera

Proc. Wash. Acad. Sci. bie us ap bestia — COVILLE, F. V. s Mes cil

rium, an Undescribed C w Mexico and Texas. Proc pes esi oc. Wash.

Vol. xiii, pp. 195-198. — DALL, W. Ps Synopsis of the Family Tellinida and of

the North American Species. Proc. U. S. Nat. Mus. Vol. xxiii, pp. 285-326. —

Dati, W. H. Contributions to the Tertiary Fauna of Florida with Especial

Reference to the Silex Beds of Tampa and the Pliocene Beds of the Caloosa-

hatchie River, etc. Pt. v, Teleodesmacea: Solen to Diplodonta. Zrans. Wagner

Free Inst. Vol. m, Pt v, pp. 950-1218, Pls. XXXVI-XLVII. — Dyar, H

Papers from the Harriman Alaska Saepeditton. XII, Entomological Résülts

(6) Lepidoptera. Proc. Wash. Acad. Sci. Wol. ii, i, pp. 487-501. Haia J. W.

A Theatrical Performance at Walpi. Proc. Wash. Acad. Sci. Vol. ii p. 605-

629, Pls. XXXII-XXXIV. — Fraps, J. S., and BizzELt. Methods H Daihen

mining Proteid Nitrogen in Vegetable pane N. C. Agr. Exp. Sta., Budi.

No. 174. Pp. 95-104. — HEIDEMANN, O. Papers from the Harriman Alaska

Expedition. XIII, Entomological Results; (7) Heteroptera. Proc. Wash. Acad.

Sci. Vol. ii, pp. 503-506. — HERRICK, C. = and JOHNSON, D. W. The Geology

of the Abano Sheet. Bull. Sci. Lab. Denison Univ. Vol. xi, pp. 173-239;

Pis. XAVU-XLVITE — Hopkins, A. D. The Periodical Cicada or Seventeen-

PUBLICATIONS RECEIVED. 243

Year Locust in West Virginia. W. Va. Agr. Exp. Sta., Bull. No. 68. Pp. 257-

330. Plates and Maps. — Hopxins, A. D. Report on Examination of Wheat

Stubble from Different Sections of the State. Supplement to Bulletin 67: The

Hessian Fly in West Virginia. W. Va. Agr. Exp. Sta., Bull. No. 69. Pp. 333,

344. — Hopkins, A. D. The Joint Worm in Wheat. W. Va. Agr. Exp. Sta.,

Bull. No. 69. Pp. 345-350, 1 plate. — Howarp, L. O. A Contribution to the

Study of the Insect Fauna of Human Excrement. With Especial Reference to

the Spread of Typhoid a e by Flies. Proc. Wash. Acad. Sci. Vol. ii, pp.

541—604, Pls. XXX-XXXI, Figs. 17-38. — JORDAN, D. S, and SNYDER, J.O. A

List of Fishes Collected in deis by Keinosuke Otaki and by the U. S. Steamer

Albatross, with Descriptions of vides New Species. Proc. U. S. Nat. Mus.

Vol. xxiii, pp. 335-380. — KINCAID, T. ers from the Harriman Alaska Expe-

dition. XIV, Entomological Rast e The Sphegoidea and Vespoidea. Proc.

Wash. Acad. Sci. Vol. ii, pp. 507-510. — LAMBE, L. M. Sponges from the Coasts

of Northeastern Canada and Greenland. Trans. Roy. Soc. Can., 2d ser. Vol. vi,

sect. iv, pp. 19-48, Pls. I-VI. — LAMBE, L. M. Catalogue of the Recent Marine

Sponges of Canada and Alaska. Ottawa Naturalist. Vol. XIV, No. 9, pp. 153-

172,1900. Dall, W. H. Synopsis of the Family Cardiida and of the North Ameri-

can Species. Proc. U. S. Nat. Mus. Vol. xxiii, pp. Pob gis URG, C. E.

The aep ic Divisions of the Acridide. Z4. Uni: E No. 1;

Pp. 73-100, Pls. XV-XVII. — McGrecor, R. C. New y jg js Con-

dor. Vol. iii, No. r, 1 page.— Maxow, W. R. Polypodium Hesperium, a New

Fern from Western North America. Proc. Biol. Soc. Wash. Vol. xiii, pp. 199,

200.— MERRIAM, C. H. Preliminary Revision of the North American Red Foxes.

Proc. Wash. Acad. Sci. Vol. ii, pp. 661-676, Pls. XXXVI, XXXVII.— MILLER,

G. S., Jr. Mammals collected by Dr. W. L. Abbott on Pulo Lankawi and the

Butang Islands. Proc. Biol. Soc. Wash. Vol. xiii, pp. 187-193. — MILLER, G.

S., Jr. A New Mouse Deer from Lower Siam. Proc. Biol. Soc. Wash. Vol.

xiii, pp. 185, 186. — MILLER, G. S., JR. A Collection a Small Mammals from

Mt. Coffee, Liberia. Proc. Wash. Acad. Sci. Vol. ii, pp. 631—649, Figs. 39-43. —

OsTERHOUT, W. LV. meres bei Batrachosperma, Flora, Bd. LX XX VII.

Heft 1, pp. 109-11 5, Taf. v. — PERGANDE, T. Papers from the Harriman Alaska

Expedition. XVI, BME gone Results: (10) Aphidæ. Proc. Wash. Acad. Sci.

ol. ii, pp. 513-517. — PERGANDE, T. Papers from the Harriman Alaska Expedi-

tion. XVII, Marie Results; (11) Formicide. Proc. Wash. Acad. Sci.

Vol. ii, pp. 519-521. — PIERCE, N. B. Peach Leaf Curl: Its Nature and Treatment.

S. Dept. Agr., Div. Vegt. Physiol. and Path., Bull. Mo. 20. 204 pp., 30 plates.

QUAINTANCE, A. L. The Brown un of Peaches, duque and Other Fruits.

Georgia Agr. Exp. Sta, Bull. Vo. 5o. Pp. 237-269, Plates. — SHuFELDT, R. W.

A Remarkable Growdr on the Bill ui a Caen iei arquatus). Ornis,

1900. Pp. 477-479, 1 fig. — SCHWARZ, E. A. Papers from the Harriman Alaska

Expedition, ue une eee ep (1 te Psyllide. Proc. Wash. Acad.

P- 539, 540. — SCHWARZ, E. A pers from the Harriman Alaska

Essi. UL A deed. ta a Proc. Wash.

d. Sci. Vol. ii, pp. 523-537. — STEARNS, R. E. C. Fossil Land Shells of the

j dia Day Region, with Notes on Related Living Species. Proc. pie Acad. Sci.

Vol. ii, pp. 651-660, PI. XXXV.— ipea B.B. A New Head-Rest for the

Removal of the Human Brain. y soc. Amer. Anatomists, 13th Session,

4 pP» 2 figs.— STROUD, B. B. If an “ ME Y niencephal, " why not an

244 THE AMERICAN NATURALIST.

“Isthmus Pursencephali,”? Proc. Amer. Assoc. Anatomists, 17th Session, pp. 2

29, 2 figs. — SUTTON, W. S. The — Divisions in Brac Hafen

Magna. Bull. Univ. Kan. Vol. ix, No. 2, pp. 135-160, Pls. XXXII-XXXV.

— WEINGIRL, J. The Bacterial fion of die Semi-Desert Region of New Mexico,

with Especial Reference to the Bacteria of the Air. Journ. Cincinnati Soc. Nat.

Hist. Vol. xix, No. 7, pp. 211-242. — WILDER, B. G. Revised Interpretation of

the Central Tissues of ` ona Suicides’ Brain Exhibited to the Association.

Journ. Nerve and Ment iseasé. October, 1900, 5 pp.

Actes de la Société puros du Chili, Tome x, Livr. 2. — American Museum

Journal, The. Vol. i, No.4. November, 19co. “Anales del Museo Nacional de

Montevideo. Tom. ii, Fasc. xv, xvi. 1900.— Bulletin Johns Hopkins Hospital.

Vol. xi, No. 117. December, 1900. — Geologischer Core iip a que für Geo-

logie, Petrographie, ipea und verwandte Wissenchaften. Herausgegeben von

K. Keilhack. Bd. i, No. January. — Z»sect. World, The. Vol. iv, No. 11.

November, 1900. — iban Marine Biological Station. Communications, I.

e. ix o i

October and November, 1900. — O. S. U. BERSE Vol. i, No. 2. December,

1900. — Revista Chilena de Historia Natural. v, Nos. 9-11. September-

November, r9oo. Rhode Island Aikaa ‘cet Station. Thirteenth

Annual Report, ix +168 — Science Gossip. New Series. Vol. vii, No. 8o.

Pp- —

January. — U. S. Commissioner of Education. Report for the Year 1898-99.

Vol. i, 1248 pp.

No. 410 was mailed February r2.

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VoL. XXXV, NO. 412 APRIL, 1901 —

THE

AMERICAN =

NATURALIST =]

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES 22

IN THEIR WIDEST SENSE ;

CONTENTS

E-A Theory of the c and Evolution of the Australian ;

Marsupialia . B, ARTHUR seNsIEY E

II. A creme study of Variation in ‘the smaller North- PEE ED

IIL A New bri for PR Te. eee Dr, aak quic

IV. Synopses of North-American Invertebrates. E Heo

—— hes

T e. v. tue 2

mies Publi ul i

Life, Notes — uisi. Traquair's | Presidential | Address, Welais

Fossils from Eastern Siberia, Tht DUM. Paleozoic c Fauna of Pium

ly R s, and De

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AMERICAN NATURALIST

Vor. XXXV. April, rgor. No. 412.

A THEORY OF THE ORIGIN AND EVOLUTION OF

THE AUSTRALIAN MARSUPIALIA.

B. ARTHUR BENSLEY.

INTRODUCTION.

ALTHOUGH considerable attention has been given to the

discussion of the family and generic affinities of the Aus-

tralian marsupials from a systematic standpoint, until very

recently the phylogenetic aspects of the question have been

generally neglected. On this account, and also from the

fact that little information has been forthcoming either from

paleontology or geographical distribution, it has not been

possible, up to the present time, to form an adequate con-

ception of the origin of the fauna and its phylogenetic

relationships.

The present paper is the result of an attempt to outline a

plan of evolution for the group by comparing the adaptive

modifications of the teeth and feet with those of placental mam-

mals, in which the course of evolution is fairly well known.

On account of the difficulties with which the writer has had to

contend in having only a small number of marsupial types at

his disposal, the present plan is advanced simply as a working

245

246 THE AMERICAN NATURALIST. [Vor. XXXV.

hypothesis. A fuller discussion of the question will be pub-

lished in the course of a few months.

. The fact was noticed long ago by Cuvier, and has been com-

mented upon by many zoólogists since, that the marsupials of

Australia assume much the same róles in nature as are taken by

placentals in other parts of the world, and that they show exten-

sive resemblances to the latter in many features of their external

organization. It is only within the last few years, however, in

fact since the evolution of the placentals has come to be under-

stood, that the exact significance of these resemblances has

been at all apparent. Knowing that during the Tertiary period

placental mammals have undergone a progressive development,

or, in other words, an adaptive radiation, from diffuse or collect-

ive types, by which not only their minor but also their ordinal

characters have been established, the question now suggests

itself, Have not the Australian marsupials undergone an entirely

similar or, in other words, a parallel radiation ?

The possibility of such a condition has already been sug-

gested by Osborn (99a) who recognizes several mammalian

radiations, including among them a marsupial radiation for

Australia, and remarks as follows: * We mark the fact that

the above radiations are all of ordinal rank, for the marsupial

radii, although termed families, are adaptively equivalent to

several placental orders.”

Examining the composition of the group somewhat in detail,

we find many indications of such a parallel radiation. Two

classes of facts may be specially noticed in this connection.

First, the Australian marsupials constitute a very homogeneous

group; although differing widely in extremes, not only the

various genera of a family but also the families themselves are

connected by almost insensible gradations of structure. Second,

the adaptive modifications of their teeth and feet are very

similar to those of placental mammals either at the present

day or during their progressive development in the Tertiary

period, and they bear the same relation as do those of placen-

tals to the general primitiveness of the animals in which they

occur.

The Australian marsupials, therefore, show no signs of a

No. 412.] THE AUSTRALIAN MARSUPIALIA. 247

composite structure such as might be expected if they repre-

sented migrated portions of the fauna of another continent, or,

again, if they were of polyphyletic origin. Except in a very

few instances, they show no signs of degeneration; as a group

they are, like the placentals, progressive throughout. In short,

the only satisfactory explanation that can be given for the

homogeneous character of the group and their adaptive resem-

blances to placentals seems to be that they have undergone an

entirely similar radiation under similar conditions, that it has

proceeded from a single sharply marked ancestral type, and that

the center of radiation has been the Australian region itself.

Assuming that a similar radiation has taken place in mar-

supials and placentals, it will be apparent that a plan of evolu-

tion may be constructed for the former by inference from that

of the latter. Thus, by noting the sequence of events in the

development of the adaptive modifications of the teeth and feet

in placentals during the Tertiary period, we may infer a similar

sequence in those of marsupials, and in this way determine

their probable course of evolution in these structures, and also

the characters of their stem form.

THE PROBABLE CHARACTERS OF THE MARSUPIAL STEM Form.

Dentition. — In the case of the placental mammals, progres-

sive evolution in dentition is accompanied by a reduction of

certain of the teeth and by an elaboration of certain others,

notably the molars. Unreduced dentitions with slightly elabo-

rated teeth are therefore primitive. But just asin formulating

such a law it is necessary to exclude those forms whose denti-

tion, while being of primitive aspect, is not typically primitive,

but primitive through degenerate specialization (Delphinide,

Otariidze), it is also necessary in applying it to the case of

the marsupials to exclude similarly conditioned forms of that

series,

The only marsupials which call for consideration in this

respect are the forms Myrmecobius and Notoryctes.! There

1 Tarsipes appears to be degenerate in dentition, but the teeth are not primi-

tive in aspect.

248 THE AMERICAN NATURALIST: [VoL. XXXV.

are two reasons for regarding these as degenerate in dentition.

First, there is a prototypal tooth pattern in marsupials, which

is shared alike by the Didelphyida and the insectivorous mem-

bers of the Dasyuridze, and of which the tooth patterns of the

carnivorous Dasyuride and the Peramelide are undoubtedly

progressive modifications, while those of Myrmecobius and

Notoryctes are aberrant. The tooth patterns of both of these

forms are more primitive in aspect than those of any other of

the Australian forms, but the animals are specialized in other

respects. Thus Myrmecobius is typically dasyurid in foot

structure and in its incisor formula of 4, while Notoryctes,

according to Dollo (99), is specialized in a phalangerine direc-

tion in foot structure. Second, both Myrmecobius and Noto-

ryctes are ant-eating forms,! such animals being usually

degenerate in dentition, judging from Echidna and the placen-

tal Myrmecophagide and Manidz, in which the teeth have

been entirely lost.

Excluding Myrmecobius and Notoryctes, we find the primi-

tive dental characters of marsupials distributed as follows :

The least reduced dentition is found in the Peramelide,

where the formula is: 7.8, ¢.4, 5.8, m.42

The least elaborated molar teeth are found in the insec-

tivorous members of the Dasyuride. The upper teeth (Pl. I,

Fig. d)? are triangular in shape, with three main cusps (proto-

cone, paracone, metacone) * and an outer row of styles. There

are no conules at the base of the protocone, such as are com-

monly met with in placental types. The metacone exceeds the

paracone in size, and its posterior border is produced into a

trenchant spur. The lower teeth (Pl. II, Fig. d) are of the

! For habits of Notoryctes see Stirling ('91, p. 158) and Spencer (96, p. 50,

Mammalia).

2 Or (m)i.3, mce4, mp. and 72.$, m.3, we Lydekker (99). The upper

incisor bey may contain teeth of both seri

is type might be more appropriately donaei by the teeth of Phascolo-

cis khi pic. or Antechinomys, Dasyurus viverrinus being more carnivorous

abit. Thomas, however, remarks that the teeth of the three former genera

worn one another very closely, and differ only from those of Dasyurus in being

more wp d cuspidate. See Thomas ('88, pp. 27 3» 298, 309).

* Following the nomenclature of the tritubercular theory. See Osborn

(88, '91)

No. 412.] THE AUSTRALIAN MARSUPIALIA. 249

tubercular sectorial pattern, with a main triangle bearing three

cusps (protoconid, paraconid, metaconid) and an antero-external

shelf, and a posterior heel bearing one outer (hypoconid) and

two inner (entoconid and hypoconulid) cusps.

Foot Structure) — In placental mammals, progressive evolu-

tion in foot structure, beautifully illustrated in Tertiary Ungu-

lata, is accompanied by a reduction of certain of the digits,

and by an elaboration of certain others, the evolution proceeding

from an ideal pentadactyl condition, such as is seen typically

in the Eocene Creodonta. Among the Australian marsupials

the nearest approach to the ideal pentadactyl type is seen

in certain members of the Dasyuridz (Pl. III, Fig. d). The

hallux is opposable but slightly reduced, while the remaining

digits are normal.

Summing up the above results, we may make the SEEN

statement that the marsupial stem form possessed the fol-

dag characters, or more primitive ones: Dental formula,

1.5, c.l, 9.8, m. 4. Upper molar teeth triangular, with three main

biis and an outer row of styles; metacone exceeding para-

cone, its posterior border produced into a trenchant spur ; no

conules at the base of the protocone. Lower teeth with ante-

rior triangle bearing three cusps and an antero-external shelf,

and posterior heel bearing one outer and two inner cusps. Foot

with hallux completely opposable, the remaining digits normal.

The fact will now be apparent that the above prototypal

characters are not found associated in any Australian form.

All of the Australian marsupials are derived types. On the

other hand, with one exception they are all found in the

American Didelphyide. The one exception refers to the pres-

ence in the latter of four instead of three lower incisors, and

in this respect the Didelphyidz are more primitive. The molar

tooth patterns of the stem form are almost exactly reproduced

in those of the Oligocene Opossums (Ferstheriums (Pls. I and II,

Fig. a)? |

! This refers dish to the pes in the case of the marsupials.

? In the specimen from which these diagrams were taken (Pirineo fugax

Cope) the outer parts of the upper molars are worn, so that, although there is a

Conspicuous external ridge, the presence of styles cannot be affirmed. Cf Cope

(84, Pl. LXII, Fig. 2a).

250 THE AMERICAN NATURALIST. [Vor. XXXV.

As regards the modifications of the teeth and feet, there-

fore, the Didelphyidz exhibit characters which are ancestral

to those of the Australian marsupials. The question now

arises as to whether or not they are ancestral in other respects.

The opportunity has not yet presented itself to the writer of

examining the case in detail, so that only a few examples will

be given here. As was pointed out in 1880 by Brass, the com-

plex modifications which result in the formation of the peculiar

median vagina in specialized marsupials are seen in their incip-

ient stages in the Didelphyide, where the two vaginz bend

toward one another in the middle line. Passing onwards from

the Didelphyidz every gradation is met with, until the final

stage is reached in the Macropodide,

Again, taking the case of the marsupium, the anteriorly

directed opening of this structure, as seen in the Phalangeridz

and Macropodidz, and the posteriorly directed opening, as seen

in the Peramelidae and Notoryctida, are obvious departures

from such a type as is seen in the Didelphyidze and Dasyuride,

in which the opening is directed vertically downwards.

Finally, taking the case of the tail, which is an adaptive

structure of the greatest importance in marsupials, it is entirely

probable that the hairy, non-prehensile condition, where it

occurs in the Australian forms, has been derived from a naked,

scaly, prehensile one, such as is seen in the Didelphyidz.

Thomas (88) describes for the Dasyuridz, in which the tail is

hairy and never prehensile, a scaly condition in various genera,

among them Phascologale wallacei, P. flavipes, and Sminthopsis

leucopus. A scaly tail, which, according to Thomas, is said to

be also prehensile, is found in Perameles broadbenti among the

Peramelida. Every grade of prehensilism is met with among

the Phalangeridz ; and, finally, among the Macropodide, a scaly

tail is found in Hypsiprymnodon moschatus, and a prehensile

one in the various species of Bettongia.

The above facts, when added to the evidence of the teeth

and feet, suggest strongly a former connection of the Didel-

phyide with the Australian marsupials, and although, in

attempting to express such a connection phylogenetically, it

is a natural procedure to regard the original members of each

No. 412.] IHE AUSTRALIAN MARSUPIALIA. 25I

group as the descendants of a common stock, the Didelphyidze

are so primitive that it is scarcely possible to regard them, as

a family, otherwise than ancestral.!

THE EVOLUTION OF THE TEETH IN AUSTRALIAN MARSUPIALS.

It has already been mentioned that progressive evolution in

dentition, in the case of the placental mammals, is accompanied

by a reduction of certain of the teeth and by an elaboration of

certain others, notably the molars. The researches of Cope,

Osborn, and others have shown that in the evolution of the

molar teeth the elaboration proceeds from a trituberculate con-

dition in the upper series, and from a quinquetuberculate or

sexituberculate condition in the lower series. These types are

exemplified in the teeth of the primitive Creodonta of the early

Eocene (Protochriacus, Oxyclzenus, etc.).

1 This involves the question of the limitations of the family Didelphyide.

Lydekker ('96, p. 109), while assigning the didelphyid incisor formula to the

ancestors of the oe pin (p. 55) of the Didelphyide and Dasyuride

as descendants of a comm ock. Spencer ('96, p. 188, Summary) speaks of

on the one hand, and to the early Australian t on the other. agito

(98, p. 16) regards the bai called by him rr uti as ancestral to the

Didelphyide and to the tralian Dasyuride. In the writer's opinion it will

not be found advisable Shans to regard the Didelphyidz as a modern derived

group, but that, when the actual ancestors of the Australian forms are identified,

it will probably be found necessary to extend the family Didelphyidz to include

the : f

about the time when the Australian fauna probably arose. Second, d

ily Didelphyidz, as at present limited, is not equivalent to any one of the four

Rinit families of Australian marsupials, the latter being much more comprehen-

sive. The diversity of organization which is met with among the Dasyuride in

the inii Phascologale and Thylacinus, eue the Phalangeride in the genera

Phalanger, Phascolarctos, and Tarsipes, among the Macropodide in the genera

Hypsiprymnodon and Macropus, is absolutely phi parallel among the Didel-

phyide. If, therefore, the integrity of the Australian families is to be retained, —

and there is every reason for r believing that they are natural groups, —there are

good grounds for extending the family Didelphyidz to include their own ances-

tors and those of the Australian forms, if the latter, as they probably did, pos-

Sessed the main didelphyid modifications of the teeth and feet, even while

differing in Prensa minor details. The Microbiotheriidae of Ameghino may be a

case in poi

252 THE AMERICAN NATURALIST. [Vor. XXXV.

From this point onwards development takes place along two

main lines. In one case the outer cusps of the teeth from

being tuberculate become laterally compressed and trenchant,

giving rise to the carnivorous modification, seen typically in

the specialized Creodonta (Oxyzna, Hyzenodon, Oxyznodon,

etc.). In the other case the primitive tooth pattern is entirely

changed; the upper teeth become quadritubercular by the

addition of a postero-internal cusp (hypocone), while the lower

teeth also become quadritubercular by the reduction of the

most anteriorly placed cusp (paraconid), and the intermediate

cusp of the heel (hypoconulid). In this way a primarily omniv-

orous modification is instituted, which is seen in its construc-

tive stages in the Eocene Condylarthra (series Protogonodon,

Euprotogonia, Phenacodus, and various separate genera).

From the typically quadrituberculate condition evolution

proceeds in two directions, both leading to the herbivorous

modifications of the Ungulata. In one series the cusps of

both upper and lower teeth become crescentic, producing the

condition characteristic of the Artiodactyla ; while in the other

they become lophoid, producing the complex modifications of

the Perissodactyla. The latter evolution is beautifully illus-

trated in passing from the Eocene genera (Hyracotherium,

Systemodon, etc.) to the modern types.

If, now, we arrange the teeth of the Australian marsupials

according to the above placental plan, we obtain such a result

as is represented diagrammatically in Pls. I and II. The primi-

tive trituberculate, tubercular sectorial type is here represented

by the teeth of Dasyurus viverrinus (Fig. d). As indicated

elsewhere in a footnote, this would be more appropriately

illustrated by the teeth of one of the purely insectivorous forms

of the Dasyuridae (Sminthopsis, Antechinomys, Phascologale).

The carnivorous line above mentioned is, in the case of the

Australian marsupials, entirely confined to the family Dasyu-

ride, and it culminates in the Tasmanian wolf (Thylacinus cyno-

cephalus). The teeth of this animal are represented in Pls.I

and II, Fig. ¢; they show all the essential characters of those

of Dasyurus, except that in the lower teeth the metaconid is

absent. The progressive carnivorous modification is, in the

No.42] THE AUSTRALIAN MARSUPIALIA. 253

upper molars, a conversion of the paracone and metacone into

cutting blades, and a reduction of the outer styles, which are

here represented by very inconspicuous tubercles. In thelower

teeth the progressive modification is a conversion of the para-

conid, protoconid, and hypoconid into cutting blades, and a

reduction of the metaconid, hypoconulid, and entoconid.

Turning to the second stem, leading to the omnivorous and

herbivorous modifications, we find the first stage represented in

the teeth of the Peramelide. The upper molars of Perameles

nasuta (Pl. I, Fig. e) show all the essential characters of those

of Dasyurus, but in addition they show an incipient hypocone.

In the third molar the hypocone is not very pronounced, and

the tooth is triangular; but in the second molar the hypocone

is well developed, and the tooth is now quadrate. Thomas

(88, p. 220) describes the triangular and quadrate modifica-

tions as characteristic of the family. The upper molars of the

Peramelidz parallel, in a general way, those of the Condylar-

thra above mentioned. The lower molars of Perameles nasuta

(Pl. II, Fig. e) resemble still more closely those of Dasyurus,

the omnivorous modification being only apparent in the posterior

heel, where the hypoconulid is reduced.

The completed omnivorous: modification is doubtless to be

met with in the Phalangerida. Unfortunately, only two exam-

ples of this family have been available to the writer, namely,

Trichosurus vulpecula, which is highly specialized, and Petau-

roides volans. The teeth of the latter animal are represented

in Pls. I and II, Fig. f. Although they illustrate the com-

pleted quadrituberculate condition, they are only approximately

transitional ; the hypocone is completely formed in the upper

molars, and the paraconid is reduced in the lower molars; but

the antero-external shelf, which is prominent in the lower teeth

of the Peramelidee and Dasyuride and‘ very conspicuous in

those of the Macropodidze (Pl. II, Fig. ï), is here absent, and

there are no traces of external styles in the upper teeth.

The selenoid modification of the cusps, which is so widely

represented in the artiodactyl Ungulata, appears to be confined

to the single form Phascolarctos (Pls. I and II, Fig. /) among the

marsupials, but the lophoid modification is widely represented

254 THE AMERICAN NATURALIST. [Vor. XXXV.

in the Macropodide (Pls. I and II, Fig. 7). The incipient

stages are seen in the teeth of the specialized Phalanger,

Trichosurus vulpecula (Pls. I and II, Fig. Z), and also in those

of Hypsiprymnodon moschatus (Macropodide). It is of interest

to note that the lophoid modification is accompanied by a

hypsodont condition of the crown in the Macropodide, as in

the perissodactyl Ungulata.

THE EVOLUTION OF THE FEET IN AUSTRALIAN MARSUPIALS.

It was pointed out by Huxley in 1880 that none of the

Australian marsupials possess a normal pentadactyl pes, that

wherever the hallux is present it presents an extensive move-

ment of adduction and abduction; in other words, that the

marsupial foot is typically prehensile.

Quite recently a careful study of the foot structure of mar-

supials has been made by Dollo (99), who endeavors to demon-

strate a derivation of all recent forms from arboreal ancestors.

Dollo has indicated all the important adaptive modifications in

Australian marsupials, although he has not discussed their

phylogenetic significance, so that all that is necessary in the

present paper is to arrange his types into a phylogenetic series,

in accordance with his interpretations and the general plan

noted above for the placentals.

Dollo has shown that the arboreal marsupials present suc-

cessive stages of arboreal adaptation, following which the foot

becomes more and more modified. Thus, beginning with the

Didelphyidz (Pl. III, Fig. æ), the foot is provided with an

opposable hallux, and the remaining digits are normal. In the

Phalangeride (Fig. 5) the hallux is again opposable, but in

addition the fourth digit is enlarged, and the second and third

are reduced and syndactylous. Finally, in one member of the

Phalangeridz, Tarsipes (Fig. c), there is, in addition to all of

these arboreal Characters, a recession of the claws of the fourth

and fifth digits. It is thus possible to recognize an arboreal

line of evolution such as is represented in Pl. III, sc.

Among the terrestrial or semi

-terrestrial marsupials, all of

the forms,

except the specialized members of the Dasyuridze,

No. 412. THE AUSTRALIAN MARSUPIALIA. 2

show some of these arboreal characters, indicating that up to

a certain stage they have been arboreal in habit. It is thus a

simple matter to arrange them in collateral groups according

to the stage at which their terrestrial evolution began. Such

an arrangement as this is represented diagrammatically in

Pl. III, in the side lines passing off from the main arboreal

line a—. The first series to be given off is that of the Dasyu-

ride. In the least modified members of this family the animals

are still arboreal, and the foot (Fig. 7) is like that of Didel-

phys (Fig. a), except that the hallux is slightly reduced. As

Dollo has pointed out, it is possible to recognize a complete

series here, following which the hallux becomes gradually

reduced and finally obliterated (e).

Two other terrestrial offshoots, which parallel one another,

and arise from the main arboreal line at the phalangerine stage,

are represented by the Peramelidze and Macropodide. In

both of these the evolution is cursorial as well as terrestrial,

and is accompanied by a reduction of the hallux and an enor-

mous enlargement of the fourth digit, so that the foot becomes

functionally monodactyl. The extremes of structure in these

lines are, as Dollo has indicated, in the Peramelidae Perameles

doreyana (Fig. g) and Peragale leucura (Fig. h) and in the Mac-

ropodida Hypsiprymnodon moschatus (Fig. 7) and Macropus

(Fig. 7).

Another terrestrial offshoot is represented by the Phasco-

lomyide (Fig. f) although the relations of this branch are

doubtful. The hallux is reduced in these animals as in other

terrestrial forms. With respect to the remaining digits, the

foot appears to be transitional between that of the Didelphy-

idæ and that of the Phalangeride, the second and third digits

not being so reduced as in the latter family. The difficulty arises

here as to whether or not the second and third digits, from being

fully reduced, as in the Phalangerida, have been secondarily

enlarged. Such a condition would appear to be not improbable

in slow-moving animals, such as the wombats; in the other

animals which have passed through a phalangerine stage the

tendency towards monodactylism has doubtless prevented the

second arid third digits from becoming secondarily functional.

256 THE AMERICAN NATURALIST. [Vor. XXXV.

For Notoryctes Dollo describes a slight syndactylism and

reduction of the second and third digits, which points out this

form as still another offshoot of the main arboreal line.

GENERAL REMARKS ON THE EVOLUTION OF AUSTRALIAN

MARSUPIALS.

When we compare the plan of evolution as determined by

the structure of the teeth with that determined by the struc-

ture of the feet, we note the fact that they practically coincide.

Thus there are two lines leading from the prototypal forms, one

of them showing an insectivorous-carnivorous and an arboreal-

terrestrial evolution leading to the Dasyuridze, the other show-

Dasyuridz Peramelidze ecu Phalangeridz :

ee rous .

ee orous Primitive Phalang-

rial eride

Myrmecobius `

y Td Phascolomyidze

Y Properamelida(hyp.)

gee tea oon

real-térres strial

Didelphyidze ?

Prototypal forms [iiem

ing an insectivorous-omnivorous and a progressive arboreal

evolution leading to all the remaining marsupial families..

Following the former line, there appears to have been a

typical evolution in all forms except Myrmecobius, which, as

noted above, is probably degenerate in dentition.

Following the omnivorous arboreal line, we find that one

of its stages has been obliterated, being unrepresented among

existing forms. The characters of other forms, however, leave .

No. 412. THE AUSTRALIAN MARSUPIALIA. 2

no doubt as to its composition. Thus the Phalangeridz present

a typical arboreal modification of the foot, combined with

advanced omnivorous and herbivorous modifications of the

teeth, while the Peramelide present a terrestrial modification

of the phalangerine foot, combined with a primitive omniv-

orous condition of the teeth. There must, therefore, have

existed at some time, if the present plan is reliable, a series of

forms such as is represented in the general diagram on p. 256

as Properamelidze (hyp.), consisting of animals which combined

a phalangerine type of foot with a primitive omnivorous denti-

tion. These animals would have given rise to the Peramelidæ

by a terrestrial modification of the foot, and to the Phalanger-

idee by a more completely omnivorous and finally herbivorous

modification of the teeth.

From the phalangerine stem, after the omnivorous modifica-

tion was established, there was apparently given off a branch

to the Macropodide, with a trend of evolution from arboreal

to terrestrial, and from omnivorous to herbivorous. The her-

bivorous modification reaches its greatest development in this

family.

The phylogenetic positions of the Phascolomyidae and Noto-

ryctidae, as indicated in the general plan, are by no means

certain. In the case of the former, the difficulty in interpret-

ing the foot structure has already been commented upon.

The unworn molar teeth of these animals are said by Krefft (75)

to be quadrituberculate. If such is the case, it would appear

probable that they arose from the omnivorous arboreal stem

after the Peramelidze were given off, and that there has been

a secondary enlargement of the second and third digits.

In the case of the Notoryctide the writer has not been able

as yet to decide definitely the cusp homologies in the upper

teeth, but the lower are undoubtedly tritubercular; as indi-

cated above, the dentition is probably degenerate. Combining

this interpretation with the evidence of arboreal characters in

the foot, it would seem probable that Notoryctes arose from the

arboreal omnivorous line somewhere near the Properamelidz.

258 THE AMERICAN NATURALIST. [Vor. XXXV.

A DIDELPHYID ORIGIN IN THE LIGHT OF GEOGRAPHICAL

DISTRIBUTION.

In discussing the question of the origin of the Australian

marsupials it is apparent that, apart from the evidence deriv-

able from the organization of existing types, any opinions

which are expressed on the subject must take into consid-

eration the present and past geographical distribution of

marsupials, the manner in which the ancestral forms gained

access to the Australian region, and the time at which their

entry was effected.

Within the last twenty-five years several attempts have

been made to connect the Australian marsupials with the

extinct types of different horizons in other countries, and to

explain in this way the conditions of their origin. Thus, in

1876, Wallace suggested a possible relation to the Jurassic

forms of the northern hemisphere. Concerning this he

speaks as follows: * As, however, no other form but that of

the Didelphyidz occurs there (in Europe) during the Tertiary

period, we must suppose that it was at a far more remote

epoch that the ancestral forms of all the other marsupials

entered Australia; and the curious little mammals of the

Oólite and Trias offer valuable indications as to the time when

this really took place"? . . . «It was probably far back in

the Secondary period that some portion of the Australian region

was in actual connection with the northern continent; and

became stocked with the ancestral forms of marsupials.’ 3

In 1882 Cope, in commenting upon the ancestry of Thyla-

coleo, placed this form in the family Plagiaulacide of the

northern hemisphere, and regarded the latter as related to the

Australian Macropodidze through a hypothetical ancestor Trito-

modon. There is, however, no valid evidence in favor of this

view.

! Haeckel has recently endeavored to express such a relation phylogenetically

by deriving all of the mo Pr a,

3 Ibid. (vol. i, p. 465)

No. 412.] THE AUSTRALIAN MARSUPIALIA. 259

By far the most complete discussion of the question, from

all points of view, is that given by Lydekker (96) in his Geo-

graphical History of Mammals. Lydekker, while agreeing

with Wallace as to the northern origin of the Australian

fauna, fixes its time of entry at a much later period, namely,

at the beginning of the Eocene, on the basis of a possible con-

nection of the ancestors of the Australian Dasyuridz with

those of the Oligocene Didelphyidz of the northern hemis-

phere. He remarks as follows (p. 55) : * Recent researches

have tended to show that the alliance between the Dasyuridz

and the Didelphyidz is much more intimate than was formerly

supposed to be the case. This being so, it is a fairly safe

assumption that both families are descended from a single

common ancestral stock." . . . “ Not improbably polyproto-

dont marsupials survived in southeastern Asia till the early

portion of the Eocene division of the Tertiary epoch, and

in this region both Dasyurida and Didelphyide were differ-

entiated. Representatives of the former family soon after-

wards found their way into Australia and New Guinea, while

the opossums would appear to have dispersed in one direction

into Europe and in the other into North America."

Objection has been taken to this view by Spencer (96) on

account of the difficulty of explaining by it the non-appearance

of Didelphyida in Australia and the paucity of polyprotodont

types in New Guinea, and also on account of the lack of

evidence of the former presence of marsupials in Asia.

For a northern origin Spencer substitutes a South-American

one taking place in the late Cretaceous through the medium of

an Antarctic continent. This idea is also favored by P. L. and

W. L. Sclater (99). Viewing the question from a phyloge-

netic standpoint alone, Lydekker has recently (99) suggested

the Prothylacinidze (Sparassodonta of Ameghino) of the South-

American Miocene as possibly ancestral to the Dasyuridz, and

Ameghino (93) regards the family called by him Microbiotherii-

dæ as ancestral to both the Dasyuridze and the Didelphyide.

Spencer, however, in attempting to derive the Australian

fauna, encounters the same difficulty which confronted Lydek-

ker, namely, how to explain the non-appearance of Didelphyidze

260 THE AMERICAN NATURALIST. (VoL. XXXV.

in Australia. Concerning this he remarks as follows (p. 188,

Summary): * The only way in which it seems possible to

account at once for the presence of forms such as Prothy-

lacinus in the Patagonian Tertiary beds, and the absence of

any of the Didelphyidze in Australia, is to suppose that on the

South-American side the connection between the Antarctic

land and what is now Patagonia was lost at a time compara-

tively soon after the early polyprotodonts had passed across,

and during which the Didelphyidz were being developed per-

haps in the more northern part of South America."

In the writer's opinion, the difficulty of explaining the

absence of the Didelphyidae from Australia is only an apparent

one, which is due to the interpretation of this family as a

modern derived group. Reasons have already been given for

the elevation of the family to an ancestral position, and if the

case is susceptible of proof, no other explanation is necessary

to account for their absence from the Australian region than

the assumption that they were formerly present and subse-

quently disappeared. If we imagine the family to have origi-

nally gained access to the region, it is not difficult to conceive

that, in establishing the foundations of an extremely compre-

hensive adaptive radiation, especially under the favorable con-

ditions of absolute freedom from competition, they should

have thrown aside their distinctive didelphyid characters.

And more especially is this conceivable when we realize that

the differences of structure separating the Didelphyidz from

the most primitive of the Australian forms (insectivorous

Dasyuridze) are very slight.!

Concerning the direction from which the ancestral Didel-

phyidz may have entered the Australian region, there is at

least some justification for the view that it was from the north-

ward. Lydekker (96, p. 57) has cailed attention to the fact

that there are marked indications of a faunal affinity of North

America and Asia, which points to an interchange of forms

between the two continents. This suggests a connection of

the Oligocene opossums of Europe with those of North

America, and, as Lydekker has pointed out, there may have

1 Cf. Thomas ('88, p. 31 5}.

No. 412.] THE AUSTRALIAN MARSUPIALIA. 261

been a migration of these animals into both countries from

a common center in Asia ; on the other hand, there may have

been simply a passage of forms via Asia. In either case the

former presence of opossums on the latter continent would

be indicated.

However this may be, the actual evidence at our disposal

is very slight, and it is equally conceivable that ancestral

Didelphyide may have entered by way of Antarctica. This

possibility has been suggested by Osborn (99b), although no

preference is indicated by him for either view. As already

indicated, the time at which the ancestral marsupials gained

access to the Australian region has been estimated by Wallace

as *far back in the Secondary," by Spencer as Cretaceous,

and by Lydekker as early Eocene. It is probable, however,

that these estimates are much too great, and that the time of

entry was considerably later. Comparing the marsupial and

placental radiations with respect to the modifications of the

teeth and feet, we notice the fact that while there is a general

correspondence between the most primitive characters of each

series, there is no such correspondence between the most

specialized ones. The most specialized characters of mar-

supials are decidedly primitive as compared with the most

specialized characters of placentals. To illustrate the case,

while the insectivorous members of the Dasyuride and the

Didelphyidae approach closely the trituberculate pentadactyl

ancestors of the placentals, the Macropodidæ, which are the

most specialized of the marsupials, are much more primi-

tive than the most specialized placentals (perissodactyl

Ungulata) whose evolution they parallel. The lophiodont

teeth of these forms are very similar to those of such per-

sistent primitive types as the Tapiride and their imme-

diate ancestors, having none of the complex characters of

those of the progressive lophiodont forms (Equide and

Rhinocerotidz). The functionally monodactyl, cursorial feet

of the Macropodidz are primitive as compared with the simi-

larly modified feet of the Equide. Further, there is the

remarkable fact that while the marsupial families have the

value of placental orders, it is barely possible to separate them

262 THE AMERICAN NATURALIST. [VoL. XXXV.

sharply as families. In fact, they show such a composition as

must have existed at some stage of the placental radiation.

Now the Macropodidz show no signs of being persistent

primitive types, and there seems to be no valid reason for

assigning to the marsupials a different potential of evolution.

The only explanation which may be given for the above facts

is, therefore, that the Australian radiation has been of com-

paratively short duration. And if the placental radiation

began in the late Cretaceous or in the early Eocene, as now

appears probable, it is unlikely that the marsupial radiation

can have begun until well on into the Tertiary. An ancestral

association with the Oligocene opossums of the North or even

with the Lower Miocene forms of South America is thus not

improbable.

As to whether or not the ancestral forms gained access

to the Australian region by means of a land connection would

appear to be a matter of little consequence if the fauna is of

didelphyid origin. Such minute animals as they must have

been, judging from the Oligocene forms of the northern

hemisphere and the diminutive opossums of South America,

might have gained access to the region by transportation in

various ways.

In conclusion, the writer wishes to acknowledge his indebt-

edness to Professor H. F. Osborn of Columbia University for

many kind suggestions during the preparation of this paper,

and to Professor Allen of the American Museum of Natural

History, New York, Professor W. B. Scott of Princeton Uni-

versity, and Professor R. Ramsay Wright of the University of

Toronto, for the privilege of examining the marsupial collec-

tions of the above-named places.

COLUMBIA UNIVERSITY, NEW YORK.

! This suggests Huxley's statement of twenty years ago: “ And I suspect that

most, if not all, of the Australian forms are of comparatively late origin." (On

the Application of the Laws of Evolution, etc., P. Z. .S. (1880), p. 656).

No. 412.] THE AUSTRALIAN MARSUPIALIA. 263

BIBLIOGRAPHY.

'98 AMEGHINO, F. Les mammifères de Patagonie. Revue Scientifique

(Jan. 7, 1893). pp. 13-17. Paris.

':80 Brass, A. Beiträge zur Kenntniss n weiblichen Urogenitalsystems

der Marsupialen. Inaug. Diss. Leipzi

':82 CoPE, E. D. The Ancestry and eae of Thylacoleo. Amer. Nat.

Vol. xvi, pp. 520-522.

'84 CoPE, E. D. Tertiary Vertebrata. Report of the U.S. Geological

Survey of the Territories. Vol. iii. Washington.

'99 DoLLo, L. Les ancétres des Marsupiaux, étaientils arboricoles?

Miscellanées biologiques. 188-203. Paris.

'95 HAECKEL, E. Katemi Piskor 3. Theil. Berlin.

'80 HuxrEYv, T. H. On the Application of the Laws of Evolution to the

Arrangement of the Vertebrata, and more particularly of the

Mammalia. Proc. Zodl. Soc. pp. 649-662.

"75 KnEFFT, G. Remarks on Professor Owen’s Arrangement of the

Fossil Kangaroos. Ann. Mag. Nat. Hist. Vol. xv, pp. 204-209.

'96 LYDEKKER, R. A Geographical History of Mammals. Cambridge.

'99 LYDEKKER, R. The Dental Formula of the Marsupial and Placental

Carnivora. Proc. Zoól. Soc. London. pp. 922-928

':88 OsBoRN, H. F. The Evolution of Mammalian Molars to and from

the Tritubercular Type. Amer. Nat. pp. 1067-1079.

'91 Osborn, H. F. Fossil Mammals of the Wahsatch and Wind River

Beds. I. Homologies and Nomenclature of the Mammalian Molar

Cusps. Bull. Amer. Mus. Nat. Hist. Vol. v, pp. 84-91.

'99a OsBORN, H. F. The Origin of Mammals. Amer. Journ. Sci. Vol.

vii, pp. 92-96.

'99b Osborn, H. F. Correlation mes Tertiary Mammal Horizons of

Europe and America. An Y. Acad. Sci. Vol. xiii, pp. 1-72.

'99 SCLATER, P. L. and W. L. The APE eei of Mammals. London.

'96 SPENCER, B. Report of the Horn Expedition to Central Australia.

Part I. September, 1896.

91 STIRLING, E. C. Description of a New Genus and Species of

Marsupialia, « Notoryctes typhlops.” Trans. amd Proc. Roy. Soc.

South Australia. Vol. xiv, Part I, pp. 154-187. 1890-1891.

'88 THomas, O. British Museum “Catalogue of the Map and

onotremata." London

76 WALLacE, A. R. The Geogtaphicál Distribution of Animals. 2 vols.

New York

264 THE AMERICAN NATURALIST. . [Vor. XXXV.

EXPLANATION OF PLATE I.

Illustrating the patterns of the upper molar teeth in marsupials and their

probable sc d relations.

Notre. — The prototypal form is atas: represented by Peratherium.

Figs. b and c are inserted for comparison

a.— Peratherium fugax Cope. f. — Petauroides volans 'Thos.

b. — Didelphys azare Temm. g.— Thylacinus cynocephalus Fisch.

— Didelphys virginiana Kerr. h. — Trichosurus vulpecula Jent.

d. — Dasyurus viverrinus Shaw. i. — Macropus sp.

e. — Perameles nasuta Geoff. j-— Phascolarctos cinereus Fisch.

Abbreviations.

Pr. — protocone. Me. — metacone.

a. — paracone. Hy. — bypocone.

St. — external styles.

Fic. b, after Giebel (Odontographie, Taf. XVII, Fig. 10, a).

No. 412.] THE AUSTRALIAN MARSUPIALIA. 265

E po

2 : Pa

Me à

PLATE I.

266 THE AMERICAN NATURALIST. [Vor. XXXV.

EXPLANATION OF PLATE II.

Illustrating the In of the lower molar teeth in marsupials. (Same

references as in Pla

Abbreviations.

Prt, — pa E^. — entoconid.

Pa?, — paraconid. y^. — hypoconid

Met, — debe H1?. — hypoconulid.

a.s. — antero-external shelf,

No.42] THE AUSTRALIAN MARSUPIALIA. 267

PLATE II.

268 THE AMERICAN NATURALIST. | [Vor. XXXV.

EXPLANATION OF PLATE III.

Illustrating the modifications of the pes in Marsupials and their probable

phylogenetic relations

a. — Didelphys nudicaudata Geoff.

e. — Antechinomys laniger Gould.

f. — Phascolomys latifrons Owen.

g. — Perameles doreyana E & Gaim.

h. — Peragale leucura Tho

i. — Hypsiprymnodon dickies Rams.

j. — Macropus. sp.

Figs. a-i after Dollo.

No. 412.] THE AUSTRALIAN MARSUPIALIA. 269

PLATE III.

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L. MARK, DIRECTOR. No. 121.

A QUANTITATIVE STUDY OF VARIATION IN

THE SMALLER NORTH-AMERICAN

SHRIKES.

R. M. STRONG.

CONTENTS. Pace.

I. Introduction 271

II. Material S 273

III. Methods 274

4. General Precautions ..... 274

B. Measurements of Variable Characters 275

1. Linear Measurements EE A E E OE a ete ow? 275

2. Curvature of s 45275

3. Color "eben Ru LUI a ec UIN ere RE E 278

C. Paci dec 2.1.1 en aig CM rM pue ee Cal eee paid cq LM c E 280

IV Rete c iced d e e N RAE . 281

A. indices of Meise POM US MUR Mq UM E . 281

i. Wing and Tail... eset tentem 281

2. oe aie and Dp of Bill .. eae S

à Color e inane PRI areari rani eremi indt Ue peret 286

B. bum of Constants of Frequency Polygons ..........——— ce 287

C. a of Results and Conclusions Se Ot EE Hee cum 295

Ve DHHIMEE Lii eqni e Eea steerer pe tob 297

Bibliography ui qu Qu DH eL i aeta iem .. 298

I. INTRODUCTION.

Tue work described in this paper was begun in the fall of

1898, and was carried on at Harvard University during the

year 1898-99 under the direction of Dr. Charles B. Davenport.

It was completed during the winter of 1899-1900.

It was my desire to employ statistical methods in the study

of variation in a group of birds, and to apply the “ Precise

271 "

272 THE AMERICAN NATURALIST. [Vor. XXXV.

Criterion of Species " of Davenport ('98) to a problem in bird

classification. The smaller American shrikes of the genus

Lanius appeared to me to offer favorable material for the appli-

cation of quantitative methods to the solution of an interesting

taxonomic problem.

The shrikes are a group of passerine birds more or less

generally distributed in northern Europe and North America,

and probably of circumboreal origin. The large northern

shrike, Lanius borealis Vieill., of North America grades into

the great gray shrike, Z. excubitor Linn., of Europe; and it

exhibits a strong tendency towards individual variation.

In the United States, Mexico, and southern Canada the

breeding! shrikes are not essentially different from the

northern shrike, Z. dorealis, in certain of its color phases,

except for smaller size and the more or less complete dis-

appearance of a conspicuous barring or mottling of the

breast in adults. This barring of the breast is persistent in

most adults of Z. borealis and characteristic of the juvenile

plumage of the southern shrikes, a fact of much phyloge-

netic interest.

According to the nomenclature of the American Ornitholo-

gists’ Union there are at present recognized three races or

subspecies of the southern or smaller shrikes, which are as

follows : Lanius ludovicianus ludovicianus Linn., L. ludovicianus

excubitorides Swains. and L. ludovicianus gambelli Ridgw.

Subspecies gaméelli includes the shrikes of California and

vicinity. The shrikes of the rest of the country are classed as

either ludovicianus or excubitorides, the former being typical

in the south central states and the latter in the vicinity of

Colorado.

The shrikes of New England and the north central states

have been variously classed, by different systematists, as either

ludovicianus or excubitorides. Palmer (98) has proposed for

the shrikes of this intermediate region a new subspecies,

migrans, whose validity, I believe, can be well tested by the

* Precise Criterion " method.

! In this paper the breeding range only is considered in discussing geographi-

cal distribution.

No. 412.] SMALLER NORTH-AMERICAN SHRIKES. 273

We have in the shrikes the following eight important variable

characters :

Length of tail.

Length of wing.

Length of bill.

Depth of bill.

. Curvature of culmen.

Color of dorsal surface of head and back.

Color of upper tail coverts.

Color of breast.

Sey An EY No

II. MATERIAL.

While the shrikes make good subjects for a study of varia-

tion, there has been an unfortunate difficulty in obtaining

material in amounts as large as is desirable. Shrikes are not

especially common birds in the regions most collected in, and

they are considered rare inmost parts of New England. They

are certainly not abundantly represented in collections of birds,

and many of the skins that were obtained were imperfect, or

did not come from the breeding areas of the birds.

I have been able to procure measurements of two hundred

and ninety-four available skins, obtained from the following

sources :

Mr. William Brewster, Cambridge, Mass. . . + + - 174 skins.

National Museum, Washington, D.C. . . + + + + 64 *

Museum of Comparative Zodlogy . . < - + + * + 18. 4

Mr. J. H. Gaut, Washington, D. C. ido one ais de Saee od esl

Mr. C. F. Batchelder, Cambridge, eg EINEN E.

Dr. A. K. Fisher, Washington, D e f ^

Mr. N. Hollister, Delavan, Wis. . MU M eg pow

Oberlin College Museum, Oberlin, Ohio y

Carnegie Museum, Pittsburg, Pa. : ) oW

Mr. William Palmer, Washington, D. c boot

The numbers given do not include skins which, for various

reasons, could not be used. I wish here to express my thanks

to the gentlemen who have secured for me the loan of the

above-mentioned material, and especialy to Mr. William

Brewster for many courtesies received.

274. THE AMERICAN NATURALIST. [VoL. XXXV.

GEOGRAPHICAL DISTRIBUTION.

Mate. FEMALE. MALE. | FEMALE.

PONE a 16 13 Montana 4 3

British Columbia. . I o evada à 2 o

Tonma o oa 15 21 New Mexico . o I

&olomdo- 6.40. >- I 2 Ne or 5 2

Connecticut .. . . I [o] North Carolina 3 o

District of Columbia 6 2 North D I o

PUO . S. n 45 23 nio 3 I

Georgi 8 6 Ontario [o] I

Illinois IO 3 Pennsylvania . I I

Indiana . I o South Carolina . 16 2

Indian Territory o I Tennessee I I

Kentucky I o Texas 7 5

Bé... o I tah o I

Massachusetts . I 2 Virginia 4 10

Mexico . 17 16 Wisconsin o I

Minnesota . I o om : 3 I

Total | 174 120

III. METHODS.

A. General Precautions.

It is hardly necessary to say that work of this kind should

be done with material in normal condition. I have made no

measurements where mutilations existed, and only adult birds

bearing good evidence of representing their breeding ranges

have been used. It has been necessary to reject a number of

skins because of lack of data as to sex. As male and female

shrikes are essentially alike in color and differ little in size, it

is not possible to verify the original determinations of sex;

however, there has been no noticeable confusion in this

respect.

No attention has been given to the subspecific distinctions

appearing on the labels of skins, for I have considered it

Important to avoid all possibility of bias of opinion which

attention to previous classification might give.

No.412.] SMALLER NORTH-AMERICAN SHRIKES. 275

B. Measurements of Variable Characters.

I. Linear Measurements. Length of Wing. — The shortest

distance between the wrist and the tip of the wing was

measured with a pair of dividers, the wing being folded natu-

rally at the side of the body. The left wing was taken in every

case for the measurement.

Length of Tail.— This measuremént was made from the

papilla of the uropygial gland to the extreme end of the tail

feathers. One arm of the dividers was placed with its point

resting on the anterior face of the papilla. On account of

mutilation and frequent difficulty in finding the papilla, the

measurement has often to be omitted. Great precision cannot

be obtained, as there is irregularity in the relative position of

the papilla in the drying of the skin, and there is also usually

some wearing away of the distal ends of the rectrices. When

the latter were much frayed no measurement was attempted.

Length of Bill.— The only satisfactory method found for

measuring the length of the bill was to take the shortest dis-

tance between the nostril and the distal apex of the upper

mandible. The point of one arm of the dividers was placed

against the most distal face of the nostril. I have had to make

allowance in some cases for the wearing away of the distal

apex of the upper mandible, though the amount of wear is too

small to materially affect the measurement.

Depth of Bill. —'The greatest dorso-ventral diameter of the

bill near its base was determined. There is considerable lia-

bility to error here on account of lack of uniformity in the

articulation of the mandibles in the dried skin, and I found it

necessary to reject quite a number of skins on this account.

2. Curvature of Bill. — This is a character of the distal half

of theculmen. Its quantitative expression is a matter of some

difficulty, for we have here a curve which is not the arc of a

circle nor a parabola, nor does it correspond to any geometrical

figure. Then, too, the form and size of the bill are such as to

render it impracticable to make precise measurements directly.

To meet the latter difficulty it was decided to try to trace on

paper the enlarged projection of the outline of the bill. After

276 THE AMERICAN NATURALLIST. [Vor. XXXV.

some experimentation, the following apparatus was devised.

Fig. 1 is a diagram of an enlarging camera with a box in front

of the lens, at the left in the figure. A circular aperture for

the passage of light was made in one side of the box, the oppo-

B C

d PR BESTE A OBERE

Fic. 1. — Longitudinal section of enlarging pparatus. B, box; C, paper cylinder;

P, photographic paper in holder.

site side being removed. At one side of the aperture (A, Fig. 2)

a simple cylinder (C, Figs. 1 and 2) of stiff paper just large

enough to admit a shrike skin was fastened to the outside of

the box in a nearly horizontal position by a single tack.

Orientation was then secured for any skin by the following

simple adjustments: (1) rotation of the skin in the cylinder

around its longitudinal axis; (2) rotation of the cylinder on the

axis formed by the tack; (3) moving the skin in the cylinder

towards or away from the aperture; (4) rotation of the box on

a vertical axis.

The apparatus was arranged so that the bill of a shrike was

about seven inches from the front of the lens and its median

plane at right angles to the axis of the camera. Strong dif-

B fused daylight was then

allowed to pass through

..4 the aperture of the box

past the bill, and into

the lens of the camera,

as shown by the arrows

in Fig. 1. In the plate-

holder at the back of the camera was placed a piece of * velox "

paper (a rapid printing photographic paper) about 2x3 inches

Fic, 2. — M Sue te (C) holding bird so that the bill

s before aperture (4).

No.412.] SMALLER NORTH-AMERICAN .SHRIKES. 277

in size. An exposure of seventy-five seconds was ordinarily

sufficient to obtain a picture, which appeared on development

as a white area on a black ground. A magnification of 3%

diameters was secured, care always being taken to have the

distance of the bill from the lens constant. An outline of

the culmen of practicable working size having been thus

obtained, the next step was the analysis of the curve.

It was highly desirable to have one simple criterion of the

curvature, if one sufficiently representative could be found.

In Fig. 3, which represents the outline in a representative case,

a great increase in the sharpness of the curvature is seen

Fic. 3. — Diag f bill showing hod of ing f cul A B equals distance

of A from nearest margin of nostril; CD, tangent to outline of culmen parallel to A B; ZF,

perpendicular at point of tangency, Z; ZAF, angle embraced between the two chords

of the culmen, 4B and AE.

towards the distal end of the culmen ; the sharpness of curva-

ture varies in different individuals. The point where the rapid

increase begins was found on inspection to be sufficiently

uniform in position to suggest the idea of comparing in dif-

ferent individuals the angle embraced between two chords of

the curve of the culmen, each of them terminating at the distal

end of the culmen. The proximal ends of the two chords were

selected by the following method: one was established at a

point on the culmen as far from its apex as the apex was from

the nearest margin of the nostril. The distance of the nostril

from the tip of the culmen, of course, could not be measured on

the photographic silhouette print; it was therefore determined

278 THE AMERICAN NATURALIST. [Vor. XXXV.

by measuring the actual distance in the specimen and multi-

plying that value by the linear magnification of the print.

The nostril distance was used because the nostril was found

to be the only point sufficiently definite and constant for the

proximal limit. This chord (AB, Fig. 3) served as a base line.

A line (CD) parallel to the base line and tangent to the cul-

men was next drawn, and at the point of tangency (E) a per-

pendicular (EF) was erected between the lines. It was then

a simple matter to measure the lines AF and £F, and with a

table of natural tangents to determine the angle EAF. This

is the angle that has been adopted as a criterion of curvature

in the shrike bill. It is easily seen that there might be an

infinite number of small variations in the outline of the culmen,

especially in the part distal to the point Æ, which would not

affect the angle ZAF; but I believe that a correlation is to be

expected between the curvatures in various portions of the

culmen, so that it is probable that any considerable variation

in outline elsewhere would affect the curvature in the region

of E and thus find expression in the angle ZAF. In order,

however, to have something more than a subjective impression

on this matter, I made a second series of measurements with

half the length of the line AB used as a chord. The result of

these measurements, and their deviations from those of the

first set, are given on page 294.

3. Color Measurements. X ds the quantitative determination

of color, the “color mixer” was suggested. A simple form of

color mixer — the Bradley Milton Color-Top — was employed.

This instrument has a graduated disk, with superimposed paper

disks of five different colors, which can be so adjusted as to

make compound disks exposing two or more colors. On being

rotated at high speed these colors are mixed into one, which

is a combination of the colors used in the proportions of the.

number of degrees of each exposed. With the aid of this

apparatus it is therefore possible to determine the proportions

of these primary colors entering into any color effect which

may be produced by various combinations of them. To deter-

mine the composition of any color in nature, it is necessary

to reproduce empirically that color in the top and note the

No.412.] SMALLER NORTH-AMERICAN SHRIKES. 279

combinations that have given the desired effect. In practice,

however, the following conditions were found essential to

accurate estimation of color: (1) Uniform strong diffused

daylight, preferably coming from a skylight or north window

and not subject to strong reflections from colored surfaces;

(2) frequent comparison with ‘previously made estimates ;

(3) careful comparison of the color obtained by the top with

that of the plumage, whose color is being measured, both

being viewed from different directions.

I found it impracticable to make color estimates for more

than two hours continuously, because of color fatigue of

the eye.

In spite of the above precautions, there are limitations to

the use of the color-top. In the estimation of color, the

personal equation plays an important part. Then, too, the sur-

face of the color disks is very different in character from that

of feathers. The peculiar luster or sheen so characteristic of

a bird's plumage cannot be imitated by the color mixer. In

estimating the color of the breast, more or less mottling, which

exists in some cases, increased the difficulty. In such cases I

have attempted to estimate the mean color. Some apparatus

for blending a complex pattern into one color would be very

desirable. I do not attempt to maintain that estimates of the

same material by another person would exactly agree with

mine; but I believe that I have a consistent classification of

individuals according to color.

After several weeks of experimentation I found that, though

there were traces of blue, red, and yellow, the colors black and

white were by far the most important elements in the areas

measured. Therefore I have taken as the color criterion the

amount of black, or the melanism, of the color area described ;

and color estimates appear in the tables as percentages of

melanism. I have found it more difficult to estimate colors

containing more than 50% of white than those in which black

predominated, and I have not attempted to make fine distinc-

tions as to melanism in the lighter color areas. The large

amount of black necessary to produce even the lightest

grays in the color mixer was a matter of surprise to me. To

280 THE AMERICAN NATURALIST. | [Vor. XXXV.

properly appreciate the estimates of color given in this paper,

a color mixer should be used to reproduce the combinations

given.

Ability in the discrimination of color I have found to be

much developed by experience. The color determinations of

my first three months’ work were rejected, as increased power

of discrimination made more accurate results possible.

C. Geographical Areas.

The material described in this paper has been collected from

a territory of great size and varying conditions, and it is there-

fore desirable to compare individuals both as parts of a single

group and in subdivisions corresponding to natural life areas,

so that correlations between individual variability and geo-

graphical variation may be made. I have adopted the life

areas employed by Allen (93) for the territory covered by my

material, which are four in number:

I. Austroriparian Subprovince, embracing North and South Carolina,

Georgia, and Florida.

2. Appalachian Subprovince, embracing Maine, Massachusetts, Con-

necticut, District of Columbia, Virginia, Ontario, New York, Pennsylvania,

Indiana, Kentucky, Ohio, Illinois, Wisconsin, Minnesota, and North

Dakota. ;

3. Campestrian Subprovince, embracing British Columbia, Montana,

Wyoming, Colorado, Indian Territory, New Mexico, Texas, Idaho, Utah,

Arizona, and Nevada.

4. Sonoran Subprovince, embracing California, Lower California, and

Mexico.

A comparison of variations for still smaller areas is desirable,

but this, to be useful, would require more material than I have

been able to obtain. The subspecies of shrikes have the dis-

tribution given by Palmer (98), excepting gambelli (Palmer

did not consider the western shrikes), which agrees very well

with the above life areas. The inhabitants of each are:

SUBPROVINCE. SUBSPECIEs.

Austroriparian — ludovicianus

Appalachian — migrans

Campestrian — excubitorides

[Sonoran — gambelli]?

No.412.] SMALLER NORTH-AMERICAN SHRIKES. 281

IV. RESULTS.

A. Indices of Variability.

1. Wing and Tail.— In Figs. 4-8, frequency polygons based

on measurements of various dimensions, the position of the mean

class is indicated by a heavy verticalline. In Fig. 4 are given

frequency polygons for the lengths of the wing (4 A) and tail (4 B)

of all available male shrikes. They show a striking absence of

variability for these characters in a series of individuals repre-

senting four subprovinces with greatly varying conditions.

In Tables I-III are given correlations between the length of

wing and the length of tail. I have used the method of

Duncker (Davenport, '99, p. 33) in determining coefficients

of correlation. The “probable error" has also been deter-

mined by the following formula (Davenport, '99, p. 34), in

which p is the coefficient of correlation.

0.6745 (1 — p?)

Van (1 +p)

On comparing Tables II and III, we find that Florida shrikes

have a greater length of tail in relation to the length of wing

than shrikes from the Appalachian subprovince.

The correlation in these characters for the series from the

Appalachian subprovince is seen to be 0.157— greater than

that of the Florida series. This difference is possibly due in

part to the skewness of the curves of frequency for these series,

Which renders precise correlations difficult.

POE qt

TABLE I.— 140 MALE SHRIKES FROM THE ENTIRE REGION.

Mean of wing = 99.06 + mm. Mean of tail = 101.571 + mm.

91 = e = 3.48 3

= 2.74 + mm. 2 mm.

i WING, SUBJECT. TArL, RELATIVE.

: —7—5-—3 ~i o 2 4 8

Rel. class 100 | roz 104. 106 108 110 mm.

Sub. class (I) 94 96 95 . 4 (II)

Doe =i 473 4

96mm. — 3 2 4. 5 3 3 I

98 mm I |! 4 9 wl HH 6 4

100 mm. o I 6 7 6 2

102 mm. 2 : ? 6 3 2

mm. a I 5 3

106 mm. (III)6 . 2 I (IV.)

p = 0.5688 + P. E. p= + 0.038 +

282 THE AMERICAN NATURALIST. | [Vor. XXXV.

TaBLE II. —41 MALE SHRIKES FROM FLORIDA.

Mean of wing = 97.463 + mm. Mean of tail = 102.292 + mm.

01 = 2.20 + mm. [23 = ^d or oH

Wine, SUBJECT. TAIL, RELATIVE.

Xa —6—4-—23 0 I 3

Rel class. . 96 98 100 102] 104 106 108 IIo mm.

Sub. class (I) (LI)

96 mm. —t I 5 I

98 mm o I I 7 4 3

100 mm 2 2 2 ki 2

mm a

104 mm. (III) 6 . 1 (IV)

p = 0.639 + P. E. p = + 0.0487

TABLE III. — 28 MALE SHRIKES FROM THE APPALACHIAN

SUBPROVINCE.

Mean of wing = 99.357 + mm. Mean of tail = 99.43 + mm.

uo ri = 23r + mm. 7 = 3.063 + mm.

WiNc, SUBJECT. Tait, RELATIVE.

Xe —-5§ —3 -! o 2 4 6

Rel. class 94 96 98] 100 102 104 106 mm.

Sub. class (I) (II)

96mm. —3.. 2 I I

98mm. —1:.. I I 5 4

100 mm. o 2 4

102 mm. ye I 2 I I

104 mm. 4. I

106 mm. (III) 6 . 1 (IV)

P = 0.796 +

P. E. p = + 0.0348 +

A general tendency towards great length of tail in southern

birds has been noted by Allen (71, pp. 230, 231). In Table

VII, p. 291, the mode of lengths of tail for Austroriparian

males is seen to be 102 mm.; whereas Appalachian males have

a mode of 100mm. The mean of Austroriparian shrikes is

101.91 +mm., while that of Appalachian shrikes is 99.43 +mm.,

283

No. 412] SMALLER NORTH-AMERICAN SHRIKES.

br Jo sje} ‘g ‘uoaa oa 891 107 sSutA 3jo[ Jo sossv[) ‘py 'suosÁ[od A»uonba1j —'* *514

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284 THE AMERICAN NATURALIST. [Vou. XXXV.

a difference of 2.48 mm. Austroriparian females show a

still greater preponderance in length of tail, having a mode

6 mm. greater and a mean 3.05 mm. greater. Not much

importance, however, can be attached to the great difference of

the modes in this case, because of the small number of females

measured.

Palmer (98) says, in referring to shrikes of the Appalachian

and Austroriparian subprovinces respectively: *In migrans

2200 See ee B 5

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Fig. 5A. b=? EHEH

16 LL] LITT LI EETTI

PEPI

E-E-EHEHEEEEEHEEEEL TET

ro EE

a ewe At LIT]

s Gt

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= ENSEEHSEEHEP V BEUBHEHHENEH \

Fig. SB. o EE AE

- 9 700 Gt On onan one SS S SN!

oe” HOHHH FPS

9 HHH pe HEBHR

o LET m —L——À

a H a EET

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Fig. sc, oL = AHL

mm, 10.6 10.9 IL.2 11.5 11,8 12.1 12.4 12.7 13.0 13.3 13.6 13.9

Fic. 5. — Frequency polygons for length of bill. 4, 164 males from entire region; 2,69 males from

Austroriparian subprovince ; C, 38 males from Appalachian subprovince.

[Appalachian] the wing is longer than the tail, due to its

migratory habit; in ludovicianus [Austrori

longest, thus indicating its fixed habitat."

I find, on the contrary,

than the tail in the Appa

and VII, pp. 290,

tail over that of t

parian] the tail is

that the wing is not absolutely longer

lachian form, though from Tables VI

291, it is clear that the excess of the length of

he wing is less in migrans than in ludovicianus ;

No. 412.] SMALLER NORTH-AMERICAN SHRIKES. 285

or, in other words, that relatively to the tail, the wing is longer in

migrans than in ludovicianus, as his theory would require.

Palmer’s explanation of length of wing in migrans as the

result of habit is plausible. However, it has not been proved

that shrikes which migrate northward actually fly any more

than those remaining in Florida; moreover, it seems to me

that there are other possible explanations. I am inclined to

think that we have here a case of the condition already noted

am FHH EFEEEEEEEELEEEEL LZ

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Fig. 6A. PAEH

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Depth in mm. 8.4 8.6 8.8 9o 9.3 9.4 9.6 9.8 10.0 10.2

Fie. 6.— Frequency polygons for depth of bill. 4, 126 males from entire region; B, 54 males from

ustroriparian subprovince; C, 21 males from Appalachian subprovince.

by Allen — a tendency towards greater length of tail in tropical

birds. Unfortunately I cannot prove this, — any more than

Palmer does his hypothesis, — for it has not been possible to

determine the actual ratios of tail length and wing length with

reference to the general size of the bird.

2. Length and Depth of Bill. — In Fig. 5 are given frequency

Polygons of variations in length of bill for the entire region and

286 THE AMERICAN NATURALIST. | [Vor. XXXV.

fortwo subprovinces. Here is shown a considerable separation

of the means (indicated by the vertical line) for the subprovince

polygons, indicating a larger bill for Austroriparian shrikes

(5.8), which are also seen in Fig. 67 to have a greater depth

of bill. The increase in size found in the bills of Florida

shrikes is also shown by comparison of Tables IV and V, in

which these characters are correlated for a series of Florida!

shrikes and a series from the Appalachian subprovince. A

striking absence of correlation and a great tendency towards

individual variation in the proportions of the bill are to be seen.

Increase in the relative size of the bill in many southern birds

has been noted by Allen (71). It would be an interesting field

for statistical investigation.

TABLE IV. — 32 MALE SHRIKES FROM FLORIDA.

Classes of Length of Bill in Millimeters.

eT of Total.

epth of Bill.

H3] In5 | 119 | tat | 13234 | 12 1.330] 1333] 15:6 | $35

8.8 mm | 1 I 2

9.0 mm. I I 2

9.2 mm. 2 I I I 5

9.4 mm. I I I 3

9.6 mm. 2 2

9.8 mm. I I I I 2 6

10.0 mm. I 5 i 4 I1

10.2 mm. I :

Total 3 10 4 3 7 4 I 32

3. Color.— Frequency polygons for variations in the melan-

ism of the head are shown in Fig. 7. The upper polygon (Fig.

7A), which includes the entire series, is seen to be distinctly

bimodal, and in the lower polygons we see that the mode of

greater melanism is due to the presence of Austroriparian

shrikes, and that the shrikes of the Campestrian subprovince

have the least melanism for the dorsal surface of the head.

1 Only Florida representatives of the Austroriparian shrikes were taken in this

case, in order to eliminate intermediate forms, for it seemed desirable to me to

compare the Appalachian shrikes with a group as nearly tropical as possible.

No. 412.] SMALLER NORTH-AMERICAN SHRIKES. 287

The Campestrian subprovince includes arid portions of the

United States, where paleness of color in the fauna is supposed

to be correlated with this condition.

TABLE V.— 22 MALE SHRIKES FROM THE APPALACHIAN

SUBPROVINCE.

Classes of Length of Bill in Millimeters.

Classes of Total.

Depth of Bill.

10.6 | 10.9 | 11.2 | 11.5 | 11.8 | 12.1 | 12-4 | 127 | 13.

8.4 mm I I

8.6 mm. I I 2

8.8 mm. I 3 I 5

9.0 mm. I I 2 I I I I 8

9.2 mm. I I 2

9.4 mm. ` I I I 3

9.6 mm o

EMEN R I 1

aa ee s.l] I 6 5 I 4 I I I 22

While such correlations are frequently suggested by the

fauna of desert regions, the relations of humidity to color

are still problems for further investigation.

Very great variations in the melanism of the upper tail

coverts are shown in Fig. 8. The shrikes of the Campestrian

subprovince have here also a mode of little melanism, 30%,

while a mode of great melanism, 70%, is found for Austrori-

parian shrikes. The upper tail coverts of Campestrian shrikes,

especially those from Colorado and Arizona, are very light

gray. This peculiarity has caused the race to be designated

as the white-rumped shrikes.

B. Tables of Constants of Frequency Polygons.

In Tables VI-XIII will be found indices of variation for all

the material studied. All computations have been made from

measurements grouped into classes. This grouping has been

adopted both in drawing the polygons and in determining the

indices of variability, in order to reduce the “probable error."

288

THE AMERICAN NATURALIST.

Percent 76 2 m 82 td 86

fa ce HHRHH FH

EEREEEEM m sum un

HHHH HHH

Pasan

EERE jas

SRT

sot —

FE HAE

EENE

Hes ic

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Fig. 7A. E

E i :

EX una HHHH

HE HH H4 ||

Fig. 7B.

EHA

eae tana

m we am omn a Saw a o7

HHHL i []

Fic. 7. — Frequency polygons for

melanism of top of head

males from entire region; B, 57

ustr rian sub-

noran subprovinc

Per cent

EAN i

- \

zanas / \ ]

10 I \

LE \

ELI x \

(eel a A Oe T ]

y y \ PE RLI

[| ya ~ -i |

pela kbd a |

Fig. 8A. H |

Fig. 8B,

-0

o ) o

LLLLIILRIRRDLLDLPDLILPSSPIUPITIIIIII

LEI

LIE

No. of individuals

7 A

EIITITT TLITTITITT.ITUTITTAEAT?,

PEOR

—

EREH |

E-LIL

Bee ]

2S Gee ee ee See /

————————

Pe BELLET ETTI TTTTA

HH SD oe ont ee oe ee ee oo a _. /

4 Fo es WA Ba O DA a REA A DA

— PRA

EN EH 4

ae a8

—— oe =a

EE] =

ee ee OA A E A

Peo tt f

eee SH

r——1—: e] a aS |] A

——- = LT A

HHH LEE] z

Ed E] LE ES] ~

— LL.

EPET =

= bt

— 1 | i

= H s

B m"

ui —

5 =

B E E

æ — SRS Oot Oh 1S ELE

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a Ela

—

"n ^" n » T ^ 4» » €» "

subprovinca; Polygons for melanism of upper tail coverts. 4, 142 males from en iin een B, 55 males

subprovince.

from Austroriparian

from

C, 34 males from Appalachian subprovince; D, 18 males from Campestrian subprovince; Z, 35 males

290

THE AMERICAN NATURALIST.

[Vor. XXXV.

The following constants of the frequency polygons have

been determined (Davenport, '99) :

1. Mode: Class of greatest frequency.

Mean: 77. RC.

3. Standard Deviation, e = Ten

4. Coefficient of variability : CV. or X 100.

TABLE VI. — LENGTH oF

Lert WING.

. Probable error of Mean: P. E. M. [sub.] = + 0.6745 wa ;

. Probable error of Standard Deviation : P. E.c [sub.] = + 0.6745 Vag

AREA.

Entire region

Austroriparian .

Appalachian .

Campestrian .

Sonoran .

MATERIAL.

168 males.

70 males.

35 males.

27 males.

36 males.

112 females.

31 females.

25 females.

24 females.

32 females.

Mope.

98 mm.

96 mm.

98-100 mm.

98 mm.

I04 mm.

102 mm.

98 mm.

MEAN. STANDARD Dev.

99.09 4- mm. 2.79 + mm.

97.98 -- mm. 2.64 + mm.

97.6 + mm. 2.04 + mm.

96.64 + mm. 2.35 + mm.

99.2 mm 2.45 + mm.

97.68 mm 2.17 + mm.

101.26 mm 2.45 + mm.

99.75 mm 2.79 + mm.

100.28 mm 2.90 + mm.

98.19 + mm. 2.36 + mm.

CV. of 168 males = 2.81 +.

P. E. M. [sub.] 168 males = + 0.145.

P. E. « [sub.] 168 males = + 0.1026 +.

CV. of 112 females = 2.69 +.

No.412.] SMALLER NORTH-AMERICAN SHRIKES.

TABLE VII. — LENGTH oF TAIL.

291

AREA. | MATERIAL. Mope. MEAN. STANDARD Dev.

Entire region 141 males. 101.55 + mm. | 3.49 + mm.

95 females. 99.55 + mm. | 3.63 + mm.

Austroriparian .| 65 males. 102 mm. IOI.9I + mm. | 3.12 + mm.

30 females. 104 mm. 100.47 -- mm. | 3.25 + mm.

Appalachian . 28 males. 100 mm. 99.43 -- mm. | 3.06 + mm.

19 females. 98 mm. 97.42 -- mm. | 3.85 + mm.

Campestrian . 20 males. 104 mm. 102.3 mm. | 3.48 4 mm.

18 females. 102 mm. 99.89 + mm. | 3.42 + mm.

Sonoran . 28 males. 100 mm. 102.21 + mm. | 3.90 + mm.

28 females. 102 mm. 99.43 + mm. | 3.88 + mm.

CV. of 141 males = 3.43 +. CV. of 95 females = 3.65 +.

P. E. M [sub.] 141 males = + 0.198 +.

P. E. ¢ [sub.] 141 males = + 0.14008 +.

TaBLE VIII. — LENGTH OF BILL.

AREA. MATERIAL, Mone.

Entire region 164 males.

112 females.

Austroriparian . males. 12.1 mm.

30 females. 12.1 mm.

Appalachian . 38 males. 11.8 mm.

29 females. 11.9 mm.

Campestrian . 25 males. 12.1 mm.

20 females. 12.1 mm.

Sonoran 32 males. 12.1 mm.

35 females. 12.1 mm.

MEAN.

STANDARD Drev.

12.01 + mm.

11.71 + mm.

12.58 + mm.

12.17 + mm.

11.63 + mm.

11.26 + mm.

11.84 + mm.

11.57 + mm.

11.96 + mm.

11.76 + mm.

0.71 + mm.

0.63 + mm.

0.59 -- mm.

O.51 + mm.

0.56 + mm.

0.54 + mm.

0.63 + mm.

0.51 + mm.

0.54 + mm.

0.53 + mm.

CV. of 164 males = 5.89 +. CV. of 112 females = 5.35 +-

P. E. M. [sub.] 164 males = + 0.0374 +-

P. E. o [sub.] 164 males = + 0.0264 +.

292 THE AMERICAN NATURALIST. [Vor. XXXV.

TABLE IX. — DEPTH or BILL.

AREA. MATERIAL. | Mops. | MEAN. STANDARD Dev.

Entire region .| 126 males. 9.27 -- mm. 0.42 + mm.

85 females. 8.95 -- mm. 0.41 + mm.

Austroriparian .| 54 males. 10.0 mm. 9.54 -- mm. 0.38 -- mm.

22 females. 9.2 mm. 9.32 -- mm. 0.37 + mm.

Appalachian. .| 21 males. 8.8-9.0 mm. 8.95 + mm. 0.30 + mm.

19 females. 8.8 mm. 8.90 + mm. 0.36 + mm.

Campestrian. .| r9 males. 9.0 mm. 9.08 4- mm. 0.28 + mm.

14 females. 8.8 mm. 8.8 + mm. 0.30 + mm.

Sonoran . . .| 32 males. 9.0 mm. 9.12 + mm. 0.44 + mm.

30 females. | 8.8-9.0 mm. 8.78 + mm. 0.36 + mm.

CV. of 126 males = 4.57 +. CV. of 85 females = 4.61 +.

P. E. M. [sub.] 126 males = + 0.0252 +.

P. E. c [sub.] 126 males = + 0.0178 +.

TABLE X. — MELANISM or Top OF HEAD.

AREA. MATERIAL. Mone. | Mean. STANDARD DEV.

Entire region . 144 males. 83.57 + 4. 3.00 + %-

99 females. 83.66 + %. 3.19 + %-

Austroriparian .| 57 males. 86%. 86.17 T5 1.69 + %-

24 females, 88%. 87.25 + %. 1.81 + %

Appalachian. . 33 males. 80%. 82.24 + %. 2.68 + f$.

25 females. 80%. 82.32 + %. ($03 + %

Campestrian. .| 21 males. 80%. 80.67 + %. 1.13 + fi

17 females. 80%. 80.94 + %. 1.78 + %-

Sonoran . . .| 33 males. 82%. 82.24 + %. 2.4 +%

33 females. 82%. 83.45 + 4. 2.38 + %

CV. of 144 males = 3-58 +. CV. of 99 females = 3.81 +.

P. E. M. [sub.] 144 males — 4- o.1686.

P.E. c [sub.] 144 males = + 0.119 +.

No. 412.] SMALLER NORTH-AMERICAN SHRIKES.

293

TABLE XI. — MELANISM OF UPPER TAIL COovERTSs.

AREA. MATERIAL. Mone. MEAN. STANDARD Dev.

Entire region .| 142 males. 53.13 + %. 15.42 + %

104 females. 47-98 + %. 18.99 + 4.

Austroriparian .| 55 males. 70%. 67.22 + 4. 5.02 + %.

25 females. 65%. 66.2 y 8.16 + %.

Appalachian. .| 34 males. 60%. 58.38 + %. 93 +%

20 females. 60%. 53-2 y 64 +%

Campestrian. .| 18 males. 30%.» 25.28 + ^. 11.4 +%

17 females. : 28.25 + % 148 +%

Sonoran. . .| 35 males. 35%. 40.43 + %. 13.01 + ^j.

i 43 females. 30%. 38.26 + %. 13.37 + %-

CV. of 142 males = 29.02 +. CV. of 104 females = 39. E T.

P. E. M. [sub.] 142 males = + 0.873 +.

P. E. v [sub.] 142 males = + 0.617 +.

TABLE XII. — MELANISM OF BREAST.

STANDARD Dev.

AREA. MATERIAL. Monk. MEAN.

Entire region .| 124 males. 31.21 + %.

98 females. 36.58 + %-

Austroriparian .| 42 males. 25%. 23.81 + %.

27 females. 35% 3241 + %

Appalachian. .| 35 males. 35% 43.28 + %

23 females. 40%. 47-61 + %

Campestrian. .| 17 males. 25%. 27.65 + %-

18 females. 35%. 31.39 + %

Sonoran . . .| 3o males. 25/5. 295 — 7

30 females. 35%: 35.9 Jor

11.07 + 4.

10.51 + %.

6.66 + %.

10.36 + %.

94 7

8.71 + %.

7.27 + %

6.63 + %.

6.65 + %.

7.30 + %

CV. of 124 males = 35.48 +. CV. of 98 females = 28.72 +.

P. E. M. [sub.] 124 males = + 0.6709 +.

P. E.¢ [sub.] 124 males = + 0.474 +.

294 THE AMERICAN NATURALIST. [Vor. XXXV.

Because of limited time I was unable to obtain measure-

ments of culmen curvature for more than forty-seven individ-

uals (partly males and partly females) a number so small that

a rather large probable error is found for coefficients of

variability.

The results obtained are given in the following table.

TABLE XIII. — CuRvATURE OF CULMEN.

23° | 24° | 25^ | 26° | 27^ | 28° | 29° | 30° | 31° | 32° | 33° | 34^ | 35° | 36° MEAN.

Austroriparian | 1 t iiio? 28.66° +

Appalachian 3 X*43419|/21311 29.32" +

Campestrian sia iar 496 bse 1 azta | ee

Sonora . rl I 32.00?

To. = : | 1 214:2160|81613|/3/2|5] 141209000

Austroriparian c = 274? +. Cl. = Dou Er i 0.534 +-

Appalachian ¢ = 2.03? +. CV.= 6.924. P.E£.¢ = + 0.206 +-

Campestrian o = 3.12° +. CV.— 10.06 +. P. E.e — + 0.372 +-

Sonoran = 2160+. CV.= 6.75. P. E. a = + 0.504 ++

Total ¢=2.74°+. CV.= 9gaig—. P. E.c- 40.191 +.

The shrikes of the Austroriparian subprovince, mostly Florida

birds, are seen to have a mean curvature 3.34°—less than

that of the Sonoran birds measured. With so few individuals

precise statements as to curvature are not very reliable, but I

believe that the figures given are approximately correct for the

four subprovinces represented. The series of measurements of

the curvature of the culmen when half the base line A B (Fig. 3)

was used as the chord gave the following constants and coeffi-

cients of variability for the forty-seven individuals measured in

the first series: Mean, 26.34° +; Mode, 27^; o = 9.35 7)

CV. =8.85; P. E.¢=+40.162 +.

The angles obtained from the second series of measurements

were never greater than those from the first series. The

greatest deviation from the first set of measurements was 7^,

which occurred in four cases, All the deviations are indicated

in the following table:

o? rÉ 29 Lu 4? 5° 6° 7?

4 2 6 to IO 7 4 4 individuals.

No. 412.) SMALLER NORTH-AMERICAN SHRIKES. 295

It was impossible to get as accurate results in the second

series, because of greater difficulty in determining the-actual

point of tangency, but the results obtained show that there is

a fairly close correlation in the curvature for various parts of

the bill; that is, where the culmen is much curved in one

place, it is likely to have correspondingly strong curvature at

other points.

The only selection exercised in the whole work was employed

in the estimation of curvature, individuals which appeared on

inspection to be typical for their respective subprovinces having

been chosen, since it was impossible, for want of time, to photo-

graph the entire series.

C. Discussion of Results and Conclusions.

Although computations have been carried out to the third

or fourth decimal place, figures beyond the second decimal

place are not given, as they would imply a degree of precision

which is not attainable in an investigation of this kind. For

example, the mean length of wing for 168 male shrikes is seen

to be (see Table VI) 99.09 -4- mm. It often happened in mak-

ing correlations that, because of general wear or some special

mutilation to a single character, some individuals could not be

included in a correlation table. This has happened especially

often in the case of the tail, so that in the correlation table for

wings and tails only 140 of the 168 male shrikes could be

included. The mean for the left wings of the series of 140

shrikes was found to be 99.06 mm. (see Table I), a result

Which differs from the mean of the whole lot (168) by 3 in

the second decimal place.

The modes and means given for the melanism of the breasts

of Appalachian-subprovince shrikes show a percentage which,

though higher than in other subprovinces, is not as high as

Would have been the case had not a large proportion of indi-

viduals come from localities intermediate between the Campes-

trian and Appalachian subprovinces. Shrikes from New

ngland were found to have 50-60% of melanism for the

breast. An analysis of material shows that only five skins

296 THE AMERICAN NATURALIST. [VOL XXXV.

were obtained from New England, whereas Illinois alone is

represented by thirteen. This lack of equalization in the

sources of material prevents certain desirable interpretations

of tendencies towards minor variations.

To my mind one of the most important results reached is the

determination of the relative variability of different characters

in a group of birds representing geographical areas of consid-

erable size. The coefficients of variability (Tables VI-VII)

indicate for the wing and tail a variability of less than 4 in.

length. The bill is somewhat more variable, as is shown by a

coefficient of 5.89 -- for the length of bill in a series of 164

males. Color, as would be expected, is much more subject to

variation. The upper tail coverts and breast are the most vari-

able; but the coverts furnish only a very small part of a bird's

coloration, and the color of the breast was the character which

it was the most difficult to measure satisfactorily, especially as

advanced age and the condition of the plumage are factors of

possible importance which I have found it difficult to consider.

Fortunately for this particular study, shrikes do not change

much after the first winter plumage is obtained.

I believe that migrans is as worthy of recognition as gam-

belli. Whether it is profitable to encumber nomenclature with

the names of these races, based on slight variations, is a ques-

tion which is worthy of further consideration.

The power of discriminating fine shades of color varies in

different persons, and it can be highly developed by education.

At the present time there is much activity among certain

systematists in the production of new subspecies for geo-

graphical varieties, which long experience and special adept-

ness enable them to distinguish. A variation, no matter how

slight, that can be correlated with geographical range is con-

sidered to warrant an addition to nomenclature; but the dis-

covery and description of geographical races can be carried on

almost ad infinitum.

Birds, because of their powers of flight, might be expected

to be less subject to the factor of isolation than non-migratory

animals, but the tendency to return in spring to the same

breeding place must, in some species at least, be conducive to

No.412.] SMALLER NORTH-AMERICAN SHRIKES. 297

the formation of numerous local variations, or family character-

istics, whose recognition is a matter of power of discrimination

on the part of the systematist.

It seems highly desirable that the question of limiting the

establishment of new subspecies or varieties by some generally

accepted criteria be considered.

I do not argue for the universal use of the method of the

* Precise Criterion," but I believe that it is both desirable and

practicable to employ it in certain problems of taxonomy, such,

for instance, as the one just discussed. The ordinary work of

classification, perhaps, does notat present require the precision

in treatment furnished by purely quantitative methods, but

problems of race distinction, I believe, need the precision of

the “Precise Criterion." The contention that quantitative

methods are less useful than those ordinarily employed because

of the large amount of material required, is mischievous, for it

argues that generalizations professing precision are possible by

methods which are not precise. The problems of finer classi-

fication can be properly settled only by the use of a large

amount of material, whatever the methods used.

V. SUMMARY.

Quantitative methods have here been applied to the study

of variation in the smaller shrikes of North America, and the

following variable characters have been measured :

. Length of wing.

. Length of tail.

. Length of bill.

. Depth of bill.

ulmen.

. Color of dorsal surface of head.

. Color of upper tail coverts.

. Color of breast.

The three color areas have been found to vary principally in

the amount of melanism present. A series of 294 shrike skins

from various parts of the United States, Mexico, and southern

Canada have been studied and measurements of these skins

have been classified. —

on Am 2 CO Wh &

298 THE AMERICAN NATURALIST.

Coefficients of variability have been determined as follows

for the whole series.

MALES FEMALES

SOB GL WANE . 5.5... XUL 2.69 +

SAME OR Me ge eS ge + 3.65 +

ween OF D Oe Boa es 689 E 5.35 +

Depth of bill : Gi. uu o CUT MB Ee 4.61 +

Melanism of top of head Pao aa s LER d 3.81 +

Melanism of upper tail coverts . . . . 29.024 39.58 +

New OF MEUM e e a . a 36548 2832 +

Curvature of culmen . . 47 males and females 9.15—

Florida shrikes were found to have relatively large bills and

long tails.

A large percentage of melanism has been found for the top

of head and the back of shrikes from the south central states,

while shrikes from the vicinity of Colorado and Arizona have

a relatively small percentage of melanism for all three color

surfaces measured.

I wish to make acknowledgments to Dr. Davenport for

supervision of the work and helpful criticisms, and to Dr. Mark

for suggestions and revision of the manuscript.

CAMBRIDGE, Mass.

BIBLIOGRAPHY.

Allen, J. A

"71. On the Mammals and Winter Birds of East Florida, with an Exam-

ination of certain assumed Specific Characters in Birds, and a

Sketch of the Bird Faunae of Eastern North America. Buli.

Mus. Comp. Zoól. Vol. ii, No. 3, pp. 161—450.

Allen, J. A

'93. The Geographical Origin and Distribution of North-American Birds,

considered in Relation to Faunal Areas of North America.

Auk. Vol. x, No. 2, pp. 97-150, 2 pls.

Davenport, C. B.

'98. A Precise Criterion of Species. Science (N.S.). Vol. vii, No. 177

pp. 685-690.

Davenport, C. B.

'99. Statistical Methods with Special Reference to Biological Variation.

59 PP., 10 tables, and 28 figs. New York, John Wiley & Sons.

er, W.

'98. Our Small Eastern Shrikes. Zhe Auk. Vol. xv, No. 3, pp. 244-258:

A NEW INSTRUMENT FOR MEASURING TORSION.!

FRANK RUSSELL.

THE apparatus that has heretofore been used to measure the

torsion of the long bones of the human skeleton has been so

difficult of manipulation as to be impracticable. The following

description of a simpler apparatus is offered in the hope that

it may prove useful to those who are interested in statistical

somatology.

The base is 7 by 30 inches; it is grooved for a distance of

14 inches from the middle to within 2 inches of the right end.

The post A, 8 inches high, is fixed to the base, and has a spur

projecting 34% inches toward the right. An ordinary brass

protractor is attached to the top of the post at right angles to

the spur. (A protractor with the figures reversed in position

would be better.) The center of the protractor is fixed at the

axis of the spur, on which is pivoted a U-shaped needle that

1 Demonstrated at the winter meeting of Section H of the — Associa-

tion for the Advancement of Science, at Baltimore, Dec. 27, 1900.

299

300 THE AMERICAN NATURALIST.

rises 3% inches above the spur and extends downward exter-

nally tothe protractor scale. Theright end of the needle should

terminate in an eye, through which the spur passes ; the figure

represents the trial needle not thus arranged.

The post B is movable ; its base slides in the groove of the

base board and is held in position by a thumbscrew. The top

is provided with a strip of steel 3 inches in length. The strip

is provided with saw teeth that engage the head and great

trochanter of the femur at the same time (or both condyles),

for it is pivoted at its center and stands in the plane of the long

axis of the bone. This strip is fixed in an exactly vertical

position, and the protractor is exactly horizontal in relation

to it.

The torsion of any long bones can be measured with this

instrument. The axis of each end is first indicated in pencil ;

the spur of the post 4 is then engaged with the lower portion

of the axial line at one end of the bone; the post B is pushed

to the left until the strip of steel is in contact with the axial

line of that end of the bone. With the left hand the needle

on post 4 is adjusted so that the arms are in the plane of the

axis of the left end of the bone, the end of the needle points

to the degree of torsion on the protractor.

The operation can be performed rapidly and with accuracy.

As the bone can be turned end for end and remeasured in

other positions, we have a ready means of making control

measurements. In practice I have not found it necessary to

use any support for even fragile bones, but to insure against

accident to such material it is advisable, perhaps, to have a

narrow table between the posts of the apparatus, which will

not support the bone but simply prevent its falling far enough

to injure it.

The apparatus can be made by any skilled mechanic. The

model figured here was made for me by a graduate student of

anthropology, Mr. W. C. Farabee, at Harvard University.

PEABODY MUSEUM OF

AMERICAN ARCH;EOLOGY AND ETHNOLOGY.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XIV. THe Hypromepus#® — Part I.

CHARLES W. HARGITT.

INTRODUCTORY.

THE following synopsis was undertaken more than a year

ago as a section of the * Synopses of North-American Inver-

tebrates " now in course of publication. Various interruptions

have delayed its completion at an earlier date.

While compiled largely from the author's notes and observa-

tions made upon the Hydrozoa of the Atlantic coast during a

period of more than ten years, the form and method of pres-

entation are patterned after the systematic works of Hincks,

Allman, Haeckel, and von Lendenfeld. For many of the

descriptive notes recourse has been had to L. Agassiz's Con-

tributions to the Natural History of the United States and to

A. Agassiz’s Catalog of the Acalephe of North America, as

well as to those of the authors just named.

The synopsis is confessedly incomplete in several of the

orders, specially upon the Campanularide and Leptomeduse.

It is, moreover, limited to a comparatively small range of hydro-

zoan life of American waters, chiefly of the northeastern

Atlantic coast. Of that of the Pacific coast our present

knowledge is still too limited to warrant even a provisional

synopsis.

The Hydromeduse comprise one of the three generally

recognized classes of Coelentera, of which the others are the

Scyphomedusze and Anthozoa. While the first two classes have

been regarded as much more intimately related phylogenetically

than has the third, it may be doubted whether after all their

relation may not be quite remote, at least so much so as to

Warrant separate consideration. Hence slight, if any, reference .

301

302 THE AMERICAN NATURALIST. [Vor. XXXV.

wil be made to the Scyphomedusz in considering possible

relationships or phylogeny among the Hydromedusz.

The Hydromedusa may be distinguished by the following

somewhat broad characteristics. In typical cases there is a

more or less well marked alternation of generations, 7.¢., a non-

sexual, hydroid stage and a sexual, medusoid, stage. The latter

are derived by a process of budding from the stem or hydranth

of the hydroid as gonophores which may become free as

meduse or may exhibit varying phases of degeneration as

medusoids or mere sporosacs, as in Clava, Campanularia, etc.

In many cases there may be exhibited proliferous medusz

from various portions of the parent medusa, as in Hybocodon.

In rare cases one or other of these phases may be entirely

lacking, as in Hydra, in which the medusa phase is wholly

absent, or as in Rhegmatodes and many others the hydroid

stage is apparently lacking.

Perhaps in no phylum of the animal world is there a more

striking exhibition of polymorphism than among the Hydro-

medusz. This seems to reach its climax in the Siphonophora,

though in such forms as Hydractinia it is also evident.

In general the hydroid exhibits a sedentary habit quite in

contrast with the free-swimming habit of the medusa. But

here again are numerous exceptions. Hydra is capable of

locomotion, as are also other hydroid forms, while as already

indicated many medusz are sessile and degenerate, and in the

Siphonophora the entire polymorphic colony is free-swimming.

In general the hydroids are colonial though with notable

exceptions, as in Hydra and many others. While in general

form the hydroid and medusoid present rather striking mor-

phological differences, they may yet be reduced to a common

and fundamental likeness. Both are of diploblastic structure,

having a definite ectoblast and entoblast separated by a middle

lamella, or mesogloea, which is a delicate, structureless mem-

brane in the hydroid and in the medusa a rather massive, gelati-

nous structure, making up the bulk of the body and giving it

the characteristic glassy appearance.

In the absence of definite knowledge concerning details of

the life history of many of the Hydromeduse, it is as yet

No. 412.] MORTH-AMERICAN INVERTEBRATES. 303

impossible to formulate any scheme of classification which

shall bring into a single view the complete ontogenetic rela-

tions of the various hydroid and medusoid phases. In the

present synopsis I have followed in the main that of Allman

and Hincks for the hydroids and that of Haeckel for the medusz,

though in each there is not a little variation both as to the order

of presentation as well as the nomenclature used.!

SYNOPSIS OF ORDERS OF HYDROMEDUS./E.

I. HvDRARLE.

Polyps solitary, never forming colonies; no medusoids; sex-cells pro-

duced in ectoderm of polyps.

Of this order only a single well-defined genus is recognized, Hydra.

The genera Protohydra and Microhydra are probably allied genera, but

their affinities are too uncertain as yet to warrant definite classification.

The former is of marine habit, the latter of fresh-water habit. Both are

devoid of tentacles, and sexual reproduction, at least in the former, seems

unknown

Of the genus Hydra there are two well-distinguished species: 77. fusca

and ZZ. viridis. Both abound in fresh waters of small lakes, ponds, and

sluggish streams, associated with various aquatic plants, notably Lemna,

various algz, pond lilies, etc.

Il. HYDROCORALLINÆ.

Colonial. Hydrosome comprising polyps of two forms, gastrozoids and

dactylozoids, supported from a network of coenosarcal hydrorhize, from the

ectoderm of which is secreted a calcareous mass which is deposited over

the spaces or meshes of the network. The colonies form incrusting, often

arborescent, masses over shells, stems of Alcyonaria, or other support,

often forming massive and fantastic shapes, as in the so-called * stag's

horn coral.” Only one genus is likely to come within the range of

the present synopsis, namely, Millepora; and of this a single species,

alcicornis.

III. TuBULARUE (Gymnoblastea).

The Tubularie are for the most part colonial hydroids, producing free

meduse, or medusoid gonophores, by budding. Hydroids devoid of

! Just as these notes were being put into final form for the press, I have been

permitted to consult the manuscript of a forthcoming Handbook of the Sear of

the Woods Hole Region, by Professor C. C. Nutting, for the privilege of Iam

under grateful obligations.

304 THE AMERICAN NATURALIST. [Vor. XXXV.

hydrothece and gonangia. Sexual individuals when set free are known

as Anthomeduse. Medusz ocellate, z.e., the sensory bodies, are visual in

character, and are located usually at the bases of the tentacles. Gonads

borne in the tissues of the manubrium.

IV. CAMPANULARLE (Calyptoblastea).

Hydroids with hydrothece and gonangia. Colonial; propagating by

budding both in development of hydrosome and in formation of gonosomes,

the latter of which may become free as medusz, or only partially develop

as medusoids, with only rudimentary medusan organs. Medusa with sen-

sory organs of the vesiculate type, otocysts, borne upon the margin of the

bell, usually between bases of tentacles. Gonads borne under the radial

canals. Medusa when free are Leptomeduse.

V. TRACHOMEDUSÆ.

Hydromeduse devoid of hydrosome (hypogenic) medusa developing

directly from the egg; no alternation of generations known. Sensory

organs chiefly tentaculocysts, containing endodermal otoliths. Gonads

borne under radial canals. Medusa generally somewhat hemispherical

in shape, with thick mesoglea. Radial canals, four, six, or eight, often

centripetal.

VI. NARCOMEDUS/E.

Hydromedusz devoid of hydrosome (hypogenic), development of medusa

being direct with no alternation of generations. Medusa rather flat in

shape, and with radial canals in form of broad gastric pouches, which vary

in number, as do also the tentacles, which are usually set at some distance

up on the outer surface of the umbrella.

VIL SIPHONOPHORA.

Hydromedusæ with free-swimming, polymorphic colonies, produced by

differential budding. The colonies of this order are characterized by an

extreme specialization of the several types of individuals which comprise

them. Reproductive products borne in gonophores which seldom become

ree.

KEY TO FAMILIES OF TUBULARIÆ.

Hydranth devoid of specialized receptacles, hydrothecæ. Sexual

products not borne in closed gonangia.

t. Hydranths with scattered, filiform tentacles . « .. CLAVE Re I

2. Hydranths with single whorl of filiform tentacles :

a. Hypostome conical, not abruptly differentiated.

^. Colony regularly branched . . - . BOUGAINVILLIDA, 3

No. 412.] MORTH-AMERICAN INVERTEBRATES. 305

&. Colony not branched. Hydrorhiza of anastomosing canals, form-

ing an incrusting base, overlaid with ectodermal ccenosarc.

c. Hydranths with sessile, fixed gonophores . HYDRACTINIDA, 5

c. Hydranths producing free meduse . . . PODOCORYNIDA, 6

a’. Hypostome trumpet-shaped or hemispherical . . EUDENDRIDA, 4

3. Hydranths with more than a single whorl of filiform tentacles :

a. Stem provided with definite sheath of hdi perisarc.

6. Distal tentacles in two whorls . . . . HYBOCODONIDA, IO

/. Distal tentacles not in two whorls . . . TUBULARIDA, 9

a’. Stem not provided with definite shéath of perisarc, more or less

definitely marked with longitudinal flutings or ccenosarcal channels

CORYMORPHID&, 8

4. Hydranths with scattered, somewhat EUM disposed, capitate tentacles

COR

only ORYNIDÆ, 2

5. CORN SR with odit AH at filiform RESI E and with distal

capitate tentacles on hypostome . . . . . . . . PENNARIDA, 7

I. CLAVIDA.

Colonial, stems simple or branching, hydranths elongate, clavate, with

numerous filiform tentacles irregularly disposed over the y. Gono-

phores borne upon hydranth, or on special branches, or occasionally arising

from the hydrorhiza. Medusoids never free.

GENERA.

I. CLAVA. Colony of simple, unbranched individuals, devoid of peri-

2. RHIZOGETON. Colony very similar to Clava. Gonophores arising

from hydrorhiza. x

3. CORDYLOPHORA. Colony profusely

branched and with definite sheath of perisarc.

Clava leptostyla Ag.

Trophosome: Hydranths simple, with slen- -

der basal portion which arises from a filiform

hydrorhiza protected by a delicate perisarcal

filiform, and scattered over the hydrant

Kondomi Gohachinesd in clusters at base Fic. 1. — Clava — Ag.

of tentacles, medusoids never becoming free. VENIET tS

Male gonads of a bright pinkish hue, similar, in general, to that of the

colony. Female gonads of a rather distinctly purple color

Habitat: Shallower waters on fucus, docks, sea wall, etc., at t Cold Te

Harbor, Woods Holl, Hadley Harbor, etc.

306 THE AMERICAN NATURALIST. [Vor. XXXV.

Rhizogeton fusiformis Ag.

(Contr. Nat. Hist. U. S., vol. iv.)

Trophosome: Colony much asin Clava. Hydranths of about X inch in

height, tentacles about twelve in number, borne on distal half of polyp.

Gonosome: Gonophores oval, arising from hydrorhiza on short pedun-

cles, the whole invested by filmy perisarc.

Habitat: Rocky pools between tide marks, Massachusetts Bay.

Cordylophora lacustris Allman.

Trophosome: Colonial, profusely branching, hydranths with scattered

filiform tentacles.

Gonosome: Gonophores borne on branches, ovate and with definite

investment of perisarc.

Habitat: Brackish, and fresh

waters in lagoons, ponds, etc.

2. CORYNIDA.

Colonial, hydranths with capitate

tentacles only, scattered over the

elongated bodies, or growing in

indefinite whorls. Gonophores usu-

ally borne among the proximal ten-

tacles, or from body of polyp and

producing medusz, which may be-

come free or remain attached.

GENERA.

I. SYNCORYNE. Stem invested

by definite perisarc; hydranths clavi-

form.

. 2. Corynitis, Stem devoid of

definite perisarc ; hydranths sessile,

with long, cylindrical bodies.

Fic, 2. — Syncoryne mirabilis Ag. Syncoryne mirabilis Ag.

(After Agassiz.) ^

Trophosome: Branched, perisarc

smooth or with only slight indication of annulations. Hydranths with

numerous capitate tentacles.

onosome: Meduse borne on hydranth body. These are of two forms,

one free and developing earlier, hemispherical, with well-developed ten-

tacles, with an ocellus at their base; the other fixed, tentacles rudimentary,

and devoid of ocelli.

No.412.]] .VORTH-AMERICAN INVERTEBRATES. 307

Corynitis agassizii McCr.

Trophosome: Colonial, not branched, hydranths with cylindrical, highly

contractile bodies, and spirally arranged, capitate tentacles.

Gonosome: Gonophores growing low on body of hydranth or among

Medusz almost spherical, the surface dotted with

the proximal tentacles.

Marginal tentacles two or four, nodulated and

clusters of nematocysts.

swollen with batteries of nematocysts.

abitat: Shells of Mytilis, usually overgrown with incrustations of

Membranopora.

3. BOUGAINVILLIDA.

Colonial, branching, with distinct perisarc. Hydranths with conical

hypostome and a single whorl of filiform tentacles. Gonophores borne

just below the hydranth. Medusz with four radial "m

canals, marginal tentacles either single or in clusters, à |

and with ocelli at their bases.

3 Wi Fi

GENERA. AX. WKF

1. BOUGAINVILLIA. Hydrocaulus with dense . Y PA L%

perisarc. Medusa with clustered mapan tentacles 4, PM.

and with branching oral tentacles. N I) TA

2. PERIGONIMUS. Stems with gelatinous peri- | y 7

sarc. Medusa with two marginal tentacles and NU

without oral tentacles. »

Bougainvillia superciliaris Ag. » i

Trophosome: Colony attaining a height of about y

two inches. Stem irregularly branched, branches

annulated proximally. Hydranths with inconspicu-

ous hypostome and from fifteen to twenty tentacles.

Gonosome : Gonophores borne mostly on pedicels fic. 3. — Bougainvillia

from ultimate branches. Mature medusæ with heavy fri iaris Ag.

manubrium and branched tentacles, those of margin

(After Agassiz.)

arising from conspicuous sensory bulbs. Colony light color with greenish

tinge, hydranth light rose tint. Medusz with yellowish manubrium tipped

with red, sensory bulbs reddish orange.

Bougainvillia (Margelis) carolinensis McCr.

Trophosome: Colony sometimes eight to twelve inches high, usually

much smaller. Stem profusely branching, with hydranths freely dis-

tributed on both stem and datura and of elongate and flexible, sub-

Conical form. Tentacles about twelv

Gonosome: Gonads borne on both stem and branches, often in clusters.

308 THE AMERICAN NATURALIST. [Vor. XXXV.

Medusz much as in previous species, but with narrower and shorter hypo-

stome. Colony light grayish tinged with dull green, hydranths with red-

dish tint. Medusa with brick-red manubrium and sensory bulbs, ocelli

ack.

Habitat: Piles of docks, occasionally on seaweed and floating timbers.

Perigonimus.

Colonial, rarely attaining a height of more than ¥% of an inch, simple or

branched, perisarc usually gelatinous and extending to base of tentacles.

Hydranths relatively large and with conical hypostome. Medusz borne on

hydranths or on stem or branches, bell-shaped and with two to four

tentacles with bulbous bases.

Perigonimus jonesii.

(American Naturalist, vol. xxviii, p. 27.)

Trophosome: Colonial, branching freely, with thick, gelatinous perisarc,

often wrinkled, extending to, or even including, the bases of tentacles.

ydranths with subconical hypostome,

with about sixteen filiform tentacles,

alternately elevated and depressed.

Gonosome: Medusz ovoid or hemi-

spherical, with four radial canals and

ocelli, but having only two tentacles,

which are often spirally coiled and dis-

posed within the subumbrellar cavity.

Habitat: Found only upon the

abdomen and legs of the spider crab,

Labinia marginata, Cold Spring

Harbor, L.i

4. EUDENDRIDJE.

Colonial, often branching with great

profusion, becoming quite arborescent.

Perisarc distinct, more or less annulated, attached by creeping hydro-

rhiza. Hydranths flask-shaped, with sharply differentiated, trumpet-shaped

hypostome. Tentacles filiform, forming a single whorl about the base

of the hydranth. Male gonophores borne in a verticil just beneath the

tentacles of hydranth, which in some species become directly metamor-

phosed into gonophores. Female gonophores not verticillate, usually borne

on body of hydranth, which often becomes transformed into gonophores

with their peculiar spadiceous, finger-like coils enclosing the ova. The

family includes a single genus, Eudendrium, fairly characterized in the

accompanying cut. The following species are designated :

Fic. 4.— Perig,

No.412.] MWORTH-AMERICAN INVERTEBRATES. 309

Eudendrium ramosum Linn.

Trophosome : Colony arborescent, much branched, attaining a height of

from four to six inches. Branches rather symmetrical, pinnate and some-

what alternate, with similar sub-branches. Hydranths somewhat ovoid,

with trumpet-shaped hypostome, and with

fo Ne single verticil of about twenty tentacles,

el j some of which are often atrophied in male.

Gonosome: Sexes distinct, though often

growing in approximate colonies. Gono-

phores of female somewhat pyriform and

scattered, springing from body of hydranth

7% % clusters, each from three to four chambered.

pete Color of male reddish, of

female orange. Abundant

on piles of docks, on racks,

etc., in shallower waters.

Eudendrium dispar Ag.

(Cont. Nat. Hist. U.S., vol. iv.)

Trophosome : Colony large,

Fic. 5.

E height, stems more slender

(After Allman.) than in former, somewhat

Fic. 5.— Colon fascicled, extensively and

Fh Goo Ry dranth with eooo p oret; variously branched and an-

nulated. Hydranths vasiform, with about twenty-eight tentacles.

Gonosome: Sexes distinct. Gonophores of female of pinkish orange

hue, variously clustered about the base of the more or less atrophied

hydranth and from distal portion of stem. Habitat in deeper waters in

Vineyard Sound, attached to rocks, shells, etc.

Eudendrium tenue A. Ag.

(No. Am. Acalephe, p. 160.)

Trophosome: Colony very small, rarely exceeding an inch in height,

branching irregularly, hydranths vasiform, borne on slender pedicels.

Gonosome: Male gonophores from two to four chambered, of pinkish

Color, clustered from bases of tentacles. Female gonophores bright orange

in color, scattered over the branches and stem. Habitat on seaweed, etc.,

in shallower waters. Not abundant.

310 THE AMERICAN NATURALIST. [Vor. XXXV.

Eudendrium capillare Alder.

Of the distinctness of this species and Æ. a/bum, listed by Professor

Nutting, I have grave doubts. It seems to me that Alder’s diagnosis of

capillare coincides so closely with that of Æ. zenue as to render their identity

highly probable. So also of Æ. album. Specimens taken at Woods Holl

seem almost certainly identical with Æ. Zezze, and therefore both should

probably be merged under Æ. capillare, whatever slight differences there

are being hardly greater than varied environment would easily explain.

5. HYDRACTINID/E.

The Hydractinide are so closely allied to the following family that it

seems unfortunate that they were not originally merged ; the only easily

distinguishable difference being in the free medusz of the latter in contrast

with the fixed sporosacs of the former. In size, general habit, and mor-

phology they are so closely identical that but for the gonosomes no differ-

ence would be recognizable, though in Podocoryne the hydrorhiza seems less

definitely covered with naked ccenosarc, — but even this differs greatly in

specimens from different localities.

In both polymorphism is a marked feature, at least three types of polyps

being distinguishable :

1. Feeding hydranths (trophopolyps); whitish in color and with numer-

ous filiform tentacles, ey appearing in alternately elevated and

depressed order.

2. Reproductive individuals (gonopolyps), more slender-bearing gono-

phores in clusters below the tentacles, which are fewer in number than in

the first and imperfectly developed.

3. Spiral polyps, elongated individuals,

wholly devoid of tentacles and with apex

of body thickly beset with nematocysts.

The entire colony arises from an in-

crusting base which is thickly beset with

jagged spines, the latter sometimes con-

sidered a fourth type of individual.

Hydractinia echinata Fleming.

(Hydractinia polyclina Ag., Cont. Wat.

Hist. U. S.)

: Trophosome: Colony composed of

ag Beraman echinata. numerous polyps, as given above.

"wn ad Gonosome: Gonads as sessile sporo-

sacs borne on distinct hydranths, gonopolyps, having but few tentacles.

Medusoids never free.

No.412.] MORTH-AMERICAN INVERTEBRATES. 311

Habitat: Usually found upon shells occupied by the hermit crab, but

occasionally found upon fucus and occasionally also upon piles of docks.

6. PoDOCORYNID.

Colony very similar to that of the Hydractinide, as given above.

Hydranths with single whorl of filiform tentacles surrounding base of the

conical hypostome.

Podocoryne carnea Sars.

Trophosome : Hydranths slender, pinkish-white in color, and with filiform

tentacles.

Gonosome: Medusz borne in clusters about the hydranth just below

the whorl of tentacles. When set free the medusa is of marked bell-shape,

with definite velum, short manubrium of reddish color, four radial canals

from the bases of which arise eight marginal tentacles.

Habitat as in Hydractinia.

Stylactis.

Under this generic name Sigerfoos describes (American Naturalist,

Vol. XXXIII) a hydroid having many points in common with the Podoco-

rynidz, and it should probably be classed under this family. He has given

to it the specific name Hooperi (cf. of. cit.). The following definitive

characters have been given of it:

Trophosome : Hydranths slender, with a length when fully expanded of

about 3% of an inch. Tentacles in single whorl, filiform, and of variable

number, eighteen to twenty-five.

Gonosome: Gonophores borne upon specialized hydranths just below

tentacles, and set free as meduse having four radial canals, eight tentacles

Which are somewhat rudimentary, devoid of ocelli. Sexual products borne

upon manubrium. Found on shells of Stygnuassa (Illyanassa) obsoleta.

7. PENNARID&.

Colony arborescent, pinnately branched, hydranths with two sets of

tentacles, one proximal composed of ten to twelve, filiform, the other borne

upon hypostome in two indefinite whorls, short and capitate.

Pennaría tiarella McCrady.

of origin. Hydranths large and flask-shaped, those terminating stem or

branches appreciably larger than others.

312 THE AMERICAN NATURALIST. [VorL. XXXV.

Gonosome: Meduse borne on hydranth body above the whorl of

proximal tentacles. Medusa liberated during early evening and dis-

charging the sex products immediately thereafter. In many cases the ova

are discharged before the liberation

yy of the medusz, as indeed are also

y Habitat: Abundant on piles of

i docks, floating timber, eelgrass, fucus,

| etc, usually in shallower waters.

| x Development from June to October.

Z 8. CORYMORPHID/.

occasionally found definite colonial

EC. E.

i » Eg

3 i amr ie

Fic. 8. Fic. 9.

| " y Usually solitary, though I have

)

Fic. 8. — Pennaria tiarella McCr.

Fic. 9. Hyd th 1 A h 1

f medusaz

buds arising direct from the hydrorhiza. Hydranths with proximal and

distal whorls of filiform tentacles. Gonophores as free medusz with four

radial canals and with one to four marginal tentacles, one of which is usually :

much the larger. i

Corymorpha pendula Ag.

(Cont. Nat. Hist. U. S., vol. iv.)

Trophosome: Hydrocaulus from two to four inches in height, the fleshy

Ceenosare traversed by longitudinal canals which ramify more or less near

the base. Hydrorhiza an indefinite root-like expansion of the base, by

which the whole is attached to the sandy substratum.

Hydranths flask-shaped, sharply distinct from stem. Proximal tentacles

large, forming a single whorl at base of hydranth. Distal tentacles very

contractile, forming alternating verticils about the base of the hypostome.

No.412]] MWORTH-AMERICAN INVERTEBRATES. 313

Gonosome: Medusz borne on branched peduncles, arising just above

the proximal tentacles, ovoid hemispherical, with single large and usually

three rudimentary tentacles.

Hydroid bright pink in color, meduse light yellowish, manubrium,

tentacles, and bulbs pinkish.

Habitat: Sandy bottom in rather deep waters at

various points in Vineyard Sound, Muskegat Chan-

nel, etc.

9. TUBULARIDA.

Hydrocaulus with definite perisarc, simple or ir-

regularly branched. Hydranths flask-shaped, with

proximal and distal whorls of filiform tentacles.

Gonophores in form of fixed sporosacs, borne on

branched peduncles.

Tubularia.

Generic description as given for family. The

following species are given:

Tubularia couthouyi Ag.

(Cont. Nat. Hist. U. S., vol. iv.)

Trophosome: Stems unbranched, attaining a height

: Fic. ro. —

of from four to six inches. Hydranths large, often ndula

expanding an inch or more in diameter, with proximal (Modified from ‘Aman )

whorl of thirty to forty filiform tentacles and a distal one of much smaller.

Gonosome : Gonophores as numerous, densely crowded racemes of

: pendulous sporosacs. Larva escaping as actinule. Hydranth and gonads

Habitat: On sandy bottoms dredged off Nobska Point, Vineyard

Sound, and other similar places in the same locality.

Tubularia larynx Elis and Solander.

Trophosome: Stems clustered, more or less branched, annulated. Height

One to two inches. Stem forming a collar-like expansion just below

hydranth, ds latter bearing sixteen to twenty proximal filiform tentacles

and a distal whorl of about the same number.

Gonosome: Gonads in pendulous clusters, similar to last. Color of

hydranth and gonads rosy. Perisarc yellowish.

Tubularia spectabilis Ag.

(Thamnocnidia spectabilis Ag., Cont. Nat. Hist. U. S.)

Trophosome : Colony irregularly branched and sparsely annulated.

Height three to four inches. Hydranths much as in former.

Gonosome: Comparable with former.

314 J THE AMERICAN NATURALIST. [Vot. XXXV.

Tubularia tenella Ag.

(Thamnocnidia tenella. Ag.)

Trophosome: Colony very small, rarely exceeding a height of one and

one-half inches. Stem loosely branched and with indefinite annulations.

Hydranths with tentacles about as in former.

Gonosome: Compare T. larynx. Color and habitat much as in last.

Fic. 12. — Single hydranth

enlarged.

(After Agassiz.)

Fic. 14.— Hydranth with pendulous

gonophores.

Fic. 13. — Tubularia crocea Ag.

No.412.] MORTH-AMERICAN INVERTEBRATES. 315

Tubularia crocea Ag.

(Parypha crocea Ag.)

Trophosome: Colonies growing in dense tufts of tangled stems of from

three to four inches in height. Stems sparingly branched, with occasional

indications of annulations. Hydranths with tentacles much as in former

species, but numbering from twenty to twenty-four in each whorl.

Gonosome: Much as in the first species. Hydranths and gonads of rosy-

red color, stem pale, whitish. Habitat. Growing in dense masses on piles

of docks, floating timbers in harbors, and shallower waters.

Hypolitis perigrinus Murbach.

Under this name Murbach describes a hydroid taken at Woods Holl

(Quar. Journ. Mic. Sci., Vol. XLII), which would seem to have some

affinities with the Tubularidæ. The fol-

lowing characters are summarized :

rophosome : Colony consisting of sim-

ple hydranths with long hypostome and

with distal and proximal whorls of filiform

tentacles. Polyp free. ;

Gonosome: Gonads borne on hypo-

stome just above proximal tentacles and

occur singly in the type specimen. Sessile

medusoids, somewhat terete in form and

devoid of tentacular processes.

10. HYBOCODONIDÆ.

Hydrocaulus 1 1 1, solitary, with

definite perisarc and hydrorhiza. Hy-

dranths large, with proximal and two

distal whorls of filiform tentacles,

A. H foy Ag.

Fic. 15. — Hybocodon z

(After Agassiz.)

Hybocodon prolifer Ag.

(Cont. Nat. Hist. U.S., vol. iv.)

Trophosome: Stems longitudinally striated, occasioned by coenosarca

canals, Perisarc enlarged and annulated just below hydranth. Hydran

similar to those of the Tubularidz, but with oral tentacles in two distinct

whorls. d

Gonosome : Gonophores closely attached to hydranth body just above

Proximal tentacles. Free meduse with four radial canals, and with a single

greatly enlarged tentacle from whose base a number of secondary medusz suc-

“essively bud, and from these still other groups of similar meduse may arise.

SYRACUSE U NIVERSITY.

REVIEWS OF RECENT LITERATURE.

ZOOLOGY.

Jordan and Snyder on Japanese Fishes. — In the Proceedings of

the United States National Museum (XIII, 335), Jordan and Snyder

give a list of the fishes collected in Japan by Professor Keinosuke

Otaki in 1895 and 1896, and by the “Albatross” in 1896, these col-

lections being in the Museum of Stanford University and the United

States National Museum. Fourteen species are described as new and

most of them figured by Mrs. Starks. These are Chimera phantasma,

Gobio biwe, Gobio mayede, Otakia rasborina, Congrellus meeki, Pseudo-

tolithus mitsukurii, Sebastodes hakodatis, Sebastodes scythropus, Scor-

pena onaria, Callionymus beniteguri, Trifissus ioturus, Blennius

Jatabet, Calorhynchus kishinouyei, and Verasper otakii. The species

last named is probably not distinct from Verasper grigorjewt, lately

described as a species of Hippoglossus from Hakodate by Herzen-

stein. The manuscript name of Lampetra mitsukurii Hatta is

adopted for the small lamprey of southern Japan.

In the list of Japanese fishes the following new genera are

indicated: Jshikauia (steenackeri), Otakia (rasborina), Konosirus

(punctatus), Bryttosus (kawamebari), Corusculus (Anthias berycoides),

Lteliscus (Etelis berycoides), Insidiator (rudis), Trifissus (ioturus),

Rhombiscus (cinnamomeus), Karcius (bicoloratus = scutifer), Usinosita

(Japonica), Zebrias (zebrina), Areliscus (joyneri).

he two species referred to Gobio belong rather to Günther’s

genus Leucogobio. The specimens called Apogon guadrifascia-

fus belong to Apogon notatus (Houttuyn) = semilineatus Schlegel.

Those recorded as Sebastodes oblongus belong to S. mitsukurii

Cramer, which differs in the larger scales. The species recorded

as Chenogobius castaneus is distinct from the latter species and is

as yet undescribed. The genus Trienophorichthys, as Steindachner

has shown, is identical with Tridentiger, and Trifissus is not differ-

ent. In fact, later investigations show the identity of Trifissus

toturus with Steindachner's Tridentiger bifasciatus. D.S Í.

Fowler on Fishes of the Tropical Pacific. — In the Proceedings of

the Academy of Natural Sciences at Philadelphia (1900, p. 493) Mr. Henry

317

318 THE AMERICAN NATURALIST. [Vor. XXXV.

W. Fowler gives a detailed account of fishes from Hawaii, Tahiti,

and Samoa, in the collection of the Academy. Most of these were

collected more than sixty years ago by the noted naturalists, Dr. John

K. Townsend and Mr. Thomas Nuttall. Others were more recently

obtained by Dr. William H. Jones and Dr. Benjamin Sharp.

The new species are the following. From Honolulu: Zycodontis

parvibranchialis, Echidna zonata, Stolephorus purpureus, Synodus sharfi,

Hemipteronotus copei, Brotula townsendi; from Samoa, Mugil caldwelli.

Mr. Fowler is very careful as to his nomenclature and synonymy,

a sure sign of good workmanship in systematic zodlogy, and his

conclusions seem everywhere tenable. The plates which illustrate

Mr. Fowler’s paper, eleven species in all, seem accurately drawn,

but are not well reproduced.

It may be here noted that the name Stolephorus cannot be used

for the great genus of anchovies, to which it has been of late years

(following Bleeker) applied. Its type, Atherina japonica of Houttuyn,

from Nagasaki, is not an anchovy, but the very common Japanese

silver-sided sardine, Kibuna-iwashi, commonly known as Spratelloides

gracilis. Stolephorus should therefore supersede Spratelloides. The

genus of anchovies called Stolephorus should probably stand as

Anchovia. The single species which Jordan and Evermann set

apart under the latter name is probably not generically distinct.

Perhaps all these species should be reunited under Engraulis, fol-

lowing Giinther’s view. The tropical anchovies have, however, 4

smaller number of vertebra: and a firmer texture of body than the

species originally called Engraulis. I may note also the necessity of

returning to Gymnothorax Bloch instead of Lycodontis. D. S. J.

Miall and Hammond's Harlequin Fly.!— This is a book about

an animal that has figured prominently in histological work for a

generation. It is a book that is intended to facilitate the study of

Chironomus, especially for inland naturalists, to whom it is so readily

available, by setting forth in detail its habits and life history, present-

ing a résumé of the studies hitherto made of it (chiefly on its salivary

glands and its embryology), and adding many new and more or less

interesting facts and observations.. The chapter headings are as

follows: Outline of Life History, and Relations of Chironomus to

Other Diptera; the Larva of Chironomus; the Fly of Chironomus ;

1 Miall, L. C., and Hammond, A. R. Zhe Structure and Life History of the

Harlequin Fly (Chironomus). Oxford, The Clarendon Press. 8vo, iv + 196 pP»

129 figures

No. 412.] REVIEWS OF RECENT LITERATURE. 319

The Development of the Pupa and Fly within the Larva; The Pupa

of Chironomus; and The Embryonic Development of Chironomus.

To this is added a brief appendix on methods, likewise intended to

promote the use of Chironomus as a laboratory subject. The numer-

ous figures are well selected and useful, but in their execution

the best of them do not rise above mediocrity.

Incidentally there is described (on page 34) and figured the larva

of Clinocera (Fam. Empidide), a new type of dipterous larva with

eight pairs of prominent abdominal prolegs. LO N.

Reactions of Protozoa. — In the Supplementband for rgoo of

the Archiv für Anatomie und Physiologie, August Pütter! presents a

most valuable contribution to our knowledge of the reactions of

unicellular organisms. The fact that the reactions of these creatures

to various stimuli are profoundly modified when the organism is in

contact with a solid, is strikingly evident to any one that has studied

the behavior of the Protozoa. Pütter has subjected to a thorough

analysis this effect of contact of solids (thigmotaxis) and its inter-

ference with the operation of other stimuli, and the results form a

contribution, an acquaintance with which is indispensable to all who

wish to obtain an understanding of the behavior of these creatures.

Exact observation of the actual movements of the organisms, close

attention to the interrelation of structure and function, and careful

analysis of the various factors involved, form the striking and valu-

able features of the paper, which stands in refreshing contrast in

this respect to some of the recent papers dealing with the reactions

of lower organisms. The paper is so full of detail, and casts

light on so many observed phenomena, that it is impossible to give

an idea of the results in a brief notice. In addition to a precise

account of the thigmotactic reaction itself, the author deals particu-

larly with the reactions to heat and to the electric current, as

modified by the thigmotactic reaction. The observations on electro-

taxis are in accord, in all essentials, with those set forth by Pearl in

the American Journal of Physiology for July, 1900, and throw some

additional light on this subject, especially on the subject of Zrazs-

verse electrotaxis. Attention may be further called to the fact that

Pütter confirms for many Infusoria the method of reaction to a

stimulus by turning toward a structurally defined side, as described

by the reviewer.

* Pütter, August. Studien über Thigmotaxis bei Protisten, Archiv für Anatomie

und Physiologie, Physiologische Abteilung, Supplementband, 1900, pp. 243-302.

320 THE AMERICAN NATURALIST. [Vor. XXXV.

By the work of Pütter another of the reactions of these organisms

— thigmotaxis — is placed on a satisfactory scientific basis, and

light is thrown on many other phenomena. HS

Notes. — Burchardt’s recent article (/enaische Zeitschrift, Bd.

XXXIV, pp. 719-882) on the body spaces and connective tissue of

Amphioxus is of general interest because of the accompanying bibli-

ography, which is intended to be complete for this important and

much studied animal. The list is arranged chronologically and

includes some six hundred references. One is reminded of Baedeker

on finding that important papers are indicated by an asterisk.

Metcalf (* Notes on the Morphology of the Tunicata,” Zoologische

Jahrbücher, Bd. XIII, 1900, pp. 495-602, Heft IV) gives us a paper

extending over a wide range of morphological and systematic topics.

A brief notice like the present one cannot mention all the results of

interest contained in such a paper as this. Among them the follow-

ing are perhaps the most important: The homology of the vertebrate

hypophysis and the neural gland of the Tunicata is a “ suggestion the

truth of which, while it may be probable, is still insufficiently estab-

lished.” Reéxamination of the structure of the interesting deep-sea

genus Octacnemus leads to the conclusion that its affinities are with

those simple ascidians that reproduce by budding, rather than with

the Salpidae, as supposed by Herdeman. It is proposed to institute

a family, the Octacnemidz, for it. A new species of the molguloid

genus Bostrichobranchus, viz., B. molguloides, is described.

A contribution to the postembryonal development of Molgula is

made by Marc de Selys-Longchamps and D. Damas (Arch. de Biologie,

Tome XVII, 1900, pp. 385-483). The development of the stigmata

is studied with special care. Six pairs of protostigmata are recog-

nized, the first two pairs forming simultaneously. The later multi-

plication and coiling up of the stigmata and the formation of the

infundibule are followed out. The larval sense organ is found to

persist into adult life in M. ampulloides. The study of the devel

opment of the sexual glands leads to the conclusion that there is no

bilateral symmetry in the disposition of the germinative epithelium.

In studies on some distomes, Jacoby describes (Arch. f. Naturges,

- I, 1900) a species, Distomum heterolecithodes Braun, which is

remarkable for the fact that the vitelline gland, which is usually

symmetrical, is developed only on one side of the body. In eleven

cases studied the organ was sinistral, while in four the author found

No412] REVIEWS OF RECENT LITERATURE. 321

a complete szfus inversus, or, as Kowalewski has termed it, sexual

amphitypy, in which the arrangement of organs is the mirror image

of the normal condition. The same anomaly was found to occur in

the following species in the ratio given: Opisthorchis crassiuscula

(Rud.) 7: 84, O. poturzycensis (Kow.) rarely; O. albida 16: 68, O. trun-

cata (Rud.) 6:50, O. felinea (Riv.) 8:100. Other cases from other

species have been recorded by Stiles and Hassall, and Kowalewski

regards it as a characteristic of the genus Opisthorchis. If this be

true, it must still be remembered that it may occur in other genera

also. Jacoby observed it in Distomum lanceolatum, which is not

related to Opisthorchis but rather perhaps to D. heterolecithodes.

The nervous system of Moniezia expansa has been studied by

Tower (Zool. Jahrb., Abt. Morph., Bd. XII, pp. 359-384, 6 pls.).

It is noteworthy that the physiological salt solution, so universally

used, is harmful if the worms remain in it more than an hour. The

author gives the formule of fluids successfully employed for trans-

porting the cestodes, for keeping them alive in the laboratory even

up to five days, and for fixing and staining the nervous tissues. In

the scolex is present an anterior nerve ring with four small ganglia

opposite the suckers, a pair of large cephalic ganglia connected with

the ganglia of the anterior ring by four nerves and surrounded by the

posterior ring which connects with the lateral lobes of the cephalic

ganglia, and also by four small nerves with the ganglia of the anterior

ring. These latter nerves are the beginning of the dorsal and ventral

longitudinal nerves, while the prominent lateral nerves spring from

the lateral lobes of the cephalic ganglia. No accessory lateral nerves

Were present. In each proglottid each lateral nerve bears an anterior

lateral ganglion near the center and a posterior lateral ganglion near

the posterior margin. From the former a transverse genital nerve

arises, and from the latter a bunch of smaller fibers, together with a

marginal nerve, which is recurrent, and the dorsal and ventral commis-

Sures which together constitute the nerve ring of each proglottid.

The Uncinarie of the Canidae and Felidz and the Sclerostominze

of the Ruminants have been subject to a careful revision by Railliet

(Arch. Parasitol., Tome III, No. 1, 1900).

The distomes of the isolated genus Rhopalias St. and H. (Rho-

Palophorus Dies) have been restudied from Rudophi’s and Diesing’s

types by Braun (Zool, Anzeiger, Bd. XXIII, pp. 27-29). Three species

are reported, of which one is new, and the genus appears to be con-

fined to the marsupials of South America. These forms are closely

322 THE AMERICAN NATURALIST. [Vor. XXXV.

related to the echinostomes, but the lateral lobes of the head, which are

supplied with spines, have been transformed into a retractile proboscis.

An Atlantic “ Palolo” is described by Mayer (Bull. Mus. Comp.

Zool., Vol. XXXVI, pp. 1-14, 3 pls.). The form, which is shown to

be distinct from the Pacific species of similar name and habits, is

named Staurocephalus gregaricus. It appeared in a dense swarm

before sunrise on a single morning and discharged its sexual prod-

ucts with the coming of the sun under contractions so violent that

the ripe segments were torn open. The author gives an interesting

discussion of the advantages of this habit in shortening the egg-

laying season, concentrating the breeding individuals and not only

insuring more perfect fertilization, but also reducing the distance

which the sperm must traverse in order to fertilize the ova; while

‘the increased struggle for food due to the production of a large

number of young larva is counterbalanced by the heavily yolk-laden

egg of this species.

The Alciopide and Tomopteride of the Plankton Expedition by

Apstein (Ergebnisse der Plankton- Expedition der Humboldt-Stiftung,

Bd. II, H. b., 61 pp., 14 pls.) opens with an extended taxonomic

account of the alciopids captured. The geographical distribution of

these forms has been much extended, as appears from the tables

given, which also show that the group belongs to warmer waters, with

the exception of strays found in the northern branches of the Gulf

Stream and of a single Antarctic species. Within the warm zones

they appear to be generally distributed without special areas and are

present everywhere in approximately equal numbers, as shown by the

hauls of the vertical net. Among the tomopterids, however, a single

species seems to occur only in the vicinity of land, though with an

extended range, while the others are all true pelagic forms. The

genus contains species found in cold waters, — those characteristic

of warm regions and such as inhabit both. While moderately equally

distributed, these forms do not manifest the uniformity noted for the

alciopids.

BOTANY.

Some Recent Forestry Publications. — The interest in the preser

PFE of the timber covering which still characterizes certain parts

of the earth, and its renewal in denuded areas, which appears to be

Nuus] AEFPIEWS OF RECENT LITERATURE. 323

spreading gradually in this country and which has long been recog-

nized as economically important in many other parts of the world,

is leading to the rapid appearance of handbooks and other publica-

tions, of varying degrees of excellence but all of utility. Notice has

been made elsewhere of Mr. Gannett's magnificent treatise on our

own forests, which stands in a class quite by itself as a work of

rare statistical and other practical value, and of the botanico-

forestal books of Professor Sargent? and Miss Keeler.* The mail

now brings an official handbook of Indian forestry, by R. Ribben-

trop! and a book by Abbot Kinney, intended to awaken and spread

popular interest in this subject on our Pacific coast.

Mr. Ribbentrop, who is Inspector-General of forests to the govern-

ment of India, traces the history of forestry development in a British

colony where, whatever else may be said, the home government has

made continuous and intelligent effort to develop and conserve the

rich endowment of nature; and the influence of this policy on the

native governments is shown to be most encouraging.

Mr. Kinney, who has had ample opportunity to observe the reck-

less waste of timber in one of the grandest existing forests, analyzes

the relations of forest preservation to the elements and the greed

or carelessness of man, and presents an outline for a forest system

for southern California. A number of specialists contribute chap-

ters to his book on irrigation and the peculiar conditions apper-

taining to sand dunes. | T

Coulter and Rose’s Umbelliferæ. — A revision of the Umbelliferæ

of our flora, by Drs. Coulter and Rose,® constitutes the opening

number of Vol. VII of the Contributions Jrom the United States

National Herbarium. Three hundred and seventy-one species,

pertaining to Seventy-eight genera, are included, with necessary

! Gannett, H. Forest Reserves, Ann. Rept. U. S. Geol. Surv., Washington,

1900, vol xx, pt. v. xviii + 498 pp., with numerous maps, charts, and process

illustrations from photographs.

2 Sargent, C. S. The Silva of North America. Boston and New York, 1890-

1900. Twelve quarto volumes, with many plates, and to be completed in one or

x additional volumes now in preparation.

4 Keeler, H.L. Our Native Trees and how to Identify Them. New York, 1900.

Ribbentrop, B. Forestry in British India. With a rainfall chart and three

t k ii + 245 pp. Calcutta, 1900. diagram

: mney, A. Forest and Water. v + 247 pp. with numerous e

alf-tone plates. Los Angeles, 1900.

6 oulter, J. M., and Rose, J. N. Monograph of the North-American Umbel-

Mere. vii + 256 pp. Pl. IX, ff. 65. Washington, 1900.

324 THE AMERICAN NATURALIST. [Vor. XXXV.

synonymy and description, and the essential fruit characters of

the genera are clearly and accurately figured. In pleasing con-

trast with the habit of some writers, the authors give a full cita-

tion of the material that they have examined in the course of

their study — on the satisfactory result of which they are to be

congratulated.

Studies of Plant Life! is a laboratory guide for high schools,

which represents the experience of three Chicago teachers. The

twenty-four studies it contains, each consisting of directions and

questions regarding easily obtainable material, are intended to sup-

plement the botanical lectures or text-book work of a school year.

Part I, made up of seventeen studies, deals mainly with Crypto-

gams; Part II is devoted to the organs of Spermaphytes and their

modifications. Much use is made of the compound microscope and

of comparatively inexpensive apparatus. In an appendix are given

directions for twenty-four experiments illustrating nutrition, metab-

olism, growth, and irritability. Following this is an outline of a

model field trip showing what may be done in the way of ecological

study. At the end of the book comes an analytical key for the

determination of one hundred seed plants of northeastern United

States, which includes a short synopsis of forty-two families of

angiosperms.

The book seems well calculated to direct young students along

profitable and interesting lines of work, following present-day

methods of teaching the * New Botany."

But few errors have been noticed, and these are for the most part

such as any competent teacher may be trusted to correct before they

are likely to become a source of trouble to pupils. Special attention

should be called to the need of amending the study on seeds and

seedlings. Here the most unfortunate misconception is the treat-

ment of the maize kernel as a seed, the funiculus and hilum of

which the student is expected to find. F. L. S.

Notes. — Professor Engler has issued the first of a series of

“Erganzungshefte” to the phanerogamic portion of Engler and

Prantl's Die Natiirlichen Pflanzenfamilien, containing additions for

the years 1897-98. The publication of the regular fascicles of this

* Pepoon, Herman S., Mitchell, Walter R., and Maxwell, Fred B. Studies of

p lant Life, a series of exercises for the study of plants. Boston, D. C. Heath &

Co. 1900. 12mo, cloth, xii + 95 pp.

No.412.] REVIEWS OF RECENT LITERATURE. 325

important work has now reached Lieferung 201, dealing with the

Hyphomycetes.

“A Taxonomic Study of North-American Ranunculacez " is the

title under which Dr. K. C. Davis publishes his thesis, submitted to

the Faculty of Cornell University last June. It consists of a series

of articles separately printed from various journals, some of which

have already been noticed in the Naturalist.

A posthumous revision of the genus Matthiola, by Pascal Conti,

accompanied by a portrait of the author, appears in No. 18 of the

Mémoires de P Herbier Boissier.

The Oxalidacee of Uruguay receive apparently careful treatment

by Arechavaleta in fascicle 14 of the third volume of Anales del

Museo Nacional de Montevideo. Thirty-five species are described.

A study of the leaf anatomy of the Melastomacez constituting

the tribe Miconiez, with reference to the classification of the plants,

forms No. 19 of Mémoires de l’ Herbier Boissier.

Bulletin 175 of the North Carolina Experiment Station contains

descriptions of twenty-one species of Crataegus and eight species of

Panicum, believed to be new.

Rubus ideus anomalus is reported from Vermont by Fernald in

Rhodora for October.

Celtis pumila, the separability of which from the arboreous species

of the Mississippi valley has long been in dispute, is critically consid-

ered, in connection with its allies, by E. J. Hill, in the Bulletin of

the Torrey Club for September.

Tig Revue Horticole for October contains an interesting article by

IS on the Mexican forms of Persea gratissima, with a colored

plate of one of the finer varieties.

n The Physiological differences between the sessile- and pedunculate-

‘Tuited English oaks are considered at some length by W. R. Fisher

In the Gardener s Chronicle of September 22.

5 No. 14 of Dr. Holm's « Studies in the Cyperacez," in Zhe Amer-

es Journal of Science for October, refers chiefly to a collection of

Carices made in Alaska by Evans in 1897-98.

Part XIV

of Kraenzlin's *Orchidacearum Genera et Species,"

recently issu

ed, reaches page 896.

326 THE AMERICAN NATURALIST. [Vor. XXXV.

A monograph of the Erysiphacez, by Ernest S. Salmon, constitutes

the ninth volume of Memoirs of the Torrey Botanical Club, bearing :

the date Oct. 4, 1900. Nine plates of details, containing 175

figures and a very full bibliography and host-index, with analytical

keys for each genus, make the book easy to use.

A preliminary report on diseases of the red cedar caused by

Polyporus juniperinus and P. carneussby Dr. Von Schrenk, constitutes

Bulletin No. 2r of the Division of Vegetable Physiology and Pathol-

ogy of the United States Department of Agriculture. The first of

these fungi is described as new. The gelatine process-reproductions

of photographic illustrations of the decayed timber are good examples

of what may be done in illustration direct from the camera.

Professor Nelson publishes, as a separate from the tenth Aor?

of the Wyoming Experiment Station, a preliminary list of the crypto-

gams of that state, exclusive of Pteridophytes.

An interesting article on the occurrence of mycorrhiza on Arctic

plants is reprinted by Hesselman from Bihang till K. Svenska Vet-

Akad. Handlingar, Vol. XXVI.

. The structure of the diatom girdle is discussed by Palmer and

Keeley, in Part II of the current volume of Proceedings of the Phila-

delphia Academy.

Five new species and one variety, chiefly in Allium and Artemisia,

are added to the Colorado flora by Osterhout in the September

number of the Bulletin of the Torrey Club.

Part III of Thomas Howell’s Flora of Northwest America, dated

August 21, 1900, reaches part way through the Composite.

: A “Botany of Southern California," by Charles Russell Orcutt,

is begun in the West American Scientist for September.

A considerable number of new species of North-American plants

are published by several writers in recent numbers of Zoe.

Professor F. Manson Bailey’s contributions to the flora of Queens-

land, New Guinea, etc., continue to appear in current numbers of

the Queensland Agricultural Journal.

Dr. Carreiro, the most active botanist now resident in the Azores

contributes a list of his recent discoveries to the October-November

number of the Buletin de | "Académie internationale de geogri aphie

botanique..

No. 412.] REVIEWS OF RECENT LITERATURE. 427

Volume VI of the * Flore de France" of Rouy and Foucaud,

devoted to Rosacez, appears as the 1899 Annales de P Académie de

La Rochelle, and is ascribed to Rouy and Camus.

The public school department of Carthage, Mo., where nature

study of the most practical kind has taken firm root through the

efforts of Professor Stevens, the superintendent, has now begun the

publication of a series of “Nature Study Leaflets,” dealing with

common objects.

Progress in American agriculture and the sciences upon which

it rests is well sketched in the voluminous Yearbook of the United

States Department of Agriculture for 1899, recently issued.

Part IX, recently published, completes the second volume of the

Bulletin of the Bussey Institution, which has been in progress since

the year 1877. A table of contents and an index make the subject-

matter of the volume accessible.

Forestry in Sweden is reported on by General Andrews, late

United States Minister to that country, in a Senate Document

recently issued.

The conditions of success in grafting are discussed by Daniel

in current numbers of the Revue générale de botanique.

Dr. P. Van Romburgh publishes an important paper on Caout-

chouc and Gutta-percha in the Dutch Indies, as No. 39 of the

Mededeelingen of the Buitenzorg Botanic Garden.

A biographical sketch, with portrait, of Klatt, whose writings on

Iris and certain Composite are familiar to all working botanists, is

reprinted by Voigt from the Jahrbuch der Hamburgischen Wissen-

schaftlichen Anstalten for 1898.

A portrait of the late Sir J. B. Lawes, whose field experiments on

the physiology of agricultural plants, extending over many years,

are known to all botanists, appears in Science Gossip for October.

PALEONTOLOGY.

Traquair’s Presidential Address, Bradford, 1900. — The Zoó-

logical Section of the British Association, in its choice of Dr.

Traquair for president, paid a graceful tribute to a scientist whose

researches during the past thirty-five years have conduced more

328 THE AMERICAN NATURALIST. — [Vor. XXXV.

to our knowledge of Paleozoic fishes than those of any other

single investigator. And the propriety of electing a paleontologist

to preside over its sessions is abundantly confirmed by the brilliant

address of sixteen closely printed pages with which its sittings were

opened.

In this essay Dr. Traquair has done for the class of fishes what

Marsh attempted for American fossil vertebrates in general, and

what Osborn accomplished for the Mammalia, in their addresses

before the sister association! in this country some years ago ; and

these three summaries, taken together, constitute a very important

chapter in the literature of vertebrate paleontology. Although not

retrospective in an historical sense, Dr. Traquair's paper is in effect

a clarified review, expressed in terms of evolution, of the leading

philosophic deductions which the science of ichthyic paleontology

has afforded up to the present time. Its significance consists in a

clearer recognition of the relationships between different groups of

fishes, together with a more precise indication of their lines of

descent; and it contains also critical observations on various dis-

puted points, such as the origin of paired fins, development of dental

plates and dermal armor from shagreen-like scales, atrophy of the

lower jaw and shoulder-girdle, modification of the caudal fin, and

similar issues. And in conclusion it is stated that *the paleon-

tology of fishes is not less emphatic in the support of the doctrine

of descent than that of any other division of the animal kingdom”;

also that “we do not and cannot know the oldest fishes, as they

would not have had hard parts for preservation, but we may hope to

come to know many more old ones, and older ones still, than we do

at present."

The author devotes special consideration to those ancient forms

whose structure he has so ably elucidated in earlier memoirs. AS

for the ostracoderms, a most enigmatical group, which he at one

— supposed were derived from the primitive elasmobranch stem

(owing to the resemblance of Thelodus scales to shagreen), an

independent origin is now admitted to be possible; but the idea is

discredited that they had any share in the evolution of more recent

types of fishes. The Ceelolepide, to which **Cephalaspis as well

as Pteraspis and its allies are traceable," were certainly shark-like |

1) i .

P Marsh, O. C. Introduction and Succession of Vertebrate Life in Am

roc. Amer. Assoc. Adv. Sci., Nashville Meeting, 1873. Madison

sborn, H. F. The Rise of the Mammalia in North America, iid.

Meeting, 1893.

No.412.] REVIEWS OF RECENT LITERATURE. 329

creatures, even if they were not offshoots from some primitive form

of shark.

A novel and somewhat surprising interpretation of the equally

problematical arthrodires is given, to the effect that they are teleo-

stomes belonging to the order Actinopterygii. Newberry, it is true,

imagined a resemblance to exist between Macropetalichthys and

recent sturgeons ; but we know now that this genus has nothing to

o with coccosteans, and Newberry totally misapprehended its

structure. The lack of a shoulder-girdle and lower jaw (properly

speaking) in all coccosteans, — an anomaly which they share in

common with ostracoderms, — and the peculiar arrangement and

mode of growth of the dermal plates, are characters which, to our

mind at least, are irreconcilable with actinopterygian affinities.

Although functional “mandibles” and “maxillaries” are often

present, yet they are evidently only modified dermal plates of the

same nature as the body armoring; and the former are suspended

in soft tissues without any articulation with the cranium or other

bones whatsoever. They are no more homologous with the jaws of

bony fishes than is the so-called *clavicular" with a pectoral arch.

Bashford Dean would exclude arthrodires from fishes altogether, as

Cope did the ostracoderms ; and elsewhere a tendency may be noticed

looking toward the revival of M'Coy's Placodermata.

Turning to the elasmobranchs, Dr. Traquair again surprises us

by declaring that their paleontological history does not throw any

definite light on the disputed origin of paired limbs or “on the

question whether the so-called archipterygium is the primary form

of paired fin in the fish, or only a secondary modification." Never-

theless, he cautiously admits that *the paired fins of the Upper

Devonian shark, Cladoselache, as described by Bashford Dean,

Smith Woodward, and others, seem to favor the lateral fold theory."

Next follows a very interesting and very learned discussion of the

Crossopterygii, the Dipnoi, and Actinopterygii, into which, however,

Space forbids us to enter. His conclusions regarding the first two

Eroups are that *the Crossopterygii were not derived /row the

Dipnoi, and that the modern representatives of the latter group are

degenerate forms; yet as to the immediate ancestry of the Dipnoi

themselves and the diphyletic origin of the so-called archipterygium,

we had best for the present keep an open mind." It need scarcely

be added that this lucid and timely essay of Dr. Traquair's is of

Prime importance, not only to paleontologists, but to zodlogists

everywhere. CRE

330 THE AMERICAN NATURALIST. [ VOL. XXXV.

Triassic Fossils from Eastern Siberia.' — In this publication, Dr.

A. Bittner has described the fragments of a pelecypod and beachiopod

fauna from the Lower Trias of the Ussuri region, near Vladivostock,

in eastern Siberia. Although the fauna is not rich, it is especially

interesting, coming from the same formation from which Dr. C.

Diener ° has already described the characteristic cephalopod fauna.

While most of the species are new, it is surprising to find among

them forms either identical or closely related to European forms,

among which may be mentioned Pecten conf. alberti Goldfuss, Pseudo-

monotis multiformis Bittner, and Myophoria conf. levigata Alberti,

of which representative forms, or geographic variations, are found

in the Werfen beds of the Tyrol.

Diener, on the other hand, found the cephalopod fauna of the

Ussuri region absolutely different from that of the Tyrol, with its

nearest relationships rather with the Triassic faunas of northern

Siberia and India. It is, therefore, remarkable that a kinship with

the European Trias should show itself in the pelecypod fauna, which

also shows decided affinities with some forms in the Lower Triassic

Meekoceras beds of southeastern Idaho, near Soda Springs. The

cephalopod fauna of the Ussuri region is almost identical with that

collected, but not yet described, by Professor A. Hyatt and the

writer from Wood’s Cafion near Soda Springs. These additional

species will aid greatly in the correlation of the Idaho beds with

those of the Ussuri region, and incidentally with the Mediterranean

Trias. TPS

The Upper Paleozoic Fauna of Russia.?— In this paper N. Jakolew

has added another link to the chain that connects the Upper Car-

boniferous faunas of Russia with those of America. Up to the pres

ent our knowledge of the gastropods of the Permo-Carboniferous,

or Artinsk, beds of Russia has been scanty, but in this work we find

à number of American species in the beds that lie above the Uralian

and below the true Permian beds, analogous to the uppermost

Missourian and possibly the lower part of the Wichita formation of

the Mississippi valley region. The analogies between these transi-

tional formations on the two continents have long been recognized,

A. Versteinerungen aus den Trias-Ablagerungen des Süd-Ussuri-

der ostsibirischen Küstenprovinz, Mém. Comité Géol. Russie, vol. Vb

2 Mém. Comité Géol. Russie, vol. xiv, No. 3, 1895.

N. Die Fauna einiger oberpalaeozoischer Ablagerungen Russlands.

alopoden und Gastropoden, Mém. Comité Géol. Russie, vol. xv, No. s

! Bittner,

Gebietes in

No. 4, 1899.

3 Jakolew,

I, Die Ceph

1899.

No. 412.] REVIEWS OF RECENT LITERATURE. 331

but every bit of additional evidence is welcomed. It all tends to

show that in the time of the uppermost Coal Measures the great

western ocean had extensions as far as the Mississippi valley on

the one side, and to eastern Russia on the other, since the brachio-

pod, gastropod, and cephalopod faunas of the two regions have many

species in common. Dr. Jakolew also announces the publication,

in the near future, by Professor Tschernyschew, of an extensive

work on the brachiopod faunas of the Upper Carboniferous of

the Ural region, in which correlations will be made with the American

formations and faunas. LPS

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS, AND DEATHS.

EDUCATIONAL GIFTS.

American Museum of Natural History, $25,000, by the will of Oswald

Ottendorfer ; $5000, by the will of Henry Villard.

Amherst College, $75,000, from D. Willis James and C. M. Platt ; $25,000,

from other sources.

Augustana College (Illinois), $30,000, from E. C. and J. A. Erickssen.

Aurora College (Illinois), $50,000, from Andrew Carnegie.

Baptist Female University of Raleigh, N. C., $20,000, by the will of Chief

Justice Faircloth.

Bates College, $10,000, by the will of Joseph Ricker.

Beloit College, a conditional gift of $200,000, from Dr. D. K. Pearson.

Bowdoin College, $20,000, by the will of Joseph Ricker.

Brown University, $250,000, from John D. Rockefeller, on condition that

the endowment be increased to $2,000,000 ; $30,000, from two anony-

u e

Carleton College, $50,000, from Dr. D. K. Pearson ; $100,000, from other

sources.

Carson and Newman College (Tennessee), a conditional gift of $15,000;

from John D. Rockefeller. :

Castine, Maine, $100,000, by the will of Charles F. Emerson, for a public

rary.

Chicago University, $1,500,000, from John D. Rockefeller ; $25,000; from

ell.

.eon Ma

Colorado College, $50,000, towards the completion of the scientific building.

Cooper Union (New York), $20,000, by the will of Oswald Ottendorfer.

Harvard University, $20,000, by the will of the late Governor Roger

Wolcott; $50,000, by the will of Henry Villard.

Johns Hopkins University, real estate valued at $750,000, from William

Wyman and Francis W. Jenks, upon condition that $1,000,000 be col-

— for the University. We understand that nearly the whole amount

as been subscribed.

Lehigh University, $5000, from Warren A. Wilbur.

Malden (Mass.) Public Library, $125,000, from Elisha H. Convers

vig Free Circulating Library, $20,000, by the will of mecs Otten-

orfer.

Northwestern University, $30,000, from Dr. D. K. Pearson.

GIFTS, APPOINTMENTS, RETIREMENTS. 333

Oberlin College, a conditional gift of $200,000, from John D. Rockefeller.

Rutgers College, $25,000, by the will of J. A. Vanderpool.

Syracuse Public Library, $200,000, from Andrew Carnegie.

Syracuse University, a conditional gift of $400,000, from John D. Archbold.

The University of Buffalo, $50,000, for the erection of the Gratwick

Research Laboratory.

University of Pennsylvania, funds yielding $400, for a prize in the school of

biology, from an anonymous friend.

University of Wisconsin, $1000, from Charles F. Pfister; $1000, from

Fred Vogel.

Upper Iowa University, $225,000, from Andrew Carnegie.

Washington and Jefferson College, $60,000, from W. J. Thompson, for

library building and equipment.

Wellesley College, $25,000, from H. H. Hunnewell, for the botanical

department.

Wesleyan University, $2000, from G. W. Quereau, for the library ; $1000,

from C. H. Buck.

Yale University, $50,000, by the will of Albert E. Kent of Chicago;

$80,000, by the death of B. D. Silliman ; a conditional gift of $500,000,

by the will of T. B. Winthrop ; $68,152, from the estate of the late

John De Koven ; a part of the estate of the late Professor E. E. Salis-

bury, estimated at $150,000; $30,000, from Alfred Vanderbilt ;

$50,000, by the will of Henry Villard. i

APPOINTMENTS.

Mr. Outram Bangs, assistant in mammalogy in the Museum of Compar-

ative Zoölogy at Cambridge, Mass. — Professor Franz Boas, curator of

ethnology in the American Museum of Natural History. — Dr. H. Borut-

tan, professor of physiology in the University at Göttingen. — Professor H. C.

Bumpus, curator of invertebrate zoölogy and assistant to the president in the

American Museum of Natural History, New York city. — Dr. A. Burian,

docent for physiology in the University at Leipzig. — Mr. E. J. Butler,

official botanist to the Indian government. — F. M. Chapman, assistant

Curator of mammals and birds in the American Museum of Natural History.

ee. F Czapek, docent for anatomy and physiology of plants in the

German University at Prag. — Dr. Paul Eisler, professor extraordinary of

anatomy in the University at Leipzig.— Dr. M. Fürbringer of Jena, pro-

fessor of anatomy in the University at Heidelberg, as successor to Gegen-

baur. — Myron L. Fuller, assistant geologist on the United States Geologi-

cal Survey, — Joaquin Gonzalez Hidalgo, director of the Natural History

Museum at Madrid.— Dr. Grabowski of Braunschweig, director of the

zoólogical gardens at Breslau. — Dr. L. P. Gratacap, curator of mineralogy

in the American Museum of Natural History. — Dr. Valentin Hacker of

Freiburg i. B., professor of zoólogy in the Stuttgart Technical School. —

334 THE AMERICAN NATURALIST. (Vor. XXXV.

. Miss Clara Hamburger, assistant in the zoólogical institute of the Univer-

sity at Breslau. — Mr. G. T. Hastings of Cornell, teacher of science in the

English Institute at Santiago, Chile. — Dr. Erich Kaiser of Bonn, geologist

of the Prussian geological survey in Berlin. — Dr. Friedrich Klein, professor

extraordinary of physiology in the University at Kiel. — A. G. Leonar

of the University of Missouri, professor of mining and mineralogy in the Uni-

versity of Idaho, at Moscow, Idaho. — Dr. G. S. Lingle, professor of experi-

mental physiology in Rush Medical College. — Dr. Rudolf Martin, professor

of physical anthropology in the University at Zürich. — Dr. Anton Nestler,

professor extraordinary of botany in the German University at Prag. — Dr.

F. Reinke, professor extraordinary of anatomy in the University at Rostock.

— Dr. Arthur Robinson, professor of anatomy in King's College, London. —

Dr. A. Sauer of Heidelberg, director of the geological survey at Stuttgart.

— Dr. Marshall H. Saville, curator of Mexican and Central American

archeology in the American Museum of Natural History. — Dr. F. Schenck

of Würzburg, professor of physiology in the University at Marburg. — O.

Schneider, geologist of the Prussian géological survey. — Dr. Oscar Schulz,

privat docent for physiology in the University at Erlangen. — Dr. H. Stech-

mann, director of the Breslau zoólogical gardens. — Dr. L. Steigert, geolo-

gist of the Prussian geological survey. — Dr. Alexander Steuer, docent for |

geology and paleontology in the Darmstadt technical school.— Dr. H.

Stille, geologist of the Prussian geological survey. — Dr. Fr. K. Studnitka,

dócent for zoólogical histology in the Bohemian University at Prag. —

Arthur Thompson of Oxford, professor of anatomy in the Royal Academy,

London. — Dr. O. Tietze, geologist of the Prussian geological survey. —

Dr. A. Tschermak, assistant in the physiological laboratory of the Uni-

versity at Leipzig. — Professor G. Vasseur, conservator of geology in the

Museum of Natural History at Marseilles. — Professor A. Vayssiere, con-

servator of zoólogy in the Museum of Natural History at Marseilles. — Dr.

Max Verworn, professor extraordinary of physiology and director of the

physiological institute at the University at Góttingen. — Dr. A. Voelzmann,

professor extraordinary of zoólogy in the University at Berlin. — Dr. Vogel

of Hamburg, bacteriologist in the agricultural experiment station in Posen.

— Dr. Charles H. Warren, instructor in mineralogy and geology in the

Massachusetts Institute of Technology. — Dr. G. Wetzel, docent for anat-

M€— in the University at Berlin. — Dr. E. Woloszczak, professor of botany

in the technical school at Lemburg, Austria. — Dr. Fred. C. Zapffe, pro-

fessor of histology in the medical school of the University of Illinois. —

Dr. Theodor Ziehen of Jena, professor of psychology in the University at

Utrecht. — Dr. W. D. Zoethout, laboratory professor of neurology in Rush

Medical College. :

No.412.] GIFTS, APPOINTMENTS, RETIREMENTS. 335

RETIRED.

Sir A. Geikie, from the directorship of the geological survey of Great

Britain. — Dr. Friedrich Goltz, professor of physiology in the University of

Strassburg. —Dr. Georg Meissner, professor of physiology in the University

at Góttingen. — Dr. E. von Mozsisovics of Vienna, from the position of

vice-director of the Austrian geological survey. — Dr. Max Reess, professor

of botany in the University at Erlangen. — John C. Smock, for ten years

state geologist of New Jersey. — Dr. Filippo Sylvestri, from the position of

chief of the zoólogical section of the National Museum of Buenos Aires.

DEATHS.

Dr. Theodor Adensamer, zoólogist, in Baden, near Vienna, November

I6, aged 33. — Dr. . von es, emeritus professor of botany in the

Stuttgart technical school, in August. — Dr. Richard Altmann, professor

extraordinary of histology in the University of Leipzig.— F. R. Bedford,

student of echinoderms, in London. — Dr. G. Boerlage, assistant director of

the botanical gardens at Buitenzorg, Java, in September, while on an expe-

dition to Ternate.— Dr. Breusing, assistant in geology in the Hannover

technical school, while on a trip in Dutch Guiana. — Mr. Philip Crowley,

entomologist and ornithologist, in London, December 20. — S. W. Egan, of

the geological survey of Ireland, at Dublin, January 6. — Dr. Gustav Hart-

laub, the celebrated ornithologist, at Bremen, November 20, aged 87.—

r. R. Hegeler, privat docent for botany at Rostock, at Stuttgart, Septem-

ber 28.— Dr. George Duryea Hulst, the well-known entomologist, in

Brooklyn, N. Y., November 5, aged 54.— Dr. William King, for thirteen

years connected with the geological survey of India. — Dr. Philip H. Kirsch,

ichthyologist, at Las Cruces, New Mexico. — Dr. S. J. Korschinski, bota-

nist, at St. Petersburg, December 1.— Mr. R. D. Lacoe, the well-known

collector of carboniferous fossils, at West Pittston, Pa., February 5. — John

Henry Leech, entomologist, at Salisbury, England, December 29, aged 38.

— Thomas Benton Marbut, geologist and mining engineer, at Esther, Mo.,

September 16, aged 29. — Dr. W. von der Marck, geologist, in Hamm,

Germany, November 22, aged 86.— Dr. Paul Marés, botanist, May 21,

1990, at Mustapha, near Algiers. — Charles Marquet, entomologist of the

Museum of Natural History at Toulouse. — John Potter Marshall, emeritus

professor of geology and mineralogy in Tufts College, February 4, aged 76.

ofessor Max von Pettenkoffer of the University dt Munich, by suicide,

in January, aged 83. — Professor Dr. Adolf Pichler, pu of the Tyrol,

at Innsbruck, November 1 5, aged 82. — E. Roze, botanist, at Chatou, France,

May 25, 1900. — Michael Edward, Baron de Selys-Longchamps, entomolo-

gist, well known for his studies of the Odonata, at Liége, Belgium, Decem-

ber 11, aged 86. — Dr. Otto Staudinger of Dresden, an eminent student of

Be THE AMERICAN NATURALIST.

Lepidoptera, at Luzern, October 13, aged 7o. — Dr. Hermann Stechm

director of the Breslau Zoö logical Gardens. — Dr. Otto Martin Torell

23, aged 35.— Dr. Friedrich Anton Zürn, a well-known investiga!

animal parasites, September 11, aged 65.

(Wo. 411 was mailed March 15.)

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CONTENTS

I. Two New Myrmecophilous Genera of Aberrant Phorids from Texas

CHARLES T. BRUES

II. The vocan a the Hexapoda and their Relation

. L. B. WALTON

III. S een bony in the Postembryonje Develogiusnk of ra e

Professor N L. KELLOGG

IV. ide squid sateen CERE.

e H. 8. JENNINGS

V. Synopses of North-American Invertebrates. XIV.

RH — Part II . unos HM. © W. HARGITT

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eere of Ferns, The Flora of Celebes, Notes — -Pilona

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Dr. LEONHARD STENEGER

un Publications Received.

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THE

AMERICAN NATURALIST

Vor. XXXV. May, rgot. No. 413.

TWO NEW MYRMECOPHILOUS GENERA OF

ABERRANT PHORIDZE FROM TEXAS!

CHARLES THOMAS BRUES.

THE discovery of two new genera of myrmecophilous diptera

allied to the Stethopathidz, in Texas, is entirely unexpected.

That a family of wingless Diptera hitherto represented by a

small group of genera in the Old World should suddenly appear

in North America is, to say the least, a very unusual occurrence.

The new genera are also very interesting from a taxonomic

point of view, as they show very clearly the affinities of the

Stethopathidze and Phoride, which have hitherto appeared

Somewhat doubtful.

Last October, while collecting ants in one of the pecan

Sroves upon the outskirts of Austin, Texas, Dr. Wheeler of

the University of Texas found in a nest of Solenopsis geminata

Fabr. several specimens of a peculiar dipteron with rudimen-

tary wings. Upon closer examination they seemed to be

Phoridze, although their habitus was extremely like that of the

Stethopathidz, from which they were easily distinguished by

their halteres and small wings.

1 E. VENUS

Contributions from the Zoblogical Laboratory of the University of Texas, No. 15.

338 THE AMERICAN NATURALIST. (VoL. XXXV.

About a month later Dr. Wheeler called my attention to a

number of somewhat similar flies in a nest of Eciton caecum

Latr. which had been in the laboratory for some time. These

also proved to be Phoridz, but of a still more degenerate type.

A more careful examination has shown that these two genera

are referable to the Stethopathidz, but at the same time are

evidently degenerate Phoridz, so that the former family must

be included in the Phorida. Such an addition does not make

the family an incongruous one. When Loew (57) placed his

African Psyllomyia in the Phoridz he made a statement equally

applicable to the present addition :

“Wenn irgend etwas geeignet ist über die verwandtschaft-

lichen Beziehungen der Phoriden eine Aufklärung zu geben, so

sind diess Arten, welche so sehr von dem Typus der in der

alten Gattung PZora vereinigten abweichen, wie die oben be-

schriebene, und welche doch der Familie der Phoriden mit so

bestimmt ausgesprochener Entschiedenheit angehóren wie sie.

* Leider muss ich bekennen, dass die oft wiederholte Ver-

gleichung der Psy//omyia testacea mit Diptern gar verschiedener

Familien mir nach dieser Richtung hin durchaus kein positives

Resultat gegeben hat, so dass ich die Familie der Phoriden von

allen andern Familien der Diptern noch so scharf getrennt und

so unvermittelt zwischen ihnen stehen sehe, wie zuvor."

With the discovery of the still more degenerate forms his

remarks lose none of their pertinence.

The family Stethopathidze was established by Wandolleck in

1898 ('98b) for the reception of several genera of Diptera which

are remarkable for the total absence of wirigs and halteres,

besides other less striking peculiarities. Two of the genera

which he describes had been previously described by Dahl (97)

as sexes of a single species which he named Puliciphora lucifera

and placed among the Phoridz. He, however, considered at

the same time that they were a connecting form between the

Aphaniptera and Diptera and accordingly announced that he

had solved the much-vexed question of the relation of the fleas

with the Diptera.

Wandolleck later showed (98a) that this idea is wholly

erroneous as far as the Aphaniptera are concerned, and that

No.413.] ABERRANT PHORID.£ FROM TEXAS.

339

they are in no way related to them, nor, on the other hand,

did he consider them to be Phorida.

His third genus, which he leaves without a name, had been

previously named in his honor by Cook ('98).

The most important characters of the Stethopathidz, as

defined by Wandolleck, are the total absence of wings and

halteres, the strongly reduced thorax, very small eyes, large

coxa, and small external genital organs of the female.!

Recently Wasmann has described a genus of termitophilous

Diptera (00a) which agrees with the Stethopathidz in all essen-

tials, except that they possess rudimentary wing-like appendages

and a slightly larger thorax. These are characters that would

not warrant the erection of another family for their reception.

Moreover, the presence of rudimentary thoracic appendages

places them still closer to the Phoride.

The two Texan genera approach the true Phoridae even more.

closely, since one of them possesses both halteres and rudi-

mentary wings, and both have the large macrochzte of the

body hairy, a character which Wasmann has pointed out as

distinguishing the Stethopathide and Phoride. In both of

these forms the abdomen is almost completely membranous.

The African genus Psyllomia, however, has an abdomen of the

true phorid type, while it agrees with one of the new genera,

Commoptera, in having rudimentary wings, as well as in the

Structure of the head. The external sexual organs of the

female are so clearly of the phorid type that they present no

important deviation.

Through these forms we can pass without any great gaps

from the Stethopathidae to the Phoride, and as such is the

case, the family Stethopathide is certainly untenable, and the

genera hitherto placed there, together with Psyllomyia and the

1I have not considered the form of the mouth-parts in Wandolleck's three

genera, which he believes to be entirely different from those of the Phoridz and

all other Diptera. He himself says, however, that Dahl, who has spent much

^ time in studying the Phoridz, considers them as phorid mouth-parts. He says:

“Ich zeigte die Zeichnungen Dahl, der sich seit Jahren mit Phoriden bescháftigt,

er erkannte sie sofort als Phoridenmundtheile an. . . .” I think Wandolleck

must have exaggerated the extent and importance of the variation in the mouth-

Parts, for the two Texan genera have typical phorid mouth-parts, often, however,

shrunken and distorted in alcoholic specimens. :

340 THE AMERICAN NATURALIST. [VoL. XXXV.

two described below, had best be considered as the subfamily |

Stethopathine of the Phoride.

It does not seem probable that the forms have had a common

origin however ; even their distribution would preclude this

in the case of such a recent group. On the other hand, there

seem to be several independent lines of descent which we can

to some extent suggest. In Psyllomyia the degeneration con-

sists in the reduction of the eyes, absence of ocelli, and reduc-

tion of wings. In Commoptera the eyes are larger, the ocelli

present ; but the abdomen is extremely degenerate in structure,

being almost wholly membranous. Ecitomyia could be more

easily derived from forms like Commoptera, as it has very

rudimentary wings, no halteres, smaller eyes, smaller thorax

(without scutellum), and about equally degenerate abdomen.

The degeneration has gone furthest in the genera from the

Bismarck Archipelago and west Africa. Here the thorax is

exceedingly small, the wings and halteres completely lost, the

eyes very small, and the abdominal segments more or less mem-

branous. Of these Stethopathus alone has retained distinct ocelli.

Wasmann’s Termitoxenia has the immensely swollen abdomen

so characteristic of all termitophiles, and a greatly modified head.

It does not approach closely to any of the other genera.

The following table, showing the order of degeneracy with

regard to different structures in the genera of the Stetho-

pathinze as compared with Phora, will serve to emphasize the

great disparity between the genera. Phora is placed at the top

in each case and the most degenerate genus at the bottom.!

Wings. |

EYES AND OCELLI. ABDOMEN. | THORAX.

no Te aa

Phora : Phora Phora Phora

Psyllomyia Commoptera Psyllomyia Commoptera

Commoptera Termitoxenia Chonocephalus | Psyllomyia

omyi Stethopathus Stethopathus Termitoxenia

Termitoxen Psyllomyia Ecitomyia Ecitomyi

Stethopathus Wandolleckia Commoptera tethopathus

Wandolleck Ecitomyia Wandolleckia Wandolleckia

Chonocephalus | Chonocephalus | Termitoxenia Chonocephalus

1 Where ocelli or wings are absent I have judged by the size of the eyes and

the thorax.

No. 413.] ABERRANT PHORIDA FROM TEXAS. 341

Still more remarkable is /Enigmatias, represented by Æ.

blattoides Meinert from Denmark, which does not seem to be

closely related to the Stethopathinz, although I have unfortu-

nately not had access to the original description of Meinert

(90). Its habitus is certainly entirely different from that of

any of the genera here mentioned. Platyphora lubbockii Verrall

has been suggested as the possible male of /Enigmatias, but

that is very problematical.

Throughout the winter we had been searching in vain for

the males of Ecitomyia, which is very common in the nests of

Eciton cecum, but not until spring (February 3) were we able

to obtain them. On that day we obtained two specimens from

different nests in which the females were abundant. A glimpse

at one of them immediately justified any surmises made as

to their phorid character, for the males, although much like

the females, possessed fully developed wings with the peculiar

phorid venation and large halteres!

Such a remarkable amount of variation in usually stable

morphological characters may be best explained by the great

tendency of all degenerate structures to vary in an unusually

high degree, and by a great power of adaptation in the

Phoridze.

The habits of only four of the genera are known with cer-

tainty. They are all myrmecophiles or termitophiles. A fifth,

Wandolleckia, lives apparently ectoparasitically upon large west

African land snails (Achatina variegata Roissy).

Their geographical distribution is extremely peculiar and is

a case of remarkable discontinuous distribution not due to the

great age of a certain type, for it does not seem possible to

regard them as an old group, but rather as several independent

and to some extent conveying lines of degeneration. This view

is strengthened by the above-mentioned impossibility of show-

ing any interrelation of the genera.

The following dichotomy will serve for the identification of

the genera thus far known.

342 THE AMERICAN NATURALIST. [Vor. XXXV.

STETHOPATHIN&.

Wingless or with rudimentary wings, eyes reduced, abdominal plates usu-

ally much reduced, coxa very long, face deeply concave, eggs usually very

large and causing an enlargement of the abdomen. Males, as far as posi-

tively known, easily correlated with the females, but much like the males of

the Phorine.

GENERA OF STETHOPATHINA (FEMALEs).!

I. Wings and halteres absent . i ; . T ; . . 5

Either wings or halteres or both present `. : i i : 2

2. Both wings and halteres present, the former abbreviated — . . 3

Either wings or halteres present, never both ; à . .

Arista of antenna dorsal, abdomen strongly chitinized throughout, pro-

boscis much longer than height of head, very slender

Psyllomyia Lw.

Arista terminal, abdominal segments small, the greater part of the abdo-

men membranous, proboscis shorter than height of head, stout

Commoptera gez. nov.

Abdomen greatly swollen, its apical segments bent forward on the ven-

tral side of abdomen. Wings absent, halteres present

: Termitoxenia Wasmann

Abdominal segments normal in position, wings present, halteres absent

Ecitomyia gez. nov.

Ocelli present s : 3 ; i : Stethopathus Wand.

Ocelli absent i à è í à i : : 6

Head longer than wide, subtriangular, palpi when viewed from above

extending far beyond anterior margin of head. Two macrochete on

posterior margin of head .- . ; ; Wandolleckia Cook

Head wider than long, trapezoidal, palpi not extending forward beyond

front margin of head : ; : Chonocephalus Wand.

If we desire to consider /Enigmatias as belonging here, it

may be separated from all the other genera by its stout blattid-

like form, with the thorax as wide as the abdomen. The

problematical male (Platyphora) differs from the male of Ecito-

myia in having no macrochzetze on the dorsum of the thorax.

1 The male of only one genus (Ecitomyia) is known and it may be recognized

by the diagnosis given farther on.

No. 413] ABERRANT PHORID4E FROM TEXAS, 343

PSYLLOMYIA LOEW.

Head swollen, lentiform, completely chitinized. Eyes extremely small,

somewhat wider than high, situated on the sides of the head. Ocelli absent.

Antenne two-jointed, only of medium size, each situated in a cavity, sepa-

rated by the front margin of the head, which extends somewhat between

them. Arista very thickly clothed with moderately long hairs. Palpi pro-

jecting, not very stout, bristly along the lower side and at the tip. Pro-

boscis very long, geniculate, without distinct labella. Thorax rounded.

Abdomen and legs as in Phora, the latter very bare, only at the tips of the

four posterior tibiae, with short, small bristles. Wings abbreviated, leathery,

resting upon the dorsum of the abdomen and having the general appearance

of short elytra (about asin Meloé). They have

indications of three very thick, rib-like longi-

tudinal veins, which are beset with small black

bristles, some of which are noticeable because

of their much greater length. Halteres be-

neath the wings, almost rudimentary.

Psyllomyia testacea Lw. (Fig. 1). Length

1.75 mm. Pale brownish yellow. Arista and

wings, with the exception of the longitudinal

veins, more yellowish white ; the entire abdo-

head finely punctured and covered with

hardly perceptible hairs ; besides these there

are two posteriorly directed black macrochete

on the extreme anterior part of the head, two

anteriorly directed ones on each side close to

the base of the proboscis, one anteriorly

directed one immediately in front of the eye,

and four posteriorly directed ones upon the

occiput, close to the rather acute margin of the

head. Thorax with scattered black bristles,

one of especially large size on the side of the thorax at the upper part of the

front coxa. On the wings two bristles are especially large, one of them

on the inner margin, the other near the apex.

Fic. 1.— Psyllomyia testacea Lw.,

Q. (After Loew.)

The specimens described by Loew were collected in Kaffir

Land, Africa. Of all the genera this most closely resembles

the Phorinz, yet it shows undoubted affinities with Commop-

tera. The long proboscis is apparently quite different from

those of the other genera.! The antenne have a dorsal arista,

1 Wasmann, however, mentions that Dorniphora Dahl, one of the Phorinz,

possesses a somewhat similar proboscis.

344 THE AMERICAN NATURALIST. | [Vor. XXXV.

which is evidently homologous to the three apical joints in the

antennz of the other genera! Such a variation in the

insertion of the arista is seen also

in the Phorine. The palpi and

eyes are very small. The wings are

shaped much as in Commoptera, but

are almost free from bristles along

the costa. The legs are of the usual

phorid type, and judging from the

figure the coxze must be exceedingly

lengthened.

Nothing whatever was known by

Loew of the habits of this peculiar

form, but Wasmann has mentioned it

('00b) as the guest of a south African

Fic. 2. — Commoftera venit doryline ant (Dorylus helvolus Linn.).

pes homes hrar He also includes in his list of the

guests of the South-American Eciton predator “ Phorid N.G.

n. sp. (prope P. testacea Lw.), S. Catharina.”

COMMOPTERA SOLENOPSIDIS (gez. et sp. nov.).

Female (Figs. 2 and 3). Length 1.5 mm. Abdomen r.or mm. Thorax

.26 mm. Head .45 mm. Halteres.1g mm. Pale yellow, head somewhat

darker, and abdomen somewhat lighter, legs concolorous with thorax. Body

everywhere more or less covered with fine hairs. Head, seen from the side,

oval Vertex gently descend-

ing, about two-thirds as long

as the mesonotum ; face as

Close to the mouth, in elongate

vertical depressions which are

quite shallow. Antenne of

the usual form, arista termi-

nal, strongly pubescent. Eyes

oval, one-half as high as the

head and two-thirds as wide as

high, with the ommatidia sep-

arated and convex. They are

Fic. 3. — Commoptera wien n.sp. Female,

ds de

! Wandolleck speaks of them as antennal joints, but such a nomenclature

seems inconsistent with the one usually adopted.

No. 413.] ABERRANT PHORID.£ FROM TEXAS. 345

placed slightly above the middle of the head. Ocelli large and well

developed, occupying their usual position in a triangle upon the vertex.

Palpi as long as eye, small and slender, with the usual macrochaetze which

are also smaller. Proboscis two

arge, downwardly directed macro-

cheta. A pair of closely approxi-

mated, anteriorly directed bristles on

anterior margin of front; another

posteriorly directed pair on the poste-

rior margin of the head ; and a third

outwardly directed pair near the

posterio-lateral corners of the head.

Mesonotum somewhat wider than long,

with four outwardly directed marginal macrochætæ on each side. Scutellum

small, convex, much rounded behind, less so in front, projecting somewhat

over the metathorax behind, with a pair of strong approximated median

macrochetz. Posterior part of metathorax produced back into the abdomen

like a very large scutellum. Seen from the side the thorax is oval, not much

larger than the head. The anterior coxa are exceedingly large and freely

movable. The other coxæ smaller, more or less connate with the thorax,

the hind pair extending back beneath the base of the abdomen. Protho-

racic stigma large and distinct. Wings short, less than one-third as long

as the body, rather narrow and pointed, subtriangular in outline (Fig. 5).

Along the costa are two rows of stout macrochætæ ; elsewhere the wings

are covered with fine, short hairs. Veins not sharply distinguishable from

the surrounding membrane, consisting of two longitudinal veins which coa-

lesce at the tip of the wing.

"Toward the base of the

wing the first furcates, as

does also the second. The

anterior branch of the

x second vein unites with

gor Se the posterior branch of the

Mur 5 first. Posterior branch of

Fic. 5.— Comntoftera solenopsidis, n. sp. second vein evanescent.

mone eme Two vestigial posterior

veins are present also. Halteres immediately behind the wings large,

consisting of three distinct joints, the basal two small, quadrate ; the third,

large, Stout, oval (Fig. 5). The third joint is almost as large as the palpi

and flattened behind, so that a sharp edge separates the plane part from

the remainder of the joint. Abdomen elongate oval, somewhat depressed,

finely hairy, membranous except for the small dorsal plates, which are

extremely rudimental. The first is a wide band, extending only one-third

Fic. 4. — Mouth-parts of Commoptera

solenofsidis,n. sp. a, ventral view;

4, lateral view.

346 THE AMERICAN NATURALIST. [Vot. XXXV.

across the abdomen; the second, four times as wide as long, oval; the

third, similar and much smaller; fourth, triangular, with a large pit in the

center. Sexual organs of the usual form. The central axis is broadly

rounded and bisetose at tip; lamella short, club-shaped, and rounded at

the tips, quite bristly. Last two abdominal segments with rows of marginal

bristles. The last two segments of the abdomen are retractile, as in Ecitomyia,

and capable of being exserted to a considerable extent. Legs moderately stout.

Hind metatarsi flattened, and with regular rows of transverse macrochete.

Three female specimens found in a nest of Solenopsis geminata Fabr.,

at Austin, Texas, Oct. 24, 1900, by Dr. Wm. M. Wheeler.

The structure of Commoptera is on the whole more degen-

erate than that of Psyllomyia. The eyes are larger and the

ocelli present, but the swollen membranous abdomen and

general habitus are at least a greater departure from the

phorid type, if not a mark of degeneracy.

The head and its appendages are much as in the genera

described by Wandolleck. The eyes are, however, less reduced

and the ocelli nearly of normal size. A most remarkable dif-

ference is seen in the proboscis, which is not long as in Psyllo-

myia. The mouth-parts do not differ to any extent from those

of some Phoras which I have examined.

The thorax and its appendages present nothing new, except

the peculiar condition of the wings. At first I thought it

possible that the wings were normally of the usual size and

had been bitten off around the edges. But this view is dis-

proved by two facts. In the first place, the wings are sym-

metrical and have the edges perfectly continuous. In only

two wings (out of six) did there seem to be any irregular or

notched places along the posterior margin. Secondly, the

extreme activity of the flies would make it impossible for

the Solenopsis, although it is quite an active ant, to gnaw off

the wings soas to present even a semblance of the perfect sym-

metry exhibited. We are then forced to conclude that such

abortion is natural and that the wings have been decreased in

size on account of the inconvenience they presented to the fly

while moving about in the galleries of the Solenopsis nest.

Wings would indeed be a great inconvenience in moving about

in the narrow galleries and quite an unnecessary burden, when

the legs are adapted to such wonderfully quick motions.

No.413.] ABERRANT PHORIDA& FROM TEXAS. 347

The condition of the abdomen is also remarkable. The seg-

ments have no doubt been reduced, independently of the

secondary swollen condition due to the immense eggs. Such

small abdominal plates fitted together in their normal position

would form an abdomen so utterly at variance with the size

required to perform its natural functions that we must consider

the segments reduced and the abdomen also enlarged, probably

by post-metamorphic growth. The difference in the size of

the abdomen in different specimens of Ecitomyia shows that

a considerable post-metamorphic enlargement occurs in that

species. The external sexual organs do not depart from those

of Phora, except that they may be slightly reduced in size.

The fourth abdominal segment differs from all the others in

having not a dorsal plate, but a chitinous ring, triangular in

shape and surrounding a membranous patch which probably

has a glandular function!

There can be no doubt that this peculiar insect is a true

myrmecophile, as the nest in which they were found contained

numerous individuals, most of which escaped on account of

their extreme activity. Although we have examined a great

number of similar Solenopsis nests, we have seen no other

Specimens, so that, in this locality at least, it is much rarer

than the genus living with Eciton.?

ECITOMYIA WHEELERI gez. et sp. nov.

Female (Figs. 6 and 7) Length 1.20 mm. Abdomen .93 mm. Thorax

16 mm. Head .14 mm. Wings .14 mm. Head and thorax yellowish

brown, much darkened above. Abdomen yellowish white, its small dorsal

plates darker, the first almost piceous. Legs concolorous with the lower

portions of the thorax. Head, seen from the side, subtrapezoidal, the front

gradually descending, nearly as long as the dorsum of the thorax. Height

of antennal cavity about equal to the front, about one-third as deep as high,

regularly arcuate. The antennz (Fig. 8) arising near the base of the cav-

Ity, of typical form : first joint small; second, large globose, obtusely pointed

attip; first joint of arista sm all, distinct; second, longer; third, nearly equal

! Owing to the limited number of specimens which we have of this species, I

could not examine its structure. It is no doubt similar to the one described at

length under Ecitomyia.

* Since writing the above, I captured another specimen in a nest of the same

ant (April 6th). ° .

348 THE AMERICAN NATURALIST. [VoL. XXXV,

to first and second together ; the terminal slender portion not very distinct

from the third joint, distinctly plumose. Eyes slightly smaller than second

antennal joint, oval, with about twenty facets. Palpi rather slender, arcuate

near base, as long as the front, bearing six

strong macrochete laterally and a few other

weaker ones. All of them, especially the

larger ones, distinctly and finely hairy.

Thorax smaller than the head when

viewed from the side and longer than high,

the suture between the pro- and mesothorax

distinct above on the Spira prothoracic

stigma distinct (Fig. 9, 7). Mesopleure

distinctly cadi. areolated below.

ings a little longer than the thorax, slen-

der, flat, and obtusely pointed at tip;

. covered on the dorsal side with short bristly

hairs. Dorsum with six macrochete : two

small humeral, two larger post-humeral, and

Metathorax small, concealed to a great

extent by the abdomen, which extends over

it almost to the meso-metathoracic suture ;

viewed from above through the abdominal

wall, it appears elongate, subtriangular, and rounded at the apex. Legs

rather stout, especially the coxze and femora. Anterior coxa two-thirds as

long as the femora and freely movable at base; the four posterior ones

more or less connate and not

solarge. Posterior metatarsi

enlarged and flattened, bear-

ing six transverse rows of

stout bristles (Fig. 9, c).

One well-developed spur on

posterior tibia. Legs every-

where covered with short

hairs

Abdomen elongate, oval,

acuminate, capable of being

exserted at the tip, so that

the last three segments may

be retracted into the abdo- Tec pos See sheet t

men or pushed out for a is

distance equal to one-half that of the remainder of the abdomen.

retracted it is about twice as long as the head and thorax taken together.

It is almost wholly membranous, only the very small dorsal plates

being chitinized. First dorsal plate trapezoidal, as wide as long, and

Fic. 6. — Ecitomyia wheeleri, n. sp

Female, dorsal view.

No. 413.] ABERRANT PHORIDA FROM TEXAS. 349

narrowed basally; second, semicircular; third, subtriangular ; fourth,

twice as wide as long, rectangular. All except the first are almost rudi-

mental. Just in front of the fourth plate there is a horseshoe-shaped piece

of chitin, enclosing a pit from which projects a papilla from

the interior of the abdomen (Fig. 9, e). Exterior sexual

organs consisting of a stout longitudinal axis, obtusely

pointed at tip, where it bears two macrochete (Fig. 9, a).

The two lamellz are attached laterally upon the sides of

the central piece near the tip. Lamella about as long as

the diameter of the central axis, elongate, gradually enlarged

toward the tip, where they are rounded ; covered with nu-

merous stout bristles. All the apical segments of the

abdomen bear several marginal macrochete.

Male (Fig. 10). Length .68 mm. ; of wing the same.

Body alutaceous. Thorax infuscated above. Abdomen

piceous on basal three-fourths above, except on the anterior

margins of the segments, where it is much paler. Antenne,

palpi, face, and legs pale testaceous. Anterior tibie black,

except at extreme base. Hypopygium more or less black.

Wings hyaline, veins pale. Head shaped much as in the

female ; eyes larger, not much smaller than in species of

Fic. 8. — Ecitomyia Phora ; ocelli present, large, in a triangle on the vertex.

n n: sP.. Head seen from the side, about twice as high as long.

; Chætotaxy the same as that of the female, except that the

most anterior pair of frontal bristles is shorter. The macrochætæ on the

in the female. Thorax

arched in front, more

than twice as long as

the head. Scutellum

well developed, bear-

with three marginal

Macrochete and a

Pair just before the

Scutellum. Legs

longer and more slen-

d d J

d bdomen, 9 :

er than those of the 8, apex of abdomen, Phora sp. (), 9: c, hind metatarsus, 9 ;

; di ttal section of ¥; 7, head; 2, pro-

at tip, hind trochan- thorax; 3, mesothorax; 4, metathorax; 5, abdomen.

Loreen d

Fic. 9. — Ecitomyia wheeleri, n. sp. a, apex of a

female, Coxæ bristly d, diagrammatic median sagi

ters each with a pair of very strong recurved spine-like macrochætæ. Wings

large, as long as the body, with a stout longitudinal vein which meets the

350 THE AMERICAN NATURALIST. [VOL. XXXV.

thickened costal margin near the middle of the wing, and three faint, oblique

longitudinal veins. Costal vein bristly along its entire length. alteres

about as long as the hind metatarsus, distinctly three-jointed. Abdomen

with six segments, the basal ones longest. It is chitinized above and mem-

branous beneath. Hypopygium large, exserted, asymmetrical.

Described from numerous female and two male specimens collected at

Austin, Texas, in the nests of Eciton cecum Latr. and Eciton schmitti

Emery, from October to

February, the males only

in February.

The head of Ecito-

myia is much as in

Commoptera, but is

sharply angled at the

posterio-lateral corners

and longer on the ver-

tex. The palpi are

flattened and appear

much thicker when

seen from above. The

eyes are considerably

smaller than in Com-

moptera. The thorax

is much wider than

long and has no scutel-

lum, while the meta-

thorax is wholly con-

cealed in the basal part

a, dorsal view; of the abdomen. The

appendages upon the

dorsum of the thorax in the female of this species do not seem

to be homologous with the similarly placed ones in Termitoxenia.

They approach more closely to wings, while those in Termi-

toxenia are, in structure at least, like the halteres of Commop-

tera. That these appendages are wings is proved by their

insertion evidently anterior to the meso-metapleural suture,

and still more positively by their structure, as the homology of

the thoracic segments is somewhat obscure. They are strap-

shaped, and not round in cross-section; the dorsal side is

Fic. 10.— Ecitomyia wheeleri, n. Sp., ó.

4, side view; c, hypopygium

No. 413.] ABERRANT PHORIDA FROM TEXAS. 351

bristly while the ventral side is bare; they show no traces of

any separate segments and articulations, whereas the halteres

of Commoptera do. The presence of wings and absence of

halteres are peculiar to this genus among all Diptera. Wings

are often absent and rarely both wings and halteres, but in no

other case are wings present without halteres. In Lvetmoptera

browni Kellogg (00) the wings are much as in Ecitomyia, but

the halteres, although somewhat reduced, are distinct. In

Termitoxenia Wasmann considers the appendages of the thorax

to be attached to the prothorax, which he believes to be greatly

enlarged and to cover the dorsum of the entire thorax. This

would certainly be an unusual development of the prothorax,

and it seems much more reasonable to suppose that they are

the halteres, or perhaps possibly reduced wings. It seems

highly improbable that a dipteron prothorax should have sud-

denly become so large and have developed wing-like appendages.

On the other hand, they are quite similar to the halteres of

Commoptera, and Wasmann’s appendices thoracicales could be

easily derived from halteres. In Termitoxenia they seem to

have taken on a new function, at least Wasmann so supposes

from their peculiar form. He suggests that they may be of

use as a means of attaching the animal to the body of the

termite in order to be carried about.

The abdomen has somewhat larger dorsal plates than Com-

moptera, but they are nevertheless very much reduced, being

scarcely visible from the side. The pit and papilla upon the

anterior part of the fourth dorsal plate are shown in sections

to be connected with a remarkable gland in the abdomen. The

dorsal plate of the segment is continued forward to form a

strongly chitinized ring which passes over into the plate pos-

teriorly Inside of this ring the integument is very thin and

delicate, and is folded in to form a pit, surrounded on all sides

by firm chitin. The bottom of the depression is swollen out

in the middle to form a papilla, which is evidently to some

extent eversible, as it shows a different form in almost every

specimen. It often appears distinctly bifurcate at the tip. The

tip of the papilla is covered with fine hairs and usually shows

Some refractive granules, most probably urates of some sort.

352 THE AMERICAN NATURALIST. [VoL. XXXV.

Internally the gland has a peculiar and complex structure

(Fig. 11). The secretion seems to be formed in two elongate

oval bodies lying in the dorsal part of the abdomen, and it is

apparently carried through some intermediate, somewhat retic-

ulately arranged cells to the surface of the papilla, which is

lined with several layers of cells of varied size and form. The

gland may perhaps supply some

pleasant secretion for the ants,

like the tufts of hairs devel-

oped in myrmecophilous Cole-

optera, although I have not

been able to decide by obser-

vation. Ican find no reference

to similar glands in any other

insect, and hope at some future

Fic. 11. — Ecitomyia wheeleri, n. sp.

Cross- : . id

g gland time to study them in detail.

hitin ring;

G oi abdomen through Med

We have found this species

papilla and extending into abdomen; i, in. g great number of times, always

re associated with species of the

ant genus Eciton. Eciton cecum Latr., a totally blind species,

which tunnels in the earth, seems to be its favorite host,

although we found it upon one occasion in a nest of £. schmitti

Emery, a species with very different habits, which lives in

compact masses under stones, making its trips in search of

food above ground.

The Ecitomyias are exceedingly quick and have the habit of

darting rapidly about in zigzag paths in the way characteristic

of many myrmecophiles. In the nests of Æ. cæcum they frequent

those parts of the nest containing the greatest number of ants,

being very often seen running along the galleries of the ants,

into which they rapidly disappear when the nest is disturbed.

Those occurring with Æ. schmitti seem to stay at a greater dis-

tance from the main body of the ants, but this species makes

! A very curious coincidence occurs in a new genus of wingless Proctotrupidz

which occurs in the nests of Eciton cecum. Zt also possesses a sharply defined

roughening of the integument at exactly the same place that the gland of Ecitomyia

has its opening! I am sure, however, that there is no gland in connection with it.

Can it be possible that this has any connection with some way these blind an

may have of recognizing their habitual nestmates ?

No.413.] ABERRANT PHORID.£ FROM TEXAS. 353

large clusters and it might be dangerous for a myrmecophile to

venture into these. It apparently prefers to move about in

the vacant galleries of the nest.!

Some females which were placed in an artificial nest con-

taining a number of Æ. cecum workers soon made themselves

at home and appeared much more at ease than the ants, which

appear to be quite stupid and slow in adapting themselves to

new conditions. Some of the flies preferred to rest upon the

glass walls of the nest away from the ants. Others darted

among the ants in the largest groups, while the ants regarded

them without the slightest animosity. Any other fly or small

insect introduced into the nest was viciously attacked by the

ants and soon killed to serve as food for a large group of ants

which had taken part in its destruction. Even dead legs and

wings were picked up and carried about. Some Ecitomyias,

however, which had presumably died a natural death, were not

molested by the ants, and remained for a long time undisturbed.

One of the Ecitomyias was apparently feeding upon some

deposit left by the ants as they moved about, and it also

approached some of the less excited ones after the manner of

Myrmecophila, but I could not see that it obtained anything

from the bodies of the ants.

Throughout the winter we had seen the females in almost

every large nest which we examined, but although probably

half a hundred nests were seen during that time, not until

February did we positively find any males. In a large flourish-

ing nest of E. cecum which extended under stones for a distance

of nearly twenty feet, we found numerous female and two male

Specimens. Although the male has ample wings, it did not

attempt to fly, but hopped about in a similar but much less

agile manner than the female, which is often exceedingly quick

and hard to catch. The male does not hold the wings flat

! We have not been able to observe how they manage to follow the ants about

as they make their regular changes of nest, for this ant does not remain in the

Same nest for any length of time, except probably during the breeding season.

Other myrmecophiles of this species (e.g., Staphylinide) march along in proces-

Sion with their hosts as they make their curious journeys. As Æ. cecum moves

only by tunneling underground, they would experience no difficulty in keeping

company with the ants. Ne

354 THE AMERICAN NATURALIST. | [Vor. XXXV.

upon the back, but keeps them in a slanting position, so that it

resembles an exceedingly small aphid or psocid.

TERMITOXENIA WASMANN.

Thorax with one pair of dorsal appendages, apparently the halteres.

Halteres styliform or hooked at tip. Abdomen much swollen, curved

downwards, anus directed forward.

This most remarkable genus is represented by four termi-

tophilous species from Africa and India (Wasmann, '00a). Only

females are known, although Wasmann considers some of his

specimens as hermaphrodites, as they apparently possessed

both ovaries and testes. He says, in speaking of T. mirabilis:

* Jetzt sehe ich an Schnittserien der letzteren, dass dieselben

Hermaphroditen sind mit noch kleinen ovarien und gut ent-

wickelten Hoden." On such evidence we must not, however,

suppose that winged males do not occur. Other cases of her-

maphroditism among insects have been noted, but in no case is

it established to be anything more than a pathological condition

of certain individuals.

STETHOPATHUS WANDOLLECK.

Both wings and halteres absent, ocelli present, epistoma not large and

prominent. Thorax rounded. Abdomen elliptical, first four dorsal seg-

ments strongly chitinized and well developed, covering the greater part of

the dorsum of the abdomen. Abdomen not greatly retractile at apex

ovipositor short. i

Only one species of this genus has been described, S. ocellatus

Wand. from the East Indies. The specimens were found upon

carrion and in the flowers of the giant Arum (Amorphophallus).

It was previously described by Dahl as the female of his

Puliciphora lucifera, but because of his poor description and

misunderstanding of its systematic position, Wandolleck

redescribed it under a new name which it is probably best

to adopt.

No. 413.] ABERRANT PHORIDÆ FROM TEXAS. 355

CHONOCEPHALUS WANDOLLECK.

Wings and halteres wanting, ocelli wanting, eyes small and sunk deepl

into the head. Front almost horizontal. Thorax in profile triangular.

Abdomen elliptical, with six strongly chitinized dorsal plates which exten

far down upon the sides. Sixth ventral plate also present, almost meeting

the sixth dorsal. Ovipositor long.

This genus is represented only by C. dorsalis Wand., from

the Bismarck Archipelago. The imagines were found upon

carrion.

In conclusion, I wish to express my greatest thanks and

gratitude to Dr. Wm. M. Wheeler for the many kind suggestions

and great help which he has given me throughout my work.

I take great pleasure in naming one of the species in his honor

as a slight token of my appreciation.

BIBLIOGRAPHY.

'94 CHILD, CHARLES M. Ein bisher wenig beachtetes antennales Sinnes-

organ der Insekten, mit besonderer Berücksichtigung der Culiciden

und Chironomiden. Zeitschr. f. wiss. Zool. Bd. lviii, Heft 3

(1894), pp. 475-528, Taf. XXX, XXXI.

97 Cook, O. F. A New Wingless Fly from Liberia. Science. Vol. vi

(Dec. 1o, 1897), p. 886. ‘

'97 DAHL, FRIEDRICH. Puliciphora, eine neue floháhnliche Fliegengat-

'00 KELLOGG, VERNON L. An Extraordinary New Maritime Fly. Bio-

logical Bulletin. Vol. i, No. 2 (January, 1900), pp. 81-87, 3 text-figs.

'57 Loew, H. Psyllomyia testacea, eine neue Gattung der Phoriden.

Wien. Ent. Monatsschr. Bd. i (1857), pp. 54-56, Taf. I,

Figs. 22-25.

'30 MEIGEN, S. W. Systematische Beschreibung der bekannten europá-

ischen zweiflügeligen Insekten. Theil vi (1830), Taf. LXIII, Fig. 13.

'90 MEINERT, F. Ænigmatias blattoides, Dipteron novum apterum.

Ent. Med. Vol. ii ( 1890), pp. 212-226, PI. IV.

VERRALL, G. H. Description of a New Genus and New Species of

Phoride parasitic on Ants. In Lubbock's Ants, Bees, and Wasps.

DP. 431—435. 1881. :

WANDOLLECK, BENNO. [st die Phylogenese der Aphaniptera ent-

deckt? Zool. Anzeiger. Nr. 553 (1898), pp. 180-182.

356 THE AMERICAN NATURALIST.

'98b WANDOLLECK, BENNO. Die Stethopathidz, eine neue flügel- und

schwingerlose Familie der Diptera. Zool. /Jahrb., Abth. f. Syst.

Bd. xi (1898), pp. 412-441, Taf. XXV, XXVI.

'98c WANDOLLECK, BENNO. On the Phylogeny of the Flea and the

Stethopathide. Naturwiss. Rundschau. Braunschweig, Nov. 16,

1898.

'00a WASMANN, E. Termitoxenia, ein neues flügelloses physogastres

Dipterengenus aus Termitennestern. Zeitschr. f. wiss. Zool. Bd.

Ixvii, Nr. 4 (1900), pp. 599-617, Taf. XXXIII.

'00b WasMANN, E. Neue Dorylinengáste aus dem neotropischen und

dem athiopischen Faunengebiet. Zool. Jahrb., Abth. f. Syst.

Bd. xiv, Heft 3 (1900), pp. 1-75, Taf. I, II

UNIVERSITY OF TEXAS, AUSTIN,

March 1, 1901

POSTSCRIPT.

After this article had gone to press, the author was so fortunate as to

obtain two other new species of Stethopathinz, even more remarkable than

the ones described above. While collecting ants in the vicinity of Austin

with Mr. A. L. Melander one afternoon during March, we found in a nest

of Eciton opacithorax Emery a small insect which by its actions = at

once recognized as a member of the Stethopathina. On carefully sifting

the earth of the nest a second specimen was obtained. On comparing them

we were exceedingly surprised to discover that not only were they different

from any described Stethopathinz, but were also quite unlike each other.

They are both females which I intend to describe in the near future as

representatives of two new genera. From this it would seem that there

must be a whole series of these peculiar Diptera living as true myrme

cophiles in the nests of various ants.

UNIVERSITY | OF TEXAS, AUSTIN,

April 10, 19or.

THE METATHORACIC PTERYGODA OF THE

HEXAPODA AND THEIR RELATION

TO THE WINGS.

L. B. WALTON.

Ow the anterior margin of the prothorax and mesothorax

of the Lepidoptera are two small sclerites known as the pata-

gium and tegula, respectively ; while in certain other orders of

Hexapoda (Hymenoptera, Neuroptera, and Trichoptera) a small

piece has been found at the base of the

mesothoracic wing which has been consid-

ered equivalent to the tegula. Further

than a few suggestions based on limited

observations, no attempt has been made

either to ascertain the value of these pieces

or to demonstrate the existence of similar

homodynamous or homologous structures

in the hexapods. :

The purpose of the present paper is to

call attention briefly to the general pres-

ence of a sclerite on the hexapod meta-

thorax which seems homodynamous with

the so-called tegula, to make some sugges-

tions concerning terminology, and espe-

cially to point out that the present view

concerning the metamerism of the anten-

nate arthropods appears worthy of recon-

sideration.

The relation of the tegula, or pterygodum

as I prefer to term it, to the pieces of the

mesothorax, which is in many respects

the most generalized of the three thoracic

segments in the insects, is shown in Fig. 1.

herda

IG. 1. — C. s lignif

Left lateral portion of mes-

ora A, pterygo-

dum (stippled); w, wing;

eps, episternum ; eft, epi-

godum extending behind

the i All

figures are

Bearing in mind

now that the mesothorax and metathorax are equivalent,

357

358 THE AMERICAN NATURALIST. (VoL. XXXV.

the various components of each being reduplicated in the

other segments, subject, however, to the factors governing the

specialization of the different groups of insects, the existence

of a corresponding piece in the metathorax would a priori

be inferred. The presence of such a piece! is represented

in the accompanying diagram (Fig. 2), while a homologous

part can generaly be demonstrated throughout the other

orders (Figs. 3, 4). Furthermore, it is to be noted that in the

typical form this is joined to the dorsal

margin of the episternum, while the wing

is articulated with the dorsal margin of the

epimeron and not, as hitherto accepted, with

the episternum.?

In connection with the facts noted above,

certain evidence is available, based on em-

bryology as well as comparative anatomy,

which adds weight to the inference that

Fic. 2.—Cossus ligni- these two pieces are rudimentary? wings

erda. Left lateral $

portion of metathorax, aNd that the thorax of the Hexapoda is com-

oe: Doar sci in posed of six somites which in the typical form

bear the fundaments of six pairs of wings.

The development of the mesothoracic pterygodum* (=tegula,

parapteron, etc.) is of extreme interest in this connection and

furnishes interesting evidence toward establishing the hypoth-

esis I have put forward regarding its relation to the wing.

This, however, is only one of a large number of facts which

corroborate such a view and which appear explainable on no

1 From comparisons made throughout the Hexapoda it is evident that this

does not correspond to the more or less chitinized part of the epimeron at the

base of the wing in most Lepidoptera.

The anterior margin of the wing is generally specialized at the base, so that

it partially extends over the dorsal portion of the episternum, while an articula-

tive process of the pterygodum may be received into a corresponding notch of

wing. (See Fig. 3) This condition, however, is secondary. A

here appears to be no evidence for regarding these as vestigial, since it ng

improbable that forms with six pairs of wings ever existed. The term “ rudi-

mentary,” as generally used, does not seem inappropriate, although the quc

* fundament" is more concise.

*Owing to its greater size, this can be more conveniently studied than the

Corresponding piece on the metathorax.

No. 413.] THE METATHORACIC PTERYGODA. 359

other basis. To one of these I have already called attention

(Walton, 1900), notably the formation of the coxa in Chilopoda

and Hexapoda from two fused pieces to which I have Appues

the name “ coxa genuina " and

* meron."

I have adopted the name v^

* pterygodum ” 1 for the pres-

ent in preference to others

which have been suggested

for the mesothoracic piece

(tegula, parapteron, squamula,

etc.), since it has priority over

terms otherwise acceptable,

and according to our present Fic. 3.—Hydrophilus seiner, Left lat-

: : era sil yardon of metathorax. x 8. a genuina

knowledge better indicates the between ante-coxal piece Ae 1 meron. Oe

function of the part. The term kir as in Fig. n pimeror

uniet

“parapteron,” which Comstock

and Needham (98), following Newport (39), have used in

reference to the mesothoracic pterygodum, appears inap-

propriate, for the reason that Audouin (24) first used it

to indicate a supposed sclerite on the anterior margin of

the mesothoracic episterna in Dy-

tiscus circumflexus, the part in ques-

tion being merely an articulative

process. Several years later, in a

» note to a translation of a paper by

MacLeay (32), Audouin stated his

belief that the piece in the Hymenop-

tera termed squamula by MacLeay

was homologous to the parapteron

- which he himself had described.

8 This supposition was not only incor-

- rect but was subsequent to the ter-

minology adopted by Latreille.

The value of the patagium on the prothorax has been more

or less discussed, but until we know more concerning its devel-

opment it is impossible fully to decide whether it is equivalent

1 (gr. mreovydóss (mrepv'yoeiófjs = mrépvyos [wing] + eldos [form].

r references as in Fig. 1

360 THE AMERICAN NATURALIST. [Vor. XXXV.

to the wing, as suggested by Cholodkowsky ('86), or to the ptery-

godum (tegula), the view adopted by Haase (86) and now so

generally accepted. It should be observed, nevertheless, that

the reasons given by Haase for reaching such a conclusion

are far from adequate, since the only evidence to which he

called attention, otherwise than a superficial resemblance, was

that (1) chitinous folds of a similar nature but of secondary

origin are present on the prothorax of certain Hymenoptera

and Diptera, while (2) the patagia do not exist during the

larval stage of the Lepidoptera, but commence their develop-

ment in the first few days of the chrysalid stage. Unfortu-

nately, however, Haase failed to demonstrate any homologous

structures in the Hymenoptera or Dip-

tera, and had he attempted to do so it is

evident, from the preceding, that proof

of their secondary nature would have

been difficult to establish. Moreover,

the Anlage of a structure must exist

in potentio, and the time during the post-

embryonic stages at which it commences

that which is known as development can

rc qu ids alone be of no particular value in deter-

relations of pterygodum, wing, mining its palingenetic or cenogenetic

episternum, and epimeron in

the thoracic segment of the Character. The question as to whether

Ta Pot References as in the patagium represents a prothoracic

pterygodum or a wing, must await a large

amount of comparative work based on embryology, with the

possibility of paleontological’ evidence affording some help

in the solution of the problem. The ratio of develop-

ment between wing and pterygodum on the other thoracic

segments allows the inference, however, that pterygoda may

te]

La!

ral

ondon) suggested. Brongiart has already pointed this out. (Note sur quelque

insectes fossiles du terrain houiller qui presentent au prothorax des appendices

aliformes, Bull. Soc. Philom., tome ii, 1890.)

No. 413.] THE METATHORACIC PTERYGODA. 361

exist in front of the patagia which then have the value of

wings.

Another interesting question in this connection is the

homology of the elytra of Coleoptera, for again further inves-

' tigation must be awaited before a logical conclusion can be

reached. The tracheation of the elytra, to which attention

has been called by Comstock and Needham (98), is not con-

clusive evidence that they are specialized wings, for in connec-

tion with the view I have here advocated this would be

expected if they were homologous with the mesothoracic

pterygoda of the Lepidoptera, and the suppressed wing was

represented by the alulet so noticeable under the Elytra in

Hydrophilus, etc.

From the preceding facts, to which I have called attention,

it appears necessary to consider that the typical thoracic seg-

ment (Fig. 5) possesses the components of both pterygodum

and wing, the former joined to the dorsal margin of the epi-

sternum, the latter articulated with the dorsal margin of the

epimeron, while furthermore the morphological position of

the pterygodum in respect to the wing indicates that it may

have an important bearing in elucidating the metamerism of

the antennate arthropods.

ANATOMICAL LABORATORY, BROWN UNIVERSITY.

BIBLIOGRAPHY.

?4 Aupourn, V. Recherches anatomiques sur le thorax des animaux

articules et celui des insectes hexapodes en particular. Ann. Sci.

Nat. Zool. Tome i 1, p. 416.

'86 CHOLODKOWsKY. Zur Morphologie der Insectenflügel. Zoo. dus.

Bd. ix, p.

'87 CuoLopxowexy. Ueber die eee bei den Lepidop-

teren. Zool. Anz. Bad. x, p.

95 Comstock, J.H. Manual for de do of Insects.

98 Comstock AND NEEDHAM. The Wings of Insects. Amer. Nat.

Vol. xxxii, p. 561.

THE AMERICAN NATURALLIST.

Haase, E. Die Prothoracalanhánge der Schmetterlinge. Zool. Anz.

Bd. ix, p. 711

HOFFBAUER. Beiträge zur Kenntniss der Insectenflügel. Zeitschr.

wiss. Zool. Bd. liv, p. 579.

KELLOGG, V. L. The Classification of the Lepidoptera. Amer. Nat.

Vol. xvii, p. 248.

KIRBY AND SPENCE. Introduction to Entomology.

KoLBE, H. J. Einfuhrung in der Kenntniss der Insecten.

LATREILLE, M. Observations nouvelles sur l'organisation des ani-

maux articulés

MAcLEAY. Explanation of the Comparative Anatomy of the Thorax

in Winged Insects. Zodl. Journ. Vol. v, p. 145.

MacLeay. French Translation, with Notes by Audouin. Ann. Sci.

Nat. Tome xxv, p. 95.

MEINERT. Sur l'homologie des élytres des Coléoptéres. xt.

Tidskr. Bd. i, p.168

Newport. Todd’s Cyclopedia of Anatomy and Physiology. Vol.

ii, p. 916.

PACKARD, A. S. Text-Book of Entomology.

RILEY. Tegule and Patagia of Lepidoptera. Proc. Ent. Sov.

Washington. Vol. ii,

SHARP. The so-called Secondary Wing of Coleoptera. Tr. Ent.

Soc. London.

SHARP. The Cambridge Natural History. Vol. v.

SHARP. The Cambridge Natural History. Vol. vi.

WALTON. The Basal Segments of the Hexapod Leg. Amer. Nat.

Vol. xxxiv, p. 267. No. 400.

PHAGOCYTOSIS IN THE POSTEMBRYONIC

DEVELOPMENT OF THE DIPTERA.

VERNON L. KELLOGG.

IN the most recent considerable! paper on the postembry-

onic development of an insect of complete metamorphosis, the

author lays much stress on the small part which phagocytes

play in the breaking down of the larval tissues during the

metamorphosis of the insect studied — the little brown ant,

Lasius flavus. In this respect the author sees in the meta-

morphosis of Lasius (belonging to the Hymenoptera) a sharp con-

trast to the metamorphosis of the Diptera, in the best-known

example of which, the much-studied Calliphora, phagocytosis

plays an all-important part. Korotneff? found in the case of

the degeneration of the larval muscles of Tinea (Lepidoptera)

that there was no phagocytosis. Rengel? in studying the

changes in the alimentary epithelium of Tenebrio and other

Coleoptera, found also no phagocytosis, and Needham,* in a

careful study of the flag weevil (Mononychus vulpeculus), simi-

larly found a complete lack of phagocytosis in the histolysis

of the larval tissues of this insect. Karawaiew strongly agrees

with Korotneff and Rengel in believing that phagocytosis is

a phenomenon of postembryonic development associated with

the length of time occupied by the metamorphosis. With the -

blowfly the metamorphosis occupies but a few days; with

Tinea, a little more than two weeks ; with Tenebrio, several

weeks, according to the temperature; and with the ants still

: Karawaiew, W. Die nachembryonale Entwicklung von Lasius flavus, Zeitschr.

J. wiss. Zool, Bd. lxiv (1898), pp. 385-478, Pls. IX-XII, and 15 figs. in text.

? Korotneff, A, Histolyse und Histogenese des Muskelgewebes bei der Meta-

während der Metamorphose, Zeitschr. J. wiss. Zool., Bd. lxii (1896).

* Needham, J. G. The Metamorphosis of the Flag Weevil (Mononychus

Vulpeculus), Bio’. Bull., vol. i (1900), pp. 179-191.

363

364 THE AMERICAN NATURALIST. [VOL. XXXV.

longer; in the case of Lasius flavus, for example, from the

first warm spring days until the beginning of June or even

longer. In the case of the insects, like the flies, with

a short time devoted to metamorphosis, there must be

space made for the new organs as quickly as possible ; that

is, the old larval organs must get out of the way as soon

as may be. The natural process, a gradual degeneration,

is a process of long duration, and on that account not suffi-

cient in the case of the flies. Hence, says Karawaiew, there

has arisen the barbaric devouring of the tissues by the

leucocytes.

It has seemed to me unfortunate that in the study of the

postembryonal development of the Diptera so much attention

should have been given to the highly specialized Muscidae and

so little to more generalized members of the order. The

metamorphosis of Coretha, Culex, and Chironomus has been

studied somewhat, but without any approach to that exhaust-

iveness which characterizes the studies of Weismann, Van

Rees, Kowalevsky, e£ al., on Calliphora. In the hope of find-

ing some new light upon these extraordinary phenomena of

histolysis and histogenesis which are a part of insect meta-

morphosis I have undertaken the study of the postembryonic

development of two flies belonging to the more generalized

Diptera, the Nematocera. One of these flies is Blepharocera

capitata, a member of the strange, small family, Blepharoce-

ride, with strangely and strongly modified immature stages,

and the other is Z7o/orusia rubiginosa, a giant crane fly

(Tipulidz), with simple immature stages. While both of these

forms are nematocerous Diptera, and to this extent allied,

there is an exceptionally wide divergence between them in

point of structure of the larval stages, and this differ-

ence has, to my mind, an allimportant influence in deter-

mining the obvious and suggestive differences in the character

of the development, which, we shall see, obtains. This

present reference to the metamorphosis of these two dip-

terous forms has to do solely with the peculiarly interesting

and suggestive conditions of the histolytic processes in the

metamorphosis. r

No. 413.] DEVELOPMENT OF THE DIPTERA. 365

The larva of Holorusia rubiginosa! is cylindrical, worm-

like, tapering slightly towards both ends, without feet or other

special organs of locomotion. It attains a length of three

inches (outstretched full-grown specimens). The head is a

retractile, strongly chitinized capsule, with biting mouth-

parts. The internal anatomy is simple. The musculation

consists, except in the head, of simple segmental, longitu-

dinal, integumental muscles and of annulate integumental

muscles. Locomotion is a simple squirming or wriggling,

caused by longitudinal contractions. The alimentary canal

is a straight tube divisible into the usual parts. The ven-

triculus has four diverticula or czca, and the large intestine

has a single forward projecting diverticulum. There are four

Malpighian tubules. There is a single pair of large salivary

glands, each gland bent double. The respiratory system con-

sists of a single pair of large spiracles situated on the posterior

aspect of the last abdominal segment, and of a pair of main

longitudinal tracheal trunks with their branches. The larval

life lasts several weeks.

The pupa is found in the same place inhabited by the larva,

and is of simple character. It is from one and one-fourth to

one and three-fourths inches long. There is a pair of slender,

short respiratory tubes on the prothorax. The pupal stage

lasts twelve days.

In the course of the postembryonic development of Holorusia

I have found no occurrence of phagocytosis. The breaking

down of the muscles and salivary glands and fat body of the -

larva (tissues in which phagocytosis most certainly occurs if

at all and in which it is most readily determinable) is accom-

plished apparently entirely by simple « selbständige Degen-

eration” (Karawaiew). The breaking down of the muscles

does not begin until after the pupal life is well started. In

fact there is no very great breaking down essential. The

musculation of the adult differs from hat of the larva more

in the addition of the wing and leg muscles of the thorax

* This is the first published reference to the immature stages of this giant

tipu - The larve were found abundantly on the banks of a small stream near

this university (Stanford), lying in mud and slime composed of decaying leaves.

366 THE AMERICAN NATURALIST. (Vou. XXXV.

and the muscles of the head than in any complete substitu-

tion of an imaginal musculation for a previous wholly different

complex larval musculation. In pupa one-third through their

existence (four days old) a great deal of the larval muscula-

tion still persists side by side with the developing new muscles

of the thorax. The nuclei of the degenerating larval muscles

show the “old-age” characters of degenerating nuclei; the

contractile substance first loses its striate appearance, then

becomes loosely fibrous, then spongy, and finally breaks up.

The degeneration of the large salivary glands is easily fol-

lowed. In the larve the nuclei of the large epithelial cells

pup y-eig ld, degenerati

Fic, 1. — Salivary gland of Holorusi bigi A cross-section of gland of larva; 2, cross-

$ p : 11 Advanced

on being

are regularly circular or elliptical (in optical plane) and sharply

delimited by a nuclear membrane. The chromatin is rather

massed together and stains strongly. The cytoplasm of the

cells is evenly granular and the cell outlines well defined

(Fig. 1, 4). Ina pupa not more than twenty-four hours old a

marked degeneration of the cells has occurred. The cytoplasm

is distinctly vacuolated, and in a pupa a day or two older the

cytoplasm is spongy, the cells have lost their shape, the nuclei

have lost their membranes and are showing other degenerative

characters (Fig. 1, B). No phagocytes appear. The degenera-

tion or histolysis of the larval tissues of Holorusia is accom-

plished thus without the interference of phagocytes. The pupal

condition is characterized by no such extensive breaking down

No. 413.] DEVELOPMENT OF THE DIPTERA. 367

of larval organs as is apparent in the pupa of Calliphora, where

the pupal body cavity is filled with * pseudo-yolk," a confused

fluid mass of degenerating tissue.

The larvae of Blepharocera capitata! are of extraordinary

external appearance, and. in their habits and structure are

widely removed from other dipterous larva. They live under

water in brooks, clinging by six elaborately developed ven-

tral suckers to the smooth rock bed or to smooth stones in

parts of the stream where

the water runs swiftly and

is shallow. The segments

of the body are greatly

modified, the three thorac-

ic segments and the head

being fused to form a single

large anterior body region.

For the control of the

suckers and for the peculiar ,

lateral swinging movement &

of the body in locomotion

an elaborate musculation is

developed, which is very

different from the muscu-

lation of the adult fly. The

Pup: are also extraordi-

nary in character and live,

like the larvze, attached to T E

the rocks in swift, shallow fic. 2,—Larval muscle of Blepharocera capitata,

Put&'of the strehimi "The. e een A pepe n der dem o

duration of the pupal stage is fifteen days. The histolysis of

the larval tissues begins three or four days before the true

pupal condition is reached. The larvae cease feeding, become

quiet, and thus remain three or four days before pupation.

The total duration, therefore, of the time devoted especially to

the change from larva to imago is about eighteen days, as

compared with twelve in the case of Holorusia.

! For an account of the structural character of the larvæ, see Kellogg, Notes on

y Life-History and Structure of Blepharocera capitata Loew, Ent. News, vol.

xi (1900), pp. 305-318. .

368 THE AMERICAN NATURALIST.

There is a great breaking down of the larval organs of

Blepharocera. With such a specialized larval life there is a

great difference between the larval organs and the imaginal

organs. The musculature, the alimentary canal, and the res-

piratory system are largely broken down and reformed. And

in all of this histolysis phagocytes are abundant and conspicu-

ous. When pupa not more than three or four days old are

dissected, the body cavity is found to be filled with “pseudo-

yolk," that is, with a lymph-like liquid containing floating bits

of degenerating tissue and hosts of phagocytes. A bit of

larval muscle (Fig. 2) in a pupa a few days old shows very

well the character and effects of the phagocytosis.

Thus in the fly Blepharocera, with its eighteen days of

prepupal and pupal condition, phagocytosis is conspicuously

present; in the fly Holorusia, with its twelve days of pupal

condition, histolysis is unaccompanied by phagocytosis. The

fly in which the histolytic phenomena occupy the longer time

is the one in which the histolysis is accompanied by phagocy-

tosis. This is a condition not at all in consonance with Kara-

waiew's conclusions, as quoted at the beginning of this paper.

What is the reason for the presence of phagocytes in the

histolysis of Blepharocera and their absence in Holorusia? To

my mind, the extent of the metamorphic changes, the degree

to which histolysis occurs, probably offers the explanation.

In Blepharocera, with its highly specialized larval form, its

peculiar and specially developed organs, the change to imago

is radical; the histolysis of larval tissues is extensive. In

Holorusia, with its generalized larval form, its less modified

organs, the change to imago is accomplished with much less

breaking down of larval organs and reformation of imaginal

ones; the histolysis is less radical and considerable. The

phagocytes are the agents or the assisting agents in the more

extended and radical histolysis.

My observations so far do not enable me to offer any evidence

regarding the moot point touching the causal agency of the

phagocytes in histolysis. Whether the phagocytes initiate his-

tolysis, or merely render effective aid after the degeneration

has been initiated independently, is a question of importance.

ON THE SIGNIFICANCE OF THE SPIRAL SWIM-

MING OF ORGANISMS:

H. S. JENNINGS.

Ir is a well-known fact that many of the lower organisms

swim in a spiral path, but the real significance of this fact has

never been pointed out, I believe, until recently. Swarm-

spores, flagellate and ciliate infusoria, rotifers, and many other

lower organisms as they pass through the water revolve on

their long axes, and thus follow a course that takes the form

(as a rule) of a spiral. Extended discussions of this fact are

to be found in many works, as in Bütschli's * Protozoen ” in

Bronn's Klassen und Ordnungen des Thierreichs, and in many

special papers. These discussions usually confine themselves

to a description of the facts, — so far as these were made

out, — and to a discussion of the mechanical factors involved

in producing the spiral movement, without any attempt to

show the biological significance of the phenomenon. To under-

stand the significance of this method of swimming was indeed

perhaps impossible until the relation between it and the

method of reaction to a stimulus in these organisms was

known, and especially until it was recognized that the body of

the organism bears a constant relation to the axis of the spiral, —

that is, that the same side of the organism is always directed

toward the outside of the spiral (as in Fig. 1). These relations

were first pointed out by the present writer in Nos. II and V

of his * Studies on Reactions to Stimuli in Unicellular Organ-

isms,” 2 where they were shown to hold for a considerable

number of Flagellata and Ciliata.

! The substance of this paper was presented at the meeting of the Western

Naturalists i in Chicago, Dec. 27, 1

? II, The Mechanism of the Motor Reactions of Paramecium, Am. Journ. of

Phys., = ii fad can P. 323; V, On the Movements and Motor Reflexes of the

d Ciliata, 4m. Journ. of Phys., vol. iii (1900).

369

379 THE AMERICAN NATURALIST. [Vor. XXXV.

The exact purpose that is served by this method of swimming

is a point deserving of further emphasis and fuller discus-

Fic. 1, — Diagram of spi-

ral course of Loxodes

and d below this plane.

sion. The Flagellata and Ciliata are as a rule

unsymmetrical in form. One of these organ-

isms, as, for example, Loxodes (Fig. 2), or

Paramecium (Fig. 3), when it leaves the bot-

tom and starts to swim freely through the

water, cannot go in a straight line, but owing

to its lack of symmetry continually swerves

toward one side, so that it tends to describe

a circle. If no method is taken of compen-

sating this deflection, the circles described

are frequently very small, and of course the

animal makes no progress by swimming in

this way. Paramecium and Loxodes thus

tend to circle toward the aboral side, Chilo-

monas (Fig. 4) toward its “lower lip,” all the

Hypotricha to the right, etc.

To obviate this difficulty, revolution on the

long axis is combined with the forward move-

ment of the organisms. By this means the

continual swerving toward one side is com-

pensated, since this side is continually turned

in a new direction. Thus, if Loxodes is

swimming (freely through the water) away

from the observer, and the aboral side is at

first to the observer's left (Fig. 1, 4), the

organism at first swerves to the left; but as

it revolves the aboral side soon comes to be

the upper side, and the animal now swerves

up (č). By continued revolution the aboral

side is brought to the right (c), so that the

animal swerves to the right. Next, of course,

it swerves down and the process is continued,

the animal swerving successively to the left,

* up, right, down, etc. These movements, 9

course, compensate each other, so that only

the forward component of the motion 1S

No.413.] SPIRAL SWIMMING OF ORGANISMS 371

effective; the animal thus moves forward as if on a straight

line, — the actual path being a spiral with a straight axis. The

principle is the same as that by which a rifle bullet is given a

straight course by making it revolve in the axis of flight.

In the Hypotricha a similar course is followed, save that the

swerving is to the right; in Chilomonas (Fig. 4) it is toward

the lower lip (a). Such a spiral path is known

to be followed by most of the free-swimming

Protista, — by swarm-spores, flagellates, and

ciliates in general; by Volvox, Eudorina,

Pandorina, Platydorina, etc. In some of these

organisms the course followed becomes almost

an actual straight line, owing to the fact that

the body is symmetrical, so that there is no

pronounced swerving toward one

side. Such is the case, for ex-

ample, in Volvox. Here

the revolution on the

T long axis probably

serves merely to com-

pensate for any acci-

dental deviations that

may occur through in-

| jury, unequal develop-

Fic. 2. Fic, 3. Fic. 4. ment, and the like.

F ee rostrum, after Biitschli, showing the Such cases are compar-

Pas Pome ae eta, 4 atively rate; however,

oral side, ` most of these organ-

Fic. dM Chil, x os 1 . *

‘nek — ATTN He INOOA e. ioma- being. markedly

ee ee ee

a Pr

i gage z

Lp pre Sa neun >

AI re p T

unsymmetrical.

The mechanical cause of the revolution on the axis.of pro-

Sression has often been discussed. In the Ciliata there seem

to be three possible factors: (1) an oblique stroke of the cilia ;

(2) the oblique position of the peristome; (3) the unsymmet-

rical form of the body, which is often of such a shape as to

favor rotation in a given direction. That the first factor is the

Primary one is indicated by the fact that the direction of revo-

lution may be reversed in many of these organisms, even when

372 THE AMERICAN NATURALIST. [Vor. XXXV.

the form of the body is such as to oppose this reversal The

unsymmetrical form seems rather an adaptation to this method

of swimming, — a consequence of it. Many of these organisms

are so shaped that the body forms part of a spiral; this is to a

certain degree the case, for example, in Paramecium (Fig. 3).

In some others this is much more marked. Phacus, for example,

is frequently strongly spiral. Some of the bacteria swim in

this same manner, and among these, Spirillum forms, as is well

known, a sort of animated corkscrew. The prevailing asym-

metry in the unicellular organisms is closely correlated with

this method of swimming.

When creeping along the bottom (as Loxodes usually does),

or when in contact with any solid object, these same organisms

exhibit no such rotation. When moving along a surface there

are, of course, only two chances to err from the straight line,

either to the right or to the left. When swimming freely

through the water, on the other hand, the chances of devia-

tion are indefinitely numerous, since the organism may swerve

to the right or left, or up or down, or in any intermediate

direction. Moreover, when in contact with a surface, this

usually presents numerous stimuli, which serve as directives

of motion, while in the free water such stimuli are lacking.

Hence the necessity of some special device for keeping the

straight course in the latter case. The movements and reactions -

of organisms differ greatly when they are moving along a gui

face from those. exhibited when passing freely through the

water. (Pütter! has recently published a valuable paper on

this subject.) Both flagellates (e2., Peranema) and ciliates

move without rotation when in contact with a surface. Yet

even then they usually cannot travel in a straight line; Col-

pidium and Oxytricha, for example, follow a much curved

course.

As the present writer has fully set forth in his * Studies on

Reactions to Stimuli” (doc. cit.), this method of locomotion is

closely related with the usual method of reaction to a stimulus.

In addition to swerving toward a structurally defined side 1n

| Pütter, August. Studien über Thigmotaxis bei Protisten, Archiv f. Har

Phys., Physiol. Abth., Supplement Band (1900), pp. 243-302-

No.415.] SPIRAL SWIMMING OF ORGANISMS. 373

their locomotion, these unsymmetrical organisms respond to a

stimulus by turning toward a structurally defined side.

It is important not to misunderstand the nature of this

spiral motion. If one of these swimming organisms is viewed

from above with the ordinary microscope, the path of the

organism seems to swerve merely first to the right, then to

the left. This is of course because the upward and downward

part of the path is lost from view with the ordinary micro-

scope which sees approximately in a single plane; with a stereo-

scopic binocular the real nature of the path is evident. If the

constant relation of the body of the organism to the axis of the

spiral is likewise overlooked, a peculiarly false conception of

the movements of these

organisms is obtained,

which seems to be

somewhat widespread.

This is the conception

that the organism

Swerves as it swims, hedcp

first toward one side, ymmetrical organism when oriented by li

then toward the other. Mijn caper n

For example, Loxodes or Paramecium, according to this view,

would swerve first toward the aboral side, then toward the

oral side. This supposed movement has even been given a

high theoretical significance, as being the natural result of the

orientation of an unsymmetrical organism by lines of force,

Such as rays of light, or the path of diffusing ions. Thus, in

Fig. 5, in the position a-a, in which the axis of the organism is

parallel with the lines of force, more lines of force impinge

on the convex side of the organism; hence the locomotor

organs on that side act more (or less) strongly than those on

the concave side. As a result of this differential action, the

organism swings (supposedly) to the position 4-2 (that is, it

Swings toward the aboral or convex side). Now more lines of

force impinge on the concave side; the locomotor organs act

more (or less) strongly on this side, and the organism swings

again (now toward the oral or concave side) into the position

a-a. This continues, and, combined with the forward motion,

ban iiladdciak: d oscillation of an

374 THE AMERICAN NATURALIST. [VoL. XXXV.

supposedly accounts for the sinuous path of these organisms.

It should be clearly stated that the actual movements of these

creatures lend no support to this account, but are, on the

contrary, quite incompatible with it. The

organisms swerve always toward the same side,

not first to one side, then to the other.

But it is well known that it is not only

unsymmetrical organisms that swim in a spiral,

but that the same is true for many bilateral

organisms also, —as, for example, the Rotifera.

Since the two sides are alike in these animals

(see Fig. 6), there is no reason for swerving

to the right rather than to the left, and the

spiral path calls for some further explanation.

The significance of the spiral path in such cases

c.6.—Dormal view 1$ clearly seen when the movements of these

parachi animals are carefully studied. When creeping

Weber) to show the On the bottom or the surface film, there is no

sadanefthetwo rotation. Here the only possibilities of devia-

tion from the straight line are either to the

right or to the left, and since the two sides are alike there is no

reason for swerving in either direction. But the dorsal and

ventral sides are not alike (see Fig. 7), and in swimming freely

through the water the animal

might err by turning toward the

dorsal or toward the ventral side,

or in any intermediate direction.

As a matter of fact, careful ob-

servation shows that most roti-

fers do swerve toward the dorsal =Z

side as they swim freely through —

the water. This tendency seems fic. 7. — Rotifer (Brachionus pala, mon

traceable to the fact that the fos fom Hoi c eet

rotifers are primitively creepers side when rising from the bottom. 4

on the bottom, and most of them 9?! 54e: ® peas a

still retain this habit. In order to rise from the bottom into the

free water, the animal must necessarily move toward the do

side (as in Fig. 7). The cilia which bring about the free

us zin

+ YF ip

'J (N

CS ene E

No. 413.] SPIRAL SWIMMING OF ORGANISMS. 375

swimming movement seem to have this tendency, to strike so

as to turn the animal toward the dorsal side, strongly ingrained.

Many of these animals cannot rise from the bottom so long as

the dorsal side is down. In such a case the dorsal side of the

head repeatedly strikes the bottom until, by revolving on

the long axis, the dorsal side is turned toward the free water ;

the animal then swerves off the bottom in that direction. Some

Fic. 8. — Diagram showing course followed by Ploesoma when swimming without revolving

on the long axis. The animal continually swerves toward the dorsal side, hence follows

a circular path.

of the rotifers, if they attempt to swim freely through the water

without revolving on the long axis, turn backward somersaults,

over and over, describing thus small circles. I have seen

Ploesoma thus describe circles (Fig. 8) for considerable periods.

But as soon as the animal begins at the same time to revolve

on the long axis, without otherwise changing its movement,

the effect is striking. The purposeless circular movement

Fic. 9. — Diagram showing the course

of Plæsoma as soon as it begins to

revolve on the long axi

higher and lower, respectively

5 and d, which lie in the same plane).

Thi 1 , with th 1

axis (a and c are

than

This spiral course the dorsa

side to the outside of the spiral, is

characteristic for many Rotifera

THE AMERICAN. NATURALIST.

[Vor. XXXV.

(Fig. 8) becomes at once a well-ordered

progression in a spiral path (Fig. 9).

No one who has seen this sudden

change from random circles to a path

having all the essential qualities of a

straight line can fail to appreciate thé

biological significance of the rotation

on the long axis in compensating the

tendency to swerve in a given direction.

This tendency to swerve toward the

dorsal side seems present in the major-

ity of the free-swimming Rotifera, and

is compensated almost universally by

the revolution on the long axis, causing

the resulting path to be a spiral with

the dorsal side directed toward the

outside of the spiral (Fig. 9). All

rotifers observed by the writer revolve

to the right, and no reversal of the

direction of revolution was ever seen.

In some of these primitively bilat-

eral animals this spiral method of

swimming has resulted in the pro-

duction of an unsymmetrical form

analogous to that of the infusoria.

In the small aberrant family of Rat-

tulidæ this adaptation to a spiral

movement is most striking. Rattulus

tigris, for example, has a Urs

body, forming actually a segment o

a spiral, and there is in WE.

high spiral ridge on one side. T is

ridge begins behind the middle, near

the mid-dorsal line, and passes for-

ward, at the same time curving onee

to the right side. The animal am

in a spiral of which this ridge 2"

its own twisted body form 4 part.

No. 413.] SPIRAL SWIMMING OF ORGANISMS. 70

Asymmetry appears sporadically in many different groups of

the Rotifera ; possibly it may in every case be brought into

relation with the spiral method of swimming.

The Rotifera are a group of organisms excessively varied in

form and movements, furnishing a most excellent opportunity

for studies on the interdependence of

structure and function. Some species

(¢.g., certain species of Diaschiza)

have the body so curved ventrally

that the tendency to turn toward the

dorsal side is more than compensated,

and the animal tends instead to curve

continualy toward the ventral side.

a swimming

: k E 5 FiG. 10. — V

This tendency is of course likewise ^ zuchianis triquetra from behind

after Eh t gt h the t

hree

corrected by the revolution on the

axis of progression, the path taking

here the form of a spiral with the ventral surface to the outside.

Some few rotifers have become so modified that revolution on

the long axis has become unnecessary for keeping a straight

course. Thus Euchlanis triquetra (a view of which from the

rear is shown in Fig. 10) has developed

three great keels, one dorsal and two

lateral, which tend to prevent swerving

in any direction; so this animal fre-

quently swims freely for long stretches

without revolving, while the closely

related Euchlanis oropha (having no

keels) almost continually revolves as it

passes through the water.

| Revolution on the long axis, with the

Fic. r.—One of the Gastrotri- resulting spiral path, is common also

after Zelinka), visae be in many other animals. In the Gas-

os cha (s on the trotricha (eg., Chaetonotus, Fig. 11)

the locomotor organs (cilia are con-

fined to a strip on the ventral side, (a) which necessarily

results in giving the organism a tendency to turn toward the

dorsal side. The revolution on the long axis is therefore of

&reat importance for producing an effective forward movement.

keels.

378 THE AMERICAN NATURALIST.

Revolution on the long axis is also to be observed in many

rhabdoceels. It is indeed one of the commonest features in

the locomotion of small fresh-water organisms, and doubtless

occurs in salt-water forms in the same way.

On the whole, then, it is clear that revolution on the long

axis, with the resulting spiral path, is of high biological

significance. Only through this device are many organisms

enabled to follow a course which is practically a straight one;

without such revolution many creatures merely describe small

circles, making no progress whatever. By means of this revo-

lution on the long axis, any organism, no matter how misshapen

and irregular, may follow a course which is, in effect, equivalent

to a straight line. The simple device of revolving in the axis

of progression is surprisingly effective, in that it compensates

with absolute precision for any tendency or combination of

tendencies to deviate from a straight course in any direction

whatsoever.

ANN ARBOR, MICH, January 5, 1901.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XIV. THe Hypromepus& — Part II.

CHAS. W: HARGITI.

THE CAMPANULARIÆ (CALYPTOBLASTEA).

Tue Campanularie are distinctively colonial Hydromedusæ,

many of them most exquisitely beautiful and graceful forms.

In size they vary from very minute forms barely visible to the

unaided eye, to forms like Halecium, measuring from twelve

to twenty inches or more in height. The hydranths are provided

with specialized receptacles, hydrothecæ, into which they are

capable of more or less complete retraction. Gonophores are pro-

duced by budding, and are provided with specialized receptacles,

‘gonangia, similar in morphological features to the hydrothecæ.

The gonophores may be liberated as free medusa, or may

remain fixed as medusoids, the sexual products maturing within

the gonangium and later escaping as free larvae or planule.

When free, the medusze are known as Leptomedusz, charac-

terized generally by a low, flat bell, marginal sense organs

usually of the vesiculate type, with the gonads usually borne

along the underside of the radial canals.

A classification of the Campanularida is almost, if not quite,

impossible without the presence of the gonosome, which in

many genera is the most distinctive differentiating feature.

In the following synopsis this feature will be in constant requi-

sition, and where it is absent in specimens the student is admon-

ished as to the doubtful character of purely morphological

determinations.

379

380 THE AMERICAN NATURALIST. [VoL. XXXV.

SYNOPSIS OF FAMILIES.

CAMPANULARID&. Hydrothecze campanulate, terminal, borne on dis-

tinct pedicels ; gonophores fixed or free-swimming. Hydranths with large

and somewhat trumpet-shaped hypostome.

LAFŒIDÆ. Hydrothece deep tubular, sessile or pedicellate ; hydranths

with conical hypostome.

HALECIDÆ. Hydrothece usually reduced to shallow, disk-like recepta-

cles (hydrophores). Hydranths with conical hypostome. Gonophores as

imperfectly developed medusoids.

SERTULARID&. Hydrothecz borne in double rows, adnate to hydro-

caulus. Gonophores sessile.

PLUMULARID;. Hydrothece arranged in single row only on side of

hydrocaulus.

CAMPANULARID&.

Synopsis of the Genera.

CLvTIA. Stems simple or rarely branched. Hydrothece deeply bell-

shaped, with toothed margins, borne on long pedicels. Gonangia pro-

ducing free meduse having four marginal tentacles. ;

OBELIA. Stems regularly branched, hydrothecæ bell-shaped, with entire

margins. Gonangia borne on stems and branches and producing free

medusæ having numerous marginal tentacles.

CAMPANULARIA. Stems simple or branched. Hydrothecæ campanu-

late, with margins entire or variously toothed. Gonangia, medusæ as mere

sporosacs, within which the sexual products develop and escape as rer

planulæ. :

GONOTHYRÆA. Stems branching ; hydrothecæ campanulate and with

toothed margins. Gonangia producing well-developed medusoids, which,

while often furnished with tentacles and capable of protruding beyond the

orifice of the gonangium, never become free, thus exhibiting an interesting

intermediate stage between the first two genera and Campanularia.

Clytia Lamx. (in part).

Generic characters : Stem usually simple, attached by creeping hydro-

rhiza. Hydrothecæ devoid of operculum. Gonangia produced from stem

or hydrorhiza and borne on pedicels which are usually beautifully annulated.

Gonosome. Medusz deeply bell-shaped and with four marginal tentacles

when first liberated. Otocysts eight, two in each interradius. Both these

and the tentacles increase in number with the age of the medusa.

No.413.] MORTH-AMERICAN INVERTEBRATES. 381

1, C. bicophora Ag. (FIG, 16).

Trophosome : Colony rarely attaining a height of more than an inch,

composed of simple or sparingly branched stems. Hydrothece deeply bell.

shaped and numerously and sharply toothed,

borne on elongate pedicels which have terminal

annulations.

Gonosome: Gonangia symmetrically annu-

lated and usually arising from the hydrorhiza.

Meduse when first liberated of hemispherical

shape and with four tentacles and eight otocysts.

Habitat: Usually on fucus, occasionally on

shells or other hydroids.

2, C. cylindrica Ag.

(Cont. Nat. Hist. U.S., vol. iv.)

Trophosome : Stems simple, hydrothece tubu-

lar, small, deep, with sharply pointed teeth. Pedi-

cels short, with proximal and distal annulations.

Gonosome : Gonangia oblong, somewhat flat-

tened, devoid of annulation, producing free

edusæ. Fic. 16, — Clytia bicophora Ag.

(After Agassiz.)

Habitat : Similar to last species.

3, C. grayi Nutting.’

Trophosome : Stem simple or irregularly branched, strongly annulated

except in middle branch. Hydrothecæ very large, cylindrical. Numerous

marginal teeth, rounded and not deeply cut. Hydranth with about twenty

tentacles

Gon : Gonangia oblong, conspicuously and regularly annulated,

attached » creping rootstocks.

Habitat : Growing on living worm tubes, composed of sand. Dredged

from Pt of 31 fathoms. The largest Clytia yet found in American

waters.

Obelia Peron and Leseur.

Generic characters : Colony often plantdike, of whitish color, attached

by M dsl hydrothecae campanulate and devoid of operculum.

angia borne on stems and branches, producing free medusæ character-

ted by numerous marginal tentacles, four radial canals, and eight otocysts

symmetrically disposed on the inner margin of each interradial quadrant.

1 Condensed from Professor Nutting’s original description.

382 THE AMERICAN NATURALIST. [Vor. XXXV.

i, O. commisuralis McCr. (Fic. 17).

Trophosome: Colony long, slender, profusely branching, branches spread-

ing in graceful curves on each side of the main stem, which may attain

a height of six to eight inches.

Gonosome : Gonangia elongate, slen-

der, obconical, opening by terminal,

circular orifice arising from the base

of the cone on a short conical neck.

Medusz when first liberated have six-

teen marginal tentacles, four radial

canals, beneath which later the gonads

develop.

2, O. dichotoma Linn.

Trophosome: Colony rather small,

stem slender, irregularly branched, an-

nulated just distal to origin of branches,

the latter annulated at irregular inter-

alisMcCr. — vals. Hydrothece large, deeply cam-

panulate, borne on annulated pedicels.

Gonosome : Gonangia axillary, slender and smooth, somewhat obconical,

and similar to those of former species. Medusz with sixteen tentacles,

manubrium somewhat trumpet-shaped.

Fio. t:5

pu ^um

(After Agassiz.)

3, O. flabellata Hincks.

Trophosome: Stem filiform, alternately branching, giving the stem 4

somewhat zigzag character. Both stem and branches variously annulated.

Hydrothecz alternate, short, widely open and with entire margins, borse

on tapering annulated pedicels.

Gonosome : Gonangia axillary, obovate, with tubular orifice. Medusz?

4, O. geniculata Linn. (Fic. 18).

Trophosome: Colony inconspicuous, rarely attaining a height of more

than an inch. Stem somewhat zigzag in form as in former species, bet

apparently jointed at each bend. Hydrothece obconical, rather short,

with plain orifice, borne on short annulated pedicels.

Gonosome : Gonangia axillary, urceolate, borne on short pedicels.

Medusz discoid, with twenty-four tentacles when liberated, greatly increas-

ing in number with age.

Habitat : Common along Massachusetts and north Atlantic coast, OP

Fucus and Laminaria.

No. 413.] MWORTH-AMERICAN INVERTEBRATES. 383

5, O. gelatinosa Pallas.

Trophosome: Stems fascicled, rising from a fibrous hydrorhiza to a

height of eight to ten inches. Branches opposite in pairs, which alternate

with each other in vertical arrange-

ment, presenting a verticillate appear-

ance. Hydrothece small, borne on

long slender, ringed pedicels, and hav-

ing notched margins of a somewhat

castellated form.

Gonosome : Gonangia axillary, ovate,

flattened at distal end and provided with

raised orifice. Medusæ with sixteen

tentacles when liberated from gonan-

gium.

6, O. longissima.

7, O. bicuspidata.

8, O. bidentata. Fic. 18.— Obelia geniculata Linn. a, go-

nangium of same enlarged.

Species 6, 7, 8 are listed from Pro-

fessor Nutting’s records, but have not been taken by the present writer.

Campanularia Lamx. (in part).

The generic characters are fairly explicit under the synopsis of genera.

1, C. caliculata Hincks.

(Clytia poterium Ag.)

Trophosome : Stem simple, of variable length, bearing a single hydro-

theca which is campanulate, with entire margin, and with a thick wall form-

ing a sort of diaphragm within the lower part, thus giving the appearance

of a double-walled cup.

Gonosome : Gonangia irregular, oval in shape, with undulating outline

and with wide circular aperture, edusoids extremely degenerate. Larva

escaping as free-swimming planule.

Habitat : Massachusetts Bay, Nahant, Nova Scotia, on seaweed, etc.

2, C. hincksii Alder.

Trophosome: Stems rather long, mostly simple; hydrothece large,

deep, almost tubular, the margins scalloped with castellated teeth.

nosome: Gonangia ovate, elongate, somewhat narrowed toward ex-

tremity, irregularly annulated throughout, borne on short, smooth pedicels.

Medusoids degenerate; ova forming a central mass within the capsule.

384 THE AMERICAN NATURALIST. [Vor. XXXV.

3, C. volubilis Linn.

Trophosome: Stems usually simple, long and somewhat twisted. Hydro-

thecz deep and sub-tubular, margins with shallow undulations.

Gonosome: Gonangia flask-shaped, smooth, with an elongate neck

borne on short pedicels.

Habitat: Frequently found growing upon other hydroids, usually in

deep water. Gulf of St. Lawrence, Massachusetts coast, etc.

4, C. neglecta Alder.

Trophosome : Stems regularly branched, delicate, filiform, branches pin-

nate, both stem and branches more or less annulated. Hydrotheca narrow,

deep, borne on annulated pedicels and with marginal teeth bimucronate.

Gonosome ; Gonangia axillary or on short pedicels which are annulated,

pear-shaped.

The colony is very minute and inconspicuous.

5, C. verticillata Linn.

Trophosome: Colony composed of erect, fascicled stems, irregularly

branched. Hydrothecz bell-shaped, rather large, deep, with from ten to

twelve teeth about the margins,

\ LI

WW borne on annulated pedicels.

y A IPA ^

: YB pa Gonosome: Gonangia flask-

if f shaped, smooth, borne on short

"c. fg. F 2 è .

f £2 pedicels and terminating in narrow

Trophosome : Colony resembling

in general aspects that of Obelia

commisuralis, attaining in some

cases a height of four to six inches.

J ; orifice.

N * 6, C. amphora Ag. (FIG. 19).

E ue M Hydrothece campanulate, with en-

AN N Em : tire margins, borne on annulated

Noll a pedicels.

~ WA Gonosome : Female gonangia elon- —

gate, somewhat obconical, borne 0?

Y short annulated pedicels and opening

by a terminal aperture. Male gonan-

T E s ad ss gia elongate, oval or spindle-shaped-

(After Agassin) . ^ pMedusoids more or less degenerate,

never becoming free ; the male, ac-

cording to Agassiz, attaining a higher stage of development than the female.

The embryo escapes from the gonangium as a free-swimming planula.

No. 413.] MWORTH-AMERICAN INVERTEBRATES. 385

7, C. angulata Hincks (Fie. 20).

Trophosome: Stems slender, slightly branched, strongly geniculate or

undulate in habit. Hydrothecz alternate, campanulate, with entire margins,

borne on long slender pedicels which arise at each flexure of the stem or

branch.

Gonosome: Gonangia somewhat ovate, obscurely wrinkled, and termi-

nated by a broad aperture. Colony small, varying from % to 3( inch.

Fic. 20. Fic. 21.

Fic. 20. — Campanularia angulata Hincks. (After Hincks.)

Fic. 21. — Campanularia flexuosa Hincks. (After Hincks.)

8, C. flexuosa Hincks (Fic. 21).

Trophosome: Stem flexuous, irregularly branched, annulated near the

io and above the origin of branches. Hydrothecz large, subcampanulate

With plain margins, borne on long annulated pedicels.

Gonangia axillary, large, elongate, smooth, and borne on

onosome:

short annulated pedicels. Male gonangia sensibly smaller.

386 THE AMERICAN NATURALIST. | [Vor. XXXV.

Professor Nutting has recorded the following species of which I have

no data, and which therefore are merely noted.

C. minuta; C. Edwardsii Nutting ; C. calceolifera.

Gonothyrea Allman.

1, G: Jovéni Allman (Fic. 22).

Trophosome: Stems erect, somewhat flexuous, irregularly branched, and

with annulations above each branch. Hydrothece deeply bell-shaped and

with toothed margins, borne on short

NJ V. annulated pedicels.

r N Gonosome : Gonangia borne on short

: annulated pedicels, axillary, broadly ob-

[^ conical in outline.

4 Habitat: On fucus and other alge,

Vy [9 rocks, etc. Cold Spring Harbor, Woods

We B d Holl, etc.

| abe ii

á Ñ M uf 7, 2, G. hyalina Hincks.

" N

Trophosome : Colony elongate, clustered,

SB fy x profusely branched, with flexuous stems

S d d^ giving off branches at each bend,

CNN } 7 4 Branches erect, very tender and hyaline,

EN y sometimes of great length and much rami-

oh fied." Hydrotheca elongate, of delicate

NN. Bip [t texture, with numerous marginal teeth and

See Y Y borne on annulated pedicels.

cw \ N Gonosome: Gonangia oval, axillary,

QN j borne on annulated pedicels.

Habitat : On various hydroids, Tubula-

b ria, Halecium, etc. (Hincks).

3, G. tenuis Clark.

Fic. 22. — Gonothyrea loveni Allman. ‘no's list.

ter Hincks.) Noted from Nutting's

LAF@IDA.

This family has been variously modified of late and by some replaced

entirely. In the present synopsis I have chosen to follow in general the

classification of Hincks, though recognizing its doubtful reliability in x

respects.

Lafca Lamx.

Stems simple or fascicled, attached by filiform hydrorhiza. Hydrothec®

tubular, with or without operculum. Gonangia oblong, often io

encrusting masses about the stem.

No. 413.] WORTH-AMERICAN INVERTEBRATES. 387

1, L. dumosa Flem. (Fic. 23).

Trophosome: Stem creeping, sometimes erect and fascicled ; hydrothece

tubular, margins devoid of teeth or operculum, usually sessile.

Gonosome: (?)

2, Z. calcarata A. Ag. (FIG. 24).

Trophosome: Stems creeping, simple; hydrothecz tubular, sessile.

Gonosome: Gonangia large, elongate, obovate or oblong, somewhat

resembling those of certain campanula-

Meduse large, transparent, with

Fic. at Trope donet Pe (After Hin

Fic. A. Ag yir bon A. Agassiz.)

gonads suspended in folds beneath the radial canals; marginal tentacles

numerous in mature specimens, only two when first set free.

Habitat: Usually parasitic upon sertularian hydroids.

3, LZ. pygmea Alder.

Trophosome: Stem creeping; — minute, tubular, elongate,

borne on very — annulated pedice

Gonosome : (?

Habitat: Parasitic on various hydroids.

HALECIDA.

Of this family a single genus comes within the range of this synopsis :

namely, the type genus, Halecium (Oken), the characters of which may be

Summarized as follows:

Trophosome : Colony more or less branched, attached by a creeping

hydrorhiza, Hydrotheca often shallow and disk-like, or funnel-shaped

(hydrophores). In many species with double or triple margins due to

388 THE AMERICAN NATURALIST. [Vor. XXXV.

subsequent secretions as the hydranth grows, leaving the old hydrophore. In

many cases the everted rim has on its inner margin a circle of small bright

dots which are rather characteristic of the genus. Hydranths imperfectly

retractile, elongate, and with conical hypostome.

Gonosome : Gonangia of varying aspects, showing distinctive differences

between male and female and affording easy means of distinguishing the

sexes. Medusoids imperfectly developed, never free.

1, H. halecinum Linn. (FIG. 25).

Trophosome: Colony erect, rather rigid, subflabellate in form. Hydro-

thecz alternate, somewhat tubular in form, and with everted rims.

Gonosome : Gonangia borne in a series on the upper side of the branches ;

those of the male elongate, slender, somewhat spindle-shaped, tapering below

e b

Fic. 25. — Halecium halecinum Linn. a, male; 4, female gonangia of same

(enlarged). (After Hincks.)

to their attachment by very short, slightly ringed pedicels. Female gonan-

gia somewhat oblong, broader toward the distal end, and with tubular

aperture nearer one margin.

2, H. beanii Johnston.

Trophosome : Colony of delicate, graceful form, somewhat dendritic,

attaining a height of about two inches. Hydrothece with everted

rims.

Gonosome : Gonangia arising from near the base of hydrothece ; male,

elongate oval; female, somewhat curved, with the aperture situated near

the middle of the upper side.

No.413.] MWORTH-AMERICAN INVERTEBRATES. 389

3, H. tenellum Hincks.

Trophosome: Colony minute, extremely delicate; stems slender, often

strongly annulated, branching irregularly. Hydrothece funnel-shaped and

with everted margins.

Gonosome : Gonangia ovate, pedicellate.

4, H. muricatum Ell and Sol.

Trophosome: Colony stout, dendritic, profusely branched, and with

joint-like divisions, alternately from below which the hydrothece arise.

onosome : Gonangia ovate, borne on short pedicels, roughly marked

with linear ridges of spinous processes.

Eastport, Me. (Verrill).

SERTULARID&.

Synopsis of Genera.

SERTULARIA. Colony plant-like, stems more or less branching, jointed,

attached by creeping hydrorhiza. Hydrothece in double rows, strictly

opposite, usually devoid of operculum. Gonangia with plain margins.

ERTULARELLA. Colony resembling somewhat the former. Hydro-

thece in double rows, but distinctly alternate, with toothed margins and

with an operculum composed of several pieces. Gonangia strongly annu-

lated throughout, slightly dissimilar in the two sexes.

Dipwasta. Colony more or less branching, stem jointed, hydrothecze

Opposite, a pair to each internode and often with a valve-like operculum.

Gonangia scattered, differing in shape in the two sexes, those of female

large, often divided into segments above, male smaller and with central

tubular aperture.

THUIARIA. Stem somewhat plant-like, jointed ; hydrothece in double

Series sub-opposite, but deeply immersed in the substance of stem and

branches.

HYDRALLMANIA. Stems flexuous or somewhat spirally inclined. Hydro-

thecæ alternate, placed on front of branches, and curved alternately to

right and left. ;

Sertularia Linn.

Generic characters given above.

1, S. pumila Linn. (FIG. 26).

Trophosome: Stems straight or slightly curved, simple or branched ;

branches opposite ; both stem and branches divided into short internodes,

each bearing a pair of hydróthecz, the latter opposite, tubular, and some-

what contracted toward the aperture, which faces outward and is more or

less cleft or notched.

390 THE AMERICAN NATURALIST. [Vor. XXXV.

Gonosome: Gonangia more or less oval, sessile, with marginal rim,

Male gonangia somewhat more slender, and regular in outline.

Habitat: One of our commonest sertularians, found attached to fucus,

etc., between tide marks and in tide pools.

2, S. cornicina McCr.

Trophosome : Colony very small, composed chiefly of unbranched stems,

which rarely attain a height of more than % inch. Hydrothece appearing

as lateral emarginations with slightly diver-

gent apertures. Hydranths slender, with

about sixteen tentacles.

Gonosome : Gonangia?

The above description is condensed and

modified from that of McCrady (Proc.

Elliott Soc., Vol. 1, p. 204).

3, S. argentea Ell. and Sol.

Trophosome : Colony of bushy and slightly

wavy stems, perisarc dark and horny; branch-

ing, alternate, and somewhat dichotomous.

Hydrothece short, urn-shaped, tapering to-

ward the free and divergent aperture, which

is small and oblique.

Gonosome: Gonangia broad, obovate,

tapering toward the base; aperture circular, and usually with two

divergent spines.

abitat: Usually from deeper waters, growing on shells, stones, etc.,

sometimes found near tide marks. Recorded from various points along the

New England coast.

Ftc. 26.

Qo 2447, > £g sp Linn.

(After Agassiz.)

4, S. cupressina Linn.

Trophosome : Colony slender, elongated. Stems rather stout and straight,

alternately branched and dichotomously sub-branching. Hydrothece tubu-

ar, transparent, somewhat alternate, and adherent throughout most of their

length, slightly divergent toward the aperture, which is wide and bilabiate-

Gonosome: Gonangia elongate, tapering toward base, and with promi

nent spine at each side of the aperture, which is slightly raised and central.

Habitat: Less abundant than the former species, though with similar

distribution.

Sertularella Gray.

Generic characters given in above synopsis.

No.413.] NORTH-AMERICAN INVERTEBRATES. 391

, oS. rugosa Linn.

Trophosome : Colony small, simple, or sparingly and irregularly branched ;

stems annulated. Hydrothece crowded, strongly annulated transversely,

and with four marginal teeth

Gonosome : Gonangia large, ovate, strongly annulated, and with a four-

toothed aperture.

2, S. gayi Lamx.

Trophosome : Stems erect, with alternate branches, somewhat obliquely

jointed. Hydrotheca somewhat urn-shaped, one to each internode, usually

wrinkled, and with narrower, diver-

gent, four-toothed aperture.

Gonosome : Gonangia elongate,

42999. ovate, tapering toward the small,

of two-toothed aperture. Usually

S strongly annulated in upper por-

tion, the lower smooth.

3, S. tricuspidata.

Trophosome: Stems slender,

Fic. 27.—Diphasia fallax Johnst. (After Hincks.) a, 9 gonangium of same (enlarged).

alternately branched, often bipinnate near the ends. Hydrothecæ cylin-

drical, prso slightly everted, with a three-toothed orifice

Gonosome : Gonangia large, with strongly transverse ridges and with a

plain, PORE saai opening.

Diphasia Ag.

, D. fallax Johnston (Fic. 27).

Trophosome : Stems thick, sparingly branched, branches alternate, often

terminating in tendril-like bodies. Hydrothece short, tubular, with upper

Part slightly divergent, and with wide, smooth orifice.

. 392 THE AMERICAN NATURALIST. [Vor. XXXV.

Gonosome : Gonangia differ in the two sexes. Male elongate, slender,

tapering toward base and expanding toward orifice, which bears four stout

spines. Female gonangium oval, deeply cleft above into four leaf-like seg-

ments, larger than male.

2, D. rosacea Linn.

Trophosome: Stems slender and delicate, branches alternate and with

internodes constricted at the base. Hydrothece long, tubular, with upper

portion free and divergent toward

the aperture, which is oblique and

entire.

N Gonosome : Gonangia slightly |

\ ri different in the sexes; female pear-

shaped, elongate, borne on short

pedicels and marked with eight lon-

gitudinal ridges, each terminating

above in a spinous process. Male

somewhat curved toward base, with

similar longitudinal ridges terminat-

ing in spinous teeth about the slender

tubular orifice.

Thuiaria Flem.

A single species of this genus

comes within the present synopsis.

Thuiaria thuja Flem.

Trophosome : Stem and branches

rather rigid, somewhat zigzag in

shape, and annulated near the base.

Perisarc black or very dark in color.

Hydrothecæ smooth, ovate at base

and tapering toward the distal end.

Gonosome: Gonangia smooth,

pyriform, and with circular slightly

emarginate aperture.

Fic. 28. — H'ydralImania Jalcata Linn,

incks.)

Hydrallmania Hincks.

Hydralimania falcata Linn. (Fics. 28, 29).

(Sertularia falcata.)

Trophosome: Stems flexuous, slender, sometimes spirally inclined.

Branches alternate, regularly pinnate and plume-like, arising just above

No. 413.] WORTH-AMERICAN INVERTEBRATES. ` 393

each joint. Hydrothecz tubular, closely appressed, arranged in rows along

the pinnz, and with plain oblique aperture.

Gonosome: Gonangia ovate, tapering toward the base, and with a

tubular orifice.

Habitat: Shells, stones, etc., generally distributed

from Grand Manan, Massachusetts Bay, and south-

PLUMULARID&.

Synopsis of genera. Modified and

condensed from Nutting’s Mono-

graph of the Plumularide.

ANTENNULARIA. Colony more or

less arbuscular, stem simple or branch-

Fic. 29. — Hydrallmania falcata Linn verticillate or scattered ; hydrothecae

eui praagi. cup-shaped ; nematophores trumpet-

shaped. Gonangia borne in axils of branches, unilateral.

Monasta&cuas. Colony dichotomously branched, stem not fascicled,

hydrocladia arising from upper sides of branches, otherwise resembling

Plumularia, from which it differs in the entire absence of cauline hydrothecz.

Gonangia oval in shape and with terminal aperture.

ScHIzoTRICHA. Colony branching, branches pinnately arranged, hydro-

cladia often forked. Gonangia borne on stem or hydrocladia.

CLADOCARPUS. Stem simple or fascicled. Nematophores not trumpet-

shaped, definitely fixed to hydrothecae or branches. Gonangia borne on

stem or hydrocladia.

Antennularia Linn.

1, Antennularia antennina Linn. (FIG. 30).

Trophosome : Colony growing in dense clusters of upright stems, often

eight to ten inches high,

Stems simple or sparingly

cai obscurely jointed,

each internode bearing a

Gonosome: Gonangia

9vate, borne singly in axils

of hydrocladia. Aperture Fic. go.— Antennularia antennina Lim. Portion of

subterminal. stem and hydrocladia (enlarged). (After Nutting) .

394 THE AMERICAN NATURALIST, [Vor. XXXV.

2, A. americana. Nutting.

Similar to former, but usually from deeper water and apparently of

exceedingly variable character.

3, A. rugosa Nutting.

Trophosome: Colony unbranched, attaining a height of six inches.

Hydrocladia in verticils of six or eight, borne on stout processes of the

stem and with proximal ends reinforced on the lower sides by a thickening

of the perisarc. Internodes long and irregular, further subdivided by

numerous irregularly disposed septal thickenings, which resemble joints,

giving the appearance of many internodes, where in reality there is but one.

Hydrothecz small, short, cylindrical, and supported below by a thickening

of the internode.

Monastechas quadridens McCr.

Trophosome : Colony subflabellate in form, dichotomously branched,

attaining a height of about six inches. Stem not fascicled, with indistinct

internodes and branching at irregular intervals, those bearing hydrocladia

being divided into long internodes, each of which bears a hydrocladium at

its distal upper side. Hydrothece large, campanulate.

Gonosome ; Gonangia sac-like, borne on short processes below hydro-

thecæ, and each protected by a pair of nematophores.

Habitat: Various stations along the North Atlantic coast, and from

Marthas Vineyard southward.

Schizotricha Allman.

Hydrocladia pinnately disposed, often branching once or more. Two

species come within the range of this synopsis.

1, S. Zene//a Verrill.

Trophosome: Colony branched dichotomously, attaining a height of

about two inches. Stems clustered or fascicled, divided into alternately

longer and shorter internodes, the latter bearing each a hydrotheca and a

hydrocladium. Hydrocladium slender, often branched, proximal intern es

short, and without hydrothecz, which are subcylindrical.

Gonosome : Gonangia of curved shape, tapering at base and gradually

enlarged toward the distal end, somewhat resembling cornucopiz.

Habitat: Gay Head, Vineyard Sound, New Haven, Greenport, R. I,

Woods Holl, Vineyard Haven

No. 413.] WORTH-AMERICAN INVERTEBRATES. 395

2, S. gracillama Sars. (FIG. 31).

Trophosome: Stem sparingly branched, having a height of about two

inches and somewhat fascicled. Branches divided into regular internodes,

each of which bears a hydrocladium on a short, stout process near its distal

. Hydrocladia alternate, branching

dichotomously twice or more beyond its

proximal internode. Hydrothe-

cæ small, cup-shaped.

Gonosome : Gonangia borne

in pairs on the stems in the

axils of the hydrocladia, and

also at the forks of the latter,

cylindrical in shape, tapering at

proximal ends, sessile.

Habitat: Shallower waters,

New England coast.

Cladocarpus Allman.

Stem simple or fascicled.

Hydrothece deep and with

smooth margins or with lateral

'sinuations, and with one or two

anterior teeth. A single species

comes within the range of this

synopsis.

Fic. 31. Fic. 32.

Fic. 31. — ScAizotricha gracillima Sars. Branched

hydrocladium (enlarged). (After Nutting.)

Fic. 32. — Cladocarpus flexilis Verrill. Portion of C. flexilis Verrill (Fic. 32).

ydrocladium (enlarged). (After Nutting.)

Trophosome: Colony long,

slender, sparsely branching, stem not fascicled, attaining a height of about

nine inches. Hydrocladia distinct, slightly sinuous, divided into rather

slender internodes, each with a number of septal ridges back of hydrothecæ,

which are deep, tubular, nearly straight, and with a single anterior tooth at

the aperture.

Gonosome : Gonangia numerous, borne on stem and bases of hydro-

cladia, oblong-ovate, with latero-terminal orifice.

REVIEWS OF RECENT LITERATURE.

ANTHROPOLOGY.

Human Spines.' — The paper is, in the words of the author, “a

description of forty-five anomalous human spines in the Warren

Museum,” and of a number of special parts of the spinal column

from the same collection ; it is also a discussion of the causes of the

spinal variations.

The author describes five classes of spinal anomalies, namely :

1. Spines in which the number of presacrals is normal, but in

which there is an irregularity at the junction of the thorax and loins,

or at the junction of the thorax and neck.

2. Spines in which the 26th is the v. fu/cralis, but in which the

25th is not quite separated from it.

3. Spines in which there are more than 24 perfectly free prasa-

crals, the extra one being thoracic, or lumbar, or there being two

extra præsacrals, one thoracic and one lumbar, the latter sacralized

on one side, the 27th being the fu/cralis.

4. Spines in which one or more praesacral vertebra are imperfectly

developed, one or more vertebrze being fused, the atlas being fused

with the occiput, or the 24th being more or less sacralized.

5. Spines in which there is a presacral too few: a vertebra being

wanting in the loins, in the back, there being 12 pairs of ribs, the

first pair being cervical and perfect on one side, the 24th being in

all the groups the Julcralis.

There are further described cervical, rudimentary first thoracic,

bicipital and tricipital ribs ; fusion of atlas and occiput, of atlas and

axis, axis and third cervical vertebra; a suppression of a cervical

and an extra half vertebra.

‘he main facts brought out by the paper are (1) a lack of relation

between the condition of the spine at one end of the thorax and that

at the other, and (2) the frequency of “concomitant” variations on

one or both sides of the spine.

_ | Dwight, Thomas. Description of the Human Spine, showing Numerical Varia-

tion, in the Warren Museum of the Harvard Medical School. Memoirs of the

Boston Society of Natural History, vol. v, No. 7 (Boston, 1901), pp- 237-312»

with figures.

397 .

398 THE AMERICAN NATURALIST. [Vor. XXXV.

(1) “If the undeveloped end of the rst thoracic rib is a step

towards the future, it would be reasonable to expect in the same

spine a corresponding advance below the thorax. Conversely, if

there is an archaic condition below the thorax, there should be an

analogous condition above it. While there are cases that fulfill these

conditions, they are quite lost in the multitude which do not, and

which even present contradictory conditions at the opposite ends

of the spine, being retrogressive at one end and progressive at

the other.”

These facts are in contradiction to Rosenberg’s theory.

(2) The author points to the cases where * we see a tendency

sometimes for the whole thorax to move forward (upward ?) by cervi-

cal ribs associated with absence of the last thoracic ones or with

their existence in a rudimentary condition.” “We also see cases in

which, when the cervical rib on one side is distinctly larger than

its fellow, the last rib on that side is either correspondingly smaller

than its fellow, or even replaced by a pretty typical transverse

process.” These concomitant variations **may extend even further, so

as to include the sacralization of one side of the last lumbar, or even

the absorption of one side of the atlas into the occiput.”

The causes of the variation: The author confesses his inability to

show the original cause of the phenomenon. “It is clear, however,

that the vertebra at the junction of regions are particularly variable,

and it seems hard to doubt that errors of segmentation may occut-

The original error having occurred, there seems to be a tendency in

the organism to reproduce the type as nearly as may be under the

changed conditions; to make as normal a series of regions as CI

cumstances will permit; and this tendency manifests itself to some

extent independently in the two halves of the spine." For this

tendency the author adopted the old and rather unsatisfactory theory

of ** the vital principle."

Professor Dwight closes his interesting work with the following

additional deductions :

1. Variations occur in two ways : (1) by irregular development of

the costal elements at and near the ends of the regions of the spine,

and (2) by irregular segmentation through which there are more OF

fewer vertebrz: than normal | :

2. Variations of both kinds are variations around a mean. It 1s

not impossible that some of them may be reversive ; that any are

progressive is mere assertion. :

3. Assuming the correctness of Rosenberg's studies in ontogenesi

No. 413.} REVIEWS OF RECENT LITERATURE. 399

his view may account for some of the variations, but even in these

cases something more is needed to explain the concomitant changes.

4. Variation of the costal elements at one end of a region is often

associated with variation of an opposite nature of those at the other

end. Several regions may be involved, and the two sides may vary

independently.

5. Variations, which separately seem either reversive or progres-

sive, generally lose that appearance when the whole spine is

considered.

6. After the occurrence of the original error in development there

is a tendency for the spine to assume as nearly as possible its nor-

mal disposition and proportions. This, as do also concomitant vari-

ations and indeed all development, implies a * vital principle."

These deductions of the author naturally invite discussion ; but it

will be of advantage if this be deferred until the material bearing on

the points in question is still more abundant and the observations

extended. Conclusions of this nature apply not only to the part

under consideration but largely to the whole skeleton. 4A B.

Notes.— Four * Cruciform Structures near Mitla" are described

by Mr. M. H. Saville in Vol. XIII of the Bulletin of the American

Museum of Natural History. After a scholarly summary of the

history of previous explorations at Mitla the author confines his

attention to the cruciform burial chambers which are unique in form

and surpass all other tombs in Mexico or Central America in size

and in beauty of stone work.

“A Bilateral Division of the Parietal Bone in a Chimpanzee ;

With a special Reference to the Oblique Sutures in the Parietal,” is

the subject of a paper by Dr. Aleš Hrdlička, appearing in the same

volume. It contains a detailed description of the skull of an adult

male chimpanzee, with a discussion of the important problems con-

nected with the abnormal parietal sutures. An oblique suture, the

author believes, can be attributed to only three possible causes, as

follows : an early fracture, a persistence of the original separation

between the two centers from which the bone is developed, and a

Coexistent difference between their relative positions ; the existence

of à supernumerary third center of ossification. The brochure is

illustrated by six outline drawings.

In the Report of the Museums Association of the United Kingdom

for 1898 Mr. Harlan I. Smith advises an “ ethnological arrangement

400 THE AMERICAN NATURALIST. [Vor. XXXV.

of archeological material and suggests a classification under

thirteen main divisions of such material, with a view to illustrating

ancient tribal life and ethnology. This method would make the

specimens aid in solving problems and would find a use for many

now discarded as unworthy of attention.

The Report for 1899 contains a paper by Mr. Smith upon “the

preservation of local archzological evidences,” showing the danger of

their obliteration by man and nature, and the necessity of systematic

explorations in this country as well as in foreign lands. More accu-

rate and complete records should be kept by means of indices and

catalogues. Mr. Smith makes suggestions for local work under the ^

auspices of state universities and historical societies.

An interesting example of *psycho-physical study" is given in

Mr. Arthur MacDonald’s “Emile Zola," reprinted from Zhe Ofen

Court, August, 1898. This study was made by a number of French

specialists, and the result published with the approval of the subject.

Among the characteristics investigated are antecedents, mental

evolution, physical peculiarities, the nervous system, ideas, senti-

ments, and will This empirical method is employed in order that

we may come to have somatology of the living as well as of the dead,

and thus gain a knowledge that will be of practical use in amelio-

rating social conditions.

In the Bulletin (Vol. VIII, No. 2) of the French Society of Anthro-

pology, M. Zaborowsky discusses at some length the problems of the

racial unity and the place of origin of the Slavs. He concludes that:

1. The Slavs of the North came from the region between the

Danube and the Adriatic. They were related to the neighboring

inhabitants of the terramare of Emilia. The migrants may have

been drawn toward the north by the trade in amber.

2. They passed the Carpathians through the valleys of the Oder

and Vistula, following especially the latter to the vicinity of the

Baltic, where they developed an independent culture.

3. The Slavs introduced the custom of cremation, until then

unknown. They brought metals and glass; iron was used for orna-

ment only.

4. They formed a branch or included the tribes of the Venedes of

the Adriatic; the name Venedes dates back in the Baltic region to

the fourth century before our era.

5. By their dominant characters they approach the French Keltic

type. The southern Slavs are dark and brachycephalic. To the

No.413.] REVIEWS OF RECENT LITERATURE. 401

northward of the Carpathians they encountered a neolithic blond

population which they absorbed and in part perpetuated in certain

regions.

6. Upon the Oder, the lower Vistula, and along the Baltic coast,

where they have pushed back the Finns toward the east, their culture

was modified by contact with the Germans from Scandinavia,

“ZOOLOGY.

Evermann and Marsh on the Fishes of Porto Rico. — One of

the most thoroughly admirable of faunal works is the report on the

Aquatic Resources and Fisheries of Porto Rico, just published by the

United States Fish Commission. The authors are Dr. Barton

Warren Evermann and Millard C. Marsh.

In this work are given full descriptions of 291 species, arranged

systematically, with analytical keys and numerous figures in the text.

A general discussion of the waters and of the geographical features of

the island is given by Dr. Evermann, and a chapter on the fish trade

and fishing methods by William A. Willcox.

The work is illustrated by 49 colored plates, by C. B. Hudson and

A. H. Baldwin. The accuracy and excellence of these plates cannot

be too highly praised. Without invidious comparisons we may doubt

if any plates of fishes ever published excel in fineness of coloration

Some of these (as the Nassau Grouper, Plate XII, and the Red

Hind, Plate XIII) by Mr. Hudson.

The nomenclature and definitions of groups are taken chiefly from

Jordan and Evermann’s Fishes of North America, and the new species

are mostly described in the final appendix to that work.

Those not thus included are the following :

Aphthalmichthys caribbeus, Gill and Sphagebranchus ophioneus, Mayaguez.

mith, San Geronimo,

Lycodontis albimentis, Culebra Island. Apogon sellicauda, Culebra Island.

ps cleroperca bowersi, Culebra. Neomenis megalophthalmus, Puerto Real.

Poi albifimbria, Culebutas. Scorpena bergi, Mayaguez.

minus beanorum, San Juan. Emblemaria pandionis, Isabel Segunda.

Citharichthys arenaceus, Mayaguez. Halieutichthys smithi, Mayaguez.

The name Peprilus is substituted for Rhombus, preoccupied in

mollusks. A few other changes in the nomenclature, adopted from

Jordan and Evermann, will be found necessary. These may be

402 THE AMERICAN NATURALIST. [Vor. XXXV.

briefly noted without statement of the reasons: Sardinia for Clupa-

nodon, which should be restricted to the group in Japan and China,

lately named Konosirus ; Anchovia for Stolephorus, which was based

on a Japanese Spratelloides; Esox (Esocidz) instead of Lucius;

(the type of a Linnzan genus, according to Linnzus," is the best

known European or officinal species") ; Syngnathus for Siphostoma,

the same rule covering these cases; Bodianus should replace Harpe,

the genus called Bodianus by Jordan and Evermann standing as

Cephalopholis; Dipterodon should replace Neomzenis, if the latter is

really distinct from Lutianus; Eupomacentrus is probably not distinct

from Pomacentrus; Tropidichthys should replace Canthigaster, the

latter a bare definition without species. Probably Carapus must

replace Fierasfer. Lepisoma must take the place of Labrisomus,

which replaces Gobioclinus. Probably Ichthycallus should be used

instead of Iridio. Det

Studies of Animal Life.'— In this new series of laboratory exer-

cises for use in high schools — the outgrowth of experience in the

schools of Chicago — the authors have aimed to make the practical

work of elementary zoólogy a study from the view-point of animal

life, interpreting structure in the light of activity. While the outlines

for the study of the activities of living animals are as extensive as is

probably practicable for most schools, by far the greater part of the

laboratory work is a study of structure. Students are not expected

to dissect, but many points of internal structure are to be demon-

strated from permanent preparations.

In the form of its outlines the book is an example of the reaction

from the older manuals, — which consisted of description to be veri-

fied by the students, — in that it contains numerous questions, along

with a minimum of description and guiding information. Some of

the questions are of doubtful value in elementary zoólogy, for exam-

ple: “Why are there no fresh-water echinoderms?” “Is there

anything about the life history of man to suggest the metamorphosis

of insects?" ‘What traces of an invertebrate exoskeleton are still

present in man?"

In order * to develop the subject of the evolution of life from sim-

ple to complex forms," the authors follow the so-called logical order

and begin with the Protozoa, because “high-school pupils are not

more familiar in any true scientific sense with higher forms." If this

1 Walter, H. E, Whitney, W., and Lucas, F.C. Boston, Heath & Co. 1999

106 pp. Teacher's Book of Suggestions, with 31 pages.

No.413.] REVIEWS OF RECENT LITERATURE. 403

be true, there is inconsistency in some questions in the first lessons ;

for example: “Is there evidence that Paramcecia can breathe? "

“Has the Amceba a stomach?” Such questions are meaningless

unless the pupil has some scientific knowledge of structure and func-

tions in higher forms.

On the whole, the spirit and plan of most of the lessons may be

commended. Many teachers will welcome this as a laboratory guide

which aims to meet the popular demand for less study of compara-

tive anatomy and more about animal life in secondary education.

M. A. B.

Human Physiology. — Dr. Wm. D. Zoethout’s translation of

Schenck and Giirber’s Human Physiology: places within reach of

the English-reading student one of the best of the shorter German

physiologies. The translation is from the second German edition

and follows the original closely. After a brief introduction on gen-

eral physiology, the subject-matter is arranged under three heads

— metabolism, the transformation and setting free of energy, and

reproduction and development. The treatment is as modern as is

consistent with general soundness. "Thus we are told that “a solu-

tion tastes the more sour the greater the number of hydrogen atoms

replaceable by metals contained in the unit of volume," a statement

which includes all that is up to date without involving the reader in

the dissociation hypothesis. Although the text of the book has been

compiled with great conciseness and care, it is to be regretted that

the illustrations are so inadequate. Thus the figure showing the

general anatomy of the ear as copied from Helmholtz, and the

positively inaccurate drawing of the cross-section of the lamina

spiralis membranacea are scarcely justifiable. Nor is there good

reason why the olfactory epithelium should be illustrated by a figure

from Max Schultze, when such work as that done by Retzius, Van

Gehuchten, and others is so readily accessible. Such defects,

however, are small compared with the merits of the volume, which

Should be in the hands of every medical student and every teacher

of elementary physiology. P.

Korschelt and Heider's Embryology of Invertebrates. The

fourth part of the English edition of Korschelt and Heider’s En-

‘Schenck, F., and Gürber, A. Outlines of Human Physiology. Translated

from the second German edition by Wm. D. Zoethout. New York, Henry Holt

& Co, 1900. viii + 339 pp.

404 THE AMERICAN NATURALIST. [Vot. XXXV.

wicklungsgeschichte der wirbellosen Thiere completes the translation

of this monumental work.! As in the second and third parts, the

translation has been done by Matilda Bernard, and the revision and

editing by Martin F. Woodward. The present part gives an account

of the embryology of the mollusks, the tunicates, and Amphioxus,

and in the groups covered agrees with the third part of the German

edition except in the omission of the chapters on the brachiopods

and the Bryozoa, which the translators had previously placed in their

second part. The third part of the German edition appeared in

1893 ; the translation, therefore, is unfortunately some seven years

late. This has put on the editor the heavy task of supplying the

more recently acquired information on the groups under considera-

tion. Mr. Woodward has wisely refrained from rewriting the third

portion of the work, and has attempted to bring it up to date by

employing footnotes and adding to the literature lists, as in the

former part. While this is perhaps the best way out of the dif

ficulty, it does not seem to have been employed very successfully in

this last part. As an example, the chapter on Amphioxus may be

cited. Our advance in the knowledge of the embryology of this

form is indicated in some seven notes, none of which give very

extensive information. The appendix to literature for this chapter

contains some fifteen new titles. As these presumably cover the

period from 1893 to 1900, the list is obviously incomplete. One

misses any reference to Lwoff’s completed paper on the germ layers

(1894), Legros’s note on the morphology of the sexual glands (1895);

MacBride's note on germ layers (1896), Garbowski's discussion of

the mesoderm (1898), Klaatsch's account of the structure and devel-

opment of the tentacles (1898), Lankester's note on the development

of the atrial chamber (1898), and Legros's description of the develop-

ment of the buccal cavity (1898), contributions which, judging from

the composition of the literature lists in the German edition, should

have been recorded. Incidentally it may be mentioned that of the

names given in this appendix Hamman is substituted for Hammar

and the capitalization of MacBride is unsteady. On the whole, the

additions made by the editor do not show the high standard of work

characteristic of the German original. The presswork, particularly

in connection with the illustrations, retains more or less of the mud-

diness of the earlier parts. Notwithstanding these shortcomings,

1 Korschelt, E., and Heider, K. Zext-Book of the Embryology of papi

vol iv. Translated by Matilda Bernard, revised and edited by Marti F.W

ward. New York, The Macmillan Company, 1900. xii + 594 PP» 3!? figs-

No. 413.] REVIEWS OF RECENT LITERATURE. 405

the translation is generally so well done that the work, now that it is

completed, cannot but be a boon to the English-reading student.

Heart-Beats in Salpa. — The pulsation of the heart in three

species of Mediterranean Salpas has been exhaustively studied by

L. S. Schultze.! As is well known, the hearts of these animals beat

first in one direction and then in the other. A complete set of

advisceral or of abvisceral beats constitutes a pulsation series. The

intervals between pulsation series are known as pauses. An advis-

ceral pulsation series and its pause, followed by an abvisceral series

and its pause, form a compound heart period.

The numbers of beats in pulsation series were so extraordinarily

variable that a normal number could not be found. The total

number of abvisceral beats may be considerably more or less than

that of the advisceral beats; thus in one case 247 abvisceral beats

corresponded to roo advisceral beats, and in another 237 abvis-

cerals to 523 adviscerals. The rates of the two sets of beats were,

however, very close; thus 100 abvisceral beats were accomplished in

175 seconds, and the same number of adviscerals in 174 seconds. As

the water in which the animal was kept lost oxygen, the rate of

beating increased; thus an individual’s heart, which at the begin-

ning of the experiment beat 100 times in 208 seconds, after six

hours beat the same number of times in 148 seconds. Of the three

species studied, the two larger ones, Salpa africana-maxima and

Cyclosalpa pinnata, had an average rate of 26 to 30 beats per

minute; the smaller, Salpa democratica-mucronata, 107 per minute.

The pauses between ad- and abvisceral series varied from 1 to 4 or

occasionally 5 seconds.

Each heart-beat is a peristaltic wave that sweeps over the heart

from one end to the other. Usually a new wave appears at one end

before the old one has passed off at the other, and sometimes as

many as seven waves may be counted on a heart at once. Kruken-

berg believed that the two ends of the heart were physiologically

very different, and that nicotine and hellebore affected the advis-

ceral pulsations only, the former diminishing, the latter increasing

€m. Schultze, however, found that these poisons influence the

ab- as well as the advisceral pulsations, and thus demonstrated that

the ends of the heart were not in this respect dissimilar.

* Schultze, L. S. Untersuchungen über den Herzschlag der Salpen, Jenaische

Zeitschr. y. Naturwissenschaften, Bd. xxxv (1901), pp. 221-328, Taf. IX-XI.

406 THE AMERICAN NATURALIST. [Vor. XXXV.

The source of the stimulus for the contraction of the heart muscle

was sought for in several ways. An isolated heart was found to be

capable of beating regularly in either direction. Stimulation of the

animal's brain had no effect on the heart-beat. Removal of the

brain reduced the rate, but this was shown to be due to the loss of

substance suffered by the animal and not to the removal of the

brain. The filling of the heart with blood was shown not to be

necessary for its contraction. As small fragments of the heart

muscle continued to contract rhythmically, and yet on examination

showed no evidence of nerve fibres or of ganglion cells, Schultze

concluded that the motor stimulus for the action of the heart muscle

must be geferated exclusively by the metabolism of that muscle

itself.

The alternating action of the heart depends on the capacity of its

muscle to transmit the stimulus to contraction directly from fibre to

fibre and on the varying rhythm of the two ends of the heart. In

moribund individuals both ends of the heart may at times give rise

to contraction waves simultaneously. These usually meet near the

middle of the heart and neutralize one another. In normal indi-

viduals the rhythm at one end is so much more rapid than that at

the other that this rhythm asserts itself for the whole heart. When,

however, the muscle tissue of the given end becomes somewhat

exhausted by continued action and thereby reduces its rate of con-

traction, the muscle substance of the opposite end, having recovered

from the effects of its own previous action, may be able to establish

a more rapid rate than its opponent, and thus the center of propaga

tion is transferred to the recuperated end. Hence the quiescence of

a given end permits that end to recuperate till its own rhythm can

supersede that of the opposing end, and its action gradually exhausts

it so that its opponent in turn will be able to gain the ascendency.

Flies as Carriers of Disease. — Dr. L. O. Howard, in a recent

paper,' has presented the possibilities of the transmission of disease

by flies in a particularly striking manner. A large number of flies,

representing many species, were bred from human excrement.

Those seen visiting the same material were collected. Then collec-

tions were made in dining-rooms and pantries, and many sheets ja

sticky fly-paper examined to see what species commonly occur in

* A Contribution to the Study of the Insect Fauna of Human Excrement, Prec.

Wash. Acad. Sci., vol. ii (1900), pp. 541—604.

No.413.] REVIEWS OF RECENT LITERATURE. 407

houses. By comparing the two series, those visiting and bred from

excrement and those found in houses, it is at once apparent what

species are liable to carry disease germs. The results show that

practically all of the house flies occasionally breed in or visit human

excrement. It is thus possible that almost any house fly may carry

the germs of typhoid fever. The next step in this process, vzz., to

find out by experiment whether flies actually carry germs on their

tarsi and labella, was not investigated.

No less than thirty-six species of flies were reared from excrement,

and forty-one other species captured visiting the same material.

Among those bred were the common house fly (Musca domestica),

and the pomace fly (Drosophila ampelophila). The latter is an

especially dangerous species, as it not only frequents houses, but

also occurs on grapes and other fruits exposed on the market.

In the course of the work, many new and interesting observations

of purely scientific value were made on the life history of various

species. A large amount of the disagreeable work was performed

by Mr. F. C. Pratt, and the determination of the flies rests on the

authority of Mr. D. W. Coquillett. N. B.

Trematode Fauna of Egypt.t— Just as the earlier works of

Looss marked a new epoch in the study of the comparative anatomy

of the distomes, so the present paper is destined to be the starting

point of a movement toward the rational dismemberment of an

ancient and honorable genus — Distomum. Not that others have

failed to recognize its heterogeneous character, or to attempt its

dissolution, but that up to the appearance of the paper under dis-

cussion no one has indicated a reasonable way to the end desired.

Many authors have recognized groups of forms whose relationship

was evident, and yet have failed to give such groups their true posi-

tion as genera, or have seized upon single and insufficient characters

to delimit them. Thus Rudolphi endeavored to employ external

features, which in a group of such uniform exterior does not

Suffice ; Dujardin selected a single feature, the character of the ali-

mentary canal, for the major part of his genera, while both Diesing

and Monticelli erred in the same direction. To be sure, certain

small groups were recognized and set off from the remainder, but

the systems proposed have never met general acceptance, probably

'Looss, A. Weitere Beitráge zur Kenntniss der Trematoden-Fauna Aegyp-

es, zugleich ein Versuch einer natiirlichen Gliederung des Genus Distomum

Retzius, Zool. Jahrb., Abt. Syst., Bd. xii, pp. 521-784, 9 pls.

408 THE AMERICAN NATURALIST. [Vor. XXXV.

because of their inadequacy and of the heterogeneous character of

the genera so formed. Then even the groups which were best made

were taken as subgenera rather than in their true place as genera,

and even subfamilies, which Looss is unquestionably the first to

recognize in any broad way. This view does not in the least under-

estimate the admirable work of Braun and Lühe, which has appeared

almost synchronously with that of Looss, and which, though dealing

with fewer forms, furnishes evidence of the naturalness of the

proposed dismemberment by the independent selection of identical

groups.

Looss discusses first the law of priority in relation to helmin-

thology, and advocates on cogent grounds the dating of generic

names in this field from Rudolphi, *the Linnzus of helminthology."

Though much to be desired, his proposal must still be regarded as

impracticable in view of the ciose relation of synonymy in all groups.

Looss protests strongly, and, most will admit, rightly too, against the

use of conjecture in restoring old generic names and cities from

Rudolphi Hemiurus and Echinostoma as recognizable and evidently

good genera, with Sphaerostoma as unrecognizable and Schisturus

which depends upon pure conjecture. The law of priority is based

upon the legal presumption that the literature is available everywhere,

but a comparison of original specimens is not called for, since they

exist, if at all, in a few places at most. The replacement of specific

names, already well fixed, by comparison of the originals is hence in

violence with the wording of the law ; if, however, it is to be carried

out, general interests demand the earliest possible revision of all

originals, since in this way the least disturbance will be produced.

Names of species which do not exist in original specimens and which

are not recognizable should be dropped at once so as not to burden

the literature further. Looss then refers to the custom of Rudolphi

in citing unknown parasites by the name of the host in genitive ;

Eg., Distoma meropis, which should be interpreted as “a distome

from Merops,” but which, as the author wrote in Latin, has the

outward form of a generic and specific combination. Such names

are pure nomina nuda, since a diagnosis is lacking and were so

regarded by the author, since he never used a genitive as a specific

name, and since he also never omitted the “R.” which is lacking

after these, from the new species actually described.

The second section of the paper on the taxonomy of the distomes

opens with a discussion of previous efforts in this direction, an

of the great disparity in form and structure between Bilharzi

No. 413.] REVIEWS OF RECENT LITERATURE. 409

Apoblema, and other forms. For the dicecious distomes a new family,

the Schistosomidz, is created, and the remainder, constituting many

subfamilies, is left in the family Distomida. Since, however, the

genus Distomum is nonexistent, this ought to have been changed to

Fasciolida. A similar change is necessary with the name Mono-

stomide, but what name shall be used in its place is not now clear.

Looss then gives the following scheme of the classification as

emended, in connection with which it should be noted that there are

numerous genera whose position even yet is a matter of doubt, and

that this is not regarded by the author as in any sense a complete

system :

A. Aspidocotylea Mont.

remains unchanged)

Metastatica Lkt.

B. jo een Mon

mily le a Ne Brds.

n7 mains unchanged)

ec Apre str. Lkt.

mily DIsTOMIDÆ Mont. (partim)

Subfamily Amphistominæ Looss (= Family AMPHISTOMIDÆ Mont.)

i epodermatinæ Lss.

goderinæ Lss.

s Brachyceeliine Lss.

i Pleurogenetinz Lss.

ephalogoniminz Lss.

M Dicroceeliine Lss.

r Syncæliinæ Lss.

y Heterolopinæ Lss.

* Urogoniminz Lss.

Family (inquir. RHOPALIADJE!

oellikeria Cobbold

T SCHISTOSOMID# Lss. Schistosomum Weinland

Bilharziella n.g.

" GASTEROSTOMID/E Braun

" DIDYMOZOONID Mont. Remain unchanged

" ONOSTOMID Mont.

! According to the recent investigations of Braun (Zow. Anz., Bd. xxiii, p. 27)

closely related to the Echinostominz and consequently not of family rank.

410 THE AMERICAN NATURALIST. [ VoL. XXXV.

According to Looss the formation of a special genus is warranted

when a certain definitely circumscribed complex of characters can be

recognized in two forms which also agree in other respects ; yet

genera may be founded on single forms of evidently isolated struc-

ture. While general appearance is of value, yet internal anatomy is

the real basis of subdivision, and just this is, in fact, little known,

partly at least owing to ignorance regarding the relative importance

of characters. Among the most weighty generic characteristics are

the copulatory organs which show the following types : (1) No mus-

cular cirrus sac closed proximally and distally about the duct and

seminal vesicle, together with the constant, if often weakly developed,

prostate which lies (2) free in the parenchyme, or (4) enclosed ina

connective-tissue covering open at both ends ; (2) a closed muscular

cirrus sac which encloses (a) the genital sinus, é.c., the more or less

elongated common terminal portion of both male and female ducts,

or (4) only the end of the male duct. Here again the cirrus sac may

enclose (a) seminal vesicle, prostate, ejaculatory duct, and protru-

sible cirrus, or (4) only the last three, the vesicle lying in the paren-

chyme, or (c) the prostate also is free, while only the duct and the

cirrus are enclosed in the sac. The course of the uterus in the body

is also an important generic character, while the size of the eggs is

uniform within narrow limits in any genus. Of specific value are the

size and form of organs in detail, the extent of the vitellaria, a very

constant feature in any species, and similar details.

In the section treating of the characters of the subfamilies and

genera one finds a great variety in manner of treatment. Most

groups are considered in extenso with full-faced headings which claim

immediate attention, but there are those which are introduced in the

middle of a topic under another heading, or even rarely one finds a

new genus thrown in parenthetically which, in the absence of key

and index, makes its discovery difficult. Still the work is generally

free from such slips, and the absence of a key is attributable to the

often repeated assertion of the author that this is a fragment and not

a finished system, having for its primary object the demonstration of

the existence of natural groups of family and generic rank within the

limits of the old genus Distomum Retzius.

From this section, as well as from the fourth and last, which con-

tains a description of the new and little known species that have

been studied by the author, it is hopeless to give here anything

regarding the wealth of descriptive and comparative matter which 3

offered. Itis not too much to say that no other helminthologist 15

No. 413.] REVIEWS OF RECENT LITERATURE. 411

the equal of Looss in deciphering, delineating, and comparing the

anatomical structure of trematodes, and it would be hard to find, save

in his own work, the equal of the nine plates he has given to illus-

trate this work. Some mention is made of a total of eighty-four

genera, including twenty-three old and sixty-one new ; of the latter,

three are clearly antedated by names proposed by Braun, five corre-

spond to groups named by Lühe in a publication of identical date,’

and four are provisional. Among the fifty-two species described

twenty-four are new. One can only regret that the author did not

give an index or table of contents, if debarred from forming a key

by the incompleteness of his system. As it is, reference to any

section or topic is not an easy matter. It may also be said that in

rare instances the author fails to apply the principles he has laid

down, without giving any reason for the exception ; but some slips

are unavoidable in a work of such magnitude, and do not detract

from its permanent value. Though Looss disclaims having formed

any complete system, his work comes nearer that than any one else has

yet reached, and will be the foundation on which such a system is

to be built. H. B. W.

BOTANY.

Primitive Algz and Flagellata.* — In reviewing Dr. Blackman's

paper, the writer has not mentioned the authorities for the arrange-

ment given therein, which may be found on reference to the paper

itself. The article is of the nature of a review of recent work, and

the following is but a condensation of its most important points.

e older arrangement of the Chlorophycez, given by Wille in

Engler and Prantl's Pfanzenfamilien, is largely an artificial one, and

consequently subject to changes. Of the three groups named by

him, the Siphonez, Confervoidez, and Protococcoidez, only the first

! It appears to me clear that both the intent and the wording of the rules

covering the choice in case of synchronous appearance of different names for the

same forms call for the preference of the extended discussion over the preliminary

notice, certainly in all cases where types are named. Under this interpretation

Looss's names stand as against Lühe's, save for Dolichosomum, which is pre-

occupied, and hence gives way to Ithyogonimus Lühe rather than to Dolicho-

desmus Looss (Zoo/. Anz., Bd. xxiii, p. 603) of later date.

2 Blackman, F. F. The Primitive Algz and the Flagellata, an Account of

Modern Work bearing on the Evolution of the Algae, Annals of Botany, vol. xiv,

No. lvi (December, 1900), p. 647.

412 THE AMERICAN NATURALIST. [Vor. XXXV.

isa natural one. According to Blackman, the green alge may be

considered to originate from two flagellate forms, Chlamydomonas

and Chlorameeba. From the first type three divergent lines of ascent

go off: one leading to Volvox, including the type of motile cell

aggregations, another ending in the so-called unicellular Siphonez.

The third, by far the largest and most important, is called the Tetra-

sporine type, including forms of non-motile cell aggregations, the main

stem of which leads through the simple filamentous forms to the

branched Coleochzte, and finally, it is to be supposed, to the higher

plants. The Ulvas are properly placed as a specially developed

side branch. The always perplexing Conjugatez are indicated as

forming a possible fourth line of ascent from the Chlamydomonas

type.

The green alge which arise from the second flagellate form,

Chlorameeba, include Ophiocytium, Conferva, and finally the Botryd-

ium forms. It is considered that the gametes of these have in

reality two cilia, not one, as previously believed. They are included

under the head of the Confervales (Borzi), a title not coincident with

that used by other authors, and, with their flagellate ancestors, form

a group called the Heterokontze. ;

The Phzophycesz are derived from a brown flagellate, Chromulina,

allied to the green Chloramceba. This is widely different from the

older view, which recognized the simplest Ectocarpus type as the

most primitive brown form. Phzocystis, a form on the border line

between alge and flagellate, is taken as the next step in the direc-

tion of plants, and is connected with the recently described un-

doubted algæ forms, Phzeothamnion and Pleurocladia, and thence with

the Ectocarpus type. The Piatones constitute a side branch from

Chromulina.

The red seaweeds, it is suggested, find their origin in a flagellate

form (Rhodomonas), which is said to possess a chromatophore of true

Floridian red color.

However much one may or may not agree with the conclusions,

the paper is a suggestive one and a valuable review of the status of

the subject. The changes suggested regarding the Chlorophyceous

algæ seem certainly to be in the right direction, except that the con-

nection of the Botrydium forms and Conferva type is not very appa"

ent. As to the brown algz the identity of the flagellate ancestor does

not seem absolutely certain and convincing. A flagellate ancestor for

the red seaweeds is a matter of such doubt that, as the author say®

it is no more than a suggestion which he makes. H. M. R.

No. 413.] REVIEWS OF RECENT LITERATURE. I

. 413

Spermatozoa of Ferns.! — This account considerably extends the

hitherto published work of Pfeffer and others, and is a valuable con-

tribution to the knowledge of chemotaxis. Buller has found that in

addition to malic acid and certain malates, experimented with by

previous authors, other substances also exert a positive chemotactic

influence on the spermatozoids of ferns. "Various tartrates, oxalates,

phosphates, and salts of potassium (all substances to be found in cell

sap) have an attractive influence. Pfeffer’s negative results with

such substances are ascribed in the case of organic salts to the

use of too dilute solutions; in inorganic salts to the fact that mix-

tures (eg. plant ash) were used. Nevertheless, the opinion of

Pfeffer that it is a malate which attracts the spermatozoid to the

archegonium in ferns is substantiated, by reason of the high degree

of concentration required by other salts. Malic acid and its salts

attract spermatozoids about fifty times more strongly than do other

substances. That it is not free malic acid which is found in the cell

sap of the archegonia seems probable in that Pfeffer determined no

acid reaction in the exudation, and from the fact that malic acid

alone is decidedly poisonous to the spermatozoids.

The fact that the diethylester of malic acid is indifferent indi-

cates an explanation of the chemotactic influence on the basis of

chemical dissociation, since the afore-named substance is undisso-

ciated in solution, whereas in malic acid and the malates the nega-

tive radicle is free as an ion. But at the same time, while it is

Shown that other undissociable substances, like cane sugar, grape

sugar, etc., do not attract the spermatozoids of ferns, it is known

that one at least (i.2., cane sugar) does attract those of the mosses,

while several attract certain bacterial forms. This fact is men-

tioned, but no explanation is attempted, beyond suggesting the

possibility that some undissociable substances may be found which

do exert a positive chemotactic influence on the spermatozoids of

ferns. It is then the case, that while certain substances (¢.g., malic

acid) may be indifferent in an undissociated form, other substances

attract in some cases, although there is no dissociation of the

molecule. This would naturally hinder at present the drawing of

any general conclusions regarding the relation of chemotaxis and

dissociation.

The fact that the spermatozoids of Gymnogramme mertensii and

; Buller, A. H. R. Contribution to our Knowledge of the Physiology of

the Spermatozoa of Ferns, Annals of Botany, vol. xiv, No. lvi (December, 1900),

P. 543.

&l4 ——. THE AMERICAN NATURALIST. [VoL. XXXV.

other ferns come to rest upon the withdrawal of water from their

contents is also considered. The swarm period of the spermato-

zoids of the above-named fern has been determined to be two hours,

— a much longer time than reported in previous cases, — during

which period the starch stored up in it disappears. H. M. R.

The Flora of Celebes. — The interest which American botanists are

likely to feel in the flora of the Pacific Islands in consequence of our

occupation of the Philippines makes Dr. Koorders' report! on Mina-

hasa, the northeastern horn of Celebes, of more than passing impor-

tance to us. In addition to physiographic and similar data this

volume, which forms one of the regular series of Mededeelingen issued

from the Botanical Garden at Buitenzorg, gives a critical review of

what had previously been written on the flora of Celebes, an anno

tated catalogue of the spermatophytes and pteridophytes known to

occur in the island, descriptions of a number of new species, and

full indices to the popular and scientific names of the plants, as

well as chapters on the economic uses of many of the species and

tabulations of plants yielding the more important useful products.

Unfortunately the text is in Dutch, but diagnoses of new species are

in Latin, and there are frequent annotations in German.

Notes. — The announcement for the Fourteenth Season of the

Department of Botany of the Marine Biological Laboratory at Woods

Holl has just been issued. The work will commence on July 3 and

continue for six weeks. Courses have been provided in Cryptogamic

Botany by Dr. Davis and Dr. Moore; in Phanerogamic Botany by Dr.

Charles H. Shaw, and in Plant Physiology by Dr. R. H. True. Plant

Cytology will be under the direction of Dr. Davis and Mr. Lawson,

and in addition to the regular class work provision will also be made

for a series of special lectures. Announcements and further informa-

tion may be obtained from Dr. Bradley M. Davis, University 9

Chicago.

The Bulletin de P Herbier Boissier, suspended for a time, appears

again, under the direction of M. Gustave Beauverd, curator of the

herbarium. The first number of the new series, bearing the pu

December 29, contains papers on African plants by de Wildeman

! Koorders, S. H. Verslag eener botanische denstreis door de Minahasa, re

eerste overzicht der flora van N. O. Celebes, uit een wetenschappelijk en e

oogpunt. Batavia, s'Gravenhage, 1898, xxvi + 716 pp., 10 charts and 3 plate

Mededeelingen van s’ Lands Plantentuin, No. xix.

No.413.] REVIEWS OF RECENT LITERATURE. 415

and Durand, ferns of the Amazon region by Christ, Brazilian fungi

by the Sydows, the vegetation of Cape Magoary, etc., by Huber,

an alpine variety of Ste//aria nemorum by Beauverd, and a valerian

new to the flora of Savoy by Briquet.

Papers of botanical interest in the Proceedings of the American

Pharmaceutical Association for 1900 are the following: Merrill and

Schlotterbeck, alkaloids of Bocconia cordata ; Gordin, alkaloids of Cea-

nothus Americanus; Kebler, notes on jalap roots; Kraemer, assay of

drugs by the use of living plants; Schneider, pharmaceutical bacteriol-

ogy; Stevens, wild-cherry bark and its preparations; Dohme and

Engelhardt, Atropa Belladonna or Scopola Carniolica; Schlotterbeck,

Adlumia cirrhosa.

Under the title of Zorreya the Torrey Botanical Club of New York

City has begun the issuance of a monthly journal of botanical notes

and news, under the editorial management of Dr. M. A. Howe.

The self-pruning of certain trees, a subject apparently first dis-

cussed in this country by Trelease, in the Report of the Wisconsin

Experiment Station for 1884, and again by Bessey in Science for 1900,

receives interesting treatment by Schaffner and Tyler in the Ohio

Naturalist for January. Figures are given illustrating the process

in Populus and Salix.

The Bulletin of the Torrey Botanical Club for December contains

Systematic papers on fungi by Peck, lichens by Zahlbruckner, mosses

by Mrs. Britton, fernworts by Maxon, and Rocky Mountain phanero-

gams by Rydberg.

Part XXIII of fittonia is largely devoted to Composite and Cru-

ciferæ, with a decade of new Gentianacez, a discussion of some

neglected generic types, and a batch of corrections in nomenclature.

Thalictrum confine is the name proposed in Rhodora for December,

by Mr. Fernald, for a plant of Ontario and Maine, which in habit

Suggests smallleaved Z Jendleri, and he shows that T. occidentale

extends eastwards so as to reach New Brunswick and Maine.

The Systematic value of tendrils in Lathyrus is discussed by

ritsch in the Oesterreichische Botanische Zeitschrift for November.

Dr.

Pan Robinson, in Riodora for December, discusses the nomen-

ure

9f Agrimonia in New England.

a g pt Contributes an article on the anatomy and morphology

genia bulbosa to the American Journal of Science for January.

416 THE AMERICAN NATURALLIST. [Vor. XXXV.

Taraxacum in North America is increased by Professor Greene, in

Pittonia for January 5, by the description of 7: Chamissonis, T. rupestre,

T. ovinum, T. lacerum, T. dumetorum, T. mutilum, T. angustifolium,

and T. ammophilum, all from the Northwest, British Columbia, or

Alaska.

Senecio, as it occurs in New England, is revised by Greenman in

Rhodora for January.

Monarda fistulosa and its allies are passed in critical review by

Fernald in odora for January.

Professor Kellerman, in O. S. U. Naturalist, No. 2, gives an

interesting plate of variations in the foliage of Smilax glauca.

Professor Scribner and his assistants publish a series of studies

on American grasses as Bulletin 24 of the Division of Agrostology

of the United States Department of Agriculture.

The specific or hybrid character of Asplenium ebenoides is discussed

by Mr. Maxon in the Botanical Gazette for December, and, unlike

Professor Underwood, he considers the suggestion of hybridity as

too patent to be ignored, though he admits the absence of positive

proof of it.

A remarkably lobed form of Asplenium ebeneum is described by.

Mr. Davenport in Rhodora for January, under the varietal name

Hortonz.

Weinzirl, in the Journal of the Cincinnati Society of Natural History

for December 28, publishes an account of the air bacteria of the ari

region of New Mexico, which leads him to the conclusion that the

rather few species found differ from those yet described from other

. regions.

A compilation of the North American Phyllostictas, with descrip-

tions of the species published up to August, 190o, by J. B. Ellis and

B. M. Everhart, has recently been distributed by the authors, and

bears the imprint of G. E. Smith, Vineland, N. J.

A paper on a spot disease of the violet, due to Alternaria Viole,

by R. H. Dorsett, forms Buletin 23 of the Division of Vegetable

Physiology and Pathology of the United States Department of

Agriculture.

No. 5 of Mr. C. G. Lloyd's Mycological Notes is largely concerned

with Collybia, as represented about Cincinnati.

No. 413.] REVIEWS OF RECENT LITERATURE. 417

Professor Cheney has published “an historical review of the work

done on the flora of the territory now included within the limits of

Wisconsin,” in the Pharmaceutical Review for December and January.

The sixth fascicle of de Wildeman and Durand’s “ Illustrations de

la flore du Congo,” in course of publication in the Annales du Musée

du Congo, of Brussels, bears date of September, 1900.

The concluding part of Vol. II, and the first part of Vol. III, of

J. Medley Wood's Wata/ Plants have recently appeared.

Dr. Henry Kraemer, in the Proceedings of the American Pharma-

ceutical Association for 1900, proposes the use of living plants in

drug assaying, to test the strength of certain toxic solutions. Ina

series of experiments, seedlings of Lupinus albus and Pisum sativum

were grown in strychnine nitrate, brucine sulphate, and tincture of nux

vomica solutions of different strengths. The growth of the radicles

was found to be inversely proportional to the toxicity of the

solutions.

In the seventeenth Annual Report of the Wisconsin Agricultural

Experiment Station, Professor Goff has a paper of interest on the

development of flower buds on a number of fruit plants, in connec-

tion with the temperature curve for March and April, in which their

development was found to lie.

Students of leaf-form and position will be interested in a paper

by Raciborski in the Annales du Jardin Botanique de Buitenzorg,

Vol. II, Part

“Open Spaces for the People” is the title of an article by Philip

MacMahon, curator of the Brisbane Botanic Gardens, in the Queens-

land Agricultural Journal for December, in which a general plan is

given of that garden and of promenades that it is suggested may be

connected with it.

Some profit and a little amusement may be obtained from an

examination of a recent Consular Report on “school gardens in

Europe,” among which Consul-General Lincoln of Antwerp includes

the Kindergartens of that city.

A compendious volume of statistics concerning the use of wood

pulp in foreign countries is published as Vol. XIX of the Special

Consular Reports of our Government.

The “Diamond Jubilee Number” of the Gardeners’ Chronicle for

January 5 contains portraits of Lindley, Paxton, Berkeley, and

418 THE AMERICAN NATURALIST. [ VoL. XXXV.

Darwin, all of whom contributed matter of botanical or horti-

cultural interest to the earlier volumes of the journal, which, though

agricultural in name, has always been of interest and value to the

botanist.

The double number of the Botanische Zeitung (Abtheilung I) for

December 1 is devoted to Brunfels, a botanist of four hundred

years ago.

The Fern Bulletin for January opens with a portrait of Professor

Underwood.

PALEONTOLOGY.

Dean's ‘t Palzontological Notes.’?! — This elaborate memoir of

upwards of forty pages, six plates, and eighteen text-figures, is one

of the most important contributions to the literature of Devonian

Arthrodires that has yet appeared in this country. It has for its

twofold object a minute description of the skeletal structures in

Mylostoma, Stenosteus, and Selenosteus (the last two being here

made known for the first time), and a discussion of the characters

and relationships of the whole group of Arthrodires, with the impro-

vision of a new system for their arrangement.

Four of the quarto plates are admirably lithographed by the

author from original specimens; the other two are from photo

graphs. Not the least valuable and suggestive feature is the inter-

spersion throughout the text of numerous diagrams showing the

arrangement of plates in the cranial, dorsal, and ventral armor of

different genera of Arthrodires. Restorations of this nature are not

only useful for the graphic information they convey, but they afford

an extremely convenient means of comparison. The interesting

modifications displayed by Mylostoma and Dean's two new genera,

especially as regards their dentition, constitute a welcome addition

to our knowledge of this group. Owing to their poor preservation,

the difficulty of deciphering the détails which the author has pains-

takingly worked out and skillfully reconstructed must have been very

great, and obviously none but an expert could have succeeded.

1 Dean, Bashford. deepen Notes. I, On Two New Arthrodires

from the Cleveland Shale of Ohio; II, On the Characters of Mylostoma, New

berry; III, Further Notes on ú Relationships of the Arthrognathi. Mem

N. Y. Acad. Sci., vol. ii (1901), pt. iii.

No.413.] REVIEWS OF RECENT LITERATURE. 419

Of more general interest, however, are the author's views as to

the relationships of the so-called * Arthrognathi" (= Arthrodira +

Anarthrodira, the latter comprising only the genera Macropetal-

ichthys and Holopetalichthys). Excluding them from both Pisces

and Ostracoderms, Dean assigns them the rank of an independent

class, which he conjectures may have been descended from primi-

tive forms like Lanarkia in the Silurian. The prime reason for

removing Arthrodires from Pisces rests upon the interpretation of

their jaw-elements, which Dean regards as merely dermal ossifica-

tions, showing **not the slightest evidence of their relation to endo-

skeletal or gill-arch jaws." There is, however, strong presumptive

evidence that a cartilaginous mandibular arch was present, the

distal portion of which ensheathed the lower dental plate (** gna-

thal Dean), and the proximal portion, or suspensorium, was

attached to the head-shield. How,.for instance, is one able to

conceive of the attenuated mandibles of Titanichthys, except as

imbedded in tissues corresponding to the Meckelian cartilage?

The author advances some ingenious theories to explain the

evolution of the articular joint between cranial and dorsal shields

characteristic of Arthrodires, and, giving free rein to his imagina-

tion, speculates on how such a joint might have arisen in the head-

shield of Macropetalichthys. A slight impediment exists, however,

in the way of accepting his conclusions, for the reason that we can-

not admit any of his premises. Dean assumes, and so, too, have

Newberry, Cope, and Eastman before him, for that matter, that a

valid basis of comparison exists between this enigmatical form and

various specified genera, where the facts prove such is not the case ;

also that the arrangement of cranial plates and sensory canals in

the genus under consideration is homologous with that pervading

Arthrodires generally, which is an egregious error.

The fact of the matter is that all writers on Macropetalichthys

have been misled by deceptive appearances, perhaps to some extent

also by preconceived ideas, and its osteology has not yet been

correctly interpreted. Eastman’s and Dean's suggestion that there

is a superficial system of investing plates arranged independently to

a distinct substratum of bony elements is unsubstantiated, if not

disproved ; and still more improbable is the notion that the “ head-

Shield" is made up of combined cranial and dorsal shields. The

transverse septum peculiar to this genus is an internal structure

situated a long distance in advance of the posterior cranial border,

and recalling in a measure the inwardly directed processes on the

420 THE AMERICAN NATURALIST. [ VoL. XXXV.

inferior aspect of the postorbital in Dinichthys. There are in

Macropetalichthys no movable ginglymoid joints, neither dorsal

nor ventral armor so far as known, and apparently no ossified lower

jaw. In short, this genus and the accompanying Asterosteus are as

far removed from Arthrodires as are the Ostracoderms, and should

perhaps be included in the same subclass with the latter. Their

canal systems and cranial plates can be no more homologized with

those of Arthrodires than with Stegocephalians, and the most we

can affirm is that their arrangement is bilaterally symmetrical, and in

a few respects similar or paralleled. Dean’s text-figures 12 and 13

are objectionable, therefore, in so far as they depict merely fanciful

conditions ; and in Figs. 15 and 16 the overlap of the clavicular is

shown, but the plate itself is omitted.

Apart from this parenthetical discussion, as it were, of extraneous

genera, the author’s generalizations respecting Arthrodires, his sub-

division of them into several new families, and summary of their

characters, reflect the advanced ideas of an investigator who has

greatly enlightened us as to their structure and interrelationships.

CRE

Jurassic Fossils from Alaska.!— In this paper Dr. Pompeckj

has given a revision of all the known Jurassic fossils of Alaska and

has described a number of new forms. According to Dr. Pompeckj

the Lias is represented at Kialagvik, or Wrangell Bay, on the Penin-

sula of Alaska, as shown by the presence of Amaltheus whiteavest

White, ZZ/a howelli White, ZL. kialagvikensis White. This fauna

was described by Dr. C. A. White as of Upper Jurassic or Lower

Cretaceous age.

The Kelloway stage of the Middle Jurassic is represented at Kat

maiskoj on the southeastern side of the Alaskan Peninsula, and on

the northwestern side of the Kadiak Island. The former locality

furnished the following fossils :

Cadoceras wosnessenski Grewingk, C. grewinghki Pompeckj, C. cato-

stoma Pompeckj, Belemnitella, sp. indet., Aucella, sp. indet., a

mus, 55. indet. The locality on Kadiak Island yielded the following

fossils: PAy/loceras subobtusiforme Pompeckj, Cadoceras conf. wosnes

senski Grewingk, C. grewingki Pompeckj, C. Schmidti Pompeckj, C.

petelini Pompeckj, C., sf. indet., C. stenoloboide Pompeckj. TE

fossils show a wide distribution of the sea during the Kelloway stage

1 Pompeckj, J. F. Jura-Fossilien Alaska, Verhandl. Kais. Russ. Mineralo:

Gesell. St. Petersbourg, Ser. 2, Bd. xxxviii (1900), Nr. 1.

No.413.] REVIEWS OF RECENT LITERATURE. 421

of the Middle Jura, but the occurrence of Lias in the same region

shows that Neumayr's hypothesis of a great transgression of the sea

in Middle Jurassic time in the arctic region will not hold good.

Marine deposits existed there even before Jurassic time, as shown

by the occurrence of Upper Triassic deposits in Alaska.

Ej DG

The Permian of Armenia.'— The strata of Djulfa in Armenia

are classic in geological literature, having long ago been assigned to

the Subcarboniferous on account of the supposed occurrence of

Goniatites striatus. But a revision of the fauna, based on a new col-

lection made by Prof. F. Frech, shows that these beds belong to the

Permian, for there is a mixture of Paleozoic and Mezozoic types.

Their Paleozoic age is shown by the occurrence of typical Permian

brachiopods, such as are known in Russia and India, and of the

goniatite genus Gastrioceras, which has never been found above the

Permian. On the other hand, the genera Hungarites and Otoceras

are known there, which elsewhere are not found below the Trias.

The evolution of Hungarites from the simple forms of Armenia into

the complex development as known in the Trias of Siberia and India

is worked out by Dr. von Arthaber in a most convincing manner,

and is a good argument against useless multiplication of generic

names in a phylogenic series. TT

Notes. — Dr. Diener (Beitr. Palacontol. Oesterreich- Ungarns, Bd.

XIII, 1900) has continued his detailed studies of the Triassic

faunas in this contribution, treating of the Muschelkalk zone of Cera-

lites trinodosus. He describes a new genus, Arthaberites, of the

family Pinacoceratide, resembling Pseudosagoceras Diener of the

Lower Trias of Siberia, and possibly descended from it. The fauna

described shows a strong resemblance to that of the Bosnian Mus-

chelkalk long since made known by the works of F. von Haner. It

consists of numerous species of Ceratites, Anolcites, Celtites, Proar-

cestes, Joannites, Procladiscites, Megaphyllites, Sageceras, Arthaber-

ites, Pinacoceras, Norites, Monophyllites, Sturia, Gymnites, Ptychites,

Nautilus, Orthoceras, Atractites. Many of these species had not before

been found in the Alps, and their discovery is of material aid in cor-

relating the Alpine strata with those of the other Triassic provinces.

! Arthaber, G. von. Das jüngere Palaeozoicum aus der Araxes-Enge bei

Djulfa, Beitráge zur Palaeontologie Oesterreich-Ungarns und des Orients, Bd. xii

(1900), Nr. 4-

422 THE AMERICAN NATURALIST. (VoL. XXXV.

Dr. Whiteaves (Geol. Survey of Canada, Vol. I, Part IV) revises

his published papers on the paleontology and stratigraphy of the

Queen Charlotte Islands, many old species being renamed, as further

studies have shown their designations to be untenable. Several new

species of brachiopods and Mollusca have been obtained by later col-

lectors and are added in this paper. Among the ammonites may be

noted the predominance of Desmoceras and Olcostephanus, also the

absence of Baculites and Pachydiseus, which are characteristic of the

Cretaceous of Vancouver Island. The faunas here described seem

to include both Knoxvile and Horsetown beds of the Californian

section, as shown by such characteristic species as Phylloceras knox-

villense Stanton, Lytoceras batesi Trask, L. sacya Forbes, Desmoceras

breweri Gabb, D. haydeni, and Aucella crassicollis Keyserling.

The revision of the nomenclature will be exceedingly acceptable

and useful to students of West-Coast stratigraphy.

Dr. Sokolow (Mem. Comité Geol. St. Petersburg, Vol. IX, No. 5,

1899) has made an interesting study of the brackish-water basin

fauna that lies immediately between the Mediterranean stage of the

lower Miocene and the Sarmatic stage of the middle Miocene Ter-

tiary of Russia. The Mediterranean stage represents the deposits

of the disappearing ancient Mediterranean Sea, and the Sarmatic

beds are the deposits of the ancient Black Sea. In this paper we

have a study of the fauna transitional between the two epochs and

the two basins. In consequence of this the fauna is a mixed one, -

showing both marine and brackish-water types, due to the rapidly

changing physical geography and the development of the extensive © a

brackish-water seas that covered southern Russia in later Miocene

time.

Most students of the Triassic paleontology of the Alps occupy —

themselves with the cephalopods, while the other groups are

neglected. But Dr. Kittl (Ann. k. k. Naturhist. Hof Museum ——

Wien, Bd. XIV, Nr. 1, 2) has given an elaborate revision of the —

gastropods of the classic St. Cassian beds of the southern Alps —

describing many new species and making known a rich fauna. — ber ue

detailed stratigraphy and correlation of these beds are taken up, xs

much new light is thrown on the relations of the various fossiliferous — '

horizons of the southern Alps. |

In the Bulletin of the Harvard Museum of Comparative ale a

Mr. C. R. Eastman gives descriptions and figures of two species of -

extinct gar pikes, Zepidosteus atrox Leidy and Z. simplex Leidy, Ë E

No. 413. REVIEWS OF RECENT LITERATURE. 2

423

the Eocene Green River shales of Wyoming. Most of the fossil gar

pikes of America have been hitherto known from bare fragments

scantily described. Mr. Eastman’s specimen of Z. atrox is especially

complete, as large as an alligator gar, and very much like it in

appearance. In fact it “lacks any positively archaic features,” and

Mr. Eastman regards it as “obviously the direct progenitor of the

alligator gar, Z. Zristechus.” Mr. Eastman finds no trace of the

earlier ancestry of Lepidosteus. The gar pikes “ blossom forth sud-

denly and fully differentiated at the dawn of the Tertiary without

the least clue to their ancestry, unheralded and unaccompanied

by any intermediate forms, and they have remained essentially

unchanged ever since.”

In the Bulletin of the Kansas University, Vol. I, No. 2, Prof. S.

W. Williston describes and figures many teeth of sharks found in

the Cretaceous rocks of Kansas, his paper being a very useful con-

tribution to this difficult branch of paleontology. In the matter of

nomenclature, apparently, Agassiz’s name, Oxyrhina, should not be

used instead of the earlier Isurus of Rafinesque, and Scylliorhinus

of Blainville has unfortunately clear priority over Scyllium Cuvier.

PETROGRAPHY.

Geology of the Black Hills. — Irving’s contribution to the geology

of the Northern Black Hills adds a great deal to our knowledge of

this interesting region, especially from the point of view of petrog-

raphy. The author agrees with Crosby, rather than with Russell,

in regarding the larger intrusives of the district as laccolites and not

as plugs. He finds also an abundance of sills and dikes. The

dikes characterize the Algonkian slates, the sheets and laccolites

the Cambrian shales. The Carboniferous limestone is almost devoid

of intrusions of any kind. The principal types of rocks recognized

are a quartz-egirite-porphyry, tinguaite, phonolite, trachytoid-pho-

nolite, quartz-porphyry, mica-diorite-porphyry, dacite, tonalite, and

augite-vogesite. The phonolites and quartz-porphyries are the

most abundant types, with the quartz-egirite-porphyries and the

diorite-porphyries fairly abundant. There is such an intimate

gtadation between the different types that they appear to be related

genetically. In the pre-Cambrian rocks, dikes and possibly plutonic

424 THE AMERICAN NATURALIST. [Vor. XXXV.

intrusions of basic igneous magmas took place before the meta-

morphism of the Algonkian sediments.

The author does not agree with Van Hise in ascribing the crys-

talline character of the schists near Deadwood to the agency of

intrusives. He regards the metamorphism as “dynamic” rather

than “ contact."

Isle Royale and Keweenaw Point Volcanics. — In Vol. VI of the

Michigan Survey, Lane? and Hubbard? give a great many interesting

details concerning the petrography of the Keweenawan eruptives.

One of the most interesting features of Lane's paper is his discus-

sion of the cause of the variation in coarseness of grain in rocks, and

the application of his conclusions to the problem of the nature of the

Isle Royale and other rock-sheets. From the fact that the Isle

Royale sheets are characterized by an increase in the size of grain

to their centers, he concludes that they are surface flows or lavas.

On the other hand, he concludes that Lawson's view as to the intru-

sive character of the diabase sheets in the Huronian beds of the

north shore of Lake Superior is confirmed by the fact that they are

characterized by a rapid variation in size of grain for the first few

feet from their contacts with the surrounding rocks, and then by a

central coarser belt of tolerably uniform grain.

Another interesting chapter in the report is that on the differences

in structure between small intrusive basic masses and their corre-

sponding effusive forms. To the already recognized distinctions

between these two forms of igneous rocks, Lane adds that the

miarolitic cavities in intrusive masses naturally become filled with

other minerals than the zeolitic and chloritic ones characterizing the

corresponding pores in effusive rocks. Among the most important

of these minerals is quartz, which often appears in diabase dikes a$

micropegmatitic intergrowths in the central portions of their masses.

The microscopic and chemical features of the Isle Royale lavas are

described in some detail.

The report by Hubbard deals mainly with the structural problems

presented by the interbedded lavas and sandstones in Keweenaw

Point. It contains a few notes in the petrography of the va various

lava beds.

1 Ann. New York Acad. Sci. s vol xh.

? Lane, A. C. Geological Report on pos dons Michigan. Geol. Survey d

Midguo, Ms Me pt. i. ]sites

* Hubbard, L. L. Keweenaw Point, with Particular Reference to the Fe

and their Feeds Rocks, iéid., vol. vi, pt. ii

No. 413.] REVIEWS. OF RECENT LITERATURE. 2

425

Occurrences of Differentiated Magmas. — As the result of a rapid

survey of the Magnet Cove, Arkansas, igneous area, Washington!

concludes that the complex described by Williams represents an

excellent though peculiar example of a highly differentiated magma,

probably in the form of a laccolite, and not a series of independent

intrusions. The rock types present in the area form a regularly

graded series, ranging from foyaite, through leucite-porphyry, shon-

kinite-porphyry, ijolite, and biotite-ijolite to jacupirangite. The

distribution of the rocks is, however, abnormal, since the basic

varieties are near the center of the supposed laccolite, and the acidic

varieties in its periphery. The author explains this abnormality by

supposing the magma to be a solution in which the solvent was in

great excess. This solidified first on the outside, leaving a more

concentrated, and consequently a more basic solution within. The

solvent continued to separate by crystallization as a more and more

basic rock, as the cooling continued inward, until, finally, at the

center the most concentrated and most basic material solidified as

jacupirangite. |

purr? has compared the order of succession of the intrusions

in the Great Basin and has found it to be in general the same

throughout the district, although at any given place certain members

of the series may be lacking. This succession as worked out is

as follows: acid rocks (rhyolites), siliceous intermediate types

(andesites), acid rocks, associated with basalts, basic intermediate

types (more basic andesites and aleutites), basic rocks (basalts),

with associated rhyolites. The Great Basin appears to have been

underlaid by a single body of molten magma which supplied at differ-

ent times lavas of similar composition to all the different parts of

the overlying surface. Since this succession is different from any

described from other regions, the author suggests that the first

rhyolite is the end member of a series of differentiates, and that the

andesites are the beginnings of two distinct cycles of differentiation.

He further suggests that during the first completely recorded cycle

beginning with the earlier andesite the siliceous differentiates of this

magma were erupted in preference to the basic differentiates, while

in the second cycle the basic members were the predominant extrava-

sations. It must be borne in mind that the processes of differentia-

tion are independent of the causes that produce expulsion of lavas,

and hence the records of the differentiation, as observed on the

1 Bull. Geol. Soc. Amer., vol. xi, p. 389.

2 Journ. of Geol., vol. viii, p. 621.

426 THE AMERICAN NATURALIST. [Vor. XXXV. z

surface, must necessarily be incomplete. For this reason the suc-

cession of lavas in any district should be studied with reference to

its general features rather than to its details, and care must be taken

to exclude from the discussion the intermediate rocks formed by mix-

ing of the different types.

The great batholite forming the height of land between the high-

land mountains south of Helena and Butte, Montana, is composed

of rocks presenting a wide variation in composition. The main

rock is a hornblende-granite (I) resembling closely a quartz

monzanite. At Butte the rock is a little more basic (II). Great

masses of aplite (III), possessing often a lenticular shape, are

associated with the granites, perhaps as a differential product of

the hornblende-granite, the Butte granite being the more basic

differentiate. At the periphery of the batholite basic dikes penetrate

the surrounding sedimentaries.

SiO, TiO, AlO Fe,0; FeO CaO MgO K,O NaO H,O Various Tot.

I. GT 48 15.00 Lr XT 53448 194 - 482 ; 2.76 DEI EU

II. 64.03 15.58 1.96 2.83 4.20 2.15 4.11 2.76

III. 77.05 12 12,84 +56 -14 57 tr 5.52 2.81 4 mcr

As the result of his studies on the igneous rocks of the Bohemian

Mittelgebirge, Hibsch? concludes that the succession is as follows :

(1) flows of basalt, preceded by phonolite in a laccolite mass, (2)

flows of tephrites, stock intrusions of essexite, dikes of camptonite, $e

monchiquite, bostonite, gauteite, and sodalite-porphyry, (3) basalts, —

trachytes, phonolites, and dikes of tinguaite and eleolite-porphyty- i

Hackman? has reéxamined the ijolite in the Parish of Kunsamo,

Finland. He finds the main portion of the several massifs to con-

sist of the normal rock (I), but in addition to this he recognizes

also the following as differentiation products — a nepheline-rich

ijolite (II), a soda sussexite (III), a magnesian essexite (IV), and a T

pyroxene-syenite which may, however, be a rock formed by the solu- — |

€ of the granite surrounding the igneous rock in the magma ofthe —

atter. - t

SiO, TiO, ALO, FeO, FeO MnO CaO MgO NaO KO Pis TUM

I 4289 10$ 1945 $34 509 ^.39 1699 374 10039 d] T LLL

ür 4402 .63 -24.63 3.59 2.17 547 1967 143: 499 M a

+ 47:43 mo sbo . 4.65 1.20 4 67 15.08 | 2.00 di

IV..45.66 2.75 1104 3.57 ‘10.61 9.11 11.08 2.60 -44 — í

1 Weed, W. H. Journ. of Geol., vol. vii, p. 737:

? Min. u. Petrog. Mitth., Bd. xix,

3 Bull. Com. Geol. d. Finlande (1

. 488.

goo), No. II.

No. 413.] REVIEWS OF RECENT LITERATURE. 427

Graphic Representation of Rock-Analyses. — Miiggs! proposes

a scheme for the graphic representation of the chemical composition

of rocks based on Brógger's and Michel-Lévy's scheme. The rela-

tive percentages of the various metallic constituents are indicated by

means of polygons drawn through points plotted on eight radii.

Within this is a second polygon which represents the silica content.

The size of the latter is determined by the percentage of this constit-

uent present, and the relative sizes of this polygon and the outer

one are an indication of the rock’s acidity. In constructing the inner

polygon the percentage of silica present is divided into eight equal

parts, and each is plotted in one of each of the radii. In plotting

for the outer polygon the Al,O; is divided into three parts determined

by the proportion borne by K,O and Na,O to one another and the

other bases.

The Origin of the Glaucophane-Schists. — Rosenbusch, as is well

known, has hitherto suspected that true glaucophane-schists are

genetically associated with sedimentary rather than with igneous

rocks, but so few analyses of these schists have been made that the

supposition has not been capable of chemical investigation. In a

recent? article, however, he shows that some of the schists have the

composition of a normal gabbro magma. In these epidote, zoisite,

lawsonite, prehnite, margasite, and garnet are usually if not always

present. Rocks of this kind are closely related to amphibolites.

Other glaucophane-schists he still believes to be metamorphosed

sediments, but analyses of these are lacking.

Washington? supplements Rosenbusch's investigations in an arti-

cle in which he records and compares fifteen analyses of these

Schists. Upon comparing their analyses he discovers that the rocks

fall into two main groups, a very basic group with a content of

SiO, varying between 46% and 49.7% and a very acid group with

SiO, between 74.5% and 82.5%. The former he believes, with

Rosenbusch, to be derived from gabbros, diabases, or their tuffs.

The acid glaucophane-schists he thinks are derived from cherts,

quartzose shales, or quartzites. The basic forms scarcely differ from

the amphibolites in chemical composition, the formation of the

one or the other kind of schists depending probably upon conditions

of metamorphism.

1 Neues Jahrb. f. Min. etc., Bd. i (1900), p. 100.

2 Sitzb. kin. preuss. Ak. Wiss. Berlin, Bd. xlv (1898), p. 716.

3 Amer. Journ. Sci., vol. xi (1901), p. 35.

CORRESPONDENCE.

Editor of the American Naturalist:

Sin, — In his valuable paper on “the Snakes of New York State,”

in the February number of the Watwra/ist, Dr. Eckel enumerates two

forms as doubtfully occurring in the state upon the authority of

Cope who, in his Crocodilians, Lizards, and Snakes of North America,

records specimens as being in the collection of the National

Museum. Dr. Eckel properly doubts the correctness of the state-

ments, and I am in a position to corroborate him.

The first is a specimen of Osceola doliata clerica, United States

National Museum, No. 1407. The value of the locality * New York"

may be inferred from the fact that the specimen was received in

1858 from the Museum d'Histoire Naturelle in Paris, together with

No. 1405, Elaps lemniscatus, and No. 1406, Dromicus cursor, — all

three said to have come from New York. It is simply a case of

French geography.

The other case is that of a Natrix fasciata erythrogaster, United

States National Museum, No. 9984, said to have come from estfield

Falls, Conn. In this case the geography is correct enough, büt the

identification is wrong. I have just examined the specimen; it is a

Natrix sipedon pure and simple! =

Thus Dr. Eckel may safely eliminate Nos. 10 and 13 from his

i ` 7

check list. LEONHARD STEJNEGER.

U. S. NATIONAL MUSEUM,

February 15, 1901.

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Professor W. M. WHEELER 431

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(Palamedea: Chauna) : . Dr. B, W.SHUFELDT 455

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THE

AMERICAN NATURALIST

Vor. XXXV. June, 1901. NO. 414.

THE COMPOUND AND MIXED NESTS OF

AMERICAN ANTS:.

WILLIAM MORTON WHEELER.

Part I. OBSERVATIONS ON A NEW GUEST ANT.

Or the many symbiotic relations known to exist between

ants and other living organisms, whether animals or plants,

few are more interesting than the relations between ants of

different species or even genera that live together in the same

nests on terms of mutual toleration or intimacy. In this

country, nevertheless, studies like those of Forel (74), Adlerz

(84, '86,'96), and Wasmann (91) on the compound and mixed

nests of European ants have scarcely been undertaken, notwith-

standing the fact that our own ant fauna is known to present

numerous cases no less remarkable than those which have been

so carefully studied abroad. Sufficient evidence of the truth

of this remark will be found in the description of a hitherto

unknown form of compound nest which is made the basis of the

Present paper. This is in many respects so unlike any of

the recorded compound nests that it seems worth while, if only

! Contributions from the Zoölogical Laboratory of the University of Texas, No. 14.

431

432 THE AMERICAN NATURALIST. [Vor. XXXV.

for the sake of comparison, to follow it with a consideration of

the other forms of mixed and compound nests that are known to

occur on our continent. While it thus becomes necessary to

review much that is well known to the myrmecologist, I hope at

the same time to call the attention of entomologists, who may

have wearied of collecting and mounting our comparatively well-

known beetles and butterflies, to an extensive subject, which is

as interesting to the collector as it is fascinating to the philo-

sophical observer. Ants’ nests of the mixed and compound

varieties are of sporadic and often very rare occurrence, so that

there is great need of many more observations extending over

large portions of our country. While reviewing the various

forms of nests in the second part of my paper, I shall include a

brief consideration of some interesting compound nests that have

recently come under my observation in Texas and Mexico.

During the late afternoon of August 1, 1900, while walking

over one of the Litchfield Hills near Colebrook, Connecticut,

I found a number of nests of the common red-brown Myrmica

(M. brevinodis Emery) under some small stones that were

rather deeply imbedded in the moss bordering the exposed gla-

ciated rock of the hilltop. In four of these nests which were

rather close together and not very populous I detected among

the Myrmica workers a few decidedly smaller and more yellowish

ants of a different species running about on the up-turned lower

surfaceof thestones. As my stay in Colebrook was at that time

limited to a few days, I carefully replaced the stones in the

moss after preserving a few of each of the two species. The.

smaller ants were sent to Professor Emery of Bologna, who pro-

nounced them to be a new species of Leptothorax allied to

L. canadensis Provencher.

Later in the month of August I returned to Colebrook and

at once revisited the Myrmica-Leptothorax nests. ae of

them had disappeared, as if in resentment of my former intru-

sion into their peaceful life on the hilltop. The third nest a

tained only a few Leptothorax workers. The fourth nest was 1n

good condition and was dug up zz £oze and carried home in xu

vas bag. For several days I searched diligently, but in vain, m

all the hills about Colebrook for more of these double nests,

No. 414.] NESTS OF AMERICAN ANTS. 433

though single nests of the Myrmica were frequently found. My

observations were therefore to be confined to only one nest.

The worker of Myrmica brevinodis, which I shall not pause

to describe, is represented in Fig. 1. The Leptothorax, however,

Fic. 1. — Myrmica brevinodis Emery. Worker.

is of greater interest in this connection and certainly merits a de-

scription before we proceed with an y observations on its behavior.

Leptothorax emersoni n. sp.!

Worker (Fig. 2). Length 2.5-3.5 mm.

Brownish yellow ; upper surface of the head except the

mandibles, clypeus, and a narrow postclypeal and occipital

! I take pleasure in dedicating this species to Mr. Ralph Emerson in memory

of the many h hich together at Rockwell Hall in Colebrook.

pigg i me x di hours Y P

434 THE AMERICAN NATURALIST. [Vor. XXXV.

strip, dark brown, as is also a very broad band across the middle

of the first abdominal segment and the basal half of each of

the succeeding segments. Hairs covering the mandibles, anten-

nze, legs and body, whitish ; those on the body rather long, coarse

and truncated, but scarcely clavate

at their tips; those on the limbs

tapering and less conspicuous.

Mandibles rather coarsely striated

longitudinally and provided with a

few setigerous punctures; cutting

edges black. Clypeus moderately

convex, smooth in the middle but

with a few delicate longitudinal

ruge on either side. Antenne

11-jointed. Head opaque, above

and on the sides coarsely and

sp. longitudinally reticulate-rugose, the

areole enclosed by the rugæ being

indistinctly and finely punctate. In the mid-dorsal region

of the head the rugz are close together and their longi-

tudinal course is very distinct, but in the antennal fovez and

on the cheeks and the sides of the head they are farther

apart and more reticulate. Some of the workers have

two or three minute ocelli which are most easily seen in alco-

holic specimens. Thorax faintly but distinctly constricted at

the meso-metanotal suture; opaque, above and on the sides

less distinctly reticulate-rugose than the head. Anterior por

tion of the pronotum, metanotum, and lower pleurz punctate ;

metanotal spines slightly longer than broad at their bases,

nearly parallel, terminating in broad, flattened, blunt points.

Petiole about twice as long as broad, subopaque, finely punctate,

with a pointed dorsal cone, the anterior and posterior surfaces

of which are of nearly equal length and inclination, the former

being very slightly concave, the latter flat when seen in profile.

Postpetiole smooth, subglabrous, hemispherical. Abdomen

rather broad, distinctly flattened dorso-ventrally, and very

glabrous.

Fic. 2.

7 Add E

Worker with ocelli.

No. 414.] NESTS OF AMERICAN ANTS. 435

Female. Length nearly 4 mm.

Thorax more opaque, more distinctly rugose, and of a darker

brown color than in the worker.

Male (Fig. 3. Length 2.75-3 mm.

Coloration brownish yellow like that of the worker; head,

thoracic dorsum, abdomen, and the middle portions of the femora

and tibia somewhat darker. Wings colorless, with very pale

veins and stigmata. Hairs almost completely absent on the

head and thorax, short and inconspicuous on the legs, long on

the abdomen, but nowhere truncated at their tips.

Mandibles striated, hairy, very small, not distinctly dentate,

and far from meeting each other with their blades. Clypeus

even in the middle with a few delicate longitudinal ruge.

Head above longitudinally reticulate-rugose; ruga radiating

backwards and laterally from the posterior ocelli as centers.

Antenne 12-jointed ; scape scarcely as long

as the three first joints of the flagellum

taken together; first joint of flagellum very

short, the others of uniform thickness but

increasing gradually in rength towards the

tip. Thorax rather smooth, indistinctly

punctate ; parapsidal and other sutures

very distinct ; metanotum with two very

short rugose projections in the place of the

spines. Dorsal projection of the petiole

rounded, its posterior slope slightly concave,

the anterior convex. Postpetiole hemispher-

ical; abdomen very glabrous, somewhat

less flattened than that of the worker. Fic. 3.— Leptothorax

Leptothorax emersoni is to be assigned to — 777^ pigri

the small group of North-American species comprising Z. mus-

corum Nyl. (also recorded from Europe), Z. canadensis Proven-

cher, with its variety yankee Emery, L. hirticornis Emery, and

L. provencheri Emery. The workers of all these forms have

I1-jointed antennz and a distinct meso-metanotal constriction.

That the new species is perfectly distinct from all of these is

436 THE AMERICAN NATURALIST. [Vor. XXXV.

quite evident from consulting Emery's table of the North-

American species of the genus (95, pp. 317,318).

The above very incomplete account of the female is drawn

from memory, as the single specimen escaped from the nest

before it could be described.

Some of the workers, especially those which approached the

queen in size, possessed minute ocelli. This is shown by the

following measurements of the four workers stil in my

collection :

I. Length 2.5 mm. No ocelli.

No. 2. Length 2.75 mm. No ocelli.

3. Length 3 mm. Two ocelli.

No. 4. Length 3.5 mm. Three ocelli.

This fact is of some interest, as I shall endeavor to show in

the sequel.

On the morning of August 25 the ants were transferred

to an extemporized Lubbock nest. This consisted of a board

surrounded by water, and a little fine, moist earth placed on

the board and covered with a pane of glass. A census of the

two species of ants gave the following :

Myrmica: 6 deálated! queens; 7 males ; about 200 workers;

a few eggs; 23 larva, both young and old (but apparently no

queen larva) ; 8 worker and male pupa. The latter hatched

in the course of a few days.

Leptothorax: 1 deálated queen; 4 recently hatched males ;

14 workers; 2 adult larvae, which soon became male pupa.

The earth with its occupants was dumped from the bag on to

the pane of the Lubbock nest, and the ants, after the first

flurry of excitement, began to seek refuge under the glass.

As usual the larvze and pupa were at once conveyed to a place

of safety. While this operation was going on, it was noticed

that some of the Myrmica workers carried the Leptothorax

larvze, and the Leptothorax workers reciprocated by occasionally

1I feel compelled to coin this term for use in the sense of the German “ent

fliigelt.” The term aféerous cannot be used without confusion, since some ants

(Eciton, Dorylus, Leptogenys, Tomognathus) have truly apterous queens, wt

=r queens of most species of Formicidæ have well-developed wings, up t° the

time of their removal after the nuptual flight.

No. 414.] NESTS OF AMERICAN ANTS. — 437

carrying some of the Myrmica larva. This action on the part

of both species was evidently the result of haste and excite-

ment, as I never saw it repeated subsequently except once,

when a Leptothorax carried a Myrmica larva a short distance

and then dropped it.

As soon as the excitement had subsided the Myrmicas pro-

ceeded to dig galleries in the soft earth between the glass

pane and the board, and the Leptothorax at once migrated into

them. A few hours later the earth that had been dumped on

the pane was carefully removed and replaced by a piece of

opaque cardboard which was only lifted from the glass when

the ants were under observation. A small dish containing a

syrup of sugar and water was placed near the nest. This

was soon found by two of the Myrmica workers, which at once

gorged themselves with the liquid and returned into the nest,

where they proceeded to dole out the store of food to their

hungry sisters. It was then that I was able to make my first

observation on the mutual relations of the two species of ants.

A Leptothorax worker was seen to follow up and to climb

on to the thorax of one of the food-distributing Myrmicas soon

after it had entered the nest. In this position the little ant

proceeded to lick the back of the head and clypeus of the

Myrmica with signs of agitation as indicated by the hastening

of the tremulous beat of its antennz and the throwing of its

abdomen and postpetiole into stridulatory oscillation. The

Myrmica paused as if spellbound by this shampooing and

occasionally folded its antennz as if in sensuous enjoyment.

The Leptothorax, after licking the Myrmica’s pate, moved its

head around to the side and began to lick the cheeks, man-

dibles, and labium of the Myrmica. Such ardent osculation

Was not bestowed in vain, for a minute drop of liquid — evi-

dently some of the recently imbibed sugar-water — appeared

on the Myrmica’s lower lip and was promptly lapped up by

the Leptothorax. The latter then dismounted, ran to another

Myrmica, climbed on to its back and repeated the very same

performance. Again it took toll and passed on to still another

Myrmica. On looking about in the nest I observed that nearly

all the Leptothorax workers were similarly employed. In one

438 THE AMERICAN NATURALIST. (Vor. XXXV.

corner of the nest a number of Myrmica workers had formed

a circle about a few of their small larvae which they were

cleansing and feeding. A Leptothorax soon found its way to

this cluster and stepped from the back of one ant to that of

another, lavishing a shampoo on each in turn and apparently

filling its crop with the liquid contributions thus solicited.

This and numerous very similar observations, which could

be made at any time on removing the cover of the nest, prove

conclusively that the Leptothorax workers demand and obtain

their food from the Myrmica workers. The method of solicit-

ing food, however, differs from that of any other myrme-

coxenous animals known to me. These animals usually request

food by tapping the ant with their antennae (many myrme-

cophilous beetles), or stroking its face with their fore feet

(Atemeles), but none of these guests are so unconventional as

to mount the backs and scratch the heads of their hosts for

the purpose of inducing the latter to regurgitate. Even the

slave-holding Polyergus and the social parasite Anergates

demand and receive food after the manner of other ants. The

Leptothorax workers are so persistent in their peculiar atten-

tions to the Myrmicas that I have come to doubt whether the

little guest ants ever really feed themselves. Once only was

a Leptothorax seen to approach the dish of syrup, lap up 4

very little of it hastily, and then return to the nest. This

happened before the ants had definitely settled under the pane

of glass. After that only the Myrmica workers visited the

manger, and the Leptothorax usually waylaid them as soon as

they had entered the nest. During my first visit to the four

natural Myrmica-Leptothorax nests I found the latter species

loitering in the outer galleries just under the stone. I am con-

vinced that they prefer this situation in order to be on hand

the very moment a food-laden Myrmica enters the nest. The

Leptothorax workers often walked on the lower surface of the

roof-pane, although they had to turn over to mount the passing

Myrmicas. They must do this also in the natural nests, for

the specimens taken August 1 were, as above stated, crawling

on the lower surface of the stones.

Although the feeding of the guest ants commonly proceeds

No. 414.] NESTS OF AMERICAN ANTS. 439

as above described, I have noticed that the Leptothorax after

mounting a Myrmica sometimes turns about and licks the

metathorax or even the abdomen of its host, as if, like Myrme-

cophila (see Wheeler, '00), it found the surface covered with

some agreeable secretion. It seems, therefore, not improbable

that the Myrmicas may derive some slight benefit from guests

which, like many tonsors, combine the occupation of the bar-

ber and bather. It is perhaps unnecessary to add that the

Myrmica does not always pay for the shampooing it receives.

But the Leptothorax is not discouraged ; it merely dismounts

and runs about in the galleries till it falls in with another

Myrmica.

The Leptothorax workers "were never seen to approach the

male Myrmicas, and only once did I see one shampooing one of

the queens. So conspicuous was this neglect of the two fertile

sexes of their hosts and their predilection for the workers, that

I was convinced that they habitually ignore the queens and

males, because these insects, like themselves, depend on the

Myrmica workers for their sustenance and are probably on

that account not in the habit of regurgitating.

On the evening of the day on which the ants moved into

the Lubbock nest and during a portion of the following day,

August 26, the queen Leptothorax wandered about outside the

nest as if seeking a more favorable retreat. By 4 p.m., how-

ever, she had entered the nest and, with eight of her workers

gathered about her and her two mature larve, was found

occupying a small earthen chamber under the very middle of

the roof-pane. This chamber, which had evidently been dug

by the Leptothorax, was surrounded on all sides by the large

galleries of the Myrmicas. Seen from above, the Leptothorax

nest had the appearance of Fig.4 a. The queen, workers, and

larvee, nearly filled the small cavity, a, which communicated

with a wide Myrmica gallery, c, by means of a passage, o, too

small to admit a Myrmica. Through this narrow passage a

few Leptothorax workers entered or passed out from time to

time, but several always remained in the nest with their queen.

Usually from one to six workers were to be seen soliciting

food among the Myrmicas.

: 440 THE AMERICAN NATURALIST. [Vor. XXXV.

Three of the six Myrmica queens, together with the Lepto-

thorax males, were preserved in alcohol, as the nest was somewhat -

over-crowded with the former species and the latter were continu-

ally escaping. The three remaining Myrmica queens stationed

themselves some distance apart in the galleries, and each was

soon surrounded by a coterie of devoted workers. The larve

and pupa were being cared for in two or three different

portions of the nest.

The Leptothorax in their small central nest passed their

time in fondling and feeding one another or in lying motion-

less as if asleep, covering the two larvae which had been placed

in the bottom of the nest. The queen was assiduously fed by

the workers and was never seen to leave the nest after once

taking possession of it. On several occasions she was observed

to throw one of the workers down on its back and to hug and

kiss it in the most animated manner. It was not easy to

decide whether this behavior signified maternal affection, the

presence of some form of the play instinct, or the more prosaic

feeling of hunger. I incline to the last possibility, although

the action certainly resembled the affectionate struggles of a

cat with her kitten.

During the remainder of this day (August 26) the Lepto-

thorax nest remained in the condition represented in Fig. 4- By

8 o'clock the following morning, however, the bottom of the

nest had been dug somewhat deeper, its narrow entrance had

been closed up and a new one, equally tenuous, had been

opened in a different position (Fig. 57). Thetwo larvae had

become male pupa. By noon the queen had laid three ellipti-

cal white eggs of rather large size. At 1 P.M. the Myrmica

workers discovered the hiding place of their little companions,

and two of them in single file shouldered their way through

the narrow passage, 7, enlarging it as they proceeded. As soon

as the head of the first Myrmica appeared in the chamber, the

Leptothorax, which had been attending to their morning toilet

and to that of their larvae and to the careful arrangement of

their eggs, turned to meet the intruders. For an instant I

fully expected to see a fierce battle, but I had misjudged the

Leptothorax character. To my surprise the Myrmica

No. 414.] NESTS OF AMERICAN ANTS. 441

her companion on entering the chamber were received with a

profusion of shampooing. The large Myrmicas, though sadly

crowding the occupants of the little chamber, let themselves

Fig 8 wy & . ANES

Fic 5 r

3S. 4-9. — Ledia ; WE ; SEU

9. — Nest of Leptoth in. sp., within the nest of Myrmica scabrinodis Nyl.

an comfortably and appeared to experience all the sensuous

Cpl of a couple of jaded roués who have dropped into a

vinim bath for the night. The naturalist of a past generation

probably have interpreted the behavior of Leptothorax

442 THE AMERICAN NATURALIST. [Vor. XXXV.

under these circumstances as a politic act of hospitality on

the part of a defenseless but intelligent creature. To-day

there would be little hesitation in interpreting it as merely a

machine-like reflex called into activity by its customary stimu-

lus, the presence of the Myrmicas. That the truth lies some-

where between these two conceptions, though possibly nearer

that of reflex than of intelligent action, was apparent from the

subsequent behavior of the Leptothorax. These ants undoubt-

edly had some dim desire to remove the Myrmicas from their

nest, for from time to time a Leptothorax was seen to pull

with her mandibles at the fore leg or antenna of one of the

intruders, as if to remind her that there are limits to polite

hospitality. This action was never performed by the Lepto-

thorax while foraging in the Myrmica galleries, but it was

regularly performed whenever, as on this and several subse-

quent occasions, any Myrmica broke into the central chamber.

The direction of the tugging was not very definite or constant.

Often when a Myrmica thrust its head through the wall, the

tugging was indeed towards the interior of the chamber, as i

to draw the intruder in. But as the small ant was not able

to move the large Myrmica, and as it could not under the

circumstances tug in any other direction, the action could

hardly be regarded as anything more than a gentle means of

persuading the Myrmica to leave. This tugging was the

only act even approaching hostility witnessed between be

two species. The Myrmicas never showed the slightest a

tation towards the Leptothorax, never seized them in their

mandibles, nor even menaced them. They seemed rather to

look upon the little creatures with gentle benevolence, much i

human adults regard little children. They never passed their

little guests without the antennal greeting, and the Leptothorax

shampooed their hosts with comical zeal.

The two Myrmica workers whose intrusion into the Lepto- —

thorax chamber led to the above observations finally departed,

only to give a second party of Myrmicas an opportunity e

make a large breach in the wall at xx. These entered the

chamber at 1.20 p.m. and were received in the same ga

manner as the first party, and in turn departed after being 3

No. 414.] NESTS OF AMERICAN ANTS. 443

politely requested to leave. The Leptothorax then at once set

to work to repair their dilapidated wall. At 1.30 a worker

went out into the adjoining gallery, picked up a pellet of earth

and placed it in the breach. Again and again she returned and

gathered earth, often going to a distance of one or two inches

from the chamber for suitable pellets. Another worker soon

began to assist in repairing the breach from the inside, tak-

ing the pellets for this purpose from the inner wall of the

chamber. Then the first worker walked around the nest,

entered it through the passageway at x and began to clean

herself, while a third worker went out through the breach

and continued the work on the outside till the wall was

completed. This was accomplished by 3 P.M.

At 4 P.M. a little water was poured under a corner of the glass

where the Myrmicas had congregated in greatest numbers.

This additional moisture induced them to move mit Kind und

Kegel to the middle of the nest. Here they soon began to

break through the walls of the Leptothorax cell in two places

(Fig. 6 ss). Two Myrmicas again settled down in the cell

and underwent the usual shampooing. As soon as they had

departed the little ants again set about repairing the walls as

before. Sometimes three or four of them worked at the same

breach. During the progress of the work they frequently went

from two to three inches into the Myrmica galleries in search

of the requisite earth. At the same time a few workers toiled

from the inside of the cell, and these were soon joined by the

queen, working as busily as any of her progeny.! Occasionally

à worker, after building for some time on the outside, would

slip through the breach, turn around and build from the inside.

Twice Myrmicas rushed up to the spot s (on the right side in

Fig. 6) and commenced tearing down the wall They easily

took out pieces of earth eight or ten times as large as those

Which the little Leptothorax workers were putting in with so

much care and difficulty. But the infraction of the Myrmicas

did not escape the attention of the Leptothorax. They

1 It is interesting to note in this connection that Forel ('74, pp. 339-341) wes

ago observed that the queens of the European species of

"um and tubero-affinis) do not shirk their share of the menial labors of the nest.

444 THE AMERICAN NATURALIST. [VoL. XXXV.

interrupted their repairs to shampoo and kiss the interlopers and

again they tugged them by a leg or an antenna, sometimes in

one direction, sometimes in another. And again I was forced

to conclude that the Leptothorax workers wished to dissuade

their big hosts from trespassing on their property. At any

rate, the Myrmicas changed their plans and retreated to another

part of the nest, just as the other parties had done on former

occasions. The Leptothorax then continued their repairs. By

6 P.M. the walls had been rebuilt and the cell had the appearance

of Fig. 7. The original entrance, 7, had been much narrowed

so as to exclude all but the slender-bodied inhabitants of the

chamber.

At 7 o'clock on the following morning (August 28) the

Leptothorax nest was found in statu quo, except that the queen

had laid three more eggs during the night. By noon, however,

the Myrmicas had again broken into the cell, so that at 4.30 P.M.

the wall was torn down in several places. Nevertheless, the

ever alert guest ants had piled up the earth so that the Myr-

micas could scarcely squeeze between it and the glass roof-

pane. The inroads of the Myrmicas had been so extensive,

however, that even as late as 7 P.M. the nest presented the

appearance of Fig. 8.

By 7 a.m. on the following day (August 29) the nest had been

almost entirely rebuilt, as shown in Fig. 9. The Leptothorax

must have labored during a large portion of the night. They

had remodeled the nest, giving it a circular form, whilst appar-

ently retaining the old opening at r. Besides this opening

they had two others at z z, which were underground passages.

The ants could be seen diving into these and anon reappearing

within the circular chamber, the bottom of which had been

sunk still deeper in the soil The neat little nest now Com

tained ten eggs. From day to day the Myrmicas had been

widening their galleries, as is readily seen by comparison of

Figs. 4 to 9, so that only small pillars of earth remained to

support the roof-pane around the Leptothorax cell.

On succeeding days essentially the same conditions as those

above described were repeated with slight differences in detai

For the sake of completing the history of the double nest, the

No. 414.] NESTS OF AMERICAN ANTS. 445

observations extending from August 30 to September 4 are

condensed in the following notes :

August 30. The round cell, which remained undisturbed all

day yesterday, was still intact at 8 o'clock this morning. By

noon, however, the Myrmicas had torn down its wall in several

places, and three of them were found in the cell, submitting to

a vigorous shampoo. By 8 o'clock in the evening the circular

nest had been rebuilt. The opening at x had been closed and a

new one opened at ;; (upper left-hand corner of cell in Fig. 9).

August 31. Almost an exact repetition of yesterday's

performance. ;

September I. .'This morning the reconstructed Leptothorax

cell is smaller. Its upper entrance, » and the two subter-

ranean entrances, z z, have not been changed. The ants have

sunk the pupz and eggs to a greater depth in the earthen floor

of the nest, so that they are almost in contact with the board.

The nest was not molested by the Myrmicas during the day.

September 2, The cell this morning has further decreased in

size but is still intact. Its cavity is not more than 7 mm. in

diameter, so that the Leptothorax family is much crowded. The

two subterranean entrances at zz are still in use but the upper

entrance has been shifted to mm (lower left-hand corner of cell in

Fig. 9). The Myrmicas still leave the cell unmolested.

September 3. To-day, too, the Leptothorax were left in peaceful

possession of their cell. Many of them went out into the gal-

leries from timeto time to shampoo the Myrmicas and solicit food,

which they then distributed to their queen and to the few workers

remaining at home. During the day the upper entrance at m

was closed so that the wall of the nest was everywhere in close

contact with the roof-pane. The ants still entered and left the

chamber through the two underground entrances at 7 7.

September 4. At 8 a.m. the Leptothorax nest was unchanged,

but by noon its circular walls had grown perceptibly thinner, as

the Myrmica workers had again taken to removing the earth

from the outer surfaces. The nest was now shaped like a volcano

with sloping sides and the guest ants inhabiting the crater. By

5 P.M. the Myrmicas had made two breaches in the walls. The

nest remained in this condition throughout the evening.

446 THE AMERICAN NATURALIST. (VoL. XXXV,

The rhythmical assaults of the Myrmicas on the Leptothorax

retreat in the afternoon, on several of the days during which

the ants were observed, is, I believe, to be explained as the

result of rising temperature. In the Litchfield Hills the nights

and mornings of late August and early September are rather

cool, while the noon hours may be very warm. The Lubbock

nest happened to be placed at a window in a room with south-

western exposure, so that the diurnal variation in temperature

must have been keenly perceived by the ants. In the warm

afternoons the activities of the Myrmicas increased; they ran

about more rapidly, became more enterprising, and indulged

their excavating instincts to a greater extent. Then in the

course of this employment they often broke into the Lepto-

thorax retreat.

The fact that the Leptothorax changed their entrances from

time to time, and, as shown by the figures, kept perfecting the

form of their cell, thereby making it easier to guard and rebuild |

and more difficult for the Myrmicas to demolish, is evidence

of the remarkable psychic plasticity of these ants. Similar

behavior on the part of ants that have been repeatedly dis-

turbed by other species are recorded by different observers.

Forel (94, p. 8) brought a large formicary of Myrmucocystus

altisquamis from Algiers and gave it an opportunity to establish

itself in a garden near Zürich. The African ants were much

annoyed by the incursions of Lasius niger and T: etramorum

cespitum, and although they at first adhered to their Algerian

custom of maintaining a large open entrance to their nest, they

learned during the course of the summer to narrow the oper-

ing gradually. Finally they plugged it up completely with

grains of earth and made only a small temporary orifice when-

ever they strolled out on sunny days. Wasmann (97, pp. 69, 7 9)

mentions a nest of Formica sanguinea that resorted to a similar

method of protecting itself from the repeated attacks of a

neighboring colony of F. pratensis. All these observations

go to show that Bethe's conception (98;'00) of ants as #7

“reflex machines” cannot be entertained.

From the persistent and strenuous efforts of the Leptothorax

to intrench themselves, and from the shape and character

No. 414.] NESTS OF AMERICAN ANTS. 447

their chamber and its entrances, we are justified in concluding

that these little guest ants must be in the habit of constructing

similar lodgings for themselves in the midst of the natural nests

of the Myrmica. In their natural environment the Leptothorax

would not be cramped for space as they were in the artificial

nest, and they would probably dig their cell where they would

not be so frequently disturbed by their inquisitive hosts.

While making the observations above recorded I was much

impressed with the poverty-stricken appearance of the Myrmica

colony. Although it originally contained no less then six fertile

queens, the number of workers, larvz, pupa, and eggs was dis-

proportionately small. Furthermore, many of the workers were

of rather diminutive size, and a few of them had crippled

abdomens. I began to suspect that the Leptothorax might

be appropriating the liquid food regurgitated by the Myrmica

workers on the mouths of their larvae. With this in mind I

closely watched the larve and the inquiline ants. Once I saw

a Leptothorax lick the mouths of two young larva that were

lying side by side and on another occasion a Leptothorax

licked the body of one of the larvæ. But finally, on September 3,

I made an observation which convinced me that the regur-

gitated food is not stolen from the mouths of the Myrmica

larvae. On the morning of that day four Myrmica workers

were found tugging at the head of a large larva. Under

the lens I could see that a mass of regurgitated syrup had

hardened over the face of the larva like a mask, and that the

four workers were trying to remove it. At last one of them

succeeded in pulling it off, and while she was moving away

with it the mass stuck to the roof-pane for a moment, and I

could see that it was quite hard and glutinous and must have

remained on the face of the larva for some time. As the

Leptothorax were continually roving about the galleries in

search of food, they could hardly have failed to appropriate so

rich a morsel if they were in the habit of obtaining their food in

this manner. If any conclusion can be drawn from the single

nest to which my observations have been confined, it would seem

to be that the poverty-stricken condition of the Myrmicas, not-

withstanding the number of fertile queens, must be brought

448 THE AMERICAN NATURALIST.

about more indirectly by the Leptothorax, vzz., by their continu-

ally pestering the Myrmicas for food and thus diverting to

their own use much of the sustenance that would, under other

conditions, benefit the Myrmicas themselves and their progeny.

If I have correctly estimated the influences which may tend

to diminish the fecundity and prosperity of the Myrmicas we

have in this double nest another striking demonstration of the

complete absence in ants of any faculty of reason. For, if the

Myrmicas possessed a glimmer of this faculty, they could easily

annihilate the gluttonous little nest mates that are forever roam-

ing about their galleries like so many animated stomach pumps.

As I was obliged to leave Colebrook I took advantage of the

cold morning of September 5, when the ants were inactive, to

transfer them all to a Cohansy jar containing some earth. From

this date till October 1 I was traveling about and was there-

fore compelled to suspend observations on the compound nest.

On returning to Austin, Texas, October 1, the ants were again

transferred to a Lubbock nest, but to my dismay I found only

the queen and a single worker of the Leptothorax remaining.

The eggs, the two male pupze, and the other workers of this ant,

together with nearly all the larvae and pupz of the Myrmica,

had disappeared. The Leptothorax queen was very uneasy and

wandered about outside the Myrmica nest. On the following

day she disappeared. She had probably contrived to cross the

moat on the bodies of some drowned Myrmicas.

The single worker remained in the nest and for several days

ran about shampooing the Myrmicas and soliciting food. On

October 7 she, too, escaped from the nest and was found strug-

gling in the moat. She was rescued from drowning and soon

recovered sufficiently to crawl into the nest, though she showed

no desire to mingle with the Myrmicas. Finally she lay motion:

less in a deserted gallery, where she was found dead the next

morning. Thus ended the last of these fascinating little ants:

(To be continued.)

THE LOUISIANA DEER.

GLOVER M. ALLEN.

Ir has long been known, in a general way, that the deer

inhabiting the lowlands of Louisiana is distinctly different

from the related races to the north and east. As long ago

as 1820, Geoffroy Saint-Hilaire and Cuvier! figured a doe

in summer pelage under the name of “Biche du Cerf de la

Louisiane.” They did not, however, consider it different from

their “Cervus virginianus” (= Odocelus virginianus virginianus

Bodd.

Through the kindness of the museum authorities, I have

been enabled to examine the series of deer skins and skulls in

the Museum of Comparative Zoólogy. In the Bangs collection

are four skins and skulls of deer from Mer Rouge, Louisiana.

These represent a well-marked race, which may be known as

Odocelus virginianus louisianz, subsp. nov.

Type: No. 9111, collection of E. A. and O. Bangs, in Museum

of Comparative Zoólogy. Adult male, from Mer Rouge,

Morehouse County, Louisiana. Collected Nov. 8, 1898, by

B. V. Lilly.

General characters : Size large; color, in winter, pale; skull

long and slender; lower row of molar teeth long; antlers

heavy and high.

Color of type —body : dorsal surface from between the eyes

to the root of the tail a grizzled cream-buff and brownish-black ;

1 Histoire Naturelle des egeta: etc., tome i, livr. xvii, , Paris, 1 1824.

UR pointed out by Dr. . Allen in the American Naturalist, vol. xxxiv.

* 400, p. 318 (April, 1900), galas did not propose the name americanus

i the Virginia deer in his sentence, * Differtne vere americanus uti Pennanto

rxle

American form, giving a description and synonymy without himself passing

judgment on its validity as a distinct species.

449

450 THE AMERICAN NATURALIST. [VoL. XXXV.

each hair has a fine black tip, below which it enlarges slightly

in diameter and presents a zone of cream-buff, then one of

brownish-black, which shades into whitish near the root. Pass-

ing ventrally from the back to the sides, this whitish base grad-

ually becomes more extensive, to the final exclusion of the

brownish-black zone, so that the sides of the belly are a nearly

uniform cream-buff to ochraceous-buff. On the lower part of

the shoulders the color of the back shades into a grizzled grayish.

The tail above is nearly unicolor with the back and has a few

pale rufous hairs at its base ; the terminal third is nearly uni-

form black. The ventral surfaces of the body and tail are

white. The chest is ochraceous-buff, with a slight intermix-

ture of long brownish-black hairs between the forelegs.

Head and neck: A small area behind and between the

nostrils and a semicircular patch ventrally on each side of the

angle of the jaw, brownish-black. Chin, upper throat, eye-ring,

and sides of the muzzle whitish; a few black bristles about the

eyes. The rest of the sides of the head are a finely grizzled

white and brownish-black, passing into the white of the chin

and the bright cream-buff of the throat. The ears on their

outer surfaces are similar in color to the top of the head and

have the edges of the tips brownish-black ; their inner surface

is lined with long, delicate white hairs.

Limbs: The inner surfaces of the axilla and thighs are

white ; the color of the outer surfaces passes from the color of

the sides of the body proximally, to an ochraceous-buff distally.

A line of white hairs is present between the toes. The meta-

tarsal gland is small, 14 mm. in length, and is surrounded by

stiff, short white hairs, forming a tuft which has a concealed

outer border of brownish-black. The hoofs and dew-claws are

as long as in the type of Odocelus virginianus borealis (Miller),

but slightly narrower.

The three other topotypes (Nos. 9112, 8622, 8623, Bangs

collection) agree closely with the type in coloration, but

No. 8623 has a darker median dorsal line, and the upper

surface of the tail is uniformly black save for a few rufous

1 Odocelus americanus borealis Miller, Bull. N. V. State Mus., vol. viii, No. 38

(October, 1900), p. 83.

No. 414.] THE LOUISIANA DEER. 451

hairs near the base ; in No. 9112, as compared with the type

specimen, the tail is more rufous, with a short black tip.

I have not seen specimens of this deer in the summer pelage,

but the doe in summer coat figured by Saint-Hilaire and Cuvier

is bright reddish, and as the nearest races to the east and west

are brighter in summer than in winter, a seasonal change of

similar nature may be looked for in the Louisiana deer.

The skull: The skull of the Louisiana deer is long and

slender. The nasals are long and narrow; the zygomatic and

Fic. 1. — Skull of Og ca

gini 5 lis (Miller), ĝ , type No. 4999, Coll. E. A, and

O. Bangs, Mus. Comp. Zoól.

the palatal breadths are narrow, being much less than those of

the northern Virginia deer (Odocelus virginianus borealis).

In the type of O. Jeuzsiame the distal end of the nasal bone

touches the proximal end of the premaxillary, and the same is

true in the skull of No. 8623, 4, topotype. In this respect

there is a resemblance to Odoca/us virginianus osceola (Bangs)!

and Odocelus texanus (Mearns)? However, in No. 9112, $, topo-

type, an upward projection of the maxillary separates these two

1 Cariacus osceola Bangs, Proc. Biol. Soc. Wash., vol. x (February, 1896), pp- 25-28.

* Dorcelaphus texanus Mearns, tbid., vol. xii (Jan. 27, 1898), pp- 23-26.

452 THE AMERICAN NATURALIST. [ VoL. XXXV.

bones for a space of about 3 mm. ; while No. 8622, also ¢, topo-

type, resembles the northern Virginia deer in that this arm of

the maxillary separates the nasal and premaxillary bones by a

larger space, in this case 15.5 mm. The lower molar row is long,

being even a trifle longer than in O. borealis, though the jaw of

the latter slightly exceeds that of O. Jouzszame in length. In

all the specimens examined the ramus of the jaw shows a slight

concavity in ventral outline between the molar region and the

tip resembling in this respect O. osceo/a and differing from

RO DIRECTIS e E di adu

gini la (B gs), $, No. 2391, Coll. E. A. and

O. Bangs, Mus. Comp. Zoól.

Skull af O4.

Fic. 2.

O. borealis, in which the ventral outline of the jaw is quoe

nearly straight. Following are some measurements in milli-

meters of the skull of the type of O. Jouisiane, and by Way

of comparison the corresponding measurements of the type of

O. borealis (No. 4999, 8, Bangs collection), the race most resem-

bling it in size, are given in parenthesis after each. (Unfor-

tunately, the occipital region has been sawed off from the

Louisiana skulls by the collector, but there seems to be no

appreciable difference in length between them and northern

Virginia deer skulls.) Zygomatic breadth, 115 (139) ; least

inter-orbital breadth, 69 (80.5); greatest length of n

No. 414.] THE LOUISIANA DEER. 453

106 (95); greatest breadth of nasals, 19 (21.5); length of

premaxillary, 81 (86); greatest palatal breadth between upper

molar rows, 45.3 (55); extreme length of lower molar row,

84 (82).

Antlers: The antlers of the Louisiana deer are heavy, and

equal in size those of the northern Virginia deer, but as com-

pared with them they arise from the head at a sharper angle

Fic. 3. — Skull of Odocælus virginianus louisiana, 3. type No. 9111, Coll. E. A.

and O. Bangs, Mus. Comp.

and bend strongly inward, so that, although the length of beam

is practically the same in the type specimens of O. /ouisiane

and O. borealis, the antlers of the former are higher and at the

same time 80 mm. less in width across the beams than those of

the latter. In these high, narrow antlers there lies another

point of resemblance between the Louisiana deer and the little

Florida deer, The antlers of the type specimen of O. /ouisiane

measure 566 mm. from the burr to the tip of the beam, following

454 THE AMERICAN NATURALIST.

the inner curves of the latter. The circumference of the

antler at the base just above the burr is 119 mm. In addition

to the basal prong (that of the right antler being single and that

of the left bearing a small additional point on its anterior face),

there are on each beam three large tines and a small point on

the inner side of the beam between the first two tines. The

second tine is, on each antler, the longest. Both of these tines

possess an extra point on the anterior face near the tip, and

that of the left antler has also one on its outer face.’

Size: The specimens from Louisiana are unaccompanied by

measurements, so that the following taken from the dry skins

can only be considered approximate. The measurements of

the four specimens are given in millimeters in order thus, —

Nos. 91 11 type, 9112, 8622, 8623: length from nose to end of

tail (exclusive of terminal hairs), 2072, 1995, 1856, 1695 ; tail,

220, 253, 285, 240; hind foot, 490, 393, 375, 382; ear from

Crown, —, 122, 130.5, 134.5. No. 9i12 is a young three-point

buck captured Dec. 10, 1898 ; No. 8623 is a spike buck, and

with No. 8622, a one-point buck, was taken March 4, 1899.

Range: The lowlands of Louisiana, perhaps found through-

out most of the Austro-riparian zone.

Remarks : The Louisiana deer is more closely related to the

little Florida deer (Odocelus virginianus osceola) than to the

more northern races. Indeed, it is in general appearance but

a larger and slightly paler representative of that animal. To

the west of its range is another small deer, Odocalus texanus

(Mearns), but, as Dr. Mearns himself states, it is not to be

confounded with the Louisiana deer. From true Odocalus vir-

Sinianus (Bodd.), the large size and proportions of O. loutsian@

are alone sufficient to distinguish it, while from Odocelus vir-

gintanus borealis, which it equals in size, it is easily separated

by the cranial and color characters here given.

1 :

dis In the accompanying photographs the skulls were all placed at the same

tance from the lens of the camera without changing its position or focus.

ON THE OSTEOLOGY AND SYSTEMATIC

POSITION OF THE SCREAMERS

(PALAMEDEA: CHAUNA).

R. W. SHUFELDT.

ProFEssor William Kitchen Parker was the first to sus-

pect that those remarkable birds known as the screamers might

in some way be related to the Anseres, and as long ago as

1863 he undertook to point out in his excellent paper on the

subject in the Proceedings of the Zovlogical Society of London

(pp. 511-518) what the relationships between the two groups

consisted in. A few years later, however, Huxley included

them among his Chenomorphz (of. cit., 1867, pp. 436-460).

Naturally such authorities carried great weight for some time,

but when Garrod came to investigate the structure of Chauna

derbiana he quickly arrived at the conclusion that the Palame-

deidee were surely no geese, whatever the affinity might be

between the two groups (P. Z. S., 1876, pp. 189-200). Dr.

Sclater in 1880 created a distinct order for them (Palame-

deze), still keeping them, however, in the neighborhood of the

Anseres. Dr. Stejneger includes them with the Chenomor-

phe as a super-family (Anhimoidez), as I have elsewhere

pointed out, while Dr. Fiirbringer makes an intermediate sub-

order for them standing between the anserine birds and the

family JEpyornithida. Mr. Seebohm kept the flamingoes,

Anseres, and the Palamedez as suborders in the order Lamel-

lirostres ; and Sharpe places only the Anseres and the scream-

ers in an order Anseriformes. Professor Beddard (P. Z. S.,

1894) makes comparisons between Palamedea and Chauna, but

expresses no decided opinion there as to the systematic posi-

tion of the group.

Of all the accounts I have read of the osteology and tax-

onomy of the screamers, none come so near to what I consider

to be the truth in the case as does the above-cited contribution

455

THE AMERICAN NA

eese.

i Vias OPE om

best

EXPLANATION OF PLATE.

Right lateral view of the skeleton of the horned screamer (Palamedea cor nuta).

Greatly reduced. P hotograph of specimen No. 18,588 of the Ost ns

Collections of th

U. S. National Museum. (Collected in Venezuela.)

No. 414.] POSITION OF THE SCREAMERS. 457

of Garrod's in the P. Z. S. of 1876. His figures illustrating

his remarks are extremely instructive, comparing as he does

the occipital aspects of the skull of a Magellaine goose (Chæ-

phaga magellanica), a Derbian screamer (Chauna derbiana), and

a razor-billed curassow (Mitua tuberosa). He also gives the

basal view of the skull of Chauna derbiana. Garrod draws

some very careful comparisons among the several forms just

named, and it is well worth the pains of the student of avian

osteology and classification to read what he has to say, in the

paper cited, in connection with the present contribution.

With the view of throwing additional light upon the affini-

ties of the screamers, I have compared the skeleton of the

horned screamer, Palamedea cornuta (see plate, opposite page),

with that of the Coscoroba swan (Coscoroba coscoroba) and the

common wild turkey (Meleagris gallopavo) and arranged the

principal osteological characters characteristic of each form in

the subjoined table in a comparative way, thus supplementing

the results already achieved by Garrod and further emphasizing

some of the opinions arrived at by him in regard to the system-

atic position of the screamers.

SKULL.

General aspect.

Superior mandi-

SCREAMER.

SWAN.

TURKEY.

C pi ly 5

Characteristically

galline.

Typically anserine.

As in other An-

seres and quite

different.

Typically galline.

Agrees almost ex-

actly the

screamer: gal-

line.

area.

Lacrymal bone. Very small but | Very large, with | As in the fowls

anserine superior portion | generally, and

elongated. very different

Interorbital sep- | Entire. Entire. Perforated

tum.

Quadrates. Ansero-galline. Anserine. Galline

Auricular open- | Galline. Anserine. Galline

ing.

Lateral processes | Anserine. Anserine. Fuse distally, and

of the cranium. thus include a

foramen.

Palate. Desmognathous. | Desmognathous. Schizognathous.

—

458 THE AMERICAN NATURALIST. [Vor. XXXV.

SKULL. SCREAMER. SWAN. TURKEY.

Palatines and | Gallranserine, or | Anserine. Galline.

Vomer ith characters

nearly equa

divide

Facets of ptery | Median, as in | Anterior. Anterior.

goids. many pigeons.

Basi-temporal

area.

Mandible.

Hyoidean appa-

VERTEBRAL

CoLu

Cervicals.

[No epipleural

Pelvis.

Subtypically gal-

line.

Strongly galline,

angular proc-

esses approach-

ing the Anseres

in character.

Galline.

18. Pair of free

ribs on last

Six pairs vertebral

ribs; all without

wie ap-

a Two

pairs pelvic ribs ;

costal ribs of

last pair do not

reach the ster-

appendages on the

Narrow and long-

ish, but on the

whole with quite

dilated.

As in other swans

and geese

As in other typical

lamellirostral

birds, and very

ifferent from

Palamedea and

the fowls.

As in the Anseres

from Palamedea.

21. Two last bear-

ing free ribs,

with epipleural

appendages on

last pair

ioe pairs ver-

ebral ribs; a//

es —

Three pairs i-e

do not reach the

sternum

pelvic ribs in any

Narrow and long.

Propubic proc-

pubic elements

Hb dilated.

As in most true

fowls.

Characteristically

galline.

Galline.

I6. Asinthe swan,

but the last cer-

vical fused with

the first dorsal

vertebra.

Four pairs vertebral

ribs; all wi

epipleural ap-

pendages. One

pair pelvic ribs;

pair do not reach

the sternum.

of these birds.]

Shorter, and broad

in postacetabular

ion. Quite dif-

ferent from swan

not dilated.

459

TURKEY.

No. 414.] POSITION OF THE SCREAMERS.

VERTEBRAL

SCREAMER. SWAN.

COLUMN.

Shoulder-girdle. Anserinecharacter | Os furculum

— pre- U-sha and

ailing. Os fur- we anser-

dea Sai ine. the

ith the arch screamer, dis co-

very broad trans- coids do not

versely and come in contact

antero-posteri- when articulated

orly flattened ; in situ

articulation wit

coracoids and

scapulz asin the

geese. Cora-

coids more an-

serine than

galline.

Sternum. Anserine char- | Agrees withall the

acters prevail- typical anserine

ing ; but the pos- fowls among the

terior pair of wans, geese,

Viens viret and some of the

lower, ducks.

lateral Fide:

processes far

roader.

Limbs. Humerus onl

Highly pneumatic.

Proportions and

characters of

pectoral limb, as

and index digits.

pneumatic (?)

Bones of pectoral

limb long and

light, as in most

large anserine

birds.

Os furculum

V-shaped, and

the elements of

the shoulder

girdle all typically

galline.

Completely galline

in characte

very different

from the sternum

of Palamedea.

Humerus pneu-

matic; femur

partially so (?)

Bones of pectoral

limb are propor-

tionately as well

con-

siderably shorter

corresponding

ones in win

of Palamedea.

460 THE AMERICAN NATURALIST. [Vor. XXXV.

VERTEBRAL

SCREAMER. SWAN. TURKEY.

COLUMN.

Limbs. Pelvic limb anser- | Hypotarsus of | Hypotarsus very

ine with long tarso-metatarsus different from

odal digits. somewhat like that process in

Cnemial. proc- that of the Palamedea.

esses of tibio- screamer. :

the swans and

From an osteological point of view the screamers are

extremely puzzling forms. Had we only the skull of Pala-

medea cornuta to judge from, there is no question but that

avian taxonomers would have placed that species close to the

gallinaceous group of birds, its desmognathous palate notwith-

standing. We obtain but little light on the subject by the

comparisons made of its vertebral column and pelvis, though

in thelatter we meet with some characters that seem to suggest

an affinity with the storks.

Had we but its shoulder-girdle and sternum to judge from, it

is quite questionable in my mind that from these bones alone

its anserine affinities, or such as it may possess, would have been

suspected, and it is only when the characters of these are taken

in connection with others now known to us, that such a belief

gains support. We should never suspect Palamedea of having

any kinship with the Anseres were we to judge from the skel-

eton of its wing alone, though it is very likely that such a

relation would have been suspected had we only the bones of

the pelvic extremities to guide us.

Taken as a whole, the anserine characters of the skeleton of

Palamedea, however, are more evident than those of any other

group of the class, but these characters are neither typical nor

are they strongly marked. Such galline characters as are

exhibited in its skeleton are far more typical and most decidedly

more evident. But neither its anserine nor its galline osteo-

logical characters are sufficiently pronounced to justify us ":

arranging the screamers either along with the Anseres or the

No. 414.] POSITION OF THE SCREAMERS. 461

gallinaceous fowls. Such other characters as it may have in

its skeleton, derived from ancestral representatives of still other

groups, are now too completely masked and too obscure for us

to decipher, especially as all of the near relatives of these birds

have long ago died out, leaving no existing forms to assist us

in the matter. Palamedea, then, is doubtless the survivor of an

extremely ancient stock of birds, and were it positively known

that the gallinaceous types and the Anseres sprang from a

common stock, and it is not at all impossible, such birds as the

screamers may have easily appeared close down at the branch-

ing and then have ascended to the present time, showing but

few structural changes in their organizations.

Palamedea is a strong, vigorous, and eminently combative

type of fowl, and one not calculated, in time, to have been

much modified by its surroundings, and so has preserved in its

morphology the major share of its archaic structure.

I agree with Fürbringer that the screamers should be placed

near the Anseres, but apart and in an independent group,

standing between the latter and the ostrich types of birds.

The Palamedez, in fact, constitute a suborder of birds

(although represented but by three existing species) coequal

in the matter of distinctness with a number of others that

have been created in the class Aves, and, as a group, even

better defined and more distinct than not a few others. It is

fortunate for us indeed that in their anatomical structure there

is so much that enables us to recognize as clearly as we can

the place occupied in the system and in the class by these

birds and to predict with such certainty what their relation-

ships are with the other main groups of avian forms.

WASHINGTON, D.C.

NORMAL RESPIRATION AND INTRAMOLECULAR

RESPIRATION.

GEORGE JAMES PEIRCE.

“ RESPIRATION is essentially the intake of oxygen and the

output of carbon-dioxide by living cells. In the higher animals

two phases of respiration are distinguished — the erternal, the

exchange of gases between the air or water and the blood; and

the zzzerzal, the exchange between the blood, lymph, and the

tissues"! In plants there is, for the most part at least, only

the one phase, the exchange of gases between the air or water

and the cells composing the tissues, an exchange which is

direct and “external,” since it takes place in most cases

between the air, whether in the intercellular spaces within the

plant, or unconfined and outside the plant body, and the indi-

vidual cells. Even in the densest tissues, within which the

intercellular spaces are small, it is likely that the cells take in

free oxygen and give out carbon-dioxide, if not directly from

intercellular spaces, then from their neighbors bordering on inter-

cellular spaces. In any case, and in every stage of the process

of respiration except the purely mechanical ones, of which only

the higher animals are capable, the exchange of gases between

the cells and the air takes place in solutions, the oxygen enter-

ing and diffusing through, the carbon-dioxide passing out from,

the cells only when these gases are dissolved in water.

The object of respiration in plants is not the maintenance

of a certain body temperature, together with the production of

energy needed for doing work, as in warm-blooded animals.

It is merely the production of energy for doing work, as in

cold-blooded animals. The average body temperature of plants

I$, in general, nearly the mean daily temperature of their envi-

ronment. It will vary within certain limits, the variation being

i Pembry, M. S. Chemistry of Respiration, Schifer’s 7ext-Book of Physiology,

vol. i (1898), P. 692.

463

464 THE AMERICAN NATURALIST. (VoL: XXXV.

large or small according to the environment. Submersed

aquatics will vary least, floating aquatics more, and terrestrial

plants most in body temperature, other things being equal. But

as the temperature of small, still bodies of water (pools, etc.) may

vary considerably, so the body temperature of the organisms

living therein will vary, being warmed by the sun and cooling

during the night. The body temperature of the larger terres-

trial plants is likely to be higher at night (except on the exposed

surfaces), and lower in the day, than that of the surrounding

air. Owing to the great extent of their surface as compared

with their mass, radiation from the larger plants is rapid, and

a body temperature independent of their environment could

be maintained only at great expense of material laboriously

collected and elaborated. Plants work economically, are com-

pelled to do so, and this extravagance is avoided.

Heat is the form in which the energy set free by respiration

usually makes itself evident, but it does not necessarily follow

that only so much energy is liberated as is recognizable as heat,

or that this is the only form in which energy is liberated.

Only that energy becomes evident as such which is not at

once used. In order to determine the amount of energy

liberated in respiration, it is necessary to know and to measure

the material products of respiration.

The substances ordinarily taking part and produced in the

process of physiological oxidation are the highly complex

nitrogenous and non-nitrogenous compounds elaborated by the

organism and carbon-dioxide, water, and various small amounts

of several ‘other. substances. Of these last, oxalic acid is the

commonest and most important. Since the production of

energy and not of any particular compounds is what is striven

for in respiration, and since the substances acted upon by free

oxygen are different in different plants and even in different

cells of the same plants, the products differ accordingly.

Although the oxidation of nitrogenous matter also takes

place, it is mainly the non-nitrogenous contents of the living

cell which are involved in physiological oxidation. In the

animal body the oxidation of organic nitrogenous compounds

(proteids, etc.) results in the production of urea and of other

NO. 414.] NORMAL RESPIRATION. 465

similar substances no longer usable and presently cast off from

the body. In plants the elimination of these products is more

economically accomplished, for they furnish the foundations

for the re-synthesis of albuminous compounds. These waste

substances are removed by transforming them synthetically

into useful compounds.

The non-nitrogenous substances which become oxidized are

the fats and oils, the starches and sugars. The oxidation may

first convert the hydrocarbons into carbohydrates, with the

liberation of energy and the formation of by-products, carbo-

hydrates and by-products then becoming still further oxidized

with the liberation of still more energy. While respiration is

going on, the other functions also in operation may involve the

use, with chemical change, of some of each substance produced

in respiration, and the formation in the cell of other substances

not the products of respiration at all. It is therefore evident

that to ascertain the material products of respiration is hardly

less difficult than to determine the amount of energy liberated.

Since each process is normal only when accompanied by all

the processes going on at the same time, it is impossible to

isolate any physiological process for purposes of study. The

products of one set of chemical activities in the living body

may enter wholly or partially, simultaneously or successively,

into other chemical activities. The end products can be recog-

nized and measured with comparative ease, but to tell exactly

where or how they are formed is much more difficult and not

now entirely possible.

Water and carbon-dioxide gas are the chief products of the

Physiological, as also of all other forms of combustion of car-

bon-containing bodies. They are formed whenever a sufficient

amount of oxygen is united with the higher carbon compounds.

In organisms living under such conditions that the air can

penetrate to all their parts, enough oxygen will always be

present for such complete decomposition. The oxygen does not

unite of itself with the combustible compound, for even if active

(nascent) oxygen is present at all, which seems improbable,’

UT daa W. Pflanzen-Physiologie, zte Auflage, Bd. i (1897), p. 554- Pyst

ants, translated by Ewart, vol. i (1900), pp. 545, 546.

466 THE AMERICAN NATURALIST. [Vor. XXXV. .

it is present in amounts insufficient to accomplish the whole

result. The union is accomplished by and in the living cell;

whether with a more readily oxidizable substance first formed

from sugar, or with sugar itself, is not now known. All that is

known is that sugar, or some similar substance, and oxygen

unite, forming as end-products mainly carbon-dioxide and water.

The following reaction, without indicating what, if any, inter-

mediate stages there may be, shows the material results :

n ((CsH1206) + 6 O2 + Aq.) = n (6 CO, + 6 H20 + Aq.).

(Ag. representing the water in which the sugar is dis-

solved in the cell, does not enter the reaction. indicates

the unknown multiple of the minimum proportional formula

C6H1206, which stands for the sugar molecule. The 6 H20

produced in the course of combustion may unite with the

solvent water (Ag.), or may pass off as vapor, diffusing the

faster from the cell by reason of the heat liberated).

Since other substances than sugar are also oxidized physio-

logically in the cell, other products will be formed, the kinds

and the quantities of the latter varying according to the former.

The commoner of these minor products are oxalic, malic, and

citric acids, which accumulate in considerable quantities in

certain plants (e.g. in the leaves of Oxralis acetocella, in the

Crassulaceg, in apples, etc., and in the citrous fruits, lemons,

limes, oranges, etc. or are converted into salts (e.g., calcic

oxalate, crystallizing out of the solutions in which it is formed

in the cell), or undergo other changes (e.g., further oxidation).

In all organisms the oxidation of nitrogenous compounds, as

well as non-nitrogenous, occurs in normal respiration. The

proportional amounts of the two groups of compounds phys-

iologically oxidized vary with different organisms. In the

majority only organic and highly complex compounds are

made to yield the needed energy, but in some much simpler

inorganic compounds suffice, and in a few organisms already

known the carbon compounds are not used at all.

The nitro-bacteria, as first shown by Winogradsky,! oxidize

! Winogradsky, S. Recherches sur les organismes de la nitrification, An

nales de Inst. Pasteur, tomes iv, v (1889-91), and other papers.

No. 414.] NORMAL RESPIRATION. 467

simpler nitrogen compounds in order to liberate energy, employ-

ing carbon compounds only in the synthesis of food to be

used in the construction of their own bodies. One set of

nitro-bacteria oxidize ammonia, or compounds of ammonia, to

nitrous acid, the first and last steps of the process being thus

indicated :

2NH,0H T 302 = 2 HNO, + 4 H20.

Another set oxidize the nitrous acid, or its salts, to nitric acid,

thus :

2 HNO, + Os = 2 HNO,

The sulphur bacteria (Beggiatoa, Chromatium, etc.) obtain

most if nót all of their kinetic energy by oxidizing sulphur

compounds. They precipitate sulphur in their own bodies by

oxidizing the sulphureted hydrogen present in the water in

which they live.! If the supply of gas remain sufficient, the

sulphur will accumulate as a reserve supply in the cells ; if it

decrease, the reserve sulphur will be oxidized and, uniting with

water, will form sulphuric acid, or its salts, thus:

| 2 H,S + 0, 2 2 H0 +S

S +0, = $0;

2 SO: + O: = 2 $0,

SO; + H,O = H5;SO,

Those bacteria (z.g., Crenothrix) which, living in water rich

In iron, deposit iron in some form in or upon their own bodies,

may obtain their kinetic energy by physiologically oxidizing

ferrous compounds, presumably ferrous oxide, to ferric oxide.?

Other bacteria may be discovered which, needing carbon and

nitrogen compounds only to supply the constructive elements

of protoplasm, obtain their needed energy by oxidizing other

substances present in solution in the waters in which they live.

: Winogradsky, S. Ueber Schwefelbakterien, Botan. Zeitung, 1887. Beiträge

zur Morph. u. Physiol. der Bakterien. Leipzig, 1888. Miyoshi, M. Studien über

de Schwefelrasen-Bildung und die Schwefelbakterien der Thermen von Gumoto

bei Nikko, Journ. Coll. Sci. Imp. Univ. Tokyo, Bd. x, pt. ii, 1897.

* Winogradsky, S. Ueber Eiseribakterien, Botan. Zeitung, 1888. Molisch, H.

Pflanze in ihren Beziehungen zum Eisen. Jena, 1892. Miyoshi, M. Ueber '

massenhafte Vorkommen von Eisenbakterien in den Thermen von Ikao,

Journ. Coll. Sci. Imp. Univ. Tokyo, pt. ii, 1897.

468 THE AMERICAN NATURALIST. [VoL. XXXV.

The essential product of respiration, the one which distin-

guishes respiration from all the other functions of the living

organism, is kinetic energy. The material products vary in

kind and in quantity according to the nature of the organism

and the substances which can be affected, these substances

being in most cases complex compounds elaborated within the

body of the respiring plant, but not in all cases, as shown by

the bacteria just mentioned.

Nor is free oxygen necessary to all organisms or to all cells.

As the haemoglobin of the blood is a complex compound from

which some of the oxygen, only loosely held, can be readily

withdrawn where oxidation for the supply of energy is needed,

so the color products of certain bacteria (e.g., Bacillus bruneus)

are reserves of oxygen which become used when there is no

longer an adequate supply of free oxygen.! From colorless

compounds also, the cells at depths in the tissues of animals

(perhaps also of plants?), to which free oxygen penetrates only

in insufficient amounts if at all, obtain by decomposition the

energy needed. These decompositions are not necessarily

effected to secure oxygen for the oxidation of other sub-

stances, for the decompositions themselves release as kinetic

the potential energy which was holding the complex substances

together.

The mutual attraction of one atom of carbon and two of

oxygen is so great as to make the molecule of carbon-dioxide

very stable as well as very simple, for the affinities of the

carbon and oxygen are satisfied. In the complex compounds

of carbon, hydrogen, and oxygen in the sugar group, the affini-

ties of the component elements are not satisfied ; the com-

pounds are much less stable, as their ability to take up more

oxygen shows. At ordinary temperatures and under ordi-

nary conditions these compounds are stable. Their stability 1s

due to the mutual affinities of their component atoms which

exert an attraction upon one another sufficiently powerful to

! Ewart, A. J. On the Evolution of Oxygen from Colored Bacteria, Journ.

Linnean Soc., vol. xxxiii (1807), p. 123.

: Pfeffer, W. Berichte d. math. phys. Klasse d. K. Sáchs. Gesells. d. Wiss. ™

Leipzig, 27 Juli, 1896, p. 383.

No. 414.] NORMAL RESPIRATION. 469

hold them together in definite form. When the atoms are

rearranged more compactly in simpler forms in space, their

bonds or affinities are more completely satisfied, they unite

more perfectly, oxidation takes place in the rearrangement,

and energy is accordingly liberated and made available for other

purposes. Energy is stored in the starch or oil or sugar

molecule; the kinetic energy (solar or other) employed in the

construction of the molecule remains in it as potential energy,

holding the atoms together. The destruction of the molecule

results in the liberation of so much kinetic energy as was

employed in constructing it from the simple compounds worked

upon. i

The complete oxidation or combustion of a gram of dextrose

(sugar), resulting in the formation of carbon-dioxide and water,

as represented by this reaction,

C6H:206 ES 60, = 6 CO, a 6 H20,

liberates 3939 small or ordinary calories,! or mechanical units

of energy in the form of heat? For the sake of obtaining these

figures in more exact terms, for use in future comparisons, we

may multiply this with the molecular weight of dextrose, thus:

atomic weight of C — r2 oft, = 92

oHe s of Hy = 12

oro = ag OFC = 96

' CgH 120. = 180 = molecular weight of dextrose.

3939 calories x 180 = 709020 calories

= 709.02 Calories

The heat of combustion or complete oxidation of 1 gram-mole-

cule, Ze, of 180 grams, of aei isst is then 709.02 great

calories (C.).

This reaction, and the production of this amount of heat,

take place only in the presence of sufficient quantities of free

oxygen. Molecules more complex than those of carbon-dioxide

1 A calorie (c.) is the heat sid to raise 1 gram of water 1? C. in tempera-

ture; a great calorie (C.) is the heat required to raise 1000 gr. (1 Kilo) of water PG:

Rechenberg, C. von. Ueber die Verbrennungswärme d Ver-

ully give

470 THE AMERICAN NATURALIST. [Vor. XXXV.

and water, though simpler than sugar, may be formed from

sugar without free oxygen or with free oxygen in smaller

proportions than 6:1. Complete oxidation (normal respiration)

yields the largest amount of energy possible ; less profound

changes yield less energy. Thus the decomposition of sugar

by yeasts, according to the following reaction, which represents

only in simplest terms the nature of the chemical changes,

C&H ;206 = 2 C; Hg O + 2 CO,

forming without oxygen two molecules of alcohol and two of

carbon-dioxide from one molecule of sear yields only 67

calories per gram-molecule.!

The decomposition of one molecule of dextrose into one

molecule of butyric acid, two of carbon-dioxide, and two of

hydrogen, which is accomplished by a considerable number of

species of bacteria, and may be represented by this reaction,

C, H,,06 = C, Hy O, + 2 CO, + 2 H,,

yields about 75 calories per gram-molecule.?

Bacteria forming acetic acid, acting on dilute solutions of

ethyl alcohol in the presence of free oxygen, partially oxidize

the alcohol and decompose it into acetic acid and water, thus:

C,H,O + O, = Ci H, O; T H20,

liberating 125 calories;? but if the alcohol were completely

oxidized, as in ordinary combustion, the reaction would be

C,H4O = Y Cu L2 CO. + 3 H20,

and the heat liberated would be nearly three times as much,

about 325 calories per gram-molecule.

In these figures we have indices of the relative values of

complete and incomplete oxidations, and of oxidations and

decompositions, as sources of energy in the form of heat.

These figures are indices, to be trusted only so far as relative,

not exactly proportional, values are concerned. The chemist

can control all the conditions under which he makes a

1 Rechenberg, Zoc. cit., p. 66. 2 Rechenberg, /oc. cit.

i OF:

3 Quoted from Berthelot in Biedermann’s Chemiker-Kalender in 1897, p. 193

of the Beilage.

No. 414.] NORMAL RESPIRATION. 471

combustion in his laboratory and determines the number of heat

units liberated ; he can so regulate the process that there shall

be no by-products, and that no other compounds are included

in the reaction than those upon which he has determined to

experiment. In the plant, on the contrary, other substances

than dextrose may become oxidized, or the oxidation may be

incomplete. In the laboratory one can deal with measured

quantities of isolated substances; in the living organism indefi-

nitely known quantities of many substances together are acted

upon. Animal physiologists have done much more in this

direction than have plant physiologists. The higher animals

are better suited to such studies than are plants. The rela-

tively high body-temperatures of warm-blooded animals permit

direct temperature determinations from weighed quantities of

known foods eaten, as well as calculations from the amounts of

oxygen needed to effect combustions or decompositions. The

animal physiologist can check the results obtained by one

method with those reached through other methods. The

results obtained by animal physiologists indicate that only about

95% of the calculated yield of energy from oxidation! appears

as heat. So we must regard these figures as somewhat too

high, but their suggestive value is great, whatever must be

admitted as to their exact numerical value.

The larger organisms demand for the normal execution of their

functions more energy than can be supplied by the rearrange-

ment of the component atoms of the molecules always at

hand. They must oxidize these molecules, and the more com-

plete the oxidation, the greater the amount of energy liberated.

Some of the smaller organisms supply themselves with adequate

amounts of energy by the destruction of complex compounds

Within their own living cells. Probably some of the cells of all

large multicellular organisms have recourse, at times at least,

to the same means of securing needed energy, and when free

TEN is not obtainable, the majority of organisms can con-

at living for a time by so doing. From this the general

né nce may safely be drawn that the ability to obtain needed

SY by the destruction of complex substances in the cells

! See Pembry in Scháfer's Physiology, vol. i, pp. 836, 837.

472 THE AMERICAN NATURALIST. [Vor. XXXV.

is inherent in all organisms, that in the majority of organisms

and of their component cells this power is little needed and

hence is practically undeveloped; but that, owing to the posi-

tion of some cells deep in the tissues of many organisms and

to the peculiar habits of some of the lowest organisms, these

are obliged to obtain energy in this way and have developed

their inherent power to a high degree.

Intramolecular respiration is the name given to this mode of

respiration, a term not explicitly descriptive and therefore not

entirely satisfactory. The German term Spaltungsathmung is

in this regard more satisfactory, but it is not concisely translat-

able. Ordinary respiration is physiological oxidation or physi-

ological combustion or aérobic respiration. It is dependent

upon free oxygen and yields needed kinetic energy only by

the union of free oxygen with combustible substances. Intra-

molecular respiration is physiological simplification of complex

compounds or physiological rearrangement of atoms or anaé-

robic respiration. It takes place only when free oxygen is

present in small quantity or is altogether absent. The results

of the two processes are the same in kind — the liberation of

the kinetic energy needed to continue living — but, as the

figures quoted above show, not the same in degree.

Intramolecular respiration was first observed somewhat more

than a hundred years ago by Rollo,! but only within the last

few years has the connection between intramolecular and ordi-

nary respiration been clearly demonstrated. Pasteur and other

bacteriologists have contributed quite as much as animal and

plant physiologists to our present knowledge of respiration.

Pasteur and his followers have shown the peculiar habit of a

large number of microórganisms of being active only when

free oxygen is absent. When free oxygen is present, they

are inactive, though they may remain alive. There is a chain

of allied process: first, physiological respiration, or what may

be called intramolecular respiration, the normal respiration

of most organisms; second, physiological rearrangement 0

atoms into simpler molecules, intramolecular respiration, the

mode of respiration to which many cells and even organisms

1 Rollo. Annales de Chimie, t. 25, 1798.

No. 414.] NORMAL RESPIRATION. 473

have recourse under stress of circumstances ; third, physiologi-

cal rearrangement of atoms into simpler molecules, also intra-

molecular respiration, but the anaérobic normal respiration of

a comparatively small number of invariably low organisms.

From experiments hitherto conducted, it would seem that

the germinating seeds are better able to survive without a

copious supply of oxygen than are the other parts of higher

plants. This is what, a priori, might be expected, for the

embryo in the seed, when it becomes active in germination, is

a very vigorous organism, usually well supplied with just such

foods as may be readily broken down into simpler compounds.

The seeds of pea, for example, stimulated to germinate by being

soaked for twelve to fifteen hours in water at room tempera-

ture, will continue to respire actively for forty-eight hours or

more, even in a vacuum, producing carbon-dioxide in nearly the

same quantity as under the same conditions of temperature, etc.,

in ordinary air. Of course some air containing free oxygen

will be carried into the vacuum by the peas, but this will very

soon be entirely exhausted in normal respiration. ^ The con-,

tinued supply of energy must be obtained by intramolecular

respiration. Comparative investigations show that different

plants and different organs vary considerably in their ability

to substitute under stress intramolecular for normal respiration,

and that in very few of the higher plants is intramolecular

respiration, as measured by the yield in carbon-dioxide, so

effective as normal respiration.

For all higher plants prolonged intramolecular respiration

is impossible. To what this is due is not wholly clear. The

substances first attacked in intramolecular respiration are the

same as in normal respiration, Z.e., the sugars, starches (after

conversion into sugar), and the fats and oils. Later the

proteid substances enclosed in the cell, and finally the living

Substance itself, are decomposed to supply needed energy.

Whether the cessation of intramolecular respiration in experi-

ments upon higher plants, and the consequent death of the

Organism, are due to the destruction of part of the living

substance, or to the production in the cells of poisonous sub-

Stances, cannot now be determined. Certain it is that for

474 THE AMERICAN NATURALIST. (Vor. XXXV.

higher organisms intramolecular respiration is a function very

limited in importance, taking place only when there is continued

need of energy in the absence of free oxygen, and capable of being

maintained for comparatively brief periods only. Like normal

respiration, it is carried on solely by the living protoplasm, more

or less actively according to the greater or lesser activity of the

protoplasm. The substances decomposed are like those oxidized

in normal respiration and differ in different species of plants.

The products differ according to the plant, the conditions under

which it acts, and the substances acted upon.

Alcohol may be produced in considerable amount. This

suggeststhat in both fermentation and intramolecular respiration

(if one may separate the two processes, for the former certainly

includes the latter as well as nutrition) much of the chemical

work may be done by enzymes produced by the respiring

organism. Organic compounds and small quantities of many

others may also be formed. In germinating peas, the alcohol

produced may equal as much as 5% the weight of the moist

seeds, enough to give some support to the idea expressed above,

that the accumulation of the poisonous products of intra-

molecular respiration, as in fermentation, may cause the ces-

sation of respiration and the death of the organism.

Between those plants for which aérobic respiration is indis-

pensable to normally active life, and for which anaérobic respi-

ration is only a means of maintaining life over unfavorable

periods, and those for which anaérobic respiration is similarly

and equally indispensable, there are all connecting stages.

These are found amang the lower plants, especially the fungi ;

but, as before stated, in all large multicellular organisms, espe-

cially among animals, there are probably cells, lying deep in

the tissues, which are forced, by the positions they occupy, to

supply themselves with needed kinetic energy by the same

means as the anaérobic organisms, ż.e. by decomposing the

complex compounds which they contain. There are then cells,

as well as organisms, which are obligate aérobic, facultative

aérobic, or obligate anaérobic. Obligate anaérobic cells and

organisms live where the access of free oxygen is impossible or

difficult and inadequate; for example, deep in living tissues,

No. 414.] NORMAL RESPIRATION. 475

either as component parts of these tissues or as parasites or

saprophytes therein ; in the deeper layers of compact soils, in

the mud of swamps and marshes, and in the ooze below bodies

of comparatively still water, fresh and salt.

As in aérobic, so also in anaérobic respiration, other proc-

esses take place simultaneously with it. These, if not directly

caused by respiration, are at all events maintained by the

energy liberated in respiration and are so closely connected

with it that to distinguish between the chemical products of

respiration and those of the processes accompanying it, is a

matter extremely difficult and still only imperfectly accom-

plished. Fermentation, decay, and disease at least accompany,

if they are not actually a part of, the respiratory processes of

certain low plants. Respiration, anaérobic as well as aérobic,

is a function of the living protoplasm, which acts upon sub-

stances enclosed within its own body, producing simpler

substances of which some remain in the respiring cell while

others diffuse out of it. Some of the latter are chemically

inactive, like carbon-dioxide and alcohol; others may act on

substances outside the cell. In higher animals and plants the

enzymes (e.g., pepsin, diastase, etc.) are produced in connec-

tion with the process of nutrition, converting the substances

upon which they act into available food compounds; but it is

also certain that, among the enzymes produced, there are some

which bring about such changes in the surrounding substances

that these become available as sources of kinetic energy.

The diastase formed in the germinating seed, dissolving the

Starch deposited in the seed as a reserve food and converting it

into sugar, makes the reserve food available for at least three

purposes: first, for the construction of nitrogenous compounds

(amides and proteids); second, for the formation of cell-wall

(cellulose); 74777, for the liberation of energy by respiration.

The production and action of this enzyme furnishes material

for respiration, nutrition, and growth. The enzymes formed

by lower plants are also useful in more than one way, not the

least important use being the conversion of irrespirable into

respirable substances.

LELAND STANFORD JR. UNIVERSITY.

ON THE SWARMING AND VARIATION IN A

MYRIAPOD (FONTARIA VIRGINIENSIS).

ABRAM VARDIMAN MAUCK.!

Durinc August, 1898, a swarm of myriapods (Fontaria vir-

giniensis) made its appearance in the fields and woods south

of the Indiana University Biological Station at Vawter Park.

They became conspicuous objects in a roadway which they

were crossing, every square foot of which held one or more

individuals. Their migration was noticed during the early

hours of several days, the swarm going in a northerly direction

towards the lake, a little over 100 feet away. No further

attempt was made to trace the migration. A few days after

the appearance of the swarm all traces of it had vanished.

Thirteen hundred and nine individuals were preserved for

purposes of statistical study.

Other swarms of myriapods have been reported, and are com-

parable with the swarms of the Palolo worm during its breeding

season. Bollman, Bull. U. S. Nat. Mus., No. 46, p. 75, 1893,

records the migration and swarming of the same species noted

bythe author. He says: * At Donaldson, Arkansas, the adults

were found crawling on the ground in company with a large

number of their young, probably one adult to 500 or 800

young, then (July 11, 1887) about half grown."

Verhoeff (Zool. Anzeiger, Bd. XXIII, p. 465) gives a detailed

account of the migration of Brachyiulus unilineatus in such

numbers in the Hungarian district of Alföld that the railroad

track became so slippery, owing to the countless numbers

crushed by the wheels, that a train was not able to proceed,

though the rails were sanded.

The specimens seen at Vawter Park were all adults and all

about the same size. Of twenty females examined, thirteen

contained eggs, about 432 in number, the eggs measuring 0.5

! Contribution from the Zoilogical Laboratory of Indiana University, No. 42.

477

478 THE AMERICAN NATURALIST.

mm.in diameter. These facts would indicate that August was

the breeding time of Fontaria virginiensis, some of the females

having deposited their eggs, while others still retained theirs.

Of the total number collected, 622, or 47.517145, are males,

and 687, or 52.482875, are females. Over 100 each of the males

and females were studied with the view of determining the

occurrence of variations; first, in regard to the number of

the segments of the body; second, in regard to the number

and bilateral symmetry of the legs; third, in regard to the

number of segments in the antenne.

First, the number of segments was found to be twenty,

with no variation whatever from this number; second, the

pairs of legs were thirty in the male and thirty-one in the

female, with no variations; as far as the number of legs was

concerned there was perfect bilateral symmetry; third, the

antenna contain seven segments, and no variation from this

number was found.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XV. THE HOLOTHURIOIDEA.

HUBERT LYMAN CLARK.

THE holothurians form a sharply defined group of marine

invertebrates, especially abundant in the tropics but well repre-

sented in both temperate and polar seas. About six hundred

species are known, varying in size from ten millimeters to

almost a meter in length, and some of the largest species are

thirty or forty centimeters in circumference. The colors vary

greatly but generally are not brilliant, some shade of brown

or gray being the most frequent. Red, however, of various

shades is not rare, and green and yellow are also not infrequent.

Wholly black or white forms are not uncommon. The shape

is often worm-like but is more commonly broader and thicker,

often resembling a cucumber, so that the name “sea cucum-

bers ” is frequently given to the group. They are found at all

depths, from high-water mark to the bottom of abysses 2900

fathoms from the surface. They are generally sluggish in

their movements, and very few forms have any means of rapid

locomotion. They feed almost wholly on the organic matter

found in the mud or sand where they live, though a few species

found among rocks seem to feed upon organic matter growing

on the rocks or brought to them in the water. Some species,

especially the large ones, live on the bottom where it is muddy

or sandy, creeping about very slowly and apparently molested by

few enemies, Others, especially the footless forms, live buried

In the mud or sand, where they burrow by means of their ten-

tacles and muscular movements of the body wall. Still others

dg under or among rocks, often snugly stowed away in some

ole or crevice, where they seem to be permanently settled and

whence they can only be dislodged by breaking the rock.

479

490 THE AMERICAN NATURALIST. [Vor. XXXV.

Holothurians are easily distinguished from any other inver-

tebrates by the anterior and usually terminal mouth, with a

circumoral ring of tentacles, and their curious combination of

radial and bilateral symmetry. Although the bilateral sym-

metry seems usually to predominate, there are always five

prominent longitudinal muscles accompanied by nerve strands

and vessels of one or more sorts. The body wall is soft or

Fics. 1-4.

1. Trochostoma boreale (Sars). Nat. size. 3. Psolus chitonoides Clark. N

2. Cauds: ¥ (G ld) at. siz

at. size.

4. Psolus phantapus (Struss.). $ nat. size.

leathery, and generally contains calcareous particles in greater

or less abundance. These particles are usually microscopic

but are occasionally in the form of large plates several milli-

meters across. Their size, shape, and arrangement differ to

an extraordinary degree in the various genera and species, and

they form one of the most satisfactory means of identification.

In their simplest condition these calcareous deposits take the

form of very minute, more or less irregular grains known 4S

No. 414.] MORTH-AMERICAN INVERTEBRATES. 481

miliary granules, or somewhat larger straight or curved rods,

which are often widened and perforated at the end. These

rods are almost invariably present in the pedicels, papilla, and

tentacles, and are then known as supporting rods. In other

parts of the body wall they often are more or less irregularly

and profusely branched and then form rosettes (Fig. 26).

Sometimes the deposits are flat, thin, and wide, and are called

plates, which may be perforated with smooth-edged holes

(Fig. 19), or the holes may have teeth on the edges (Fig. 13).

If a perforated plate has a projection rising perpendicularly from

the middle, made up of several rods more or less joined together,

it is called a able (Figs. 15-17, 23-25); the perforated plate

being called the disk and the projection the spire. Sometimes

tables occur with the disk reduced or wholly wanting, and

sometimes the spire is greatly reduced. Very often the per-

forated plates are oval or elliptical in shape, with the perfora-

tions regularly arranged in pairs, and they are then called

buttons (Fig. 27); these may be smooth, knobbed, or spiny.

Quite often the calcareous particles assume the shape of some

familiar object, from which they receive names; as anchors

(Fig. 12), wheels (Fig. 14), cups (Figs. 20-22), etc. In a few

species these calcareous deposits seem to be wholly wanting ;

In some they are confined to special regions of the body ; in

many, two or more different kinds are found, usually in

Separate layers of the body wall; and in a few they are so

abundant, or so large and close together, as to make the body

wall stiff, and sometimes firm, or even rigid. Beside these

colorless calcareous deposits, we find in a few genera (Trocho-

stoma and its allies) reddish-brown or claret-colored concretions

(Fig. 18), which are often so abundant as to give their color to

the whole animal, or at least to large spots and patches. In

Such species the calcareous deposits themselves are often

iota tinged with the same coloring matter as the concretions,

un! they are free from it. In a few genera prominent

Bias ` bits of lime, usually five in. number, are grouped

e anus. These are called anal teeth.

— the cesophagus is a ring made up of plates of lime,

as the calcareous ring. There are usually ten of these

482 THE AMERICAN NATURALIST. [Vor. XXXV.

plates; five serve as points of attachment for the radial muscles

and are called radial pieces, while the five which alternate with

them are called zwzerradial. The size and shape of these

pieces differ greatly, the most important difference being

whether they have posterior prolongations (Fig. 7) or not

(Fig. 8). The tentacles around the mouth vary greatly in

number and appearance. There are usually ten or twenty, but

species are known which have normally eight, eleven, twelve,

fifteen, eighteen, and twenty-five, and in individual cases the

number runs up to thirty. In some species the number is very

constant, and individual diversity is quite rare ; while in other

species the amount of diversity is extraordinary, ranging in

some species of Holothuria which have normally twenty tenta-

cles, from twelve to twenty-nine. In appearance the tentacles

may be simple, finger-like, and unbranched, but this condition

is very uncommon. Usually each tentacle has several to many

branches ; when these all arise in a tuft from near or at the

tip and are more or less subdivided, the tentacle is called

peltate (Fig. 10); when the branches occur regularly along the

sides in two opposite series and without subdivisions, the ten-

tacle is pinnate, or, if the branches are very few and all arise

from the tip, digitate (Fig. 11); when the branches are sub-

divided and irregularly arranged the tentacle is said to be

dendroid (Fig. 9). In some species these tentacles are used

for locomotion, but they are more commonly used as feelers

and to carry food to the mouth. Locomotion is generally

accomplished by means of ambulacral- or tube-feet, as in the

other echinoderms. These feet, when present, may be

arranged in regular rows along the five radii (ambulacra), or

scattered more or less irregularly over the whole body surface.

Frequently those on the upper side of the animal are very

different in size, appearance, and structure from those on the

under or ventral side, and no longer serve for locomotion, the

terminal sucking disk being generally absent. They are then

called papille, the true-feet being pedicels. Rarely, all the

tube-feet are papilla and no pedicels normally occur. The

papilla are often situated on warts or tubercles, and these are

sometimes of considerable size. The pedicels are frequently

No.44] ANORTH-AMERICAN INVERTEBRATES. 483

confined to the ventral surface and are there often arranged in

three broad longitudinal series. Many holothurians have a

pair of long, much-branched outgrowths of the wall of the

cloaca lying in the body cavity, one on each side of the intes-

tine. These serve for excretion, and perhaps for respiration,

and are called the respiratory trees. Their presence or absence

is of importance in classification. The stomach and anterior

Fics. 5-11

4 A radial and interradial piece of the calcareous ring of Synafta inhærens (O. F. Müll). x 4s.

e same of Syzapta roseola (Verr.). x 45.

7. À radial and two interra dial pieces of a “calcareous ring with posterior prolongations.”

one briareus (Less. NS.

8. The same of a “ calcareous ring without posterior prolongations." Thyone suspecta Ludw. x 3.

9- A dendroid tentacle. Cuca ia punctata Ludw. x 3.

10. A > ss 3

Agent. hence.

Portion of the intestine are held in position by a thin membrane

attached to the body wall in the mid-dorsal line, and called the

orsal mesentery. In this lies the genital duct, and the geni-

tal glands lie in groups or tufts either on one or both sides

of it. In some holothurians the long radial muscles are divided

anteriorly, one-half continuing attached to the body wall and

Serving merely as a longitudinal muscle, while the other is free

484 THE AMERICAN NATURALIST. [Vor. XXXV.

from the body wall and is attached at the end to the radial

piece of the calcareous ring. By their contraction these five

muscles serve to draw the tentacles and whole anterior end of

the animal back into the body cavity for protection, and they

are therefore called the retractor muscles.

Owing to the fact that the early describers of holothurians

paid little attention to internal characters, and none whatever

to the microscopic deposits in the skin, it is difficult, if not

impossible, to determine from one of their descriptions what

species is meant. Even within the past forty or fifty years so

much importance has been attached to those characters in

which there is the greatest individual diversity (size, shape,

color, number of tentacles, arrangement of pedicels, etc.) that

the same species has been described under a number of dif-

ferent names. Many of our common species have two or three

synonyms, some have four or five, and one has no less than

eleven! As very few types of the species described by the

earlier writers are now in existence, the untangling of this

snarl of synonyms is no easy task. No less than eighty-eight

different specific names have been given to holothurians from

thé shallow waters of North America; but these seem to be

reducible to at most forty-seven valid species, and very probably

the correct number now known is under forty-five. Since

many of the names in most common use in this country are

incorrect, they will be looked for in vain in this key. In order

that such forms may be properly placed, the following list of

synonyms is given. It is not intended as a complete synonymy

of the species given in the key, but is simply an alphabetical

list of names incorrectly applied, or no longer in use by the

best authorities.

There are a number of such generic names, as follows :

Actinopyga Brown = Miilleria of the key.

Anaperus Troschel = Thyone "um

Botryodactyla Ayres = Cucumaria “ “ «“

Cladodactyla Brandt = Cucumaria “ “ +

Embolus Selenka = Trochostoma * * «+

Echinosoma Semper = Eupyrgus à uc

Leptosynapta Verrill = Synapta 64 d “

No.414.] MORTH-AMERICAN INVERTEBRATES.

Liosoma Brandt

Liosoma Stimpson

Lophothuria Verrill

Ocnus Forbes

Sclerodactyla Ayres

Semperta Lampert

Stereoderma Ayres

Synaptula Oerstedt

Thyonella Verrill

Trochinus Ayres

= Chiridota

= Trochostoma

= Psolus

= Cucumaria

= Thyone

= Cucumaria

Lyone

= Synapta

= Thyone

= Chiridota

of th

“ee

e key

485

Below are the specific names the use of which is no longer

correct :

affinis (Botryodactyla) Ayres

albicans (Synapta) Selenka

albida (Cladodactyla) Brandt

albida (Cucumaria) Ludwig

ayresii (Ocnus) Stimpson

ayresii (Synapta) Selenka

botellus (Holothuria) Selenka

musculosum (Thyonidium) Ayres

oolitica (Molpadia) Selenka

pallidus (Trochinus) Ayres

pauper (Embolus) Selenka

rtz

C C ladodactyla Brandt

Bentacta 3 Holothuria

na (Holothuria) Pourtales = Holothuria atra

= Cucumaria frondosa

= Synapta inherens

= Cucumaria populifer “

= Cucumaria populifer *

= Cucumaria minuta “

= Synapta inherens

= Holothuria impatiens “

= Myriotrochus rinkii *

= Thyone briareus

= Eupyrgus scaber

Psolus phantapus

= Thyone elongata

of the key.

partim Zrochostoma boreale of the

key.

= Chiridota levis

= Trochostoma ooliticum *

7 Sti

Pellucida (Sy napta)4 ieu] = Synapta inherens

partim Zvochostoma ooliticum of

the key

of the key.

O. F. } =Cucumaria frondosa “

= Cucumaria nigricans *

= Synapta vivipara

= Orcula barthii

= Cucumaria miniata “

Lii

486 THE AMERICAN NATURALIST. [Vor. XXXV.

fusiformis (Cucumaria) Desor = Thyone unisemita of the key

fusus (Thyone) Verrill = Thyone scabra RT

girardii (Synapta) Pourtales = Synapta inhaerens 6 4 €

glabrum (Thyonidium) Ayres = Thyone briareus uen

gracilis (Synapta) Selenka = Synapta inhaerens d xr

grandis (Botryodactyla) Ayres = Cucumaria frondosa. “ “ “

granulatus (Psolus) Ayres = Psolus regalis A d

sitchense (Liosoma) Brandt = Chiridota discolor Ww. 0

sitchensis (Cuvieria) Brandt = Psolus fabricii E 0€

tenella (Thyone) Selenka = Thyone briareus E ae

tenuis (Synapta) Ayres = Synapta inherens ww

tigillum (Chiridota) Selenka = Chiridota levis w Ww

turgidum (Molpadia) Verrill = Trochostoma boreale “« “ *

typicum (Chiridota) Selenka = Chiridota levis eue ee

viridis (Synapta) Pourtales = Synapta vivipara «c un

The following species are so very poorly described and so

little known it is impossible even to assign them as synonyms:

Holothuria tentaculata Forster, Synapta bachei Ayres.

The artificial key given on pp. 487, 488 is intended to include

and make possible the identification of every known species of

holothurian which occurs on the coasts of America, north of

Mexico, in less than 100 fathoms of water. There are a few

species which probably occur within these limits (as they have

been taken just outside), but they are not included in this key.

The holothurians of Florida and the Gulf coast and those of

the Pacific coast are almost unknown, and the geographical

range of even our most common species is very imperfectly

known. It has therefore been thought best to give as exactly

as possible the localities where the various species have been

taken. There are five species included in the key which are

new to science and the descriptions of which have not yet been

published; but descriptions and figures are now in press of

two of these forms, which were collected in Puget Sound in

1896 and 1897 by the zoólogists of Columbia University.

These are Psolus chitonoides Clark and Cucumaria lubrica

Clark. The third species is Cucumaria curata Cowles, also

from the Pacific coast. Mr. H. P. Cowles is now engaged in a

study of the life history of this species, and in advance of his

own publication he has very kindly given me the name he has

487

ARTIFICIAL. KEv TO THE GENERA OF NORTH-AMERICAN HOLOTHURIANS.

Calcareous deposits in skin, DUET of anchors and, plates

Sigs. 32 and P4) Se aa pun Synapta

Without respiratory trees | Calcareous deposits in skin, wheels with 6 a bs Hh col-

(Synaptidz) | lected in little heaps .. . Chiridota

Calcareous d cn in skin, scattered wheels with about 19

| spokes . Ps cos ls eda ue. v... V Mirobodhis

( Tentacles simple, unbranched; calcareous ring without pos-

Without pedicels or papilla 4 terior prolongafions o = GR e 4 1.5 ee. Eupyrgus

Caudal portion of body short,

With respiratory trees

(Molpadiidz) Tentacles digitate (Fig. 11) ;

calcareous ring with pos-

terior prolongations.

brown ellipses, or both . . . Trochostoma

Caudal portion of body long,

tapering (Fig. 2); calcareous

deposits, perforated disks or

tables with a narrow spire . . Caudina

NORTH-AMERICAN INVERTEBRATES.

No. 414.]

THE AMERICAN NATURALIST. [Vor. XXXV.

488

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No. 414] MORTH-AMERICAN INVERTEBRATES. 489

adopted and the diagnostic characters of this interesting little

holothurian. The remaining two species are very curious and

noteworthy ones from the coast of California, collected during

the summer of 1900 by Mr. Benjamin Thomas. They are

Thyone rubra Clark and Thyonepsolus nutrians Clark. Descrip-

tions of these two species are now in press. Each of these

five species bears at present only a nomen nudum, but it has

seemed best to include them, nevertheless, in order to make

the key as complete as possible.

KEY TO THE SPECIES UNDER EACH GENUS.

Synapta.

( Viviparous; mm. long; reddish-brown or

greenish, odi ud white; lives in seaweed

S. vivipara (Oerst.)

Radial pieces of cal- Biscayne Bay, Florida ; Bermuda, Jamaica; Brazil.

careous ring perfo-

SP N B pd larger, 250 mm.; white, yel-

rated for passage 4

of the nerve ish, or kn — lives buried in sand

(Fie. s) . . S. inherens (O. F. Müll)

Widely distributed in Northern Hemisphere; Maine to South

Carolina in Western Atlantic; Puget Sound, southward on

Pacific coast. Also Mediterranean, Eastern Atlantic, North Sea,

L and Arctic Ocean.

Radial pieces of calcareous ring, not iea but me notched

(Fig. 9; 1 100 mm.; rosy red. . . . à . roseola (Verr.)

Bay, Tete d idea Sound, and Bermuda.

Chiridota.

Wheel Papille very numerous, scattered over all the interambulac

besides wheels there are many small slightly curved rods UC sola (Pourt.)

Viviparous; roo mm. long; deep flesh color, wheel papille white; under stones and

loose rocks, and in sand; Key West, Biscayne Bay, Bermuda, Jamaica, and Brazi

Wheel papillæ in rows of 20-30 in each dorsal interambulacrum an

3-12 in each ventral one; North Atlantic coast . . . C. but (Fabr.)

Wheel papilla in only three interambulacra; Alaskan species C. discolor Esch.

two saar are practically PEAP and further study will probably

the i to be ide ccur under rocks, o comparatively shallow

Wa:

ter; may reach a soap m size; color very bner from gray to pink.

490 THE AMERICAN NATURALIST. [Vor. XXXV.

Myriotrochus.

Small, 50-60 mm. ; blackish or dusky; tentacles light . M. rinkii Steens.

Circumpolar; northeastern America as far south as Labrador. Occurs in water not over

300 fathoms deep.

Eupyrgus.

Very small, 10 mm.; grayish . E. Scaber Ltk.

Labrador, ohne Sp RR quo]: Kara Sea. At i addi of 4-240 fathoms, generally

on muddy

Trochostoma (Fig. 1).

Deposits wanting; ee smooth and leathery; body o

€ 100 mm. aSa or reddish-white with reddish-brow

atc T oan Stimp.

Coast of southern California.

With colorless calcareous tables, more or less

imperfect (Fig. 17), in addition to the ellipses

T. boreale (Sars)

An Arctic species, reported from the coast of Florida, Lesser

Antilles, and Maine. Verrill’s Molpadia itm probably

this species, was taken at various points on the lan

coast up to the Gulf of St. Lawrence, in 40-100 nied

Deposits, reddish-

brow

the animal a dark

red color; skin Without tables . . . . . . T. ooliticum (Pourt.)

grayish.

Boston, Block Island, Boon Island. Very possibly identical

L with "as Pom eding.

Caudina (Fig. 2).

Medium size, 50-100 mm.; about oce duet plea of the long,

C. arenata (Gould)

New England coast, Revere Beach, — Manan d, SEEN and Vineyard Sound.

Occurs buried in the sand in shallow wa

Psolus (Figs. 3 and 4).

Body subcylindrical; ( Dorsal

perisome without large plates; with

anal region elon-

numerous minute da plates and roun:

edly conical granules . . . . . P regalis Veral

caudiform; pedi- Banks of Newfoundland, in [x fathoms.

cels in three j

complete rows on Dorsal perisome with a mr scal

ventral surface imbricated P, Alenteon: "(Struss.)

i North Atlantic and Arctic oceans; on American coast as far

L south as Massachusetts ; occurs from low water to 130 fathoms.

No. 414.]

NORTH-AMERICAN INVERTEBRATES.

Fics. 12-18.

12. Anchor of Synafta scd (O. F. Müll). x 225.

13. Plate of the same species. x 22 .*

14. Wheel of Chiridota rotifera | (Pourt. J)e X235.

15 ig sade (Gould). Seen from above. X 105.

16. The seen from + & ros.

17. Table of t Tr ochostoma boreale (Sars). 105.

18. Reddish-brown deposits of chimie, iie boreale (Sars). X 105.

f Dorsal scales with fine granulations; sole with

Body depressed oval, lates ; 130

Scattered, irregular, reticulate plate:

chiton-shaped xe abr (D. & K.)

anal re- l

E us d North Atlantic; common on New England coast in shallow

Er ers Leer. Paaie Grove, Cal. (?)

pedicels forming

complete rows

color generally

Some shade of

Orange or red.

Size small, 20 mm. long by 8 wid

very — perforated -—

On along shore at Pacific Grove, Cal.

Dorsal scales with coarse granules; sole wi

reticulate cups or spheres P. peo in & K)

maller than preceding, but poesie identical with it; eastern

North votes from Massachusetts to Greenland; also on

Alaskan c

Dorsal scales, without evident granules; sole

with crowded k obbed, perforated plates

65m ET T : P. P diasin Clark

"iit "noe

T iere

; red; deposits, reticulated cups and

; th Mg os the bet

carries the TE T. nutrians Clark

492 THE AMERICAN NATURALIST. | [Vor. XXXV.

Pedicels scattered

ore or less on the

interambulacra. 4

also C. miniata

in the next section.

Pedicels present

Cucumaria.

( Size large, up to 600 mm. ; deposits, when present,

irregular smooth perforated plates C. frondosa (Gun.)

Brown; practically circumpolar; on Atlantic coast of America

)

south to Florida; on Pacific coast, to Lower California (?).

Size small, 40-50 mm.; Nae irregular

knobbed buttons, sometimes with four symmet-

rically placed holes, but more often wth several

scattered ones . ^ lubrica Clark

Pale brownish gray; n Sound.

Size very small 15-20 mm.; deposits rather

and, in the ventral wall, a few much larger

plates with much larger holes . . C. curata Cowles

Black; “eggs laid and cared for by mother; brooded under

ventral surface,” “ FAM Grove, Cal, and possibly in region

\ near San Francisco.’

( Deposits minute, reticulate cups and roundish or

oval buttons with four holes C. quinquesemita Sel.

Mendocino, Cal.. (Other specimens were labeled * Charleston,”

but this is probably a mistake.)

Deposits scattered irregular perforated plates,

with one end drawn out as a spire or dentate

process = = == =: . + C. albida Sel

Yellowish (white in ae with brown spots; 80 mm.

Mendocino and Pacific Grove, Cal.

Deposits polygonal PM with 2 Sofi: holes

a central sharp spin * jene (Stimp.)

130 mm.; Sitka and oh Sound.

Deposits an inner layer of narrow oblong plates

lacra, and there - with two or three rows of holes and an outer

or two rows

layer of tables . . . . . . C. caleigera (Stimp.)

20 mm.; Massachusetts, Labrador, Greenland, and Arctic

Ocean. Pacific Grove, Cal.

Deposits irregular spinous rods or spicules

C. nigricans (Brandt)

Dusky yellowish-brown ; 60 mm.; Sitka and Puget Sound.

Deposits a very crowded inner layer of hollow

fenestrated ellipses and an outer Py of small

reticulate cups or spheres . . Lbs Théel

White; 7o mm.; Vancouver Island and Puget Sound. Pacific

Grove, Cal.

Deposits roundish perforated plates . C. minuta (Fabr.)

- Clear brownish; 10 mm.; Grand Manan to Arctic Ocean.

No. 414.) MORTH-AMERICAN INVERTEBRATES. 493

Pedicels present

generally in

or two rows.

í Deposits “in numerous etie €

layers, their true shape not fully c

C. k (Ayres)

Eastern United States from Vineyard Sound south to South

Carolina, This species needs a reéxaminatio

Deposits crowded, thick, knobbed buttons or

plates with few or many perforations, and one

end drawn out into a thorny spine or proces

miniata s (Brandi)

Pinkish-white ; 40 mm.; Alaska and Pacific Grove, CaL (n.

C. albida Sel, but there are two species at Pacific Grove, Cal.,

one of which is apparently albida, and the other is clearly dis-

tinct from it, but answers to the description of miniata. There

are pedicels, however, on all of the interambulacra in the speci-

| mens from Pacific Grove

Fics. 19-27.

19. Knobbed plate of Thyone suspecta Ludw. X 450.

20.

ies; seen from T X 450.

21. The same; seen A € —

22. The

23. Table

same:

n fro 450-

of Holothuria atra a Jaeger? seen from the side. X 450.

25. The top of the sam

so.

26. Branched rods and peek of same species. X 450.

27. Buttons of Holothuria captiva Ludw. X 450.

494 THE AMERICAN NATURALIST. [Vor. XXXV.

Thyone.

Deposits in body wall wanting . . . . . . . . . . TZ briareus (Less.)

Dark brown, almost black; 150 mm. ; in mud, Cape Cod to Texas.

Deposits in the form of tables . . eo. LD. seabra Verr

Brownish; 5o mm.; coast of New Kp to i Clasp Bay.

Deposits small reticulate cups and uneven oval buttons with four holes

T. gemmata (Pourt.)

Body spindle-shaped, dusky brown ; North Carolina to Brazil (?).

Deposits unperforated rods . . . . T. elongata (Ayres)

Banks of Newfoundland, in anes fathoms; Massachusetts Bay (?).

Deposits crowded oval buttons with four holes . . . . Z. unisemita (Stimp.)

Brownish-gray ; 60-75 mm.; coast of New England from Nantucket northward.

Deposits buttons with four holes, and very numerous ~ complex

tables T. rubra Clark

Red above, whitish below; 20 mm. ; viviparous; Pacific Grove, Cal.

Orcula.

Size medium, 65 mm.; brownish ; deposits wanting . . . O. darthii Troschel

Labrador, Greenland, and the Arctic Ocean.

Thyonidium.

Size small, 15 mm.; pale yellowish; deposits few . . . 7: productum (Ayres)

Eastport and Grand Manan.

Miilleria.

Tentacles 20; deposits tables and buttons . . . . . . . M. parvula Sel.

Chestnut-brown; 5o mm.; Florida

Tentacles 2 TES tables and buttons wanting; yes small branched

rods M. agassizii Sel.

Light brown mottled with darker; 300 mm.; Florida and West Indies.

Stichopus.

Tables with round perforated disk; no buttons . . . . . S. dadionotus Sel.

Olive green or olive brown; 200 mm. ; Florida

Tables without disk; no buttons. . . 20. S. sitchensis (Brandt)

9o mm.; Alaska. The status of this species is doubtful.

Tables with well- rds disks; numerous buttons with nine pairs ©:

holes . S. cali Veri oe

Brown; 300 mm.; coast of ries: Pet Sound.

No. 414.] MORTH-AMERICAN INVERTEBRATES. 495

Holothuria.

Deposits tables and very numerous small irreg-

ular perforated plates . . . H. atra Jäger

Dark brown, 300 mm.; shanty: distributed in the tropics ;

Papille on dorsal, Finds

pedicels on ven- -

tral surface. Deposits tables and two kinds of ait mm

buttons H. rigida Sel.

Olive Mis 7o mm.; Florida "e Zanzibar and Society

|. Islands).

Papillz on both dor- ( Disk of tables with a smooth margin

sal and ventral H. impatiens (Forsk.)

surfaces; no pedi- Dark purplish or grayish, more or less blotched with darker ;

cels; apes j 150 mm.; very widely distributed in the warmer seas; Flori:

kind of perforated Disk of tables with twelve prominent horizontal

button teeth on margin o. TI E Prines Bel

L Dark brown; 120 mm.; Florida.

THE MOST IMPORTANT PAPERS DEALING WITH THE

SHALLOW-WATER HOLOTHURIANS OF NORTH

AMERICA ARE AS FOLLOWS.

AYRES, W. O. Notices of Holothurie. Proc. Bost. Soc. Nat. Hist.

Vol. iv. 1851-54

BRANDT, J. F. Prodrosds descriptionis animalium ab H. Mertensio

observatorum. Fasc. i .

CLARK, H. L. The S nMptes of the New England Coast. Bull. U..S.

Fish Comm. 18

Duncan, P. M., and MOORS W. P. A Memoir on the Echinoderms of

the Arctic Sea to the West of Greenland. 1881

GEROULD, J. H. Anatomy and Histology of Caudina arenata. Proc.

Bost. Soc. Nat. Hist. Vol. xxvii. 18

KINGSLEY, J. S. Contributions to the Anatomy ‘of Holothurians. Mem.

Peabody Acad. OF O0. Vol L No. s.

Lampert, K. Die Seewalzen. 1885.

Lupwic, H. Revision der Mertens-Brandt'schen Holothurien. Zeitschr.

SJ. wiss. Zool. Bd. xxxv. 1881.

PouRTALEs, L. F. On the Holothurie of the Atlantic Coast of the

United States. Proc. Am. Ass. Adv. Sci. 5th meeting. 1851.

SELENKA, F. -Beitráge zur Anatomie und Systematik der Holothurien.

Zeitschr. f. wiss. Zool. Vol. xvii. 1867.

496 THE AMERICAN NATURALIST.

SEMPER, C. Die Holothurien. 1868.

Stimpson, W. (a) Descriptions of New Species of Holothurians. Proc.

Bost. Soc. Nat. Hist. Vol. iv.

(4) Synopsis of the Marine Invertebrates of Grand

nan. 1853.

shores of North America. Bost. Journ. of

Nat. Hist. Vol. vi. 1857.

THEEL, H. Report on the Holothurioidea. Challenger Reports. Vol.

xiv, pt. xxxix. 1886.

VERRILL, A. E. (a) Report on the Invertebrate Animals of Vineyard

= ound. Report of the Comm. of Fish and

Fisheries. 1874.

(^) Preliminary Check-List of Marine Invertebrates of

e Atlantic Coast from Cape Cod to the Gulf

of St. Lawrence. 1879.

Recent Papers on Algz, N cr

IX. Publications Received . * . * Ld r . e se . *

BOSTON, U.S.A.

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AMERICAN NATURALIST

S WD XXXV. August, Igor. No. 416.

THE TEXAN KCENENIA.

AUGUSTA RUCKER.

Tuis most interesting microthelyphonid was discovered last

May in the neighborhood of Shoal Creek, near Austin, Texas,

under stones in company with Campodea, Japyx, and Scolopen-

drella. A short description of it was given by Dr. W. M.

Wheeler in the November Naturalist, under the title ** A Singu-

lar Arachnid (Kenenia mirabilis Grassi) occurring in Texas.”

The writer believed the Texas species to be identical with the

Sicilian one figured and described by Drs. Hansen and Sorensen,

and therefore wrote of it under that name. A number of speci-

mens have since been examined zz fofo and in sections, and

some interesting results have been obtained. It did not seem

Probable that the Texan Koenenia had been imported, yet it

Seemed less probable, though our climate is much like that of

Southern Europe, that the two species were identical. Close

‘Microscopic examination has shown them to be two distinct

. Species of the family Kæneniidæ.

Kenenia has since been found in Siam and Paraguay and,

like all archaic types, it will doubtless be found to be cosmopolitan

1 Contributions Jrom the Zovlogical Laboratory of the University of TS a S

615 ; | ie anne

616 THE AMERICAN NATURALIST. [VoL. XXXV.

in its distribution. A young Danish zoólogist has recently

. found in Siam a distinct species of Koenenia which Dr. Hansen

is to describe. Dr. Silvestri, the discoverer of the species from

Paraguay (K. grassii), has promised me a few of his specimens

in return for the Texan species, which I hoped to have for com-

parison before this paper was finished. I have likewise been

unable to collect any K. parvula,! of which species Dr. Wheeler

has found a single specimen, which he has briefly described

in his paper (p. 233). When descriptions of these species are

published a more correct idea can be framed of the valuable

taxonomic characters of the hitherto unknown order.

Koenenia wheeleri n. sp.

I desire now to name our principal Texan form after its

discoverer, Dr. Wheeler, and to give along with its internal

structure a short description of the characteristics which set

it apart as a new and distinct species.

In the beginning I may say we have been more fortunate

than Drs. Hansen and Sorensen in being able to distinguish

the two sexes. It hardly seems possible that the males of

Grassi’s species could be so rare when they are so abundant in

our species. In fact, in the material collected in the fall, the

males predominated. Very few females were to be had then,

and those few were so small and insignificant that it was

thought they were the males, The criterion taken for dis

tinguishing the sexes in this material was the opaque glisten

ing body in the second abdominal segment of the female, ag

receptaculum seminis of Hansen. Unfortunately the seminal

vesicles of the male are situated in the same region and have

much the same appearance; hence Dr. Wheeler, in his inter

esting paper, was misled into thinking that the male was F

the other sex. It was not until fresh material was collected

this spring and sections made that the mistake was discovered:

In size and general form the sexes are alike, and it was pei

after examining sections that one could say for the first we

that the animal possessing the more complex reproduce”

appendages is the male.

1 I have the pleasure also of giving-this species its name

No. 416.] .THE TEXAN KG NEMNIA. 617

External Anatomy.

On comparing K. wheeleri with the figures and descriptions

of Drs. Hansen and Sorensen for X. mirabilis the following

differences are apparent: (1) the number and position of the

hairs of the body; (2) the appendages of the reproductive

orifice situated between the second and third segments of the

abdomen; (3) the three pairs of prominent orifices on the

ventral surface of segments four, five, and six, through: which

the lung sacs are everted; (4) the lateral

sense organs of the cephalothorax ; (5) the s ^ zi oN

number of teeth on the two last joints of the /- à

chelicerze. EONS

The hairs over the ventral surface of the ü a 9n (UMP và

abdomen of the Texan species are more

numerous and are distributed in a manner palio m

quite different from those over the abdomen

of the old-world form. In the second seg-

ment of the female there are three rows of me

plumulose seta. The first row of six runs |

across the ventral surface anterior to the e y

: i : : ^ Mini

middle line and is continuous with the hairs (ub.

of the dorsal surface. These form a belt j

around the second segment. The second i

r 4 . Fic. 1. — Ventral surface

row of six arises along the base of the tri- of the abdomen of male

angular flap of the reproductive orifice. The

third row is so irregular that it can scarcely be called a row.

Here the hairs follow in a fashion the other two sides of the

triangular appendage. Fig. 2 is a camera drawing of this

portion of a specimen which has been treated with potassium

hydrate. The figure clearly shows the arrangement of hairs

over this region of the female. The dotted lines of the figure

represent the portion of the organ which can be seen only by

focusing through the triangular appendage.

In segment two of the male there are three primary rows of

plumulose hairs. The first row, consisting of from four to six

small setze, runs across the ventral surface at about the middle

of the segment and continues dorsally to form the circular row

618 THE AMERICAN NATURALIST. [Vor. XXXV.

of setz. This second row of four small setae extends across

the middle of the shield-shaped appendage of the reproductive

organs. The third set consists of two secondary groups, the

first of which, containing eight hairs, runs across a sort of hem

attached to the lower margin of the shield-shaped organ. The

second set is made up of large plumulose spines arising from

the tips of ten papilla, four of which spring from the edge and

six from under the edge of the hem. Fig. 3 is a careful camera

drawing of this region of the male. The first row of setze not

on the reproductive appendage is not figured. ;

Segment three in both male and female has on the two blades

of the reproductive appendages irregularly placed hairs which

are of varying lengths. In a line posterior to the middle of the ~

segment are eight other setze in a row continuing with those of

the dorsal surface to form the setigerous belt of segment three.

Most remarkable are the differences between segments four,

five, and six of X. wheeleri and the corresponding segments of

K. mirabilis. Concerning the fourth segment of Grassi's species,

Drs. Hansen and Sorensen have written, * Provided on the ven-

tral side in front of the middle with a wart-like protuberance

which bears six stiff setze, almost spines (acu/ez), arranged in

two transverse rows, the foremost of which is arched and has

four spines. Grassi seems to consider these spines sen

organs, an opinion we by no means share.” In K. wheeleri

segment four is provided on its ventral side with three pairs of

hair groups. The first pair consists of three long backward-

pointing setæ on each side of and near the midventral line.

These setze are situated on delicate chitinous flaps of the integu-

ment which arise in front of the middle of the segment. T

second pair is made up of four much shorter backward-curvins

hairs arranged in an arched transverse row near the middle "

the segment to the right and left, respectively, of the first pal

These hairs are most evidently for the protection of the ai

cate lung sacs when they are ejected, and for protection 0

their orifices when they are drawn in. The third pair conse”

of two small setae in the same line and lateral to those - yt

second group. The former are continuous with the belt o

sete encircling the whole segment. |

No. 416.] THE TEXAN K(ENENIA. 619

Of the fifth segment nothing is said concerning K. mirabilis,

while the figures show it to be like segment seven. This is

decidedly not the case in our species. The fifth segment is —

as far as the setze are concerned — a facsimile of the fourth.

Again, concerning the sixth segment of the Sicilian species,

the Danish investigators write, “It is furnished on its ventral

side with a rather strongly protruding wart, bearing a somewhat

arched transverse row of six forward-curving setae. We do not

think these to be sensory organs either." In K. wheeleri the

ventral surface of the sixth segment is in reality less prominent

than that of the two preceding segments, for it is lacking in

the first group of large hairs situated on the flaps on each side

of the midventral line; the flaps, however, are retained. Fig. 1

is a ventral view of the abdominal exoskeleton of a male

Keenenia, which has been thoroughly cleaned of its cellular

contents in KOH. In this specimen all six of the sacs are

thrown out, and the protective function of the groups of four

hairs is quite evident. All the setae shown in this figure are

plumulose; but under a magnification of about 120 diameters

they barely appear to be so, and I did not attempt to represent

this condition in the drawing.

It was only after repeated attempts that I succeeded in obtain-

ing clear views of the reproductive appendages. In both sexes,

segments two and three are con-

spicuous for their relatively enormous

appendages. In the female the mid-

ventral surface of the second segment

projects downward and backward into

a triangular appendage, which is not

emarginate at its apex, as in K. mira-

bilis. This projection almost covers Ven o Bo caer wel —

a couple of heavily chitinized down- sac in the center of Sern

ward projections of the third segment. ei

The receptaculum seminis opens between these two appendages,

while the reproductive organs open further forward into the

vagina formed by the three appendages. Fig. 2 shows what

I take to be the receptaculum seminis, while the reproductive

orifice only shows in section. The male appendages are very

pi yu

620 THE AMERICAN NATURALIST. (Vor. XXXV.

complex and difficult to explain. Fig. 3 will aid in making the

account clear. The appendage of the second segment repre-

sents in surface view a truncated conical flap having at its base

a hem which is notched in the midventralline. The hem near

the outer sides of the flap projects into two papilla, each of

which terminates in a heavy spine.

Projecting from the under surface

of the hem are two pairs of large

and one pair of small papillze, with

their corresponding spines. This

papillate appendage partially con-

ceals two trowel-shaped, strongly

chitinized, downward and backward

directed projections of the third

segment. The accessory glands

| and vasa deferentia open near to-

Frc.3.—Reproduciveappendages gether, in the median line, where

of male K. wheeleri.

the posterior surface of the un-

paired appendage is continuous with the anterior surfaces

of the paired appendages.

Another important specific difference seems to be in respect

to the lung sacs, organs which are evidently much more promr

nent in the Texan species. These organs must, in fact, be

entirely lacking in K. mirabilis, for such careful observers as

Drs. Hansen and Sorensen could not have entirely overlooked

them. On examination of a few of our specimens, one cannot

fail to observe peculiar little sacs projecting from the ventral

surface sometimes of the fourth, sometimes of the fifth, some

times of the sixth, and occasionally from all three segments.

Again a specimen may be found in which all six sacs are inva

nated, giving it the appearance of possessing three pairs of

stigmatic apertures. Of these organs Dr. Wheeler writes: " In

many specimens a delicate sac may be found evaginated from

under a flap on all three segments. These sacs are in all prob-

ability lung books. They appear to be the only pease

organs of Koenenia apart from the delicate integument, whic

` in so. small an animal must of itself nearly suffice for pP

tory purposes. If I am correct in regarding the above-descri

No. 416.] THE TEXAN KG@NENIA. 621

sacs as lung books, they must represent those organs in an

extremely simple form, in a form, moreover, which strongly sug-

gests their origin from invaginated appendages serially homol-

ogous with those of the cephalic and thoracic segments."

The two anterior sensory organs do not appear different in

any respect from the same organs of K. mirabilis. The lateral

organs, howevet, though situated in about the same place,

consist of three sensory rods each, instead of the two blades.

These are short-pointed rods pressed close siti

together and projecting, when at rest, for-

wards and outward. Fig. 4 is a camera

drawing of these lateral sensory hairs, under

a high magnification. In cross-section the

hairs appear as three rings in contact with Fıs. 4 — Lateral sense

one another. I am unable to make out on me

the surface of these organs anything more than the minute

projections which are found over the entire surface of the

animal's body.

The second and third joints of the chelicere, which form

pinchers, in specimens examined for this special purpose, were

found to be each provided with eight teeth. The teeth of the

fixed portion are long and very acute, with barbs at the base ;

while the teeth of the movable joint are short, broad, and blunt.

The above are the most evident differences of the two species

of Kcenenia, unless it be that there are more segments in the

caudal flagellum of one than of the other. I have examined a

number of complete specimens of both males and females, and

find in every case that the tail is made up of fifteen segments.

Grassi states for his species, which has been redescribed for all

other points except this, that it possesses thirteen or fourteen

joints.

Internal Anatomy.

In considering the internal anatomy of this minute animal,

several difficulties have arisen which I fear I have not entirely

surmounted. The extreme minuteness of the cells of the very

delicate tissue enclosed in the comparatively heavy chitinous

case makes microscopic study rather unsatisfactory.

622 THE AMERICAN NATURALIST. (VoL. XXXV.

Integument. — Drs. Hansen and Sorensen state, “As a

peculiarity in Koenenia, we think right to emphasize at once

that its skin is but slightly chitinized, especially on the abdo-

men, where, consequently, there is no distinction between the

(dorsal and ventral) plates and the pleura; so the expansion,

which the abdomen must be capable of allowing, probably

depends on the elasticity of this thin chitin itself." I think

had these gentlemen attempted to section Koenenia they would

not have been so emphatic about the thinness of its exoskele-

ton. The chitinous cuticle, which rests on a delicate hypo-

dermis, corresponds more nearly to that of most spiders, in

that over the expanded abdomen it does not appear to form

special plates. It also corresponds to spiders, in that it is a

flexible or accordion-plaited covering, the folds of which run

parallel with the long axis of the body. This arrangement

thus allows of great expansion of the abdomen, its function

being evidently the same as the folds in the late integument

of the abdomen of Thelyphonus. Over the chelicerze labrum-

hypostome and reproductive appendages the chitin is thick

and yellow, while between the joints it is very thin. In the

floor of the mouth the chitin is thrown into folds, running at

right angles to the long axis of the body. This, in sagittal

sections of the animal, gives it the appearance of possessing

teeth. The entire chitinous surface of the animal is not

smooth, but under high magnification appears to be covered

with small dot-like elevations.

Kenenia wheeleri is remarkable for the comparatively thick

covering of hairs arising from its flexible cuticle. The smallest

hairs are like down, covering the anterior surface of the labrum

and the under surface of the hypostome. The longest and most

delicate are the tactile hairs of the sixth, seventh, and eighth

segments of the third pair of appendages. The broadest and

heaviest spines are situated on the underside of the proxim

joint of the chelicerz ; otherwise the setze are distributed as

Dr. Wheeler has already shown. All the hairs over the body

of Koenenia, with the exception of the tactile and the vor

minute ones on the mouth appendages which are too delicate

to be made out, are microscopically plumulose.

No. 416.] THE TEXAN KQGENENIA. 623

The muscles of Koenenia are decidedly striated, like those

of insects. They represent a condition of musculature which

would be expected in so small and primitive an animal. Worthy

of note are the two pairs of simple dorsal and ventral muscles

of the abdomen and thorax. Other important muscles are those

running from the roof and side of the thorax to the chelicere.

The muscles of the appendages need not be described, with

the exception of those extending from the side of the thorax to

be inserted on the thoracic appendages. These muscles arise

on the sides of the thorax, opposite to their corresponding legs,

and, crossing over just above the suboesophageal ganglion,

become inserted on their proximal joints. The only other

muscles that need to be mentioned aré the primitive dorso-

ventral muscles of the thorax and abdomen. These are

decidedly a very striking feature in the abdomens of

arachnids.

Nervous System. — One of the most singular things about

Koenenia is its large proportion of concentrated nervous sub-

stance. The concentration of the ganglia is almost equal to

that of the Araneidz. In this respect Koenenia is even more

specialized than Thelyphonus. There seems to be no reduc-

tion of nervous element here due to the absence of eyes, but,

like all primitive types, it retains its cephalothoracic ganglia

unmodified. The brain and subcesophageal ganglion unite to

form one large mass perforated by the small cesophagus. The

brain is enormous, occupying the entire dorsal portion of

the head above the level of the cesophagus. It innervates

the median and lateral sense organs, the labrum, and cheli-

cere, The subcesophageal ganglion covers the entire floor of

the head and thorax, and shows in section swellings corre-

sponding to each of the five pairs of appendages. With a

slight constriction at the waist, the subcesophageal ganglion

connects with a single abdominal ganglion which is situated

in segments two and three, dorsal to the reproductive orifice.

From this ganglion, nerves run to all parts of the abdomen and

tail. In its nervous system Koenenia is thus very unlike the

Scorpionidea and differs from Thelyphonus in the relative size

of the cephalothoracic ganglion and in the situation of the

THE AMERICAN NATURALIST. [Vor. XXXV.

624

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No. 416.] THE TEXAN KG NENIA. 625

abdominal ganglion, which has not been drawn up toward the

head in the latter, but remains in the eighth segment.

Digestive System. — The downward-curved, crescentic mouth

leads into a strongly chitin-lined pharynx. This in turn runs

into a very delicate cesophagus which penetrates the cepha-

lothoracic nerve mass, only to dilate immediately into a

pouch-like sucking stomach. This stomach is roofed over

by the brain, while underneath it the subcesophageal ganglion

extends. At the sides the brain is not continuous with the

suboesophageal ganglion, thus leaving a passageway for mus-

cles arising from the sides of the cephalothorax to enter and

attach themselves to the stomach. When expanded to its

utmost the stomach fits snugly in between the two ganglia, but

when pulled on by the muscles it is flattened dorso-ventrally.

The comparatively thick-walled stomach opens through a valve-

like constriction into the exceedingly thin-walled intestine.

The intestine, before it leaves the thorax, gives off a pair of

small diverticula. It then passes into the abdomen, becoming

much dilated, and giving off five shallow metameric pairs of

diverticula, from the third to the seventh segments inclusive.

These diverticula are very diagrammatically represented in

Figs. 5 and 6, as are all the other organs, save the brain

and the anterior portion of the digestive tract as far back as

to the first pair of diverticula. At about the eighth segment

the thick-walled large intestine begins. Unlike Thelyphonus,

no Malpighian tubules are present, opening into the hind gut,

before it terminates at the anus. On this point Kcenenia is

most primitive, since it seems not yet to have reached the

stage in which intestinal diverticula become modified as excre-

tory organs. There are also no salivary glands present ; these

would hardly be of any use to an animal living under such

simple conditions. The intestine and diverticula are invari-

ably filled with food particles, which have the appearance of

yolk granules. Strange to say, — because of the conditions

under which Koenenia is found, — throughout the entire digest-

ive tract no dirt ever appears. This goes to prove that the

food is probably derived, as Dr. Wheeler has already suggested,

from the eggs of animals with which it associates. The

626 THE AMERICAN NATURALIST. [Vor. XXXV.

digestive tract is thus admirably constructed for such an

illegitimate practice as egg-sucking.

Excretory System. — I have succeeded in tracing the pair of

tubular glands, **tappezzata d' un semplice strato di cellule epi-

teliali," of which Grassi speaks. According to him, these

glands extend through a large part of the cephalothorax, and

perhaps have their orifice in front of the third pair of limbs.

These excretory organs in reality arise in the second segment

of the abdomen, and after forming one or two convolutions run

into and straight through the thorax, to terminate between the

second and third pair of appendages. There being no Mal-

pighian tubules in the small animal, this simple pair of coxal

glands would seem to represent the only excretory organs,

unless, indeed, the glandular cells around the respiratory sacs

can be considered as possessing excretory functions. If this

be the case, the eversible sacs will then have a double func-

tion of respiration and excretion like the vertebrate allantois.

These cells are not represented in the drawings.

Respiratory Organs. — Respiration in so small an animal as

Koenenia must necessarily be very simple, and, if I have rightly

interpreted the facts, we have in this minute Palpigrade the

most primitive form of respiratory organs. These organs con

sist of the three pairs of lung sacs which are situated in seg-

ments four, five, and six, with their corresponding orifices on

the ventral surface. They are evidently evaginated through

the internal blood pressure. For each pair of sacs there 1$

a pair of dorso-ventral muscles, corresponding to the dors

ventral muscles of Thelyphonus, which have the function in

Keenenia of drawing in the everted sac appendages. These

lung sacs possess on their inner surface (inner when they are

evaginated) granular bodies which stain a deep blue with alco-

holic carmen if they happen to be invaginated, but which take

on a normal red stain when the sacs are thrown out. Often,

in examining sections through the inverted sacs, one can i

refrain from calling them tracheze, so very much do they loo

like simple tubes. In truth, according to whether the sacs are

pulled in by muscles, remaining contracted, or whether they

have been pulled in by muscles that have immediately become

No. 416.] THE TEXAN KÆNENIA. 627

relaxed, allowing the sac to flatten dorso-ventrally and wrinkle,

do we obtain diminutive trachez or simple lung books. After

examining a great number of sections one cannot refrain from

believing that simple sac trachez like those of Koenenia may

have given rise to both lung books and trachez in other arach-

nids. If this be the case, we may hold that Koenenia, which

possesses the simplest phase of these organs, the lung trachez

(which are in reality abdominal appendages belonging to distinct

body segments), is the most primitive of all Arachnoidea.

Circulatory System. — As to the circulatory system, the

simplest condition possible exists. A definite heart has not

yet made its appearance. The blood can have no regular

course through the lacunae and sinuses, and it probably makes

its exchange of gases in the neighborhood of the lung sacs.

There must be some definite region for the interchange of

oxygen and carbon dioxide, for though Koenenia is small, its

exoskeleton is rather too thick to allow of a general surface

respiration. I do not think that the dots over the entire sur-

face of the chitin can be minute pores, which the spiders alone

of all the arachnids possess, over the skin of the abdomen.

Reproductive System. — In the female the unpaired ovary

begins as a blind tube in the seventh or eighth segment and

extends into the third. From each side near the anterior end

it is prolonged into two oviducts, consisting, for the greater

part of their length, of large glandular cells — the largest cells,

in fact, of the body. These ducts run forward and upward,

becoming very small and thin-walled ; probably the last por-

tion, for a short distance, being chitin-lined, as in Galeodes.

In the second abdominal segment they become very much

swollen, forming a sort of pouch on each side, filled with a

gelatinous secretion, evidently derived from the gland cells

of the oviduct. The duct continuing from each pouch or

vesicle runs backward and downward to meet at the place of

entrance in the vagina. It is this portion of the reproductive

organ, the vitelline vesicles and their terminal ducts, of which

Hansen and Sorensen write: “In the second abdominal seg-

ment there is an organ which shows the same peculiar luster

and refraction of light which one of us knows so well for the

THE AMERICAN NATURALIST. [Vor. XXXV.

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No. 416.] THE TEXAN KQNENIA. 62

r3 9

receptaculum seminis of small crustacea." In all of my sec-

tions I have never been able to see any sign of spermatozoa in

the vesicle, which is always full of a non-granular, gelatinous

secretion. Just posterior to the outlet of the oviducts, and

between the two lateral appendages of this region, is the

outlet of the small flask-shaped receptaculum seminis. The

accessory glands are small and insignificant, and in the lateral

anterior portion of the abdomen they empty into the vagina.

The ovary, which fills almost the entire lower portion of

the abdomen, does not show its: primitive paired condition.

Although the muscular walls of the pouch-like ovary are

bulged out with what appear to be cells, only a few of these

become the eggs, while the remaining seem to be the nurse

cells, and are consumed by the growing ova, which early in the

spring lie in the upper portion of the organ. Later in the

season a few of the eggs fill the entire ovary, while most of

the small bodies have disappeared, and in their place a few oil

drops remain. These oil drops are seen at this stage in the

lumen of the oviducts. Fig. 5 is, for the most part, a diagram-

matic sagittal section through a female Konenia. The ovary,

however, is an exact camera drawing which shows the propor-

tion between the true eggs and the food bodies, and their

corresponding position in the ovary of an animal taken early

in the spring. In Koenenia the egg is evidently not fertilized

until it reaches the vagina, and all the food that it receives

before its fertilization is a product of the ovary and oviduct.

The latter must also necessarily furnish the membrane of the

egg. Just as there is a pair of dorso-ventral muscles for each

pair of lung sacs, there is also a corresponding pair for the

reproductive appendages of each segment. These, together

with special muscles, undoubtedly cause à slight protrusion

and retraction of these organs. :

In the male the primitive paired condition of the generative

organs is retained. The testes consist of two equally swollen

tubes, beginning in the seventh or eighth segment and extend-

ing along the floor of the abdomen, to be continued anteriorly

into the vas deferens of each side. The vasa deferentia are

very much coiled, and fill — at certain times of the year — all

630 THE AMERICAN NATURALIST.

the space in segments two and three of the abdomen, before

descending on each side to swell and form the seminal vesicles.

From these vesicles the ducts run slightly up and back to a

common opening situated between the points of attachment of

the two pairs of the dorso-ventral muscles of these appendages.

The accessory glands, which are larger in the male than in the

female, appear to open into the atrium at a point just posterior

to the orifice of the vasa deferentia. The exact courses of the

vasa deferentia and the accessory glands are extremely difficult

to follow. In places the walls of the ducts entirely disappear, .

leaving only as a guide, in the case of the vasa deferentia,

the contained spermatophores. In some specimens the whole

anterior portion of the abdomen is packed with spermatophores.

Fig. 6 is a diagrammatic drawing of a longitudinal section

through a male Koenenia, taken to one side of the sagittal

plane. Only the testis and vas deferens, with the accompany-

ing accessory glands of one side, are shown. There are, evi-

dently, delicate muscles in the walls of the testes. In the

posterior ends of the testes are numerous cells which are

undoubtedly sperm mother-cells, while the anterior portion

appears to be crowded with small dotted packets. These

dots, which must necessarily be the spermatozoa, glisten under

transmitted light and rarely show a stain, even with iron-

haematoxylin. Nowhere throughout the whole course of these

organs is there any trace of spermatozoa possessing flagella.

It must be that, in Koenenia, the condition is retained which

is found in most crustacea, which possess non-motal sperma-

tozoa often imbedded in gelatinous spermatophores. It 15

probably due to this spermatophore secretion that the sperm

cells almost entirely refuse to take on the stain.

UNIVERSITY oF TEXas, AUSTIN, TEXAS,

May 23, 1901.

THE MAMMALS COLLECTED IN SAN MIGUEL

ISLAND, PANAMA, BY W. W. BROWN, JR.

OUTRAM BANGS.

IN the spring of 1900 Mr. W. W. Brown, Jr., made a short

collecting trip to San Miguel Island, the largest of the islands

in the Bay of Panama, known as the Archipelago de las Perlas.

Here he stayed for nearly a month, — middle of April to middle

of May, — and in this time secured, he believes, representa-

tives of all the species of land mammals that occur on the

island, with the possible exception of some bats.

San Miguel Island is about fifteen miles in length and lies

twenty miles distant from the nearest point on the mainland,

and sixty miles from Panama. It is composed of low-lying hills

clothed in heavy tropical forest, hot, damp, and unhealthy.

The collection of birds made on this trip has already been

reported upon.! The birds of the island are for the most

part like those of the adjacent mainland, and five species only

were considered different enough to receive distinctive names,

Most of the mammals, on the other hand, seem to be well dif-

ferentiated island species. I am much indebted to Gerrit S.

Miller, Jr., for identifying the bats for me, and to Dr. J. A.

Allen for comparing the large opossum with the various forms

he has lately described from different parts of northern South

America.

One hundred and fifty-seven mammals, belonging to twelve

Species, were collected by Mr. Brown in San Miguel Island,

as follows

Marmosa Sulviventer nob. : ‘ 4 specimens

Didelphis karkinophaga cauce An X

l Bangs, Outram. Birds of San Miguel Island, Panama, Awé, vol. xviii

(January, 1901), No. remi

angs, Outram. A New Honey Creeper from San Miguel Island, Panama,

Proc. New Eng. Zoil. Club, vol. ii = 8, 1901), pp. 51, 52-

631

632 THE AMERICAN NATURALIST. [VoL. XXXV.

Lepus incitatus nob. ; i ; i ; I specimen

Dasyprocta callida nob. . i ; > 6 specimens

Loncheres labilis nob ; 3 I

Proechimys burrus nob. . ‘ : à ; 51 «

Zygodontomys seorsus nob. : i : ; 68 ks

Mus musculus Linn. à : i A : I Specimen

Mus rattus rattus Linn. . : A

Mus rattus alexandrinus (Geoft.)

Vampyrops helleri Peters i

Hemiderma brevicaudum (Wied.)

Marmosa fulviventer sp. nov.

Four specimens, taken between April 28 and May 8.

Type: No. 8435, adult 2, Bangs Collection, Museum of

Comparative Zoólogy, collected April 28, 1900.

Characters: Nearest to the form of the Marmosa murina

series from the Santa Marta region of Colombia, — M. mitis

Bangs. Differing in being smaller; ear smaller; fur shorter,

harsher, less silky; color, above more fulvous, less olivaceous,

brown; under parts wholly fulvous, darkest on the sides (in

AM. mitis the under parts are yellowish-white) ; skull similar.

Color: The usual black facial ‘markings, rest of upper parts

between cinnamon and tawny-ochraceous; upper surface of

arms, sides of neck, and sides, much brighter, more -ochra-

ceous-rufous ; under parts strong buff, shading into ochraceous-

buff on lower sides and on under surfaces of arms and legs,

the fur unicolor except on lower sides, where it is pale gray

basally ; tail dusky, paler below than above; feet and hands

dull grayish-white.

MEASUREMENTS.

Bec

EAR FROM

ToTaL AI no

NUMBER. SEx. TRES VERTEBRÆ.| FooT.

8435 | d adult type 330 175

8436 | Q9 “ topotype 325 ie

8437 | “cc i 340 17 5

8438 | ĝ old adult Pere head & body, 170

No. 416.] MAMMALS IN SAN MIGUEL ISLAND. 633

Skull, type, adult 4, basal length, 34.4 ; occipito-nasal length,

37.4; zygomatic width, 20.4; least interorbital width, 6.2;

length of nasals, 17.6; width of nasals, 5; length of palate,

18.8; upper tooth row, from front of canine to back of last

molar, 15.2; length of single half of mandible, 27.8.

Didelphis karkinophaga cauce Allen.

Two specimens, 4 and 9, were taken May 8. These have

been compared by Dr. J. A. Allen with extensive material

from South and Central America ; and it is Dr. Allen's opinion

that they are best referred to this form, though they do not

represent it in its extremes.

Lepus (Tapeti) incitatus! sp. nov.

Type (and only specimen): No. 8441, old adult ?, Bangs

Collection, Museum of Comparative Zoólogy. Collected April

30, 1900.

Characters: An island form of Lepus gabbi Allen. Distin-

guished by its larger size, paler, more reddish-brown coloring,

and larger, heavier skull, with much heavier, more arched

rostrum.

Color: Upper parts tawny-ferruginous ; brightest on top of

head, palest — shading to dull ochraceous — on sides ; middle

of back and top of head varied by the admixture of black-

tipped hairs; nuchal patch, arms, and outer surface of legs

clear, bright, tawny-ferruginous ; superciliary stripe dull buffy-

white ; outer surface of ear colored like back, with short, dusky

tip and narrow, yellowish-white outer border, inside of ear very

sparsely haired — the hairs colorless ; under parts soiled white,

except underside of neck, which is Isabella-color, shading into

dull, tawny-ferruginous on sides of neck.

Skull: Compared with skulls of true Z. gabbi from Panama

and Chiriqui, the skull of Z. incitatus is larger and heavier

throughout ; rostrum much wider and heavier, and more

rounded and arched; bony palate wider and longer; molar and

incisor teeth heavier.

1 Jncitatus ; swiftly running; rapid, quick.

634 THE AMERICAN NATURALIST. [Vor. XXXV.

Measurements: Type, old adult 9, total length, 420; tail

vertebra, 20 ; hind foot (with claw), 80; ear from notch, 45.!

A. Type of Lepus incitatus, adult 9.

Lepus gabbi from Loma del Leon, Panama, adult e .

occipito-nasal

h, 23.8;

width of

Skull, type, old adult 9, basal length, 57;

length, 74.6; zygomatic width, 35.4 ; mastoid widt

least interorbital width, 16.2 ; length of nasals, 30.4 5 W™

nasals, 13 ; length of palatal bridge (from incisive foramina to

1 Specimens of true Z. gabbi taken by Mr. Brown at Loma del Leon, Panama

and Divala, Chiriqui, measured as follows :

| EaR

: | Tar Hinp Foot

NuMBER. | SHE. | LocaLrTY. iin Varrusam.| (CLAW) | SS

| ENGTH. ut. Ms

; | 38

8424 9 | Loma del Leon 340 18 r1

ts fbn Qoo be 345 20 ie

10065 3 | Divala 370 20 77 | `

en $1 E pr i d rs | $

I 0

10067 9 3 370 18 | 78 :

I er

No. 416.] MAMMALS IN SAN MIGUEL ISLAND. 635

palatal notch), 9 ; length of incisive foramina, 17 ; width of

incisive foramina, 6.4 ; upper tooth row, alveola, 14.6; length

of single half of mandible, 56 ; lower tooth row, alveola, 15.4.

Remarks. — The hare was not at all common in San Miguel

Island, and Mr. Brown saw but one other during his stay.

Mr. Brown tells me that Lepus gabbi and L. incitatus are ex-

traordinarily swift of foot and. are seldom seen except for an

instant as they dart like a flash through the undergrowth.

The form found in San Miguel Island differs from true Lepus

gabbi of the Isthmus in its larger size and heavier, slightly dif-

ferent skull. The color is also slightly different, the island

race being less marked along the back with black-tipped hairs,

and the general coloring is decidedly redder.

Dasyprocta callida! sp. nov.

Six specimens, April and May.

Type: No. 8443, adult 2, Bangs Collection, Museum of Com-

parative Zoólogy. Collected May 8, 1900.

Characters: A well-marked island species of the variegata-

isthmica group. Color pale yellowish ; rump hairs black, with

white tips. Skull very slender, with long, light rostrum and

narrow nasals. Molar-form teeth small.

From D. isthmica, the new species can be told by its white-

tipped rump hairs (yellow-tipped in isthmica), and from D. co/om-

biana (the form of this group inhabiting the Santa Marta

region of Colombia), which also has white-tipped rump hairs,

by much paler and more yellowish coloration. From either it

is easily known by its very different skull (see figure on next

page).

. From the Central American D. punctata, D. callida and its

allies differ chiefly in the character of the hairs of the rump.

In D. punctata these are not so elongated and are annulated

and colored like the hairs of the rest of the upper parts.

Color: Hairs of upper parts annulated with yellowish and

black, giving a slightly variegated look, though the yellowish

much predominates, and the general color is yellowish clay-

color, brighter, more ochraceous in middle of back anterior to

! Callidus : that is, taught wisdom by experience; shrewd, cunning.

636 THE AMERICAN NATURALIST. (VoL. XXXV.

rump ; rump hairs elongate, black with white tips ; under parts

soiled white, the hairs annulated with drab ; feet and hands

brownish-black.

The type and No. 8447 are in fresh pelage and are similar.

No. 8445, adult 2, is rather darker and more nearly approaches

D. colombiana in color. The three other specimens, of which

A. Dasyfrocta isthmica from Divala, Chiriqui, adult 9.

B. Type of Dasyprocta callida, adult á.

two are nursing females, are in worn, shabby pelage, in which

the black of the rump has faded to dull brown and the w white

tips of the hairs are worn down and discolored. In No. 8446

new, normally colored rump hairs are growing in again among

the old faded ones.

No. 416.] MAMMALS IN SAN MIGUEL ISLAND. 637

Skull: The skull of D. callida is very distinctive, and can

at once be told from that of any of its allies by its slenderness,

its long, narrow rostrum and long, narrow nasals. The molar-

form teeth are very small.

MEASUREMENTS.

Mn ula Sex DI 'ToTAL TAIL Hinp EAR FROM

: i y LENGTH. | VERTEBRÆ.| Foor, Norcu.

8443, type | ĝ adult | May 8 480 25 102 38

8444, topotype | QM April 22 460 22 100 38

, e: | e BELA 465 22 102 37

8446. “ |,g ^ “ 21| 510 3o 105 38

$45, Y | ii i 26 485 20 97 35

Bo. .* | d dnd Uc og 420 20 96 33

NOTE. — All external measurements are the collector's. Hind foot is measured

with the claw.

Skull, basal length, 85.4 ; occipito-nasal length, 98.6 ; zygo-

matic width, 44 ; mastoid width, 32 ; least interorbital width,

26.2; length of nasals, 38 ; width of nasals, 15.4; length of

palate, to palatal notch, 39 ; to end of pterygoid, 55.4; upper

tooth row (with four molar-form teeth in place), 17.2 ; greatest

width of rostrum, 24 ; length of single half of mandible, 58 ;

lower tooth row (with four molar-form teeth in place), 18.8.

Remarks. — The six specimens were all shot by Mr. Brown

among mangroves, the leaves of which they are very fond of.

The animal is much hunted by the negro pearl divers, and is

exceedingly shy and wary, and for some time Mr. Brown was

unable to secure one. One day during a storm he noticed

that when a mangrove blew over it was at once stripped of its

leaves by the agoutis. Acting upon a plan that this habit of

the animal suggested to him, he took several large stones with

him, and concealed himself in a tree. After a little he sent a

stone crashing through the mangroves and presently saw an

agouti cautiously approach the spot, thinking a mangrove

had fallen over. The first day he shot two specimens in this

way, and afterwards four more.

638 IHE AMERICAN NATURALIST. [VoL. XXXV.

Loncheres labilis! sp. nov.

Nineteen specimens, April and May.

Type: No. 8480, old adult ¢, Bangs Collection, Museum

of Comparative Zoólogy. Collected April 26, 1900.

Characters: Most nearly related to L. caniceps Günther of

Medellin, Colombia. Pelage long and stiff, but without spines.

Judged by description alone, the new species is brighter and

redder in color and its head is not gray. The skull compared

with Giinther’s figures (P. Z. S., 1876, p. 746) is more

slender, the palate and post-palatal regions are narrower, and

in profile the skull of the island animal is much flatter, the

supraorbital ridge not rising in a curve but lying flat and straight.

Type of Z A labilis, adult é (three views).

Color and Pelage: Pelage moderately long and harsh, but

wholly without spines. Whiskers very long, black, as also a

patch of whisker-like hairs behind the eye and others in front

of and on edge of ear. Top of head, nose, and cheeks black,

the hairs somewhat annulated with yellowish, giving à pepper

and-salt appearance ; yellowish-white patches at base of whs

kers, above eye, and behind ear ; rest of upper parts’ bright

ferruginous, varied by black-tipped hairs along middle of back,

varying in amount in different individuals, — some specimens

1 Labilis, gliding, slipping.

No. 416.) MAMMALS IN SAN MIGUEL ISLAND. 639

being wholly of an intense ferruginous color from nape to tail,

others having the rump only ferruginous, the rest of back

being darkened by a copious sprinkling of black-tipped hairs ;

chin dull grayish-white ; rest of under parts varying from

strong buff to clear ferruginous, some examples having both

colors irregularly distributed in patches; upper surfaces of

hands and feet yellowish-brown ; nails white; tail colored like

the body for a short distance, then black, sometimes yellowish-

white at tip, thickly clothed with stiff hairs, which, however,

do not hide the scales (except at base, where the tail is colored

like the body) ; ears blackish, small and low, nearly naked except

for the long black whisker-like hairs that surmount their edges.

Skull: Compared with the figures of that of Z. caniceps,

the skull of Z. /abilis is much more slender, with narrower

palatal and post-palatal regions; viewed in profile it is much

flatter, with the supraorbital ridge lying flat and straight and

not rising in a curve.

MEASUREMENTS.

Yun maw: | Sex Dirk |. Torar | "TArL HiND EAR FROM

| s | LENGTH. ratte Foor. NoTcnu.

| i

8480, type | & adult | April 26 | 540 | 240 47 15

8468, topotype gf “ ‘ 540 220 48 15

8471, * 5 ue “ 26 540 220 47 15

8470, * pa 4 “26 535 2 48 16

8469, “ Le ^ " 26| $30 210 48 15

MIL ^4 D? « e 500 240 45 15

8474 * | IC * 23| 470 220 47 15

8472, oe | g s "* 198 470 230 45 16

8475 * | d young; “ 2 440 215 45 14

MS. o. | 2 o! May 4 420 175 42 15

8476 “ | i * sg 420 195 42 14

Skull, type, adult 4, basal length, 47.8; occipito-nasal length,

56.6; zygomatic width, 27.4; mastoid width, 22.4 ; least inter-

orbital width, 12; length of nasals, 16; width of nasals, 7.2;

length of palate, to palatal notch, 21.2; to end of pterygoid,

36.4; upper tooth row, 13; length of single half of mandible,

34 ; lower tooth row, 13.2.!

! Some skulls that appear to be of about the same age are smaller, others larger.

640 THE AMERICAN NATURALIST. (Vor. XXXV.

Remarks. — This island species of Loncheres belongs to a

group quite different from either true Loncheres or Isothrix.

The skull is, in important characters, similar to that of true

Loncheres, but the pelage is very different in wholly lacking

spines. Isothrix has still softer pelage and more hairy tail,

and also a different skull. I hope some mammalogist, with

more material at his command than I now have, will before

long readjust and properly divide the different groups of this

series of “ spiny rats."

Loncheres labilis was abundant in San Miguel Island, but

was wholly arboreal,' Mr. Brown catching all his specimens in

traps set on the branches of large trees. It appears to be

diurnal, and on one or two occasions Mr. Brown saw the

animal proceeding along the branches with a curious gliding .

gait, his account suggesting the name I have used for the

species. It is the “Raton Marenero”’ of the islanders.

Proechimys burrus! sp. nov.

Fifty-one specimens, April and May.

Type: No. 8458, adult 3, Bangs Collection, Museum of

Comparative Zoólogy. Collected April 30, 1900.

Characters: A large, bright-colored, insular form of the

Proechimys centralis series. Differing from its nearest main-

land ally, P. centralis panamensis Thomas, by its larger size

and richer red color.

From P. centralis chiriquinus Thomas it differs in less spiny

back and redder colors. In color it is nearest to P. centralis

centralis (Thomas), but its feet are much darker.

The skull is slightly different from that of any of the other

subspecies. The rostrum is heavier, even, than in chiriquanus.

The nasals are long, broad, and bluntly truncate posteriorly.

(In centralis and panamensis the nasals are pointed posteri-

orly, and in chiriguinus they are shorter and less bluntly

truncate.) Hamular process broad and spatulate, as in the

other subspecies. Palatal foramina shorter than in chirigui-

nus, and less parallel-sided and wider open than in centralis

and panamensis,

1 Burrus, red, rufous.

No. 416] MAMMALS IN SAN MIGUEL ISLAND. 641

Color and Pelage: Spines confined to anterior two-thirds of

back (as in all members of this group of the genus); spines

not particularly numerous (as is the

case in cAzrzquzmus), but well covered

by the hair ; whole upper parts rich,

deep, ferruginous, slightly varied along

back by the brownish-black tips of the

spines showing through the hair; top

of nose, cheeks, and lower sides a

little paler, more yellowish; under

parts pure white, except anal region

and base of tail, which are colored

like the back ; hands and feet dusky

brown, in some specimens slightly

grayer along inner edge of metatar-

sus; tail bicolored, black above, gray-

ish below, well clothed with short, tyre ot Proechimys burrus, adult d -

stiff hairs ; ears dusky, nearly naked.

Young examples, half grown, are much darker, less reddish

above ; but among the adults there is little variation in color.

MEASUREMENTS.

| | Bas FROM

NuMBER. Sex. DATE. Neal | Gina bi. | Norcu.

8458, type $ adult | April 3o | 470 185 58 23

8448, topotype | d. « u 21| 490 205 et

8455, “ Pow "9 470 165 54 as

8449, “ ee “24 470 165 58 21

8451, “ 4 on “ 22 460 180 58 21

8450, « A ow “ 27 440 160 58 ne

8459, « 9 « “ 21 425 150 52 "

8454, “ erik “ay 438 165 2 22

8456, “ 9 “ “ 24 410 145 50 | 20

8460, « $ May 5 | 400 140 ow"

Skull, type, adult 4, basal length, 52; occipito-nasal length,

1.2; zygomatic width, 29; mastoid width, 22.2; least inter-

orbital width, 13.4; length of nasals, 24.2; width of nasals,

642 THE AMERICAN NATURALIST. (Vor. XXXV,

7.6; length of palate to palatal notch, 21; to end of ptery-

goid, 34; length of palatal foramina, 5.2; width of palatal

foramina, 3; upper tooth row, 9.8; length of single half of

mandible, 33.8; lower tooth row, 10.2.

~ Remarks.— The San Miguel spiny rat is a slightly differ-

entiated island form of the centralis series. It was very

common in the island, and Mr. Brown easily took as many

specimens as he wanted. It is known to the islanders as

* Raton mockungay." They, however, believe the tailless

individuals are a different animal. About one-third of the

specimens taken were tailless. The animal was generally

distributed throughout the island, and was often found living

in the huts and sheds of the negroes, like the common rat.

Mr. Brown took P. panamensis at Loma del Leon, Panama,

and P. chiriqguinus at Divala, Chiriqui.

Zygodontomys seorsus! sp. nov.

Sixty-eight specimens, April and May.

Type: No. 8490, old adult 4, Bangs Collection, Museum

of Comparative Zoólogy. Collected May 5, 1900.

Characters: Similar in color and general appearance to

Z. brevicauda (Allen and Chapman) of Trinidad, but much

larger, hind foot much bigger, tail less distinctly bicolored

and more coarsely scaly. Skull larger, rougher, supraorbital

heading heavier, transverse ridge where squamosal and supra-

occipital meet more pronounced, forward end of zygoma and '

zygomatic plate standing farther out from skull.

From Z. brunneus Thomas, Z. seorsus differs by its larger

size, larger foot, and richer, more reddish coloration.

Color and Pelage: Upper parts russet-brown, shaded with

dull ferruginous, the latter color more intense on the rump;

whole dorsal region thickly set with brownish-black-tipped

hairs; sides paler, more yellowish; under parts dull buffy-

gray, strongly shaded with dull ferruginous in anal region;

line of demarcation between colors of upper and under parts

indistinct ; upper surfaces of hands and feet yellowish-brown ;

1 Seorsus, isolated, living apart.

No.416.]] MAMMALS IN SAN MIGUEL ISLAND. 643

soles naked ; ears sparsely haired, dusky ; tail very sparsely

haired, coarsely scaly, indistinctly bicolored, blackish, rather

paler towards base below.

Young examples are rather darker, less reddish, brown above,

and the upper surfaces of feet and hands darker brown, less

yellowish.

The adults vary but little in color; some specimens are

yellower above and others more strongly ferruginous.

MEASUREMENTS.

| | | ČAR FROM

Huse, gae | Dem d ae aa room | Wro

| | HRS gE |

| d adult | May 5 | .320 |. 140 34 | 18

8485, topotype | [de m 315 | 140 34 17

8492, " Eze soia April 2 6 . 195 32 17

8495 * |3 H ecu 305 | 42 33 17

8487,“ | k ah May 2 300 | 10 k 17

8486, “ Le s April 25 w | DE 1 17

84906, * La A ay 6 295 | 120 | 32 17

B, *" |9.« « 8 |. 290 | neu s 16

8483, « Le April 30 290 | 435 | 30 18

8484, « E ON ir ad 390 ^o Bids *» 18

8489, “ TI May 5 285 | 120 | 30 15

8488, “ P$." “ 3 285 | 30 [uode 16

8481, ^ « pe m April 26 285 IIO 39 16

Bon ^ pa M a ik 280 120 30 17

8495, o“ Se 5 * 22 275 110 30 17

Skull, type, old adult 2, basal length, 32.8; occipito-nasal

length, 35.2; zygomatic width, 18.4; mastoid width, 12.8;

length of nasals, 15; width of nasals, 4.4; least interorbital

width, 5.4; width across zygomatic plates, 11.2 ; length of inci-

sive foramina, 7.4; width of incisive foramina, 3.2 ; length of

palate, to palatal notch, 16; to end of pterygoid, 22; upper

tooth row, 5 2; length of single half of mandible, 21 ; lower

tooth row, 5.2. _

_ Remarks. — The San Miguel vesper rat is a strongly marked

island species, most nearly related to Z. brevicauda, of Trinidad,

which it precisely resembles in color and character of pelage.

Its much greater size, bigger foot, and different tail distinguish

644 THE AMERICAN NATURALIST.

it, externally, from the Trinidad species, and the skulls of the

two can easily be distinguished.

Z. seorsus was an abundant animal in San Miguel Island,

inhabiting the dense, swampy woods, and Mr. Brown found no

difficulty in trapping it in numbers.

That the vesper rat of San Miguel Island should be so like

the Trinidad species is interesting, especially so as the yellow

honey creeper of the island (Cwreba cerinoclunis Bangs) is

much more nearly related to C. /uteola of Venezuela, Trinidad,

and northeastern Colombia than to C. mexicana of the neigh-

boring coasts of Panama and Colombia.

Mus musculus Linn.

One adult ¢, taken April 20, 1900.

Mus rattus rattus Linn.

One adult 2, taken April 20, 1900.

Mus rattus alexandrinus (Geoff.).

One adult 2, taken April 23, 1900.

The three introduced species of Mus could not have been

very numerous in San Miguel, as one individual of each was

all that fell into Mr. Brown's traps in over three weeks of

collecting.

Vampyrops helleri Peters.

One adult 9, taken April 25, 1900.

Mr. Gerrit S. Miller, Jr., tells me that this is the only exam-

ple of this rare bat that he has seen, but the specimen agrees

so perfectly with the description that he has little doubt of its

being true V. kelleri.

Hemiderma brevicaudum (Wied.).

One adult 4, taken April 25.

SOME PROTOZOA OF ESPECIAL INTEREST FROM

VAN CORTLANDT PARK, NEW YORK.

GARY N. CALKINS.

BroLocisrs in the vicinity of New York are particularly for-

tunate in having in Van Cortlandt Park a pond which yields a

great variety of interesting forms belonging to the lowest classes

of animal life. For the casual passer-by the pond is an

unsightly spot, but the green scum which covers parts of its

surface, and the healthy water-plants around its edges indicate

to the microscopist a rich fauna.

Van Cortlandt Park is about nine miles from the Grand Cen-

tral Station at Forty-second Street, New York, and is reached

by way of the Putnam Division of the New York Central Rail-

road. The trains on this division leave New York at the

terminal of the Elevated Railroad at Eighth Avenue and One

Hundred and Fifty-fifth Street at least twice each hour, and

from this point Van Cortlandt is only four miles. The pond

lies in front of the station at Van Cortlandt, extending away

to the northeast. Following the railroad up to the northern

end of the pond, beyond the spot where an offshoot of the

pond passes below the track to the left, a thick bed of lily

pads on the right indicates the spot where Amba proteus can

be invariably found. A quantity of the superficial slime cover-

ing the bottom should be scooped up in a Mason jar, or other

collecting jar which has been used only for living things, and

with a small amount of water carried back to the laboratory.

Immediate examination shows isolated forms of different genera

of Protozoa, which will be seen again later in considerable

numbers, Among these will be an occasional Amoeba, Actino-

spherium, Stentor, etc., — forms commonly found in similar

Pond water. If the material be emptied into flat dishes about .

z deep and 12” to 15” in diameter, and covered with about an

Inch and a half of tap water, the majority of the forms originally

645

646 THE AMERICAN NATURALIST. [Vor. XXXV.

observed will disappear within a few days, the tap water

apparently not agreeing with them. Examination of the mate-

rial from time to time during the ensuing month, however,

shows the gradual return of the original organisms, and with

this return the animals are much more numerous than before,

and with the advantage that only one type of organism appears

to occupy the stage at a time. Amada proteus comes along

from four to six weeks after collection, and so constant is

this appearance that I can plan in September for my classes

in November.

The addition of new tap water has a retarding effect upon

the cultures, and a too rapid culture can be checked by this

means. A slow culture, on the other hand, should be kept

from evaporating and should not be replenished.

In the present contribution I wish to offer a few notes in

regard to some forms of particular interest that have appeared

during the year, — forms which, for the most part, are rarely

observed and but littleknown. Among theseare Nuclearia, Mas-

tigamoeba, Multicilia, Lacrymaria, and a species of Actinobolus.

It may be of interest, however, to enumerate some of the other

forms that are found every year in this water. Of the Rhizo-

poda there are four species of Amceba (not counting A. radiosa,

the young form of A. proteus); two species of Pelomyxa, three

of Difflugia, one of Centropyxis, Arcella, and Euglypha ; and

of the Heliozoa, Actinophrys, Actinospherium, Acanthocystis,

Pinacocystis, and the dainty stalked Clathrulina are frequently

found. Of the Mastigophora there are several species of

Euglena, Phacus, Astasia, Trachelomonas, Peranema, Chilo-

monas, Cryptomonas, Petalomonas ; the beautiful colony form

Poteriodendron, and the more common colonies Volvox, Pan-

dorina, Eudorina, Gonium,andSynura. The minute choanoflagel-

lates Monosiga and Codosiga, and the dinoflagellates Peridinium

and Glenodinium, with their actively vibrating transverse fla-

gella, occur in greater or less abundance. The more common

forms of Ciliata in the pond are : Paramoecium, Dileptus, Bur-

saria, Stentor, Spirostomum, Enchelys, Chilodon, Prorodon,

Coleps, Urocentrum, Loxophyllum, Loxodes, Colpoda, Coeno-

morpha, Gastrostyla, Urostyla, Euplotes, Kerona, Trichodina

No.416.] PROTOZOA OF ESPECIAL INTEREST. 647

(the last two parasites on Hydra fusca), Vorticella, Zoótham-

nium, Epistylis. Finally, the Suctoria are represented by the

two genera Spharophrya and Podophrya.

Turning now to the less frequently observed forms, there is,

in some cases, apart from the fact that they may be found at

Van Cortlandt, little to be said in addition to the existing well-

known descriptions. A few remarks as to their mode of life

and relationships, however, may not be out of place.

Nuclearia delicatula Cienk.

This form, which was first described by Cienkowsky, has

occupied rather a varied position in classification. Placed by

some writers with the Heliozoa, by others with the Rhizopoda,

it belongs to those forms which, in any artificial classification,

must be considered as individuals rather than in groups. The

peculiarity of Nuclearia lies in the fact that its pseudopodia,

while long and needle-like, and in this respect similar to those

of the Heliozoa and the Radiolaria, are at the same time very

changeable, like those of the Rhizopoda. When the needle-

form pseudopodia are drawn in, the organism moves along like

an Amoeba and changes its place much more rapidly than the

Majority of the Heliozoa. When not moving, the animal

thrusts out its fine ray-like pseudopodia to a distance equal to

one and a half times its body diameter. The animal is rest-

less, however, and retains this condition only a short time,

moving off soon with an elongate form and by means of its

ameeboid processes. Unlike the fine ray-like pseudopodia of

the common Heliozoa, the appendages of this form have no

axial filaments to give them rigidity, and they frequently branch

and sub-branch, while the animal is moving (Fig. 1, B). In

this condition it surrounds and ingests food particles of various

kinds, especially plant cells, euglenoids, etc. I had a very

good culture of Nuclearia last year, and had studied the little

Organism rather carefully for several days, devoting many

hours to their mode of life, etc., but I had not seen one divide.

One day I had a very fine specimen under an immersion lens,

when a colleague, who was born under a lucky star, came in.

€ had not watched the specimen five minutes before he

648 THE AMERICAN NATURALIST. [Vor. XXXV.

remarked, ** This specimen is beginning to divide." He was

generous enough, however, to give me several views of the

animal during the process, which lasted only less than a minute

Fic. 1. — Nuclearia delicatula Cien A, th R 3 with ray-like pseudopodia :

B, the motile ameebold condition. x 1200.

and was not unlike the cleavage of a sea-urchin egg. After

division the daughter-individuals moved apart with the char-

acteristic amoeboid motion.

No: 416.) PROTOZOA. OF ESPECIAL INTEREST. 649

It is not an easy matter to interpret Nuclearia, for it may be

regarded as a degenerate heliozoan which has lost its axial

filaments, or as a primitive form which has never had these

peculiar axial structures. The interpretation involves the old

dispute over the most primitive forms of Protozoa. Are we,

with Klebs, to hold that the flagellates were the most primi-

tive forms ; or with Lankester, that the rhizopods are to be so

considered ; or with Bütschli, that the flagellated amoeboid

forms were the-ancestors of both rhizopods and flagellates?

The problem, to my mind, has no great importance, for, it is

quite probable that the Protozoa have not come down without

change from that unknown period of the primitive animal forms,

while even in historic times they may have been adapted and

readapted many times over to changing conditions of environ-

ment. The fact that the majority of Rhizopoda have flagel-

lated swarm-spores is not sufficient proof that their ancestors

were flagellated, while on the other hand the frequent amceboid

condition of the flagellates is equally valid evidence that the

thizopod form was the older. Flagella and pseudopodia are

not far removed from one another, and frequent observations

have confirmed Dujardin's early view that flagella may become

pseudopodia and pseudopodia flagella. The transitional forms

such as Dimorpha, Actinomonas, etc., only strengthen the con-

nection, and it will be shown, I believe, that many of the pseudo-

podia of Heliozoa are similar to the flagella of the Mastigo-

phora. Blochmann has shown, for example, that in Dimorpha

the pseudopodia have axial filaments, which, as in Actinophrys,

focus in the nucleus or its vicinity, The flagellum, too, is

focused at the same point, so that in Blochmann's figure the

axial filaments and the flagellum appear to be the same in struc-

ture, Artodiscus, according to Pénard, is a heliozoón which

dances about the field upon the tips of its ray-like pseudopodia.

In this case the pseudopodia are like so many flagella, for there

can be no doubt that the motion is due to the contraction and

elasticity of the pseudopodia and probably of the axial filaments

in them, Acanthocystis, also, has a slow, rolling motion, by

means of which it covers a distance equal to twelve times its

Own diameter in one minute (Pénard). Here, also, although

650 THE AMERICAN NATURALIST. [Vor. XXXV.

less than in Artodiscus, the pseudopodia are probably con-

tractile.

There thus appears to be some morphological ground in

favor of the hypothesis that Rhizopoda may have originated

from the Mastigophora and through the Heliozoa. Artodiscus

and Acanthocystis may be regarded as having many flagella of

modified form, while Dimorpha, which has a typical flagellate

stage, has one true flagellum and at the same time pseudo-

podia of the heliozoan type. In Actinophrys, Sphzrastrum,

Heterophrys, etc., the contractile element, whatever it is, is

lost, and the pseudopodia appear rigid and apparently serve no

purpose in movement. They are of the same type, however,

as in Artodiscus and Dimorpha, and, as in the latter, they

center at a common point to form a system of rays very much

like a centrosphere in Metazoa. In Actinosphzerium the rays

no longer focus in a common center, although each ends in the.

vicinity of some nucleus.

In respect of the axial filaments the Foraminifera apparently

represent an intermediate stage between the Heliozoa and the

fresh-water Rhizopoda. It has been frequently pointed out that

the central portion of the pseudopodia of these forms is dense

and stiff as compared with the outer plasm, which has the typi-

cal flowing motion of a rhizopod, thus reversing the conditions

of one of the lobose forms.

If the relations as outlined above would hold good between

the Heliozoa and the Mastigophora, then Nuclearia, having lost

its axial filaments, finds its real position as an intermediate form

between Heliozoa and fresh-water rhizopods, retaining, however,

the fine and needle-like pseudopodia of the Heliozoa. Considered

in this connection it would be a degenerate form, The entire

argument, however, might work equally well the other way,

and it might be held that an Amoeba such as A. radiosa, OF

A. tentaculata, or Nuclearia, develops vibratile pseudopodia hav-

ing an axial filament of especial contractile substance. The

foraminiferan pseudopodia would be a first step in such a

development, Acanthocystis, Artodiscus, and Dimorpha succes-

sive following steps, until a flagellate would be the outcome. It

would be difficult only to place Actinophrys, Actinospheerium,

No. 416.] PROTOZOA OF ESPECIAL INTEREST. 651

Heterophrys, Sphzerastrum, etc., according to such an hypoth-

esis. In either hypothesis Nuclearia would form an important

intermediate link.

Mastigameeba verrucosa S. K.

A number of specimens of this interesting form appeared in

one of my cultures during October, and I had a good oppor-

tunity to study it. The species is different from that described

by F. E. Schultze (Mastigameba aspera), and belongs appar-

ently to that which Kent described as M. verrucosa, although

in size and general form, etc., it agrees with the description of

Mastigameba invertens of Klebs. I could not make out, how-

Fic. 2. — Mastigamæba verrucosa S. K. A, spherical form assumed when swimming with the

ellum in advance; 2, flattened form with lobose pseudopodia. x 1250

ever, that the organism reverses its axes when it comes to rest,

— à point upon which Klebs placed considerable importance.

When first observed this species of Mastigamoeba appears

like a monad with an irregular outline, which makes its way

through the water with a peculiar irregular course, vibrating

the while as though shaken vigorously. The cause of the

vibration is the action of the rather long flagellum, which is

directed in advance. On all sides of the minute body are

lobose processes of variable length (Fig. 2). After pro-

Sressing in this way for some time it finally comes to rest

and the pseudopodia are stretched out in all directions, like

those of a small Amoeba. The flagellum, in the mean time,

vibrates slowly at one end. In its usual motile condition

the organism is round, but when quiet and with outstretched

652 THE AMERICAN NATURALIST. [VOL. XXXV.

pseudopodia it assumes a flattened shape, and a distinct ecto-

plasm and endoplasm can be made out. After a shorter or

longer period of rest the action of the flagellum becomes

more and more rapid, the body becomes detached from the

substratum and assumes a spherical form, after which it

again moves through the water with the peculiar vibrating

motion which first attracted my attention. A minute con-

tractile vacuole pulsates at irregular intervals and cannot be

made out easily when the organism is moving. A small

spherical nucleus lies near the base of the flagellum, but no

structure could be made out.

In the resting condition the pseudopodia surround and take

into the body small objects of different kinds, which are stored

up in the endoplasm until digested. The process is apparently

no different from that in Ame@ba proteus.

Mastigameeba belongs to a group of flagellated amoeboid

forms, the Rhizomastigide which Bütschli regarded as ances-

tral to both flagellates and rhizopods. Klebs, on the other

hand, placed the beginnings of both groups farther back, and

considered the Rhizomastigidze as flagellates which have become

amoeboid.

Multicilia

Only one individual of this interesting genus was seen, and

no details were made out save the presence of about twenty

long flagella-like cilia distributed evenly about the body. By

means of these appendages the organism moved along with a

rolling motion.

The search for the beginnings of the ciliate stem from some

more generalized organisms has been singularly vain. The con-

servative structure which the infusorian body presents gives

but little clue to their ancestry. In only one other known case

(Polykrikos among the Dinoflagellidia) is there a similar nuclear

dimorphism, while the almost universal reproduction by trans-

verse division is met with among the Mastigophora but rarely.

It is generally conceded that the infusorian stem must have

arisen from the flagellate stem at an early period, and must

have become progressively differentiated until the present

"EE

xo T

No. 416.] PROTOZOA OF ESPECIAL INTEREST. 653

striking peculiarities were well established. It is assumed by

Bütschli and by the majority of others who adopt the hypoth-

esis that the ciliates were derived from the Mastigophora,

that the original forms may have been similar to the present

genus Multicilia, whose motile organs are more like flagella

than they are like cilia. There is no close connection between

the cilia and flagella, as there is between the latter and pseudo-

podia, but Bütschli holds that the Ciliata might have been

derived from such a form as Multicilia, in which the motile

organs became progressively shortened and increased in

number.

There is another hypothesis which has been maintained by

Entz and Maupas amongst others, in which the Ciliata are

derived from the Rhizopoda through such forms as Actinobolus

and the Enchelinidæ in general, two types of which have been

found in Van Cortlandt water, Lacrymaria and Actinobolus.

Before speaking of this hypothesis in detail I will add a few

notes on these two genera.

Lacrymaria olar Ehr.

Of all known micro-organisms, Lacrymaria olar is the most

remarkable for its elasticity, and manner in which it stretches

Out a neck-like portion bearing the mouth. The wonderful

variability of this portion of the body cannot be represented in

a figure, but when one has witnessed the contortions which it

undergoes he will never forget the organism. Sometimes the

“neck” is stretched forwards in a straight line to a distance

equal to six times the diameter of the body ; again it may be

wound around the body of the animal from the insertion point

to the posterior end of the animal, and forward again on the

other side ; and even then it may stretch out to a considerable

distance in advance of the forward end ; or, on the other hand,

it may be retracted until the mouth appears to be merely the

extremity of an ovoid body.

Lacrymaria is not an uncommon form, and in the present

Connection it is mentioned chiefly for its bearing upon the

hypothesis as to the origin of the Suctoria.

654 i THE AMERICAN NATURALIST. [Vor. XXXV.

Actinobolus radians St.

This infusorian is much more rare than Lacrymaria, and from

its rarity perhaps more interesting. It was first described by

Stein, then by Entz, who studied it very carefully, and later

by Von Erlanger, who made out a number of minor points of

structure. It is almost always found in the company of certain

forms of Suctoria, Von Erlanger alone not finding these asso-

ciates. In the Van Cortlandt water the organism was over-

looked for some time, because of its supposed identity with the

suctorian Sphzrophrya, which was present in considerable

numbers. Actinobolus is a small spherical organism not much

larger than a Urocentrum,

ae and when at rest resem-

‘Nal Wi) bles a Sphzrophrya,

S NS NN i Hd Vy which in turn resembles

: WSS Mg A an Actinophrys. It is

only when this form is

studied with a medium

high power that a fringe

of vibrating cilia can be

* made out around the

^ 7 i uem NÑ NN profile of the périu

/ DN M a AN mA NN `N, It may then be seen that

B M i il C there are groups of long

| cilia (ten or twelve to a

Fic. 3.~ Actinobolus radians St. The tentacles are group) arranged about -

th side withthe small mouthinadvance, Themes the base of each of the

is elongate and constricted. Six Halteria are faintly long tentacle-like proc-

represented in the endoplasm. x 1200.

esses, which give to the

organism, when at rest, the aspect of a heliozoón (Fig. 3).

The tentacles are the most interesting feature of the ciliate,

and their function has been in considerable doubt. Stein was

unable to assign any function, but described them as retractile,

like the tentacles of the Suctoria, to which he compared them,

observing at the same time that they might be entirely with-

drawn into the body, leaving no trace of their former presence.

Entz thought that they might assist the organism in getting

No. 416.] PROTOZOA OF ESPECIAL INTEREST. 655

food, and observed that the ends of the tentacles were attached

to threads of algae which appeared to be ruptured at the points

touched, and he concluded that a secretion of some sort is poured

out upon the alga by the tentacle. He also maintained that the

organism is exclusively a plant-eater. Von Erlanger described

the tentacles as possessing a stiff dart or needle-like trichocyst at

the end ; these, he said, are drawn in with the tentacle until,

when the tentacles are entirely withdrawn, they form a peripheral

layer of trichocysts as in a Prorodon. He also observed that the

organism may swallow other living forms and described the

seizure of a small flagellate by the cilia about the mouth opening.

There are a few points which I may be able to make in

addition to the above. The organism, when feeding, lies

attached to the bottom of the vessel by its oral tentacles

(.e., those about the mouth, which are no different from the

others on the animal) and mouth downwards. This brings

the posterior end of the organism upward, and the anal spot

and contractile vacuole can be easily seen and studied while

the animal is in this position. The vacuole is not in the center

of the dorsal surface, but lies a little to one side (Fig. 4). The

tentacles are almost as distensible as the neck of Lacrymaria,

and may stretch out a distance of two and a half diameters

of the organism, or retract until they disappear in the body.

They arise from the substance of the cortical plasm, which is

covered by a very thin cuticle or outer membrane, but they

cannot be traced into the inner plasm. The cilia are more

like flagella than cilia, and move with a vibratory rather than a

stroking motion ; while resting they are easy to follow, but

when the tentacles are all withdrawn the motion becomes

much more rapid, and only the general waving surface can be

made out.

It is when the animal is quiet that it takes in food. It

remains for hours, sometimes, without giving a sign of feed-

ing, but the tentacles are stretched out to their greatest

length and it appears to be merely waiting for something to

come. Things do come, especially flagellates like Chilomonas,

Trachelomonas, Euglena, etc., and ciliates such as Urocen-

trum, Parameecium, Bursaria, Colpoda, etc., but no attention is

656 THE AMERICAN NATURALIST. [Vow. XXXV.

paid to them. Finally, however, a particular kind of ciliate

comes along, Halteria grandinella, which, sailing along with its

graceful but spasmodic motion, suddenly finds itself on the end

of one of the long tentacles of Actinobolus. The captive whirls

7 ty Bf ese X < 3 RA SSS N

Ti Pye ae RASS

Fi. 4. — Actinobolus radians St. at rest, with 1 tstretched, waiting for Halterias,

one of which has been captured. The mouth is on | the eee x 12

round and round for a brief second or so, and then becomes

quiet, just as it does when captured in a similar way upon the

tentacles of a suctorian. The tentacle then begins to shorten

and as it shortens the victim is drawn towards the capto

No. 416.] PROTOZOA OF ESPECIAL INTEREST. 6

perhaps to the side, perhaps on top ; wherever the prey may

be drawn, however, it is gradually worked downwards by the

tentacles to the underside toward the mouth, and then with a

single swallow it is transferred to the waiting endoplasm in

which it is digested. So insatiable is the appetite of this form

that, after adding water containing Halteria in abundance, I

have seen it swallow in the manner described above no less

than ten individuals within half an hour, after which its endo-

plasm appeared filled with foreign bodies, as Von Erlanger

described in the form he observed.

This then is the function of the tentacle, a food-catching

organ or set of organs ; the trichocyst which Von Erlanger

described, or its analogue (I was unable to make it out as

described by Erlanger), brings down the prey, and the tentacle,

by shortening, fetches it to the mouth. The tentacle itself is

inserted in the Halteria, for it can be seen with the greatest

distinctness when the victim becomes quiet, and I believe that

the dart at the end as described is not discharged into the

prey but is driven into the soft body of the victim as a minute

spear, with shaft attached.

When fully fed or when warmed by artificial means the

animal detaches itself and draws in its tentacles. It then

swims mouth first in a rambling sort of way, turning the while

on its longer axis (Fig. 3). !

Apart from the general biological interest which such an

organism has, there is a considerable interest from a theoret-

ical point of view. Maupas maintained in 1881 that the Ciliata

have been derived from the Rhizopoda through the Suctoria,

and that the tentacles are to be compared with pseudopodia,

While the forms with both cilia and tentacles were considered

intermediate. So important was the group of tentacle-bearing

forms considered that Mereschowsky formed a special group —

the Suctociliata — for their reception. Neither Maupas, Entz,

nor Mereschowsky could regard the tentacles in these forms as

food-taking organs similar to those of the Suctoria, and could

assign them no function other than that of assisting inthe capture

of aliments. As well known, the tentacles in the Suctoria are

for the most part hollow, and the food is taken in some manner

658 THE AMERICAN NATURALIST.

through the open ends. Maupas regarded them, together with

the tentacles of Actinobolus, Ileonema, and Mesodinium, as

pseudopodia, and thought that through them the ciliates came

from the Suctoria and the latter from the Rhizopoda. This

theory of the origin of the Ciliata from Rhizopoda has but an

historical interest, and it is now very generally if not almost

universally held that the Suctoria came from the Ciliata ; while

the tentacled forms of ciliates are merely independent modifi-

cations for special ends and with no phylogenetic significance.

A number of minor points suggest the close relation of the

forms like Actinobolus to the Suctoria, but more careful con-

sideration leaves little reason to doubt the view that they

represent merely parallel evolution, being adaptive features for

special ends.

THE MOLLUSCAN FAUNA OF THE GENESEE

RIVER.

FRANK COLLINS BAKER.

Tue study of faunal distribution has always been a favorite

occupation of zoólogists, and particularly of those interested in

the study of the Mollusca, as in this branch we find a very

large number of species, covering wide areas and subject to

every variation of environment. In no department of the Mol-

lusca is this of such absorbing interest as in the fresh-water

forms (unless we except, perhaps, the air-breathing pulmonates),

especially those inhabiting a large river wherethere are several

barriers to the homogeneous distribution of its shell fauna.

During the past summer the writer spent several months in

Such a study of the Genesee River, where the environments

are quite different in several parts of the stream, with a corre-

sponding difference in the mollusk fauna. The river was care-

fully surveyed from near its mouth on Lake Ontario to beyond

South Park, a distance of ten miles. A large collection was

made, which is now in the museum of the Chicago Academy

of Sciences. My thanks are due to Miss Edna E. Hall for

valuable assistance in collecting, and to Rev. John Walton

for many notes,

The Genesee River rises in Potter County, Pennsylvania,

and flows in a generally northward direction for about 120

miles, emptying into Lake Ontario, seven miles north of

Rochester, N. Y. The Genesee valley is very fertile, and the

river flows between low banks rich in vegetation. Before

passing Rochester the river is deep, the banks muddy, and the

current steady but not very swift. From a point a little north

of Genesee Valley Park (or South Park) the bottom of the river

“comes very rocky, the current swift, and at Rochester the

"VT drops to the valley below in three series of falls of

considerable magnitude.

659

660 THE AMERICAN NATURALIST. [Vor. XXXV.

The geography of the river from South Park to the lake

may be thus described: From south of Genesee Park to

within a short distance of a small dam it is broad, deep on

one side and affording a shelving, shallow bank on the

other, the bottom being composed of a clayey mud. Unios,

Campelomz, Goniobases, Sphzria, and Physz are here very

common. Just before reaching the dam the river shallows

very perceptibly, flowing rapidly over great ledges of lime-

stone. Lampsilis luteolus and L. iris are very common

here, and also Goniobasis, while Physa is not quite so

common.

Below the dam, which is now broken and very much out of

repair, the river flows rapidly over a rocky bed, in many places

the water being so shallow that persons can wade across. The

right bank is very rocky, more so than the left, and forms little

sheltered bays here and there, in which Goniobasis and Physa

live by thousands, the former making a veritable pavement in

some places. Lampsilis luteolus is also common, although

apparently preferring the middle of the river. Along the

bank here the Unios lie in great windrows, their dead shells

piled up like the débris on an ocean shore. In this pile of

shells such species as Lampsilis iris, L. luteolus, Quadrula

rubiginosa, Lampsilis complanatus, and Alasmodonta rugosa are

common. The water near the rocky shore also affords good

retreats for the mollusks.

Near the Erie Railroad bridge the water deepens, and the

Unios begin to thin out, although Lampsilis /uteolus seems to

be as common as ever. Goniobasis also flourishes, while Physa

almost dies out. Near Clarissa Street bridge the water

becomes still deeper, and the only species found are Lam-

psilis luteolus, L. iris, L. complanatus, and Goniobasis, with à

stray Campeloma or Spherium. - Physa seems to have disap-

peared completely. Below Clarissa Street bridge the water

deepens to the state dam, beyond which it is very shallow and

rocky, so far as known affording no mollusks save an occa

sional Goniobasis or Campeloma. At the dam the water 1s

deflected in a large stream for factory uses and again reaches

the river at the first falls.

BRE NEIN NAT

No. 416.] MOLLUSCAN FAUNA OF THE GENESEE. 661

At Rochester, below the bridge of the New York Central

Railroad, the river drops to the valley below in a fall 96 feet

in height, cutting through Niagara limestone and Clinton

shale. Below this fall the current is rapid, the river flowing in

a sinuous course, leaving on either side at intervals a low, flat

piece of ground with a shallow, shelving shore, the opposite

side, however, being very deep. The upper part of the river

is so contaminated with oil and gas products from the gas

works and a number of manufactories that mollusks cannot

live here. A short distance down the river, where the water

is not so badly contaminated, a few Physz, Planorbis, and Lim-

nea live. These grow more abundant as the purer waters are

reached. About two miles farther north a second fall occurs,

26 feet in height, cutting through the Clinton shale. Below

this fall the river runs swiftly, is very deep, and forms a circu-

lar pool just beneath the fall. The walls are very steep and

stony, the river having cut away the soft shale, leaving great

projecting ledges. Physa, Planorbis, and Limnza are here

notably abundant, living on ledges of rock where the water is

a few inches deep.

A few hundred feet north of this fall a third one occurs,

cutting through Medina sandstone 83 feet in height, and the

river drops to the valley below, flowing at first rather rapidly,

but after a mile or so rather quietly, between well-wooded

banks 210 feet in height. The riveris here bordered by many

Swampy flats covered with rank vegetation, and the shores are

muddy. -Near the falls the river is somewhat shallow, flow-

ing over fallen rocks, but it soon becomes quite deep. The

senera present are Limnzea, Physa, Planorbis, Bythinia, and Caly-

culina ; Limnzea and Planorbis predominating. Inthe mouth of

the Genesee there are some eight or ten species of Unios, with

Some gastropods, which have come in from Lake Ontario, but

they have not as yet reached far enough up the river to form

a factor in this discussion.

The following table shows the comparative distribution of

the species in each section of the river. An asterisk (*) indi-

cates that the species is very abundant, while the dagger (T)

Shows that the species is more rarely found.

THE AMERICAN NATURALIST. [VoL. XXXV. —

662

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No.416.] MOLLUSCAN FAUNA OF THE GENESEE. 663

A study of the table will show a few interesting facts. The

first will be the abundance of Unios above the falls and their

total absence below, showing that the upper falls afford an

insurmountable barrier to their further distribution in this

direction. Another fact of equal note is the abundance of

certain gastropods in all four sections of the river (Planorbis,

Physa). The general absence of ctenobranchiates below the

falls (Bythinia excepted) is noteworthy. Bythinia is a recent

addition to the fauna of the lower river, appearing in immense

numbers in 1898. It was very common at the mouth of the

river, in Lake Ontario, in 1895, and in these three years it

became one of the most abundant mollusks in the lower river.

The distribution of Physa is also peculiar. Physa integra

is the dominant and (so far as I know) the only form above

the falls, and Physa gyrina below. Physa sayii is the only

species between falls 1 and 2 and is very abundant only between

the upper and lower falls, a very peculiar distribution, for which

I cannot account. P. integra does not occur between the upper

river and falls 2 and 3, and I was not able to find it between

falls 1 and 2, which, if borne out by future observations, offers

an interesting phase of distribution. Why P. integra should

be so interrupted in its distribution may be accounted for by

two hypotheses : (1) that specimens may have been carried over

the first and second falls and found lodgment above the lower

falls ; or (2) specimens might have been carried by birds or

other animals. Why Physa sayii is found everywhere except

In the upper river is also an interesting question.

Limnza seems to be evenly distributed, and yet in this

senus we find the same peculiar distribution as in Physa.

L. falustris has been collected in small numbers in the upper

Diver, but Z. catascopium and L. caperata take its place in the

other three regions, the former being very abundant. Ancylus

has been found only in the upper river, in the dead valves of

Unios, but it may be that it inhabits all parts of the river. As

in the case of Physa integra, we find the distribution of Lim-

"à palustris to be peculiar, jumping, as it does, the first fall and

occurring sparingly between falls 2 and 3. P/anorbis trivolvis

's the only evenly distributed species. Among the bivalves,

664 THE AMERICAN NATURALIST.

Calyculina transversa is interrupted in its distribution, but, like

Physa gyrina, P. heterostropha, and Bythinia tentaculata, may

have come up the river from Lake Ontario. This, however,

will hardly account for Calyculina partumeta, which is found in

the lower river and between falls 2 and 3.

As one glances over the table the fact presents itself that

there is a marked division in the faunas between the upper

river and the series of falls, and the lower river. This may be

accounted for by the fact that for a distance of about two miles

above the first fall the river is shallow and very rocky, in: fact

flowing over great ledges of Niagara limestone, and the Unios

all seem to prefer the deeper, more quiet waters above this

section of the river, only a few stragglers, like Lampsilis luteo-

lus, L. iris, and Alasmodonta rugosa, being found in this rocky

region. The writer was unable to find Unios within a half mile

of the upper falls, showing conclusively that this environment

is unsuitable for them. Another reason why Unios are not

found below the falls is probably that their heavy shells and

also their habit of burying themselves in the mud prevent

them from rising to the surface and being swept over the falls,

as might be the case with Planorbis, Limnzea, or Physa, which

come periodically to the surface for air.

The foregoing discussion indicates that a series of falls like

those at Rochester will prove an effective barrier to the dis-

tribution of some mollusks (as the pelecypods with mud-

burrowing habits and the ctenobranchs, which cling to the

rocks and do not come to the surface), while to others (like the

fresh-water pulmonates, which come to the surface frequently

and hence could be swept over the falls) it is not a barrier.

Future studies and collections, however, may modify the

above conclusions.

THE CONCHOMETER.

HAROLD S. CONANT.

In connection with a recent study of variation in Purpura

lapillus, it seemed necessary to measure the apical angle of the

shell, among other features, and since no apparatus adapted to

this purpose was known, an instrument, which I have called a

conchometer, was constructed.

About 4000 measurements have been taken by this means ;

and so well did they correspond with theoretical considera-

tions that the writer believes the value of the instrument to

have been proven. With the possibility that those who are

using the statistical method for studying variation may find it

suited to their purpose, a description is here given.

It consists of two parts : (1) a device for measuring angles

at the ends of the shell, and (2) a device for measuring the

long axis of the shell and the distance from the apex of the

shell to the aperture. The first consists of an ordinary draw-

ing compass, of which one arm, ~, is firmly attached to a

board, and the other, M, may be moved over a graduated arc,

4, which is also attached to the board. The arc reads to half-

degrees and has its zero-point at the fixed arm. Attached to

each arm of the compass is a plate of metal some 2 inches

long and ł of an inch wide, standing vertically to the board, so

that only the edge is visible in the figure. The two plates are

so placed on the arms that they just touch each other at the pivot

of the compass. When the board is set in position, vertically,

the plates form a V whose angle may be made larger or smaller

by the adjustment of the movable arm by means of a suitable

handle, Æ. It is with this part of the apparatus that the apical

angle of the shell may be measured, as described later.

The second part of the apparatus, by means of which linear

measurements are taken, consists of two scales graduated to

millimeters by means of concentric lines. One is etched upon

665

666 THE AMERICAN NATURALIST. [Vor. XXXV.

a glass plate, P, placed in front of the V, parallel to the board,

and secured to it by means of a block, W, and the other is a

ruled scale attached to the board. The lines on the second

scale are directly behind those on the glass plate and, there-

fore, are not seen in the figure. The two scales are exactly

alike, except that one is etched on glass and the other ruled on

No. 416.] IHE CONCHOMETER. 667

paper. The object of having two scales is to avoid parallax

errors in reading. The paper scale is protected from damp-

ness (and consequent warping) by a sheet of transparent

celluloid.

A few words may be necessary to explain the working of

the instrument. A shell of Purpura lapillus is lowered into

the V, and the arms are adjusted to fit the apex. It is not a

difficult matter to do this, and the adjustment may be made

more accurately by having the plates polished to mirror-like

brightness and using then the well-known method of determin-

ing contact by reflection. The shell having been well fitted,

the position of the point of the movable arm may be read on

the before-mentioned arc, A, and the apical angle be thereby

determined. A simple bob, B, is used to lower the shell into

the V and to withdraw it. This is made of stiff copper wire,

with a small ball of cotton wool dangling at the end of a piece

of finer wire. The cotton wool is thrust into the aperture

of the shell, and in this way the latter is held firmly, apex

downwards.

The total length of the shell and the distance from the apex

to the aperture are both measured while the shell is in the.

position just described and shown in the figure. It is important

to note that the zero-point of the linear scale lies at the apex of

the V, and that the apex of the introduced shell must neces-

sarily be at that point. Hence the total length of the shell

will be determined by observation of the part of the shell lying

farthest from the zero of the scale, and when found it may be

read directly by means of the concentric lines. In the same

way the distance from the apex to the aperture may be read

directly by noting where the aperture comes on the scale.

Care should, of course, be taken always to present the ventral

side for measurement. After some practice the measurements

may be taken rapidly and accurately.

Dow ACADEMY, FRANCONIA, N. H.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XVI. THE PHALANGIDA.

NATHAN BANKS.

THE Phalangida, or the Opiliones as they have been called

by some authors, form a very distinct order of the Arachnida.

The cephalothorax is of one piece, but often presenting one or

two incomplete transverse furrows. There is also a furrow

on each side, which is supposed to indicate that the portion

beyond is a part of the pleura. On this part on each anterior

side there is a small, somewhat circular depression, with a

membranous bottom; these are called the lateral pores. On

the middle line of the cephalothorax, and frequently near the

anterior margin, is a rounded elevation, with a simple eye on

each lateral face ; this is the eye-tubercle, or eminence. The

abdomen is united to the cephalothorax by the whole of its

breadth, so that there is no trace of a pedicel. It is always

quite broad when compared to its length, never slender, and

usually quite high; in the males it is often tapering behind.

There are considered to be eight segments in the abdomen,

but in many forms the basal and median ones are poorly

defined on the dorsum ; on the venter there are usually six

distinct segments. The underside of the cephalothorax is

occupied by the coxze and a median piece similar to and often

called a sternum. It is, however, something quite different ;

It is the basal ventral segments advanced forward over the

bases of the coxze ; it carries with it the opening of the genital

organs, which thus, in certain forms, appears to issue close to

the mouth. This advancement of the abdomen is so pro-

nounced in many of our common forms that the coxa are

crowded at base and so appear to radiate from a central point.

The legs are usually very long and slender, though tarsi and

670 THE AMERICAN NATURALIST. [Vor. XXXV.

ofttimes several of the other joints are divided into smaller

portions by what are termed false articulations. Each tarsus

is terminated by one or two claws.

The habits of the Phalangida are rather simple. With our

common forms the length of life is but one year. The eggs are

laid in the ground, or in crevices of wet or decaying wood,

during the summer or fall; the young hatch in late fall or

early spring. At birth they have the general appearance of

Fic. 1. Fic. 2.

Fic. 1. — Dorsal view of a phalangid. a, palpus; 5, mandible; c, lateral pore; d, eye-tubercle;

' e, cephalothorax; /, abdomen.

Fic. 2. — Ventral view of a phalangid. 4, mandible; 4, palpus; c, maxilla; 4, pedal lobe;

e, coxa I; f, coxa II; g, coxa III; 4,coxa IV; z, advancement of abdomen; 4, spiracle ;

4, 2d ventral segment; 7, 3d ventral seg.; », 4th ventral seg.: o, sth ventral seg.; £^ 6th

ventral seg. ; s, anal plate.

the adult, often, however, with slight modifications. Some

species having short palpi when adult have long palpi when

young. In several of our species the young have the patella

of the palpus with a long branch, whereas it is absent or

nearly so in the adults.

They spin no web and make no retreat of any kind. They

usually move slowly, but a few can run rapidly. Some species

occasionally congregate in numbers on trees. They feed mostly

on living insects, but sometimes on decaying animal matter.

They appear to have few enemies, but their long legs or hard

No. 416.) NORTH-AMERICAN INVERTEBRATES. 671

bodies are a serviceable protection. When handled they often

. exude a whitish fluid, which in some species has a disagreeable

odor.

The points for the separation of species are often best

observed in the males; this is especially true in the large

genus Liobunum, in which the females are not known for

several species. The males, as a rule, have more slender legs

and palpi, and a more tapering abdomen ; but in some species

the abdomen of the male is very short and broadly truncate.

In Liobunum the last joint of the male palpus shows a row of

small denticles beneath. In a few forms the male has the

hind coxa enlarged. In several cases there are colorational

differences between the sexes. In many forms the male is

more spinose than the female. The order Phalangida is

usually divided into two suborders: Mecostethi (Laniatores

of Koch) and Plagiostethi (Palpatores of Koch):

First ventral segment of abdomen not reaching in front of hind coxz;

hind tarsi with two claws at tip or else with a compound claw; palpus

with tibia and tarsus more or less depressed; hind legs usually the

ongest MECOSTETHI

First ventral Mcd of PNE eid ik in n froid of hind coxa;

all tarsi with but one simple claw ; pts with tibia and tarsus cylin-

drical, second legs thelongest . . . . .' PLAGIOSTETHI

MECOSTETHI.

This suborder is represented in our fauna by two families:

1. Hind pair of coxe free at apex; spiracles obscure . PHALANGODID/E

Hind coxz wholly united to the venter; spiracles distinct COSMETID/E

The Cosmetide have but one genus, Cynorta, of three species, in the

United States ; they can be distinguished as follows :

1. Two acute spines near end of dorsal shield. . C. ornata Say (Fla.)

Two rounded tubercles instead of spines . C. sayéi Sim.’ (S. States)

Neither spines nor tubercles ; body with two pale spots above

- bimaculata Banks (Calif.)

Our Phalangodide are arranged in four genera:

1. One compound claw to each of the four sper tarsi; palpi not as

long as the body; eyes present : Sclerobunus

Two simple claws to each sphere: tarsus . eee Suy

1 Includes C. albolineatus Scerensen.

672 THE AMERICAN NATURALIST. (Vor. XXXV.

2. Eye-tubercle arising from anterior — of the cephalothorax; palpi

shorter than the body . . Sitalces

Eye-tubercle arising back ie he margin; dM ipe tan body... 3

3. Tarsus II about twice as long as body, its joints very slender; no eyes

Phalangodes

Tarsus II about length of body; eyes often present . . Scotolemon

SCOTOLEMON Lucas:

1. Eye-tubercle ends in a spine; second joint of Vise three times as long

as wide; body with short, stout spines . S. spinifera Pack. (Fla.)

Eyetibercle without spine ; second joint of T about twice as long

as broad "UY

2. Tarsus II with short joints ; ue stouter

S. flavescens Pack. (Ky., Ind., Va.)

Tarsus II with joints slender; palpi more slender

S. californica Bks. (Calif.)

SCLEROBUNUS Banks : À

Color red, tips of legs black. . . . S. robustus Pack. (W. States)

Color brown, tips of legs yellowish .S. órunneus Bks. (Wash., Alaska)

SITALCES Simon ; `. (005 9c. as Se californica Bka (Ci)

PHALANGODES Tellkampf. . . . P. armata Tellk. (Mammoth Cave)

PLAGIOSTETHI.

Three families occur in the United States:

. Last joint of palpus with a claw at end, this joint longer than pre-

ceding one (except in male of Protolophus) . . . PHALANGID/E

Last joint of palpus much shorter than penultimate, no clawatitstip. 2

2. Palpi very short, concealed under the projection of eye-tubercle

TROGULIDE

Palpi long and prominent. . . . . . . . NEMASTOMATID/E

Our genera of Phalangidz are separated as below:

1. Male with palpus enlarged, last joint shorter than penultimate ; female

with patella of palpus provided with a long branch; palpal claw

smooth ; a row of teeth on sides of coxe . (Protolophini) Protolophus

Last joint of Urbe Ex than cene ; aE without an

in adult

2. Ga of enormous size, three ae spines on femur of pipe

CH Caddo

Eye-tubercle of normal size : x.

3. A group of spinules on anterior margin of the | copia iod the

eye-tubercle spinose; palpal claw smooth, rarely if ever with late

rows of teeth on cox, frequently with spines on femur of palpus

(Phalangini) 19

—

.416] NORTH-AMERICAN INVERTEBRATES. 673

Eye-tubercle smooth, if spinose then no group of te on anterior

margin of the cephalothorax

Palpal claw denticulate, a row of teeth (adult) on i hides oF coxa, pidlatinct

at least on anterior side of coxa I, legs usually long . (Liobunini) 5

Palpal claw smooth, no such rows of teeth on coxa, legs usually

shorter . . . (Leptobunini) 7

Body very hard MP diciate abo ind below: legs very short, fourth

pair nearly as long as second pair. . Mesosoma

Body softer, although often mboi, w jiii below; legs

longer, fourth pair much shorter than second pair . iow

Fic. 3. — Protolophus singularis.

Femur I longer than body or in some females a little shorter, but

longer than width of body . Liobunum

Femur I shorter than body, in pater er not as uA as width of body

Hadrobunus

- Inner margin of patella extended; body very hard and rough; eye-

tubercle spinose: coxa III and IV in males enlarged. Trachyrhinus

Inner margin of patella not extended, or at least the >I quite

soft . 8

Legs iada i be as in ITUR coxa n not sisi out pus I

and III. Le

i uronychus

Legs much aier, i I dii or not as jong as width of

y; eye-tubercle smooth rae

Femora and tibia I and III thickened: coxa ati shorter’ than 1

urybunus

Femora and tibie I and III not thickened; coxa II about as

arge as I : v 5 c Laptobanus

Femur of pip oed s with prominent spines . . . . . - H

9 such prominent spines . . a

No false articulations in metatarsus E eyetberce m more peiie from

anterior margin. . Lacinius

At least one false ietibabiton Í in metatarsus I; : _eyetuberle nearer to

Magn o o a ae Vw . Oligolophus

674 THE AMERICAN NATURALIST. [Vor. XXXV.

12. Femora and tibiz I and III thickened; coxa II prape shut out

byIandIII . . Globipes

Femora Bax .tibize I dd I ditta; coxa qr not abit out by I

and II j Mery

13. Two prominent prioiaddibulas teeth ; [WES puncta: ,

—

No such teeth . . E ny oe ee

14. Femur I longer than width of ret nimus nti D cae Phalangium

Femur I shorter than width of body. in . |... 2 MEM

PROTOLOPHUS Banks:

Abdominal tubercles unarmed . P. tuberculatus Bks. dece, Tex.)

Abdominal tubercles with a few spinules, z^ palpi lar

P. singularis b (Calif.)

CADDO abe — o. evo 0C. agilis Bks. (N.Y, B C]

PHALANGIUM Linné :

Palpi very long, in male longer than body, in female longer than width

of body; the second joint of. male mandible is prolonged above in a

pur. n P. longipalpis Weed (Ark.)

Palpi uii Mite. in i fexise scarcely as long as width of body; no

spur on mandibles of male . . P. cinereum Wood (N. States)

MrroPus Thorell :

I. Tibia II is much longer than metatarsus II; one false articulation in

metatarsus I ; eye-tubercle about its daei from anterior margin

M. californicus Bks. idus /

Tibia II is subequal to or shorter than metatarsus 11

2. Gray and white, no false articulation in metatarsus I

M. dorsalis Bks. (Alaska)

Brown or black and White, two false articulations in metatarsus I

M. montanus Bks. (N.H.)

LaciNIUS Thorell : :

Metatarsi I and III -banded in middle; spines at tips of femora and

patella very prominent . . . L. texanus Bks. (Tex.)

Metatarsi I and III unbanded ; des joues spines at tips of femora

and patella less prominent . L. ohioensis Weed (E. States)

OniGOLOPHOR Koch. — . . —— — O. pictus Wood (N. E. States)

HoMoLopnus Banks . . , . . . gy biceps Thor.! (Colo. to Wash.)

GLOBES Banks... V. 27. . . G. spinulatus Bks. (Calif.)

LEPTOBUNUs Banks : ;

potes of coxz dark, tibia I with one dark band, palpus lineate with

brow L. borealis. Bks. (Alaska)

Apices wi coxae pale, tibia I with two dar bands; palpus not lineate

L. californicus Bks. (Calif)

1 Includes Z. punctatus Bks.

No.416] NORTH-AMERICAN INVERTEBRATES. 675

EunvBUNUS Banks

Body smooth ; ies IV nearly as long as II. E. drunneus Bks. (Calif.)

Body with transverse rows of spinules; leg IV much shorter than II

E. spinosus Bks. (Calif.)

TRACHYRHINUS Weed:

Coxe pale, maculate with brown, legs mostly pale

T. marmoratus Bks. (N. Mex.)

Coxe unicolorous, legs mostly black. . . T. favosus Wood (Colo.)

LEvuRONYCHUS Banks:

Body with a brown dorsal vitta, eges of legs brown lineate with

white . . L. pacificus Bks. (Calif., Wash.)

Body without vitta. puteis not t lineate. . L. parvulus Bks. (Wash.)

Fic. 4. — Lacinius texanus.

LioBuNUM Koch. Mates:

I. A distinct spur on the femur of palpus, dorsum yellowish, without

stripe, trochanters concolorous with coxa

L. calcar Wood (N. States)

No such spur . : mf c

2. Palpus, except PER wani black. 3a ro eter OS

Palpus yellowish or brownish =

3. Dorsum dark, often with two liegi pale pem behind, rochanters

usually pale. . L. exilipes Wood (W. Coast)

Dorsum pale yellowish, no stipe trochantes and bases of legs

gen igripalpi Wood (E. States)

4. Femur, ae xe tibia M palpis pus incrassate, dorsum with a

lack stripe, trochanters dark brown . Z. crassipalpis Bks. (D. C. ;

Palpal joints not incrassate .

5. Femur of palpus very long, cred adik much above surface oi

M LUE. dorsum with a distinct black stripe, trochanters

black L. vittatum Say ! (U. S.)

1 Includes Ziobunum dorsatum Say.

676 THE AMERICAN NATURALIST. [Vor. XXXV.

p»

-—-

Femur of palpus shorter, dorsum without distinct black stripe. he

Dorsum dark, with two large tig spots on union of cephalo-

thorax and abdomen. . . . L. bimaculatum Bks. d

Not so marked . * oy

Apex of tibia II kite, Kodimi blike ody aor

ME Weed (E. —

Apex of tibia II not white .

Apex of femur I white, E aw a black bani: coxa Be

outside . . . L. townsendi Weed (N. Mex., -

Femur I not so ar i

. Trochanters dark, Md in edk with the cox o uU

Trochanters pale, concolorous with coxæ . (CUBE

. Legs black, dorsum dark brown, eyotubercle slightly spinala

L. nigripes Weed (Ohio)

Legs pale, sometimes marked with black . «82 0 0n AD RN

d&n AS

Fic. 5. — 71 palliZ

. Femur I barely as long as body, tips of femora and tibiae dark brown

or black, eye-tubercle nearly smooth Z. formosum Wood (E. States)

Femur I plainly longer than body, eye-tubercle spinulate, legs not so

distinctly marked with black e

- Abdomen tapering behind, dorsum piden biis g (ini not yt black,

trochanters dark brown, of moderate size

L. verrucosum Wood (N. Y.)

Abdomen rather short and broad, legs very long, small species . 13

. Dorsum and trochanters light brown, bases of femora not black

L. bicolor Wood (S. States)

Dorsum yellowish, trochanters and bases of uie black

L. speciosum Bks. (Ala)

- Body short and broad, femur IV often black at base, small species

olitum Weed (E. States)

Body tapering behind, femur IV not black a at base, larger species "B

. Extremely spinose beneath, an TERUS line on last ventral segment,

large species. à 20.05 . L. flavum Bks. (S. States)

No. 416.) MORTH-AMERICAN INVERTEBRATES. 677

Moderately spinose, rather granulate, no impressed line, moderate

ee. 8 ; . . 5. . L. ventricosum Wood (E. States)

HapRnoBUNUS Banks :

Dorsum finely iE ; ek Bor in female not much marked with

brown at tips of join : H. grande Say (E. States)

Dorsum more einig wid: many itid round, pale spots; legs

shorter, more marked with brown

H. maculosum Wood (S. States)

MESOSOMA Weed. . . . . . . MM. niger Say (S. States, Neb., Dak.)

The Nemastomatida include three genera, which can be separated as

below :

PHandibles longer than the body, be Sia forward

(Ischyropsalinz) Taracus

Mandibles shorter than body, direcind downward . (Nemastomina) 2

Fic. 6. — Dendrol. "abili

2. Fourth joint of palpus much thickened . . . . . . Phlegmacera

Fourth joint of palpus scarcely thickened. . . . . . Nemastoma

hee eam Packard :

- Several false articulations in femora III and IV, eye-tubercle very

Bind . cordiam E gra Bks.! (Alaska

No false arcatons 1 in Telnet .

2. A pair of. ee erect spines on de anterior jäk of ‘the

Sidon : . P. occidentalis Bks. (Wash.)

No ih a spines . d ru p P. avian Pack? (Ky., N. Y., N. H)

1 Includes Zvmicomerus bispinosus Pavesi.

2 Includes Sabacon spinosus Weed.

678 THE AMERICAN NATURALIST. | [VOL. XXXV.

NEMASTOMA Latreille :

. Fourth joint of palpus less than twice as long as ” bs

bird Pack. -

Fourth joint of palpus twice as long as the fifth . .

2. Dorsum with some spines . . . . W. modesta Bks. “Calif, Woe

Dorsum without spines . . . . . . W. troglodytes Pack. (Utah)

TARACUS Simon:

1. Dorsum of abdomen spinose, mandibles pale in mae

mE Bks. aw

Dorsum of abdomen smooth, mandibles dark .

2. Mandibles smooth . . RU UE po Bks. Er

Mandibles rough, very Giai i spines on basal joint

T. packardi Sim. (Colo., N. Mex.)

The Trogulide in our fauna is represented by but two genera, both

restricted to the Pacific slope:

i. Eye-tubercle projecting in the form of a spoon, two spines at each side

on the anterior margin . . . Ortholasma

Eye-tubercle branched, a sng club at each side on the anterior

Waren oe. SR ate M LI Dendrolasma

ORTHOLASMA Banks . . . . . . . . . . O. rugosa Bka. (Wash)

DENDROLASMA Banks. . . . . . D. mirabilis Bks. (Calif. Wash.)

PRINCIPAL WORKS ON PHALANGIDA.

BANKS, N. The Phalangida Mecostethi of the United States. Trans.

Amer. Ent. Soc. Vol. xx (1893), pp. 149-152.

The Phalangine of the United States. Can. Entom. (1893),

pp. 205-211.

The Nemastomatide and Trogulidz of the United States. Psyche

(1894), pp. 11, 12; 51, 52.

WEED, C. M. A Descriptive Catalogue of the Harvest Spiders (Phalan-

giide) of Ohio. Proc. U. S. Nat. Mus. Vol. xvi (1893). PP- 543-

563, 12 pls.

New or Little-Known North-American Harvest Spiders. Trans.

Amer. Ent. Soc. Vol. xix (1892), pp. 187-194, 7 Pls.

Woop, H. C. On the Phalangez of the United States of America.

Commun. Essex Inst. Vol. vi (1868), pp. 10-40.

No.416]] MORTH-AMERICAN INVERTEBRATES. 679

PRINCIPAL WORKS ON SCORPIONS, SOLPUGIDS, AND

PEDIPALPI;

KRAEPELIN, K. Revision der Scorpione. /ahré. Hamburg. Teil I, Bd. viii

(1891), pp. 1-144; Teil II, Bd. xi (1894), pp. 1-248.

— —— Scorpiones und Pedipalpi. Das Tierreich, 8. Lieferung (1899),

Pp.

Putnam, J. D. The Solpugide of America. Proc. Davenport Acad.

Nat. Sci. Vol. iii (1882), pp. 1-149, 4 pls.

Woop, H. C. Descriptions of New Species of North-American Pedi-

palpi. Proc. Acad. Nat. Sci. Philadelphia (1863), pp. 107-112.

On the Pedipalpi of North America. Journ. Acad. Nat. Sci.

Philadelphia. (2.) Vol. v (1863), pp. 357-376.

1 This list was omitted in Synopsis IX.

Het.

VARIATION NOTES. — Nos. 4-6.

4. A Case of Polydactylism. — A number of years since, a case

of polydactylism attracted my attention, and in view of the value of

records of such cases I have decided to offer an account of the facts

as known to me. The case was that of a young man twenty-five

Years of age, a student in Hamline University. The accompanying

; : 681

682 THE AMERICAN NATURALIST. | [Vor. XXXV.

drawings are tracings made by running a pencil around the outlines

of the feet and hands. The right hand is entirely normal. The

left hand varies as to the little fingers, there being two digits. A

careful manipulation of the palm showed that there is a single meta-

carpal at the base, and distally two digits, each with three bones ;

the bones are entirely separate, though the skin grows across between

them on the level of the proximal joint. Both of the feet are six-

toed, the ulnar digit in each being doubled. This doubling was

found, by feeling through the skin, to begin with the digit, the meta-

carpal being single. The left foot bears a lump on the outside at

the base of the outer digit, seemingly due to an irregularity connected

with the variation presented by the toes. I was told that it had

been there from before the time of fitting the first pair of shoes.

There was no knowledge of a similar condition of the digits in the

family ; the grandparents were personally known to the young man

and nothing like it had been remarked among his relatives.

The case appears to fall in the class of cases noted by Bateson

(“ Materials for the Study of Variation," p. 345) as among the com-

monest forms of polydactylism, in which there is an extra minimus,

the metacarpals and metatarsals being normal. H. L. OSBORN.

s. Pedigree Mouse Breeding. — V. Guaita has bred white and

walzing mice through seven generations. There has been a loss of

fecundity, due to too close inbreeding. Tables are given showing

the ancestry of about three hundred mice of varying colors. The

author does not make sufficient use of his data. The most striking

fact is that when pure-bred white mice were crossed with pure-bred

walzers (chiefly black and white), all of the twenty-eight progeny

were gray, or like a house mouse, and none were walzers. When

these were bred together, nine color classes at once appeared, includ-

ing albinos and walzers, as well as grays and gray-blacks. If the

results of breeding in the later generations are compared with what

one should expect from Galton's law of ancestral inheritance, it

results that the albinos appear unusually stable and prepotent. On

the other hand, the walzing condition seems to be unstable and to

be less potent than normal.

! Von Guaita, G. Zweite Mittheilung über Versuche mit Kreuzungen von vet

schiedenen Hausmausrassen, Berichte d. maturf. Ges. Freiburg, Bd. xi, Heft 2,

pp. 131-138, 1899.

No. 416.] VARIATION NOTES. 683

6. The Long-Tailed Field Mouse! (Mus sylvaticus) has a distribu-

tion which is almost coterminous with the limits of the Palearctic

Region ; hence is a “ wide-ranging species” in Darwin’s sense. It

is also an old species, for its bones are found in the cave deposits.

It is much less variable than many other small mammals, and sports

especially are rare. Thus it is a case against Darwin's law of the

great variability of wide-ranging species and for Sedgwick's law of loss

of variability in an old species. However, albinos (with pink eyes)

have been noted as rare sports. The prevailing color is white below

and more or less rufous above. Since the young are more like the

house mouse in color, this color may be considered ancestral. The

adult mouse becomes small and dark in certain isolated maritime

localities (Lewis, Skye, Galway, and Kerry); a variation paralleled

by that of squirrels, cattle, birds, the slugs, Limax, and butterflies.

The mouse becomes rich-colored as it approaches the Oriental

Region, — the home of rich-colored birds, — for example, at Kuatun.

It is bright, clear-colored in the cold, drier regions of central Europe,

just as the squirrel and red-backed vole are.

! Barrett-Hamilton, G. E. H. On Geographical and Individual Variation in

Mus sylvaticus and its Allies, Proc. Zool. Jour., pt. ii, pp. 387-428, London,

August I, 1900

REVIEWS OF RECENT LITERATURE.

PHYSIOLOGY.

Heliotropism. — In this paper Holt and Lee! are chiefly concerned

with a theoretic discussion of the nature of the responses of organisms

to light ; but they present, also, in support of their theory, the results

of some experiments with Stentor Cceruleus, a species of Lynceus,

and some fresh-water planarians. The title of the paper, “ The

Theoty of Phototactic Response," is misleading, since the term

* phototactic" is used, not in the ordinary sense, but as including

those responses which have been called photopathic. In the present

state of our terminology **heliotropic " would be a better term.

Intensity of light and direction of ray have been championed for

half a century as the important factors of light as a. stimulus: at

one moment * direction" has been emphasized, at another “ inten-

sity" The present writers maintain that light acts only by its

intensity, the direction of the ray being a secondary factor and one,

furthermore, of an entirely different nature. Their theoretic discus-

sion centers about the views of two prominent physiologists, Loeb

and Verworn; and Verworn's theory is accepted as adequate for the

explanation of all responses to light. Loeb is characterized as a

believer in the importance of “direction,” and Verworn in that of

"intensity," but it is observed that their theories are not contra-

dictory. So far as the exposition of these two theories, as given

in the text, is concerned one seems just as applicable to the facts

in question as the other, the only difference being that Loeb has

sometimes used the term * direction " when intensity is evidently

the determining factor, thus in appearance giving emphasis to the

Importance of direction, while Verworn has in all cases expressed

himself in terms of intensity. i

The facts of response to light, admitted as such by Holt and Lee,

are thrée : (1) orientation in the axis of the ray, (2) positive or

negative movement, (3) random movement. In explanation it is

assumed that animals which show these three phases of reaction have

‘Holt, Edwin B, and Lee, Frederic S. The Theory of Phototactic Response,

Amer. Journ. Physiol., vol. vi, pp. 460-481, January, 1901.

685

686 THE AMERICAN NATURALIST. [VoL. XXXV.

an optimum intensity in which they are not directed by the light

but move at random. Below this is the sub-optimal and above it

the supra-optimal. An animal in either the sub- or supra-optimal

intensity is oriented by the rays so that symmetrical points of the

body are equally stimulated ; this is, of course, accomplished by

the placing of the longitudinal axis of the organism in the axis

of the ray. And whether an animal orients itself with head toward

or away from the source of light (7.4, whether it is positively or

negatively phototactic) depends upon the physiological condition

of the animal and the intensity of the light. By sub-optimal inten-

sities organisms are supposed to be directed toward the light through

the expansion of those motion-producing elements which are on the

side most strongly stimulated ; hence there results from this kind of

orientation a positive reaction. The same organism, if in a supra-

optimal intenSity, will be oriented with head away from the light,

because in this case contraction instead of expansion is caused, and

the reaction will be negative. The orientation theory has been very

clearly stated by both Loeb and Verworn. ;

Several different kinds of reactions, representatively selected, are

explained by the writers by their theory ; among them is the case

a positively phototactic animal moving toward the source of light

into a less intensely illuminated region. This reaction has been

taken heretofore as evidence of the independent influence of “ direc

tion" of ray. It is clearly shown in the present paper, however,

that such a contention is probably false, for difference in intensity and

the angle at which the organism strikes the side of the vessel are

sufficient to explain the observed courses taken under such conditions.

That light acts through intensity alone is a conclusion which this

paper makes plausible, but it scarcely justifies the unconditioned

statement that it does not act by the course which the rays take

through the organism. Such a reaction as the reversal of response

observed by Towle in Cypridopsis and by Yerkes in Daphnia and

Cypris is not easily explained by the hypothesis under consideration.

The paper is valuable in that it makes clear the importance of

intensity and at the same time indicates the danger of confusion

in using direction as a causal term, although intensity does in part

depend upon it. ; : R. M. Y.

Notes. — The chief defects of laboratory guides arise from the

difficulty of giving sufficient directions to the student without sup-

plying him with information that he should get from laboratory

No. 416.] REVIEWS OF RECENT LITERATURE. 687

study. This is the principal failing in Brown’s Physiology for the

Laboratory (Boston, Ginn & Co., 1900, viii + 167 pp.), which, how-

ever, is so well balanced in other respects that it deserves to be in

the hands of the teacher if not in those of the pupil.

Kelly Vena. Zeitschr., Bd. XXXV, p. 429) has pointed out that

calcic carbonate occurs in nature in five forms: calcite, aragonite,

ktypeite, conchite, and amorphous calcium carbonate. Of these

calcite and conchite are the only ones found abundantly in organ-

isms. Conchite is probably slightly more soluble, harder, and has

a higher specific weight than calcite. Both forms occur through-

out the animal series ; thus calcite is the mineral component of the

shells of echinoderms, brachiopods, crustacea, bryozoa, of the cal-

careous spicules of sponges, of the ear stones of fish and amphibia,

and of the eggshells of mollusks, most reptiles, and birds ; conchite

is characteristic of the skeletons of most stone corals, and the shells

of many mollusks.

Steinach and later Magnus have shown that the iris of a frog’s

eye will contract when stimulated by light, even after the eye has

been removed from the animal. Steinach believed this to be due to

the direct action of light on the sphincter muscle ; Magnus attributed

it to a short nervous reflex arc within the eye. Guth (Archiv ges.

Physiol., Bd. LXXXV, p. 119) finds that frog eyes show this reaction

fully two weeks after their removal from the animal,—a period much

longer than that during which other organs containing reflex arcs,

like the intestine, remain active. Moreover, pieces of the edge of

the iris as well as minute isolated groups of muscle fibres from the

Sphincter pupille contract on illumination. As the latter were

shown on microscopical examination to contain no ganglion cells, it

must be admitted that Steinach's contention that the muscles of the

ms are capable of being stimulated directly by light is correct.

BOTANY.

Recent Papers on Algae. — (Comére, Joseph. Les Desmidées

sal v Paris, 1901, 222 pp., r6 pls.) In the introduction the

Mis States his doüble purpose in writing this work ; first, to give as

account as possible of our present knowledge of the desmids ;

Second, a manua] for the study and determination of the French

688 THE AMERICAN NATURALIST. | [Vor. XXXV.

species. The first chapter deals with structure and physiology, the

second with methods of collection, preparation, and study ; then is

a historical sketch, with bibliography, dealing with France only.

Chapter 4 occupies the greater part of the book, giving the general

principles of classification, followed by description of all known

French species and forms. A table showing the distribution in the

different provinces of France and a list of all works cited complete

the text. There are sixteen plates, containing over eight hundred

figures.

It is somewhat singular, considering how much of our best algo-

logical work is done by French writers, that there has never been

any general systematic work on French algæ, such as Harvey's

Manual or Phycologia in Great Britain, Hauck's Deutsches Meeres-

algen in Germany, and similar works in other countries. This seems

to be the first work treating of alge, covering the whole of France,

and giving descriptions of all the species mentioned. It must

certainly be of much use to French students.

The classification follows in general that used by De Toni in the

Sylloge Algarum, but with some modifications. The tribe Docidez

is united with the Closteriz, while the Micrasteriz retain only the

forms with chromatophores as parietal disks, the forms having radial

chromatophores forming the tribe Cosmaria. The form of the

chromatophores is recognized as of value in classification, but the

author does not give it the value attributed by Gay.

A scientific work written by a Frenchman always has one great

advantage, the “inexorable clearness” of the language ; moreover,

a French author apparently does not feel that a good style of writ-

ing will cast doubts on his scientific thoroughness or soundness.

These merits the work in question has, and also the French pre

dilection for symmetry ; the description is a description, the note à

note, and one does not run into the other ; history does not intrude

into physiological discussion, nor bibliography into directions for

collecting.

The plates are clear, every species described being represented,

mostly by camera drawings by the author. In nearly every case à

colored figure is given to show the appearance of the living plant,

and a line drawing to show the markings of the empty cell. Only

two scales of magnification are used, 200 and 300, and the ne

scale is used in all the species of a genus, which is very convenient

for comparison. There is an index, both of accepted names and of

synonyms.

No.416.] REVIEWS OF RECENT LITERATURE. 689

The descriptions are sufficiently full, and appear to be accurate ,

occasionally a word is used in a sense that might be misleading, as

on p. 21, where certain desmids are said to occur *en parasites,"

on Sphagnum, etc. Full references are given to French authors,

and a considerable number to foreign works, but the English lan-

guage-seems to offer great difficulties to M. Comére, as it has to

many others of his nation. There are many references to Wolle's

Desmids of the United States, and this work, with three papers pub-

lished in the Zorrey Bulletin, appears in the bibliography. It is fair

to assume that M. Comére must have seen the book, but both in

references and bibliography the name is * Woole." Quite curiously,

at p. 98 there is a reference to another Torrey Bulletin article,

which does not appear in the bibliography, and in this one place the

name is properly spelled, * Wolle." “ Dysphynctium TÀwatesiz," with

synonym “ Calocylindrus Twatesii,” at p. 92, and * Twaites," p. 130,

represent three attempts on the name of a well-known botanist.

bs Barcker in Quartely Journal, p. 208,” is an interesting combina-

tion. A certain amount of carelessness in proofreading is noticeable

here and there: in the bibliography Reinsch is followed by a period,

as if an abbreviation, while Schaarschm has no period, though it

represents Schaarschmidt. Sutarastrum, Closterium monoliferum are

similar cases. But let him that never overlooked an error ona proof-

sheet cast the first stone at M. Comére.

The work makes no innovations and does not claim to add

largely to previous knowledge, but it puts in small compass, and in

shape convenient for the student, what otherwise would have cost

him much time and trouble. Any work doing this is valuable. The

nomenclature is eminently conservative; no new species is proposed,

nor 1s any one transferred from one genus to another; the author

must have nobly resisted’ the temptation to attach his name to

Something,

oS F. R. d Om Floridé-Slägtet Galaxaura dess Organo-

p ystematik," Kongl. Svenska Vetenkaps Akademiens Hand-

Sar, Bandet XXXIII. Stockholm, 1900. 109 pp., 20 pls.) The

_ of which this paper treats, has its home in tropical

Ma Cin waters, the broad belt occupied by it extending

in : m world, its northern limits being near the straits of

Sieg ins ^ the coast of Florida in the Atlantic, Mexico and

Misa: € Pacific. In the southern hemisphere it reaches similar

690 THE AMERICAN NATURALIST. [Vor. XXXV.

In Engler and Prantl’s aturfamilien Pflanzen the genus is noted

as having about twenty species, many of them little understood.

In examining the forms from Japan, collected by the Vega expedi-

tion and subsequently by Petersen, Dr. Kjellman found many that

could not be placed under known species, and was led to make a

thorough study of the genus. The result of this study may be seen

in the fact that, of the sixty-two species here recorded, only sixteen

are of previous authors; the remaining forty-six are new.

Habit characters, on which the previous classification was almost

entirely based, are here considered as of little value, and structural

characters are used instead, the descriptions being supplemented

by nineteen quarto plates of anatomical drawings by the author,

while a double plate gives habit figures, photographically produced

from herbarium specimens. The rich collections of. Areschoug,

Agardh, and the author himself, with those in the Swedish museums,

formed the material on which the work is based, and the fact that it

was all dried material in herbaria constitutes the one possibly weak

spotin the results. The ideal method of study would be to see all

the species in their homes, to notice the range of variation of a spe-

cies in different local conditions and at different seasons, and then,

with the results of this study clearly in mind, to refer to the original

types. But when this study of living material must be made over

half the surface of the globe, or at least the seacoast within that

area, it is evident that this ideal is hardly likely to be attained.

In the tables showing the distribution of the species, only three

are assigned to the coast of the United States, G. flagelliforme, G. stel-

lifera, G. umbellata. This number will have to be increased, as

on the shores of Florida there occur G. obtusata, G. rugosa, G. lapi-

descens, and G. marginata. The last two species, in the older sense,

are divided by Kjellman into several species each, so that the figure

for our coast must be seven, and may be one or two more.

The descriptions of the new species and the explanations of the

plates are in Latin ; the rest of the work, including all the historical

notes and the anatomical studies, are in Swedish, which is unfortu-

nate in a work of such general scope and interest.

(Svedelius, Nils. “Studier öfver Östersjöns Hafsalgflora,” Akade-

misk Afhandling: Upsala, 1go1.) The author has made a careful

study of the red, brown, and green algæ of the eastern and northern

part of the Baltic Sea. The flora is quite a limited one, only fifty-

six species being enumerated, there being a steady falling off in the

No. 416.] REVIEWS OF RECENT LITERATURE. 691

number of species to be found as we pass from the German Ocean

and along the Baltic to the Gulf of Bothnia. There is a steady

reduction in the saltness of the water in this direction until at

last it is almost imperceptible. Moreover, the east and west sides

of the Baltic have differing degrees of salinity, and sudden changes

in this, as also in the temperature, are produced by winds, conditions

quite unfavorable to the existence of most marine algz; hence the

poverty of the flora.

Two elements appear to compose the flora, an Atlantic and an

Arctic, but without any sharp division line. Four species, one of

them new, Ascocyclus affinis, are considered endemic, thirty-five are

. characteristically Atlantic, seventeen characteristically Arctic. Most

of the species appear in somewhat reduced or depauperate forms, in

comparison with normal conditions. This is not the case with the

green alge, however, which are fully as well developed as elsewhere :

not unnaturally, as the green algz are largely fresh-water plants, the

red and the brown being specially marine. Asin the Arctic regions,

Some species cover considerable areas of bottom in loose-lying

Panes, not attached to any substratum, new growth continually

forming as the old decays. These detached individuals are uni-

formly sterile, as is the case in the Arctic Sea, and also with the

s bna of By ie in the Atlantic. Green alga: are

Niel Sai littora region, brown in the lower littoral,

à al, in about the same proportions as in the

Atlantic.

i mai a number of illustrations, mostly habit illustrations,

iiio 2 a "ber of forms of Fucus vesiculosus, and transitions

fie site since names have been given these forms, it is

eS to illustrate them, but it is doubtful if they can be

Saini s all well defined. It is usual to include under

mile d all dicecious forms with vesicles, as well as those that

he M ht to have vesicles. It is not unlikely that more

ae pecies may be here included, but we are not yet in a

Position to draw the lines.

s : in Swedish, which is perhaps natural under the cir-

S ot its issue, but like Kjellman's work previously noted,

easy of use by the majority of botanists.

cumsta

itis n

(Børgesen, F, Fr

of the Feröes.

Sen's paper on th

eshwater Algæ of the Færöes. From the Botany

Copenhagen, 1901.) Together with Mr. Børge-

€ algz, there are printed E. Warming's * Historical

692 THE AMERICAN NATURALIST. [VoL. XXXV.

Notes on the Botanical Investigation of the Fzróes," and C. H.

Ostenfeld's * Notes on Geography, Climate, Topography, Geology,

and Industrial Conditions." Mr. Ostenfeld's notes are furnished

with excellent illustrations, so that we reach Mr. Bérgesen’s list

of the alga with a good understanding of the conditions that

have made the flora what it is. This flora must have been intro-

duced since the last glacial epoch, and, according to the writer, it

is probably more than anything else the birds that have gradually

contributed the various fresh-water algae found on these eighteen

rocky islands, only five of which have an area of over roo kilo-

meters each. In their migrations through Great Britain to the

Arctic region the birds make a stop on these islands, the distance

from the northernmost British isles being only a two hours' flight.

So we find two elements in the Ferée flora, a southern, agreeing

with that of Great Britain, and including some forms supposed to

be peculiar to the latter, and an Arctic element; the one brought by

the spring migration, the other by the return in autumn.

forms. The desmids, with 175 species, constitute more than half

of the whole, — the other -green algze number 103 ; there are forty-

one Cyanophycez ; one Nitella, one brown alga, Hydrurus fetidus,

and one red alga, Chantransia hermanni, complete the list.

Two species. of Enteromorpha, Æ. compressa and Æ. anicrococca

Jorma subsalsa, were found in brooks at a height of 200-300 meters.

The former has been considered exclusively marine, the latter has

been found in.brackish pools in the Arctic regions ; their occur-

rence in running fresh water is quite noteworthy. Beside several

new varieties and forms, two new species are described, Zuastrum

lyngbyei and Cladophora lyngbyei; a plate of the latter, however,

shows a form of branching which hardly accords with Cladophora;

it would seem as if the plant belonged rather in Siphonocladus.

There are four good plates and an excellent map, and the whole

is in unexceptionable English.

(Hirn, Karl E. « ‘Monographie und Iconographie der Oedogonia-.

cien," Acta Societatis Scientiarum Fennice. Helsingfors, 1900.) This

volume of 400 quarto pages and sixty-four plates is a notable addition

to the working tools of the systematic algologist. One hundred

and ninety-nine species of CEdogonium, forty-four of Bulbochete,

and one of (Edocladium are composed in the family. Full diag-

noses of all the species, with their many varieties and forms, M

No.416.] REVIEWS OF RECENT LITERATURE. 693

given in Latin, and there is an elaborate key to the species in

the same. The detailed notes under the species, as well as the

general chapters, are in German. These general chapters give a

full résumé, with illustrations in the text, of the structure, develop-

ment, and physiology of the (Edogoniacezm. Description, measure-

ments, synonyms, and localities are given in ample detail, and the

plates represent all the species and forms. The figures are in out-

line, and were drawn by the author from nature; all are of the same

scale, 300/1, except a few details requiring a higher scale, and one

habit figure of CEdocladium. This uniformity of scale is. a great

convenience in actual use of the work.

The standard work on this family has been Wittrock's Prodromus

Monographie GEdogoniearum, published in 1874. In this are repre-

sented 140 species, but only a single plate, of general types, accom-

panied the work. In a family like this, where the vegetative

Characters are of great simplicity, the need of accurate figures

increases even more rapidly than the number of species ; the best

of descriptions are insufficient. So that, though De Toni’s Sy//oge

Algarum gives descriptions of all species published up to 1889, it is

extremely difficult to determine species*by it. Dr. Hirn had the

advantage of study with Wittrock and Nordstedt, had opportunity to

examine type specimens of nearly all species, and had been receiv-

ing material from collectors in all parts of the world for a number of

years. As a result, the list of habitats includes all parts of the

earth; even South America, generally an algological zerra incognita,

being quite well represented. North America, though with few

exclusive species, shows quite a rich flora, but the author wisely

places a (?) against those references in Wolle which he has not been

able to verify by authentic specimens. Of the 244 species, forty-

five are described here for the first time. F. S. COLLINS.

Notes. — Biltmore Botanical Studies is the title of a new journal of

botany embracing papers by the director and associates of the Bilt-

more herbarium, the first number of which was issued on the 8th of

April Itis published at Biltmore, N. C. The first number contains

a revision of the species of Marshallia and descriptions of a consid-

able number of new species of other genera.

Fascicle 1 of Vol. II of Plante Bakeriane, by Professor E. L.

Greene and others, covering fungi to grasses of Mr. Baker’s collec-

tions of 1899, has been issued under date of March 11.

694 THE AMERICAN NATURALIST. [Vou. XXXV.

The first part of the “ Illustrations of the botany of Captain Cook's

voyage round the world in H.M.S. Zzd4eavour, in 1768-71,” by Banks

and Solander, with determinations by James Britten, recently issued

by order of the Trustees of the British Museum, contains one hun-

dred folio plates representing Australian plants.

Parts II and III of Mr. F. Manson Bailey's Queensland Flora, pub-

lished under the authority of the colonial government, have been

issued and cover the orders Connaracez to Gentianacez, inclusive.

A new edition of Pond and Clements's Phytogeography of Nebraska

has been brought out. Only one hundred copies are offered for

sale, and as the work possesses real and general merit, though its

purpose is local, the edition is likely to be soon exhausted.

Dr. Holm publishes a fifth list of additions to the flora of Wash-

ington, D. C., in the Proceedings of the Biological Society of that city,

under date of April 2.

A supplement to Zhe Flora of the Upper Susquehanna and its Trib-

utaries, by Willard N. Clute, is published by the author at Bingham-

ton, N. Y., under date of February rs.

The opening part of Vol. XV of the Zrazsactions of the Linnean

Society consists of a report on botanical collections from Mount

Roraima, in British Guiana.

An article on the determination of the type in composite genera

and species, by President Jordan, appears in „Science for March 29.

A paper on the Ranunculacee of Iowa, by T. J. and M. F. L.

Fitzpatrick, is reprinted from the Zu//etiz of the laboratory of natural

history of the University of Iowa.

The germination of Bertholletia excelsa is described in a well-illus-

trated article by William Watson, of Kew, in the Annals of Botany

for March.

A paper dealing with American plants that is likely to be over-

looked is E. H. L. Krause’s * Nova Synopsis Ruborum Germanic

et Virginiz," the first part of which was published by the author in

1899 at Saarlouis.

In an account of a collection of Crategus from near Montreal,

made by Mr. Jack, Professor Sargent, in Rhodora for April, describes

six additional new species.

No.416] REVIEWS OF RECENT LITERATURE. 695

Dr. Rydberg describes further new Potentillas in the Bulletin of the

Torrey Botanical Club for March.

Eupatorium boreale is the name proposed in Rhodora for April by

Professor Greene for what commonly passes in New England as

E. ageratoides.

The genus Teucrium, as it is represented in the eastern United

States, is passed in review by Bicknell in the Bulletin of the Torrey

Botanical Club for March. :

A supposedly new horse gentian is described from the northern

states by Bicknell, under the name 77iosteum aurantiacum, in Torreya

for March.

Miss Eastwood describes some small-flowered Nemophilas from

the Pacific coast in the Bulletin of the Torrey Botanical Club for

arch

Engler's Botanische Jahrbiicher of March 12 contains a paper by

Lopriore on the geographical distribution of Amarantacee with

reference to their relationships.

Pentstemon heterophyllus is illustrated in the Revue Horticole of

April 1.

Ale Lynchii, an artificial hybrid between Ale striata and Gasteria

verrucosa, is figured in the Gardeners’ Chronicle of March 30. As

with most bigeneric hybrids, this is a cross between representatives

of genera which, though logically separable, are capable of treatment

as sections of a single genus. :

A number of new or little known grasses are described in Circular

Wo. 30 of the Division of Agrostology of the United States Depart-

ment of Agriculture.

A portrait of Mr. George E. Davenport appears as the frontispiece

to The Fern Bulletin for April.

Several American species of Cypripedium are photographically

illustrated in Die Gartenwelt of March 16, in an article on their

cultivation.

Poisoning by the pileate fungi, which has recently been exhaust-

ively treated by Gillot in a thick volume published by the house of

P Klincksieck of Paris, is the subject of a number of recent notes

In the Bulletin de la Société des Naturalistes de ? Ain. Of 222 cases

of Poisoning, the records of which were examined by Gillot, 86

resulted fatally, and of these 2 were doubtful and the other 84 were

due to Amanitas or Volvarias.

696 THE AMERICAN NATURALIST. — [Vor. XXXV.

Professor Peck's quarto plates illustrating the edible fungi of New

York, with a number of additional plates and revised letter-press,

have been issued as Vol. IV, No. 3, of the Memoirs of the New York

State Museum.

A study of Boletus luteus, deformed by the parasitic Hypomyces

chrysospermus, by Van Bambeke, appears in the recently issued

Bulletin de la Société Royale de Botanique de Belgique for 1900.

A new species each of Tolyposporium and Ustilago in the ovaries

of Eriocaulon septangulare, from Massachusetts, is described by

Clinton in Rhodora for April.

The cambial slime diseases of trees are being rediscussed by

Dr. Holtz in current numbers of the second Abteilung of the Central-

blatt fiir Bakteriologie, etc.

Dr. E. F. Smith has published an extended and well-illustrated

résumé of what is known of bacterial plant diseases, in recent

numbers of the Centralblatt für Bakteriologie, Abteilung 2.

A paper on the organography and taxonomy of Galaxaura, a genus

of red alge, by Kjellman, is separately issued from the Æ. Svenska

Vetenskaps- Akademiens Handlingar, Nol. XXXIII, No. 1.

Nowhere have ecological facts found so good exposition in garden-

ing as at the Berlin botanical garden, where, from the moment of

assuming the direction of the establishment, Dr. Engler has devoted

himself to a development of this most important museum feature of

a botanical garden. One of the most instructive of recent treatises

on the ecological distribution of plants is issued as Appendix No. 7

to the current volume of the (Vofizb/att of the Berlin garden, and con-.

sists in a terse logical analysis of the plant -formations of the Alps

as exemplified in the newly established garden at Dahlem.

An extensive and well-illustrated treatise on the dissemination

ecology of Scandinavian plants, by Dr. Rutger Sernander, is dis-

tributed from the University of Upsala, and forms an octavo of over

450 pages, the Swedish text being accompanied by a German summary.

A good example of what Huth has called stem-fruiting plants ux

given in a plate of Artocarpus integrifolia, published by Dr. Wilcox

in the Proceedings of the Columbus Horticultural Society for 1900.

R. E. B. McKenney publishes some illustrated ecological notes 07

plant distribution in the Beihefte zum Botanischen Centralblatt, Bd. X,

eit 3

No. 416. REVIEWS OF RECENT LITERATURE. 6

Dr. Clements, of the University of Nebraska, is to conduct a class

in field ecology in the higher mountains of Colorado next summer,

extending through the months of July and August. The class should

be large and enthusiastic.

The photosynthetic activity of chlorophyll occurring below corky

tissue in the stems of several plants is discussed by Mlle. Goldflus

in the Revue Générale de Botanique of February 15.

The apparatus of transpiration, or sudation, as the author prefers

to call it, is discussed by Goffart in Vol, XXXIX of the Bulletin dela

Société Royale de Botanique de Belgique.

“The Indian doctor's dispensatory, being Father Smith's advice

respecting diseases and their cure, by Peter Smith of the Miami

country," published in Cincinnati in 1812, is reprinted as No. 2 of

the “reproduction series” of the Buletin of the Lloyd Library. It

is accompanied by a biography by John Uri Lloyd.

A neat little book on alpine plants and their cultivation, by W. A.

Clark, has been brought out for the author by L. Upcott Gill of

London and Charles Scribner’s Sons of New York. It is illustrated

by a number of exquisite half-tones showing some of the choicer

species as grown in artificial rockeries.

Portraits of a number of the American botanists of the last two

centuries are published in the recently issued second part of the

Report of the U. S. National Museum for 1896-7.

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846. "gestu A. The Study of Children. Everybody's Magazine. Jem

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5 pls. —Mason, O. T. A Primitive Frame for hune. Narrow hie Rept.

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Variations of a newly arisen Species of Medusa. Bull Mus. Brooklyn Inst. A.

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Contribution to the Geology of the John Day Basin. Univ. Cal., Dept. Geol.,

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rex. Journ. Anat. and Phys. Vol. xxxv, pp. 405-412, Pl. XXXIX. — SMITH,

H. S. Prehistoric Michigan. Pop. Sci. News. May, 1901. Pp. 110, 111, 6 figs.

— STEJNEGER, L. Description of a New Species of Snake from Clarion Island,

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THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

L The Compound and Mixed Nests of American Ants. Part II (Continued)

Professor W. M. —

II. Synopses of North-American Invertebrates. XVII. The

Rotatoria . ‘ . . ^. ^. . Professor H. 8. JENNINGS

III. Reviews of Recent Literature: Zod/ogy, Reptilian Affinities of Primitive

Mammals, Anatomical Miscellanies, The Position of the Centrosome in

— Botany, Bergen’s Foundations of Botany, Notes = - + + is

IV. Correspondence: Corrections in Nomenclature of Hydroids

Professor

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VOL. XXXV, NO. 417 ^^ Se SEPTEMBER, Igor

701

The American Naturalist.

ASSOCIATE EDITORS:

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E. A. ANDREWS, PH. D., Johns Hopkins University, Baltimore.

WILLIAM S. BAYLEY, PH.D., Colby st Neel

CHARLES E. BEECHER, Pu.D., Yale University, New Haven.

DOUGLAS H. CAMPBELL, PH.D., Stanford rH

J. H. COMSTOCK, S.B., Cornell University, Itha

WILLIAM M. DAVIS, M. E., Harvard Uem site Conil

ALES HRDLICKA, M.D., New York Ci

D. S. JORDAN, LL.D., Stanford Universi n:

CHARLES A. KOFOID, Pu.D., University of Illinois, Urbana.

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H. M. RICHARDS, S.D., Columbia University, New York.

ISRAEL C. RUSSELL, LL.D., University of Michigan, Ann :

ERWIN F. SMITH, S.D., U. S. Department of Agriculture, Washington.

LEONHARD STEJNEGER, Smithsonian Institution, Washington.

W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, Pu.D., University of Nebraska, Lincoln.

WILLIAM M, WHEELER, PH.D., University of Texas, Austin.

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THE

BMERICAN NATURALIST

Vor., XXXV. September, 1901. No. 417.

THE COMPOUND AND MIXED NESTS OF

AMERICAN ANTS.

WILLIAM MORTON WHEELER.

PART II (continued).

V. Du tosis.

UNDER this heading we may include all those remarkable

mixed nests which owe their origin to the enslavement of one

species of ant by another. This condition is characterized by

Wasmann (91, p. 43) as follows: ‘Here ants of different

Species dwell together, not only on the same spot, but coalesce

to form one colony, a single social whole. In such communi-

ties the unity of the colony is of paramount importance, and

the specific differences between the various components of the

Colony lapse so far into abeyance that they appear to be non-

existent ; the consociating ants, belonging originally to differ-

ent nests, behave towards each other as if they were kith and

kin, and carry on in common the construction of the nest, the

Acquisition of food, the education of the offspring, the defense of

the nest, etc., so far as this is permitted by their physical and

. PSychical endowments and the law of the physiological division

701

702 THE AMERICAN NATURALIST. (Vor. XXXV.

of labor. Hence the term ‘slaves’ is much less appropriate than |

the term * helpmates ' (auxiliaries)."

While the various forms of social symbiosis hitherto con-

sidered may exist between ants belonging to very different

taxonomic subfamilies, dulosis is known to occur only between

species of the same subfamily. This rule is based on but few

cases, for only four genera, two belonging to the Camponotinz

and two to the Myrmicinz, vzz., Formica, Polyergus, Strongy-

lognathus, and Tomognathus, are known to contain dulotic

species. Still it seems obvious that such close symbiotic

relationships as those under consideration could be entered |

into only by species of very similar habits and phylogenetic

derivation.

With the exception of Strongylognathus, the above-men-

tioned genera are all known to occur on our continent, the

dulotic species of Formica (F. sanguinea Latr.) and Polyergus

(P. rufescens Latr.) being represented by distinct races, or sub-

species, the genus Tomognathus by a distinct species (7. ameri-

canus Emery). So little attention, however, has been devoted

to our ants, that we may yet look forward to the discovery of

an American Strongylognathus, for there are in America sev-

eral species of the genus Tetramorium (including the sub-

genus Xiphomyrmex) which are allied to the Tetramorium

cæspitum auxiliary of the European Strongylognathus.

The meagre work which has been done on our American

dulotic ants is barely sufficient to show that their behavior 1$

essentially like that of their European allies. Since these ants

in America select their auxiliaries, or slaves, from à slightly

different though allied ant fauna, we may yet expect to find

some interesting differences in the details of habit and

behavior. ;

Before enumerating the American species, together with

their auxiliaries, it will be convenient to present a much con-

densed résumé of the splendid accounts of the European

observers, Forel, Wasmann, and Adlerz. :

Formica sanguinea Latr.— This species is a true Formic

which is sometimes found living without auxiliaries. It has

broad, toothed mandibles, of the type characteristic of

No. 417.] NESTS OF AMERICAN ANTS. 703

genus, and is naturally carnivorous, though it has been

observed to attend aphides for the purpose of collecting their

sugary excrement. Although this ant is, therefore, quite able

to exist alone, it nevertheless has a very pronounced penchant

for robbing the larvæ and cocoons of other species of Formica,

eating great numbers of them but allowing others to develop

and to function as its slaves, or auxiliaries. The latter feel

themselves to be members of the colony in which they emerge

from their cocoons, and direct all their activities to maintaining

and defending their foster nest and its occupants. In Europe,

as a general rule, the normal slaves of F. sanguinea are the

workers of F. fusca, less frequently the workers of F. rufibarbis.

Sometimes both species of auxiliaries may be found in the

same mixed nest. In extremely rare instances the workers of

F. rufa and F. pratensis may serve as slaves. The expeditions

for robbing cocoons are usually carried out during July and

August, but they seem to be rather infrequent or irregular and

are not often observed. The tactics of F. sanguinea, like those

of other dulotic ants, consist in surprising the colony they

wish to rob and in carrying away the pupa as rapidly as pos-

sible without engaging in unnecessary slaughter. Only the

ants that offer active resistance are dispatched.

F. fusca is most frequently enslaved because it is a weaker

and more tractable species and forms smaller colonies than

F. rufibarbis. The rare occurrence of F. rufa and pratensis in

sanguinea nests is due to the more savage nature of these

Species, which are enslaved only when they belong to small

colonies or when they are of small size individually.

The number of auxiliaries in nests of sanguinea varies

greatly. In Holland, in more than 100 nests, Wasmann (91)

found the ratio of sanguinea to slaves varying between 1:0 and

1:3. Most frequently the sanguinea are from 2 to 5 times

as numerous as their slaves. The number of the latter de-

pends on various circumstances, such as the abundance or

Scarcity of nests of the auxiliary species in the vicinity. It is

à singular fact that the weakest colonies of sanguinea contain

the greatest number of slaves, so that it would seem as if the

dominant species tried to make good the deficiency in the

794 THE AMERICAN NATURALIST. [VoL. XXXV.

number of its workers by importing and employing foreign labor.

This may result naturally from the fact that in weak colonies

on an average a larger percentage of the stolen pupæ are per-

mitted to develop into slaves. In populous sanguinea colonies,

on the other hand, a considerable portion of the prey is de-

voured even when thereis plenty of other insect food within

reach. ;

The relations implied by the terms “slave” and * master"

do not adequately express the conditions existing in these

mixed nests, since sanguinea works side by side with its auxil-

iaries, which are neither a mere luxury nor an absolute neces-

sity. Still, although sanguinea is capable of excavating and

maintaining its own nest, the auxiliaries appear to be more

enthusiastic and skillful workers in the earth. And although

sanguinea looks after its own brood and the hatching of the

cocoons of the auxiliary species, it must, nevertheless, derive

some advantage from the assistance of its slaves. The latter,

moreover, bring into the nest a good deal of food from the

aphides, which they assiduously attend.

F. sanguinea, on moving to a new nest, usually carries its

slaves, and is rarely carried by them. This is probably due to

the fact that the sanguinea are of a more excitable tempera-

ment and therefore have a greater tendency to take the initia-

tive in a change of dwelling than their more stolid auxiliaries.

Continental authorities uniformly maintain that the n

guinea-fusca nests contain only workers of the auxiliary species.

In England, however, Rev. T. D. Morice (00, p. 98) recently

found a nest which contained also fusca males and queens in

addition to the workers of this species. This very exceptional

condition would seem to have arisen either from the failure of

the sanguinea to consume all the pupz of the fertile sexes of

fusca, or less probably from the formation of an alliance colony

between fertile queens of sanguinea and fusca.

Polyergus rufescens Latr. — The * amazon," as the paragon

of dulotic ants, has been observed with great care by à num

ber of investigators, among whom Pierre Huber (10) and Forel

(74) hold the first place, It is a rather large, brown-red ant,

allied to Formica, but characterized by the possession of slender,

No. 417.] NESTS OF AMERICAN ANTS. 705

sickle-shaped mandibles, the cutting edges of which are fur-

nished with minute serrate teeth. Such mandibles are beauti-

fully adapted to fighting, but scarcely fitted for the many other

uses to which these organs are put by most ants. Polyergus is

therefore a warrior, and on this account its life presents two

very different phases, one replete with the brilliant tactics

whereby it gains possession of the larvae and cocoons of its

Fic. 15. — a., Polyergus rufescens Latr., subsp. dreviceps Emery, worker ; $., mandible of same ;

c., Formica fusca L., var. subsericea Say, subvar.; d., mandible of same.

Slaves, the other characterized by abject helplessness and com-

plete dependence on these same auxiliaries. -

The auxiliaries of Polyergus are furnished by the very same

Species as in the case of F. sanguinea. In this case, also,

F. fusca is most often victimized, zufibarbis less frequently.

Occasionally, too, both species are found in the same nest.

he number of slaves, however, is much greater than in san-

guinea nests, being about seven-eighths of the entire colony.

The dulotic expeditions of Polyergus have been often observed

706 THE AMERICAN NATURALIST. | [Vor. XXXV.

since the days of Pierre Huber (10, p. 210 eż seg.). They have

been admirably described by Forel (74), who has also estimated

(pp. 320, 321) the number of expeditions undertaken by a

single powerful colony of these ants during a single summer.

In thirty days (from June 29 to August 18, 1873) he witnessed

forty-four expeditions of the amazons. These usually occurred

between 2 and 5 o'clock P.M., the time limits being from 1.30 to

6p.M. Among the forty-four expeditions there were forty-one

attacks, nineteen on fusca and nineteen on rufibarbis, and three

of which only the return was observed. The total number of

cocoons robbed was estimated at 29,300 (14,000 fusca, 13,000

rufibarbis, and 2300 of unknown origin but probably fusca).

Counting in the expeditions after August 18, which he was

unable to witness, Forel concludes that not far from 40,000

larvae and pupa of the auxiliary species were appropriated

during the summer of 1873 by a single Polyergus colony ! Most

of the pupze were consumed, so that few of them ever hatched

and became auxiliaries. And although two species were pillaged

the colony later became almost entirely F. fusca.

Wasmann (91, pp. 61, 62) has observed that the fusca auxilia-

ries are noticeably: fiercer and more courageous than when

nesting alone. The same is true of fusca in sanguinea nests.

This is explained by Wasmann as merely a special case of the

general rule that all ants are more courageous when they feel

themselves backed by numbers.

The shadow side of the life of Polyergus is seen within its

nest, where it is abjectly dependent on its slaves. Here e:

spends most of its time preening its legs and antennz, as it is

quite unable to excavate. On this account the character of

the nest architecture is entirely determined by the auxiliary

species. Moreover, the conformation of its mandibles is such

that Polyergus cannot care for its own young or the pup® of

its slaves, though it sometimes licks the newborn callows.

fter a minute investigation of the question as to whether

Polyergus is able to feed itself, Wasmann concludes that it can

lap up liquids but is usually fed by the slaves. This mode of

obtaining its food is, in fact, so essential, that it dies of starva-

tion when deprived of its helpmates.

No. 417.] NESTS OF AMERICAN ANTS. 707

Polyergus goes on its cocoon-robbing expeditions unattended

by its auxiliaries. When the colony moves to a new nest the

Polyergus are nearly always carried by their slaves (cf. F. san-

guinea!) In this case the slaves commonly initiate the

change of dwelling. At home the Polyergus appear to be

under the guardianship of their slaves and to be treated

like helpless dependents. They are sometimes even held

back from their sorties by the auxiliaries.

The way in which the fertilized Polyergus queen starts her

colony has not been observed. Forel and Wasmann have

demonstrated that a friendly alliance may be easily effected in

artificial nests between Polyergus queens and strange workers

of F. fusca, and Wasmann concludes from this fact that new

mixed colonies may be started by such consociations under

natural conditions. But it does not appear to be necessary to

accept this inference. The fertilized Polyergus queen may be

quite as well able as other queen ants to raise unaided an

incipient colony of small workers which could then pillage

adjacent nests of fusca and provide themselves with the neces-

sary auxiliaries. It has, moreover, been observed that Polyer-

gus queens occasionally accompany the workers on their raids,

and this habit may be still more pronounced in the queens of

incipient colonies. I deem this probable because the young

queens of other species very generally perform nearly all the

functions which are later delegated more or less completely to

the workers alone.

Tomognathus sublevis Mayr (Fig. 16). — This is a small,

rather hairy ant, with broad and edentulous mandibles. It

occurs only in northern Europe (Finland, Sweden, and Den-

mark). Two very careful studies of its habits have been pub-

lished by Adlerz (86 and '96). The auxiliaries are furnished

by Leptothorax acervorum or L. muscorum, more rarely by

L. tuberum. | Adlerz's observations show that the Tomognathus

Secure these auxiliaries by attacking a Leptothorax colony,

driving away the ants, and taking possession of the nest,

together with the larvae. The latter are then reared as help-

mates, It is probable, however, that the T omognathus may

occasionally recruit the number of their slaves by making

708 THE AMERICAN NATURALIST. [VOL. XXXV.

sorties like Polyergus, for Adlerz succeeded in finding a nest

of Tomognathus with two species of auxiliaries (L. acervorum

and muscorum).

The mixed nests of Tomognathus-Leptothorax may contain

males, queens, and workers of both the dominant and victim-

ized species, a condition not known to occur in the case of

other dulotic nests. The males of Tomognathus (Fig. 16, a)

resemble the males of Leptothorax so closely that Adlerz failed

Fic. 16.— T'omognatA blevis Mayr (after Adlerz); æ., male; 3., female (ergatoid).

to distinguish them till he published his second study (96).

The female is also of such a remarkable character that it, too,

was originally overlooked. This sex is apterous (Fig. 16, 4)

and resembles the worker except in possessing ocelli and à

receptaculum seminis. :

The industrial instincts of Tomognathus are very rudi-

mentary. It rarely or never excavates. It is able to feed

itself if food is within reach, but it does not go in quest of

provisions. This it leaves to the Leptothorax auxiliaries, by

whom it is usually fed. Occasionally it may be seen caring

for the larvæ. A number of Tomognathus which were isolated

with larvæ and some food managed to live for 135 days, but

the larvæ died or shriveled up. It seems probable, therefore,

that Tomognathus depends on its slaves to a certain extent

even for the care of its larvæ. When the colony is compelled

to move to a new nest, the Tomognathus are usually deported

by the Leptothorax ; only rarely are the rôles reversed.

No. 417.] NESTS OF AMERICAN ANTS. 709

Sometimes when they desire to leave the nest, the Tomognathus

are detained by the auxiliaries in much the same manner as

Polyergus.

The males of Tomognathus do not mate with the females of

the same nest, but do so readily with the virgin queens of other

nests. The larva are so similar to those of the Leptothorax

that Adlerz was unable to distinguish them. They are nour-

ished with both liquid and solid food. Adlerz's description of

the manner in which the larve are fed with pieces of flies

tallies exactly with my observations on the primitive method

of feeding adopted by the Ponerinz and the Myrmicinz of the

genera Stenamma (Aphzenogaster) and Pheidole (00 and '00^).

Strongylognathus. —'The species of Strongylognathus have

perfectly edentulous, falcate mandibles, and more or less pro-

jecting postero-lateral corners to the head. Four species of

the genus are known to occur in Europe: S. huberi Forel,

S. testaceus Schenk, S. christophi Emery, and S. cecilie Forel.

The habits only of the first two have been observed (Forel,

74 and '00*; Wasmann, '91) and these present interesting

differences. Both species form mixed nests with Tetramorium

cespitum: S. huberi in southern Europe and northern Africa,

and S. ¢estaceus in southern and central Europe.

S. huberi seems like a diminutive and feeble caricature of

Polyergus. Forel (74) found by experiment that it would rob

the larvae and pupz of Tetramorium and fight with the rightful

owners after the manner of Polyergus. But it is not known

whether S. Zuberi under natural conditions really provides its

nests with auxiliaries by carrying on regular marauding expedi-

tions. Forel, in a more recent paper on this species ('00%,

P- 275), expresses the opinion that it may not make sorties but

keep up the mixed colony by alliance with the Tetramoriums

Instead. The workers of Strongylognathus are able to exca-

vate, but they are fed by the Tetramorium workers. The

latter are present in considerable numbers in the mixed nests,

but up to the present time fertile queens of Tetramorium have

not been found with them, though from what is known of

S. testaceus one or more of these queens may perhaps be

Present in some cases.

710 THE AMERICAN NATURALIST. [Vor. XXXV.

S. testaceus is a commoner and better known form than

S. huberi, and is supposed to represent a further advance

towards a condition of social parasitism. The number of workers

of S. testaceus is decidedly smaller in proportion to the number

of Tetramoriums. On this account Forel maintains that the

worker cast of S. ¢estaceus is on the road to disappearing

(cf. Anergates !). As fighters these workers, though provided

with sabre-like mandibles, are indeed but sorry caricatures of

Polyergus and decidedly less valiant than the workers of

S. huberi. They do not kill the Tetramoriums, but seem to

frighten them into deserting their larvae and pupz. Their

weakness is further shown by the fact that they do not under-

take their pillaging expeditions alone, but accompanied by their

Tetramorium auxiliaries, and it is these latter that determine

the success of the enterprise undertaken for the sake of rob-

bing their own species. The workers of S. Zeszaceus are even

awkward in their attempts to carry away the conquered larva

and pupe. Although the Tetramorium auxiliaries commonly

do all the work within the nest, such as excavating the galleries,

caring for the larvae and pupz, and feeding the Strongylog-

nathus, the latter are, nevertheless, able to feed themselves

and to dig the nest, but they are apparently unable to care

for the young.

Forel and Wasmann have succeeded in throwing consider-

able light on the obscure problem of the origin of the S. tes-

taceus-Tetramorium colonies. The former found a single

fertile queen of the Strongylognathus living amicably in the

midst of a colony of Leptothorax acervorum ; and Wasmann

made the significant discovery of a fertile queen of the Stron-

gylognathus and a fertile queen of Tetramorium living side by

side in the same nest. This nest contained workers of both

species (15,000-20,000 Tetramoriums and some thousands of

Strongylognathus), and pupz, about 70% of which were males

and females of Strongylognathus. The remainder included

two large male pupz of Tetramorium. From this discovery

Wasmann infers that the mixed nests of .S. testaceus-Tetra-

morium are alliance colonies brought about by the adoption of

fertilized queens of Strongylognathus by Tetramorium colonies.

No. 417.] NESTS OF AMERICAN ANTS. pet

The fact that these mixed nests rarely contain male Tetramo-

riums and never, so far as known, queen pupz of this species,

is explained by Forel (00%, p. 273) as the result of a general

regulative instinct : * The females and males of Strongylog-

nathus are smaller and less troublesome to nourish. This is

obviously sufficient to induce the Tetramorium workers to rear

them in the place of their own enormous queens and males,

the larvae of which they therefore undoubtedly devour or

neglect, as they do in the case of all that seems to be super-

fluous."'

After this brief review of the European species we may turn

to our American dulotic ants.

16. Formica sanguinea Latr., subsp. rubicunda Emery.

Although the typical F. sanguinea is not known to occur in

America, the species is, as Emery has shown (932, p. 647), far

more variable on this continent than it is in Europe. We

should therefore be prepared to find a corresponding variability

in its instincts, though this may not be commensurate with its

taxonomic variation.

F. sanguinea is also occasionally found without slaves in

America, but far more frequently it is attended by ants belong-

ing to the great group of forms which centers about F. fusca.

The best known subspecies of F. sanguinea in the Northern

and Atlantic States is undoubtedly rubicunda. This is usually

found with slaves belonging to F. fusca, var. subsericea Say, but

one colony which I observed near Rockford, Ill, Sept. 16,

1900, contained about equal numbers of auxiliaries belonging

to two species, viz. F. pallide.fulva Latr., subsp. nitidiventris

Emery, and F. fusca, var. subenescens Emery.

The above assumption that the habits of /. sanguinea in

America may differ to some extent from those of the European

form seems to be borne out by some recent observations of

Forel (00v, pp. 11-12). Owing to Forel's long and very inti-

mate acquaintance with the European sanguinea, these obser-

vations on our American form are of great value. He had

Occasion at Cromwell, Conn., to witness the attack of a very

712 THE AMERICAN NATURALLIST. [Vor. XXXV.

small troop of F. sanguinea (probably rubicunda) on a large

formicary of F. subsericea. “ There were scarcely thirty F. san-

guinea, and a third of these were recently hatched workers,

still immature. The troop was evidently from an incipient

colony. The subsericea had their nest about the roots of a

great mullein (Verbascum). Their numbers were at least ten

times as great as that of their assailants, and it may be

admitted that each of them was fully as well armed and on the

average larger and more robust than the sanguinea. Well, the

mere arrival of the little troop of sanguinea sufficed to spread

consternation through the nest of the subsericea, which betook

themselves to flight with their larvae and pupæ, but permitted

the sanguinea to snatch these away and to conquer their nest

without even making a serious show of defending themselves.

Not more than one or two small sanguinea were killed in the

fray. This fact is of importance, for in this instance we can-

not allege the redoubtable weapons, hard integument, or even

the impetuosity of the analogous attacks of the little troops of

Polyergus rufescens which I have described in my * Fourmis de

la Suisse.” The bold and courageous tactics of the sanguinea

were even less noticeable than in the European form of this

species, which wages war on smaller and more feeble species

than itself. I have never yet seen such complete and absurd

cowardice as that of the American subsericea, a cowardice

which brings clearly into prominence the instinctive adaptation

to attack on the part of the enslaving, and to flight on the part

of the enslaved species.”’

At Colebrook, Conn., during August, 1900, I had an oppor

tunity to see a colony of rubicunda moving to a new nest.

Each of the ants was carrying a motionless, curled-up F. sub-

sericea in its jaws. The rather open phalanx of ants presented

a very striking appearance as it moved from a shady hedge

where the old nest was located, across a dusty road and dis-

appeared in the undergrowth of a wood on the opposite side.

No. 417.] NESTS OF AMERICAN ANTS. 713

17. Formica sanguinea Latr., subsp. rubicunda Emery, .

var. subintegra Emery.

This variety, originally found by Mr. Pergande in the Dis-

trict of Columbia, has the same slave as the preceding, viz.,

F. subsericea. During July, 1900, I found a very large nest of

this variety on Naushon Island, Mass. It contained the usual

auxiliaries and was compounded with a large nest of Solenopsis

molesta. |

18. Formica sanguinea Latr., subsp. rubicunda Emery,

var. subnuda Emery.

This form was discovered by Mr. Dieck near Yale, D. C.

Its auxiliaries are also furnished by F. subsericea.

19. Formica sanguinea Latr., subsp. puberula Emery.

This small form, which occurs in Colorado, is probably the

one observed by McCook (82, pp. 152—153). Its auxiliaries,

according to this observer, belong to F. schaufusst and to a

small black species (probably one of the western varieties of

F. fusca).

20. Formica sanguinea Latr., subsp. obtusopilosa Emery.

Emery described this subspecies from New Mexico. Its

auxiliary is not recorded, but is probably furnished by some

variety of F. fusca, like neorufibarbis Emery or neoclara

Emery.

21. Polyergus rufescens Latr., subsp. lucidus Mayr.

P. lucidus, the “shining slave-maker ” of McCook, is the

best known of the three American subspecies. It has been

taken in several of the Atlantic States from Cape Cod to North

Carolina and westward into Pennsylvania, but its exact geo-

graphical distribution has not yet been determined. Rev. P.

J. Schmitt has sent me specimens from New Jersey, Maryland,

and North Carolina. The specimens from the last-mentioned

locality are decidedly opaque, thus resembling the European

714 THE AMERICAN NATURALIST. [Vor. XXXV.

form much more closely than do the typical specimens from

other localities.

The habits of P. lucidus were first observed by McCook

fully twenty years ago (80). His account is fragmentary and

barely sufficient to show that the habits are essentially like

those of the European form. In the nest which he observed

the slaves belonged to F. pa//ide-fulva, subsp. schaufusst. The

same slaves were observed in a mixed nest taken on Cape Cod

by Mrs. Mary Treat (Mayr, '86, p. 424). According to Per-

gande's observations cited by Wasmann (94, p. 164), F. pallide-

fulva, subsp. nitidiventris, is the auxiliary species in the District

of Columbia. In the case observed by McCook it was, of

course, the F. schaufussi which determined the character of the

nest, since Polyergus does not excavate. Hence the title of

McCook's paper is misleading. His observations on the feed-

ing habits of P. /ucidus are, as he remarks, “ chiefly confirmatory

of those recorded by Huber, Forel, and others in the European

Polyergus.” These, and a few additional notes on the belli-

gerent disposition of this ant, are not, however, sufficient to

leave no doubts in our minds that * our American species has

precisely the same habit " of carrying on its dulotic expeditions,

which he did not observe. Its slaves are certainly somewhat

different in this country, and it is therefore to be presumed

that the military tactics of the dulotic species may also be

different.!

To McCook's inapposite criticism of Darwin's

ing the phylogenetic origin of dulosis, I shall

to return in the sequel.

views concern-

have occasion

22. Polyergus rufescens Latr., subsp. breviceps Emery.

This subspecies (Fig. 15, 2, 4), founded on specimens from

South Dakota and Colorado, resembles the European rufescens

still more closely than does /ucidus, since it has the "T

sculpturing, opacity, and pilosity. It is, however, somewha

smaller, with a relatively shorter head, and its antennal scape

is distinctly enlarged towards their tip. . breviceps 18 ae

dently the form observed by McCook (82, P. 384) in! pu

American Faq

1 Cf. the above-quoted observations of Forel on the

No. 417.] NESTS OF AMERICAN ANTS. 715

Garden of the Gods. The slaves, according to McCook,

belonged to F. schaufussi, Recently Rev. P. J. Schmitt has

sent me specimens of P. óreviceps from Breckenridge, Col.

In this case the slaves accompanying the specimens belong

to a rather small, monticolous subvariety of Z subsericea

(Fig. 15, c, a).

23. Polyergus rufescens Latr., subsp. mexicanus Forel.

This Mexican subspecies (Forel, '99, p. 129) is related to

breviceps, but is larger, and without pubescence on the upper

surface of the body. The exact locality of Forel’s specimens

is not given. It is safe to say that they must have been taken

somewhere on the high plateau of central or northern Mexico.

The auxiliary Species is not recorded, but it is probable that

the varieties of F Jusca (F. subsericea and F. neorufibarbts)

recorded from the Mexican plateau (Forel, '99, p. 128) furnish

the requisite slaves.2

24. Tomognathus americanus Emery.

This Species (Fig. 17), which is both smaller and in other

respects quite distinct from the European sub/evis, appears

to be very rare. The type specimens were taken by Mr.

Pergande at Washington, D.C., in a nest of Leptothorax curvt-

Spinosus Mayr (Fig. 18), but no observations on the mutual

relations of the two species were recorded. Rev. P. J. Schmitt

of tty, Pa., writes me: “I have taken this species on but

one occasion, — when I carried home a bushel of sifted vege-

table matter from the woods. On examining this carefully

* While this article was going to press I discovered a fine large colony of P.

breviceps in some woods near Rockford, Ill. The ants with their slaves — in this

ance Formica fusca L., var. subenescens Emery — were living in a rotten stump

m Cavities excavated and long since abandoned by Camponotus pennsylvanicus.

The Polyergus workers, like the Colorado specimens, are of small size with dis

unctly club-shaped antennal scapes. The head and thorax are opaque and the

abdomen are long and projecting. The abdomen is shining and

nearly black in color. : OE |

xs Buckley C66, p. 170) describes a black female ant from Texas as Polyergus

ana, but no one has since succeeded in recognizing this species. Itis probably

not a Polyergus at all

716 THE AMERICAN NATURALIST. [VoL. XXXV.

I found about a dozen of the ants, which were readily recog-

nized as Tomognathus. There may have been a few Lepto-

thorax in the material, — certainly very few, if any, — but when

Fic. 17. — 71 th canus Emery. Worker.

collecting with the sieve it is hazardous to affirm that any

ants that are found belong to mixed or to independent

colonies.”

VI. CoracoBiosis.

It is very difficult to establish a clear distinction between the

ants of this and the preceding category, since Strongylognathus

is obviously transitional. Forel even includes this genus among

the social parasites, while Wasmann includes the whole of

Forel’s category among the forms which I have designated as

dulotic. I believe, however, that I am justified in erecting *

special category for Anergates, which is the only well-known

No. 417.] NESTS OF AMERICAN ANTS. 717

social parasite, and for the American Zpaews pergandei, since

these forms have become so extremely dependent on ants of

other species that they have even lost the worker caste, thus

leaving the species to be represented only by the fertile sexes

like the vast majority of living organisms. The following con-

densed account of the work of European observers on Anergates

atratulus is translated from Janet (97, p. 58 e seg.), who inci-

dentally adds to it some valuable observations of his own :

Anergates atratulus is a very bizarre ant, which inhabits cen-

tral and northern Europe. It has been studied by Schenck (52),

Fic. 18. — Leptothorax curvispinosus Mayr. Worker.

von Hagens (67), Forel (74), Adlerz ('86), and Wasmann (91).

As indicated by its name, it is a species which possesses no

Worker form. At the time of hatching from the pupa the

718 THE AMERICAN NATURALIST. (Vor. XXXV.

female presents very nearly the normal shape of queen ants

and possesses wings. After fecundation, however, owing to an

extraordinary development of the ovaries, her abdomen takes

on the appearance. of a sphere 4 mm. in diameter (Fig. 19, b),

on which are seen in the form of little plates, isolated by the

distention of the articular membranes, the strongly chitinized

rings which constitute the whole external surface of the abdo-

men in the young individual (Fig. 19, c). The male (Fig. 19, a)

is apterous, and its abdomen is strongly curved downwards.

He has a dawdling gait. The strigil is well developed in the

Fic. 19. — Anergates atratulus Schenck. a., male (after Adlerz) ; 4., fertile female (after Forel);

¢., abdomen of virgin female (after Adlerz).

female, while in the male it is very small, but nevertheless

pectiniform in certain specimens (Switzerland, Forel), although

in others it lacks the teeth and is quite rudimental or even

almost obsolete (Sweden, Adlerz; Holland, Wasmann ; Beau-

vais, Janet).

In some young male specimens collected at Beau

preserved in alcohol, I observed by transmitted light, in the

head, near the eyes, and of about the same size as these, the

two mandibular glands and their excretory ducts opening at

the base of the mandibles. These glands are therefore well

developed, notwithstanding the fact that the mandibles, which

are rounded at their tips, are much reduced. The male an

female are both provided with well-developed ocelli. The

antennz are rr-jointed in both sexes.

vais and

No. 417.] NESTS OF AMERICAN ANTS. 719

Owing to the absence of wings in the males, mating takes

place within the nests. This can be easily observed both in

the natural and in the artificial nests. The couples may be

killed without separating, by immersion in warm alcohol.

The nuptial flight of the females was observed by von Hagens

(67) on the 12th of August in the Rhine province. Some of

the queens may perhaps fly to other nests and there be ferti-

lized, and although there is usually only one fertile queen to

a colony, it is possible that there may occasionally be several

originating from different nests. If this were not the case we

should have the condition to which Forel has called attention

(74, p. 343), viz., that all mating must necessarily take place

between brothers and sisters of the same colony.

The missing workers of Anergates atratulus are replaced in

the mixed colonies by the workers of Tetramorium cespitum.

Whatever progeny is found in these colonies belongs exclu-

sively to the Anergates. The Anergates of both sexes are

nourished with food regurgitated from the mouths of their

Tetramorium auxiliaries. They appear to be incapable of

obtaining their food in any other manner.

Adlerz (86, p. 231) and Wasmann (91, p. 136) have ascer-

tained that the Tetramorium auxiliaries of the Anergates pay

relatively little attention to the young queens, while, on the

other hand, they very frequently carry the males about and

lick them long and assiduously. During this operation the

males assume a characteristic motionless attitude. The two

authors compare the attention thus bestowed on the male

Anergates by the Tetramorium auxiliaries to that bestowed

on myrmecophilous beetles that secrete certain substances of

Which the ants appear to be fond; e.g., the attention bestowed

on Claviger testaceus by Lasius flavus.

Adlerz and Wasmann have made experiments with a view to

determining the method whereby a new mixed colony is formed,

t.e., by the association of the female Anergates with the Tetra-

morium workers. Adlerz (86) in Sweden placed several unfer-

tilized Anergates queens in a strange nest of Tetramorium.

They moved about among the Tetramorium as if unperceived.

He obtained nearly the same results on placing unfertilized

720 THE AMERICAN NATURALIST. [Vor. XXXV.

queens of Anergates in a normal colony of Tetramorium com-

prising a queen and her progeny. He also placed a consider-

able number of the larvae, pupze, and male and female imagines

of Anergates in a normal colony of Tetramorium which were

living in an artificial nest. In all cases the strangers were

almost at once amicably received. Wasmann (91, p. 142)

obtained similar results in Holland. He observed that strange

Tetramoriums did not in the least injure the male or female

Anergates which he gave them, whereas they killed without

mercy the Strongylognathus testaceus males or females that

were placed in their nest. I have reported an experiment

made on the same subject (96, p. 27). I have also performed

the following experiment: A normal colony of Tetramorium

cespitum provided with a deálated queen, and a normal colony

of Anergates comprising an obese queen, some slender young

queens, some males and some Tetramorium workers — both

colonies comprising about the same number of individuals —

were put together in an artificial nest. There ensued some

struggles of relatively little importance, but some days later the

obese queen was found lying dead in the midst of a cluster of

Tetramoriums which seemed to be caring for her assiduously.

Some weeks later all the Anergates males and females had dis-

appeared, so that the colony again became a normal colony of

Tetramorium. Von Hagens (67) kept a single formicary of

Anergates under observation during several consecutive years

in the same place. It is difficult to assume that the number

of Tetramoriums may be maintained in an Anergates colony by

the introduction in one way or another of newcomers, $0 that

I am inclined to believe with Wasmann (91, p. 143) that the

duration of such a colony is limited to the duration of the life

of the Tetramoriums.

25. Epecus pergandei Emery.

Emery (94, p. 274) believes that this species, like Ane

gates, has no worker forms. Up to the present time it has

been taken only once, when Mr. Pergande found it in a nes:

of Monomorium minutum Mayr, var. minimum Buckley, ye

Washington, D.C. This nest contained not only the wing

No. 417.] NESTS OF AMERICAN ANTS. 721

males and females of the parasitic species, but was also pro-

vided with the winged sexes of the Monomorium. When both

species were put together in the same vial the Epoecus queens

attacked and killed some of the males of Monomorium. These

meagre data constitute all the forthcoming evidence for sup-

posing that the habits of Epoecus are analogous to those of

Fic. 20. — Epæcus pergandei Emery (after Emery). æ., male; b., female.

the European Anergates. I may add that I have examined

many dozens of Monomorium minimum nests in Texas in the

hope of finding their rare parasite, but up to the present time

my search has been in vain.

VII. SvNCLEROBIOSIS.

The mixed nests of uncertain origin and significance are of

considerable interest, but unfortunately they are very rare, and

as their origin has never been observed in any single instance

either in Europe or America, it is possible to do little more in

the present state of our knowledge than to catalogue the dif-

ferent cases. Mixed nests of this character are formed by the

union of dominant species with unusual auxiliaries or vice

versa, or by the close consociation of species which normally

inhabit independent colonies. It is generally agreed that such

nests must be either predatory unions, established after the

manner of dulotic species by robbing the larvæ and pupæ of

Species which never function as normal auxiliaries, or by alli-

ances between queens of different species before or soon after

Starting their colonies. . Experiment may be expected to throw

722 THE AMERICAN NATURALIST. [VoL. XXXV.

considerable light on the extent to which such unions are

possible. Forel (74) and Wasmann (91) have recorded a num-

ber of interesting observations, some of which are very similar

to the following cases observed in America.

26. Formica pergandei Emery and F. pallide-fulva Latr.

Mr. Pergande (Emery, '933, p. 646) found near Washington,

D.C., a mixed colony of Formica pergandei and the typical

F. pallide-fulva, but the nature of the consociation was not

determined. Emery suggests that the former species may be

a true dulotic ant and F. pallide-fulva its auxiliary species.

Mr. Pergande informed him that he had known of the exist-

ence of this colony for several years, but had seen only pallide-

Julva in the nest till the summer of 1892, when the 7. pergandet

made their appearance in the colony. This observation would

seem to favor an explanation by alliance rather than dulosis, or,

at any rate, on the suppositon of dulosis the róles of the two

species would seem to be the reverse of that suggested by

Emery, pallidefulva being the dominant and fergandei the

auxiliary species.

27. Formica exsectoides Forel and F. subsericea Say.

Forel (00°, p. 12) found a small mixed formicary of these

species at Hartford, Conn. “There could be no doubt con-

cerning the intimate life in common of the two species in the

same nest. They entered and went out through the same

doors, etc." Rev. P. J. Schmitt writes me that he has found

at different times five different nests of F. exsectoides-sub-

sericea, These invariably contained females of the exsectoides

only. All these colonies were, moreover, obviously incipient,

as shown by the fact that they contained scarcely more than

fifty ants, including both species. These six cases observed

by Forel and Schmitt are probably of the same nature as the

very similar cases of F. ersecta-fusca, F. truncicola-fustt, F.

exsecta-pressilabris-fusca, F. pratensis-fusca, and F. —

t 1$

rufa-fusca described by Forel (74) and Wasmann (91).

probable that the American ersectoides, like the Europea"

No. 417.] NESTS OF AMERICAN ANTS. 723

exsecta, has dulotic proclivities which are shown only while the

colony is young. In this connection the above-recorded fact,

that weak colonies of F. sanguinea are found to have the most

slaves, is perhaps significant.

28. Dorymyrmex pyramicus Roger, vars. niger Forel and flavus

McCoo

A peculiar mixed colony containing both the common vari-

eties of Dorymyrmex pyramicus was found by Forel at Faisons

N.C. (00°, p. 5). * There were two or three nests situated

several meters apart. The yellow workers and the black

workers entered and went out peaceably side by side, worked

together and treated one another with every show of friend-

ship. The two forms were, nevertheless, perfectly distinct,

without presenting any transitional varieties. I completely

demolished one of the nests and had the good fortune to find

the females and males of niger and the male of flavus, the

latter being larger and paler." Forel believes that this nest

arose “without doubt by the fortuitous association of two

fertile females, one of each variety." He therefore regards it

as of the same nature as the peculiar mixed nest of Tapinoma-

Bothriomyrmex which he described in the * Fourmis de la

Suisse ” (74, p. 372).

29. Pogonomyrmex barbatus Smith and its var. molifaciens Buckley.

At Aguas Calientes, Mexico, Dec. 31, 1900, I happened on

a huge gravel cone nest of the agricultural ant (Pogonomyrmex

barbatus), containing about equal numbers of the typical species

(with black head and thorax) and the entirely red var. molifa-

ciens. There were no transitional varieties. Both forms were

living together on the most amicable terms. As I entertained

little hope of finding the queens, since this would, in all proba-

bility, have required a very careful excavation of the soil to a

depth of from five to eight feet, I had to rest satisfied with dig-

ging into the nest a short distance and examining the hosts

of belligerent workers that swarmed forth. As this was obvi-

ously an old and very flourishing colony, there can be little

724 THE AMERICAN NATURALIST.

doubt that it must have been formed by alliance between.two

or more queens representing the two distinct color varieties.

The whole country about Aguas Calientes is covered with the

most flourishing nests of these two forms, often very close to

each other, so that it is not at allimprobable that an occasional

mixed nest should arise in this manner.

30. Stenamma tennesseense Mayr and S. fulvum Roger, subsp. aquia

Buckley, var. piceum Emery.

Rev. P. J. Schmitt found this singular mixed nest near

Beatty, Pa. The queen of the colony belonged to S. tennes-

seense. Rev. Mr. Schmitt was impressed by the fact that the

nest was under a stone, whereas /ennesseense normally occurs

only in dead wood at Beatty. This seems to be generally true

of the species. In Illinois and Wisconsin I have never taken

it except in the old logs in the rather open forests. It is a

singular fact that in this and in nearly all the other cases of

synclerobiosis the two consociating species or varieties repre-

sent a light and a dark colored form. This can scarcely be a

mere coincidence, but I am unable to suggest any explanation

of this peculiar phenomenon.

I cannot conclude this portion of my paper without express-

ing my indebtedness to Rev. P. J. Schmitt and to Professor

Auguste Forel. These gentlemen have most generously sent

me specimens of several of the rare and peculiar Formicide

which I have figured.

(To be continued.)

! A somewhat similar mixed nest appears to have been found by Mapi

(78, p. 37, footnote) at Mentone. This colony consisted of nearly equal aper ;

Stenamma [s. gen. Messor] structor, barbara, and the red-headed variety 9

barbara.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XVII. Tue RorATORiA.

H. S. JENNINGS.

Tue Rotatoria, or wheel animalcules, consist of minute,

chiefly microscopic animals, which are everywhere abundant in

fresh water. Along with the Protozoa they constitute by far

the largest number of species and individuals among the ani-

mals to be found in pools, ponds, rivers, and lakes. A few are

found in the ocean, but the rotifers are typically fresh-water

organisms. :

The most characteristic feature in the organization of the

Rotatoria is the ciliated area at (or near) the anterior end of

the body, serving as a locomotor organ or to bring food to the

mouth. Coupled with the.lack of cilia elsewhere on the body,

this constitutes a character by which a rotifer may as a rule be

recognized at once. This ciliated area is usually known as the

corona. It varies excessively in form and structure, and in a

few rare aberrant species is lacking.

Male and female differ in form, the males being, as a rule,

much smaller than the females, and reduced in structure, —

usually lacking the alimentary canal. The males are little

known, many species existing in which they have never been

Observed. Discussions of the structure and classification of the

rotifers are therefore based generally on the females alone.

The form of the body is exceedingly varied in the different

representatives of the group, — ranging from spherical in Tro-

chosphzera to the excessively attenuated form of Rotifer neptu-

nius (Fig. 29) or the spiny, turtle-like form of Polychztus

(Fig. 111). As a rule, however, the body is somewhat elon-

gated, and is extended at the posterior end, behind the cloaca,

to form a long stalk or a tail-like appendage, called the foot.

725

726 THE AMERICAN NATURALIST. [Vor. XXXV.

This frequently ends in two small points, called the toes.

There may or may not be a distinct head, set off from the

body by a neck. Dorsal and ventral surfaces are usually (not

always) markedly differentiated. In many rotifers (Loricata)

the cuticula or outer covering of the body is hardened to form

a shell or Zorica ; this may bear elevations, spines, teeth, etc.

The simplest and probably primitive form of the corona is

that of a plane disk covered with equal cilia, on the ventral side

of the animal at the anterior end, as in Proales. From this the

varied forms found have probably been derived, chiefly by the

following steps : (1) the outer cilia became longer and stronger,

forming a distinct marginal wreath; (2) the cilia within this

‘wreath were partly or entirely lost; (3) a second wreath of

cilia was developed (or has remained from the original cilia)

within or without the above-mentioned wreath. Thus we

obtain the condition, which has often been considered the

primitive one, of two concentric wreaths, with the mouth

between them. This condition is found only in much special-

ized forms.

In the alimentary canal can be distinguished a mouth, usu-

ally situated excentrically in the coronal disk and leading into

a muscular pharynx, or mastax, which is furnished with chiti-

nous jaws or ¢rophi; a narrow cesophagus; a large stomach

furnished with a pair of gastric glands, and an intestine leading

to the anus. The latter is lacking in some groups. The jaws

or trophi are very various in form and furnish a most important

systematic character. Certain types of jaws are distinguished ;

of these, the more important are the malleate (Fig. 170) and

the forcipate (Fig. 171). The other forms (ramate, Fig. 34:

incudate, Fig. 54; uncinate, Fig. 10, etc.) may be related to

one of these two types.

Points of especial practical importance for distinguishing

species are the following : form, structure, and position of the

corona; presence or absence, form and position, of the foot ;

form of the toes; presence, number, and position of the red

eye-spots ; structure of the jaws or trophi ; presence or absence

of the intestine and anus ; nature of the cuticula, — whether

soft and flexible or hardened to form a shell orlorica ; presence

No.417.] WORTH-AMERICAN INVERTEBRATES. 747

or absence of appendages on the body; and mode of life, —

whether fixed or free-swimming.

The Rotatoria are a cosmopolitan group, so that the same

species may be, and as a matter of fact frequently are, found

in America, Europe, India, China, and Australia. Our rotifer

fauna is, on the whole, so far as known, almost identical with

that of the one other well-known continent, — Europe. From

this it results that the only entirely satisfactory key for

America or any other country would be one including all

the species of the group. Two hundred and forty species have

thus far been recorded from America; it is probable that

nearly as many more will be found before the list can be con-

sidered to approach completeness. Under these circumstances

the following key to known American species can be considered

to have merely provisional value. Moreover, the species of

many of the larger genera can be determined only from full

descriptions and detailed figures; the key should in such cases

be used in connection with fuller accounts.

The classification of the Rotatoria is in an unsatisfactory

condition. The system employed by Hudson and Gosse in

their monograph of the Rotifera is used almost exclusively,

and is therefore adopted in essentials in the present key.

But it is undoubtedly unsatisfactory in many ways, separating

widely many closely related species, and bringing together

some that are widely divergent in essential structure. A

better classification has been outlined by Wesenberg-Lund, but

this has not yet been worked out in sufficient detail to make

its use practically satisfactory in such a key.

Owing to the large number of species of the Rotatoria and

the frequent difficulty of assigning definite distinguishing

Characteristics even to the genera, it will be necessary for the

key to the genera and species to be purely artificial in char-

acter. I give first, therefore, a systematic synopsis of the

orders, suborders, and families. This synopsis is based on

that given by Hudson and Gosse in the monograph of the

Rotifera. I have introduced certain divisions not given by

Hudson and Gosse, — such as the general division into Mono-

£ononta and Digononta ; this division is one which is employed

728 THE AMERICAN NATURALIST. [VoL. XXXV.

by almost all later authors. Certain families, as the Apsilide,

Ploesomadze, Gastropodidz, and Anapodidz, are added to those

given by Hudson and Gosse; this has become necessary through

the progress of investigation since the monograph was written.

Many of the definitions are taken from Hudson and Gosse.

The system is based throughout on the structure of the

females.

SYNOPSIS OF THE ROTATORIA, WITH CHARACTERS OF

THE CLASS, SUBCLASSES, AND ORDERS.

Crass RoTATORIA OR ROTIFERA (WHEEL ANIMALCULES).

Small microscopic organisms, living chiefly in fresh water, bearing at the

anterior end a ciliated area which takes various forms. Body often extended

backward to form a stalk or foot. Pharynx with chitinous jaws ; cloacal

opening, when present, on the dorsal side, at the boundary between body

oot. Excretory organs in the form of fine tubes bearing “ flame cells "

or “ vibratile tags,” and opening into a contractile vacuole near the cloaca.

Sexes separate ; males usually minute, .degenerate, lacking the alimentary

canal

SuBCLASS 1, DicoNoNTA (HAVING Two OVARIES).

Order 1. Bdelloida. Swimming with the ciliary wreath and creeping like

a leech (or parasitic). Jaws ramate (Fig. 34).

Family i. Philodinadz.

Genera. Philodina, Rotifer, Callidina, Discopus.

Family 2. Adinetade.

Genus. Adineta.

Order 2. Seisonacea. Marine ; parasitic on Nebalia ; no corona. Males

Genera. Seison, Paraseison, Saccobdella.

SUBCLASS 2, MONOGONONTA (HAVING ONLY ONE OVARY).

Order 4. , Rhizota. Fixed forms ; foot ending in a disk or cup.

Genera. Floscularia, Stephanoceros.

Family 5. Apsilide.

Genera. ae E (Atrochus).

Family 6. Melice

Genera. Me viii: Gadi Limnias, Limnioides, Cephalosiphon,

CEcistes, Pseudcecistes, Lacinularia, Megalotrocha, Conochilus.

No.4175.] NWORTH-AMERICAN INVERTEBRATES. 729

Order 5. Ploima. Not fixed ; swimming with the ciliary wreath; not

creeping like a leech. Jaws never ramate.

Suborder 1. Illoricata. Cuticula flexible, not hardened to form a shell

or Zorica.

Family 7. Microcodontide.

Genera. Microcodon, Microcodides.

Family 8. Asplanchnade.

Genera. Asplanchna, Asplanchnopus, Ascomorpha, (Hertwigia ?).

Family 9. Synchetadze

Genera. Syncheta, (Anartani, (Polyarthra).

Family 10. Triarthradæ.

Genera. Triarthra, Pedetes, Pteroessa, (Polyarthra and Anarthra).

Family 11. Hydatinadæ.

Genera. Hydatina, Rhinops, Notops, Triphylus, Cyrtonia.

Family 12. Notommatadæ

Genera. Albertia, Taphrocampa, Pleurotrocha, Notommata, Copeus,

gray Furcularia, Triophthalmus, Eosphora, Diglena, Dis-

a, Notostemma, Arthroglena, (Drilophaga) (Monommata ?).

Shri 2 gibus Cuticula stiffened to form a distinct armor or lorica.

Family 13. Rattulide.

Genera. Rattulus, Mastigocerca, Elosa, (Ccelopus?), (Diurella ?),

Genera. MN Polychatus, Scaridium, (Stephanops).

Family 15. Salpina

Genera. Salpina, Diglax Diaschiza, Diplois.

i idx

Genera. Cathypna, Dis Monostyla.

Family 18. Coluride. .

Genera. Colurus, Metopidia, Monura, Mytilia, Cochleare, (Steph-

anops).

Family rg. Pterodinadz.

Genera. Pterodina, Pompholyx.

Family 20. Brachionide

Genera, Brachionus, Scblebci Noteus.

ureade.

Genera. Anurza, Notholca, Eretmia.

Family 22. Plæsomadæ.

Family 24. Ana apodide. .

Genera. Anapus, (Hertwigia).

730 THE AMERICAN NATURALIST. [Vor. XXXV.

Order 6. Scirtopoda. Swimming by means of branched appendages

resembling in some respects those of the Crustacea. (The ordi-

nal value of this character is decidedly doubtful.)

Family 25. Pedalionade.

Genera. Pedalion, Hexarthra.

Of uncertain ‘sytematic position — Trochosphera, Atrochus, Adactyla,

Balatro, Cypridicola.

ARTIFICIAL KEY FOR THE DETERMINATION OF THE

GENUS AND SPECIES OF AMERICAN ROTATORIA.

Ar. Adult animals attached or united in colonies, usually dwelling in

tubes; or if separate and free-swimming, then carrying the transparent

tube with them. Foot ending in a flat disk or cup, which is attached

to the substratum or to the bottom of the tube (or foot absent in

Nos. 15-17) (free-swimming when young) Order RHIZOTA, H. & G.

Br. Not forming colonies ; corona with long slender setze and usually

produced into a varying number of lobes bearing the sete ; mouth in

the center of the large corona ; cilia few, about the mouth, scarcely

noticeable. Trophi uncinate (Fig. 10) . . Family FLOSCULARIAD/E

ar. Sete not arranged in whorls or parallel rows on the lobes of the

corona, but scattered or in groups . . . . Floscularia

br. Free-swimming, carrying the trakopii tbe

cz. Corona two-lobed ; two eyes on the dorsal lobe. 1, Floscularia

mutabilis Bolton

c2. Corona circular, ciliated, with five short prominences bearing

sete . P3: 2, F. pelagica Rousselet (Fig. 1)

ó2. Not ires eiim

cz. Corona without distinct lobes, but pratend va set

short, chiefly on the dorsal and ventral parts of the

3, F. edentata pni (Fig. 2)

c2. Lobes of the corona three

dr. Lobes large, separated by iid curved depressions ; set On

the entire rim of the corona. . 4, F. trilobata Collins

(Fig. 3)

d2. Lobes HOCUNE but three — one large dorsal and two small

ventral ones ; but really with two minute additional lobes between

the dorsal and ventral. See cj.

£3. Lobes five

dr. A long flexible process on the back of the dorsal lobe ; lobes

knobbed . 5, F. cornuta Dobie (Fig. 4)

42. No dorsal process ; lobes kaobbed

er. Lobes rather long and slender. . . . 6,4. onetta

Cubitt (Fig. 5)

e2 Lobes shorter and thicker . . 7, F. ornata Ehr. (Fig. 6)

No.4175.] MORTH-AMERICAN INVERTEBRATES. 731

43. Lobes broad, not knobbed, all five well marked ; sete arising

both from the summit of the lobes and from the intervening

depressions. 8, F. campanulata Dobie (Fig. 7)

d4. The two lateral iei very small, at first view bardly notice-

able; dorsal lobe much larger than ventra

ez. Corona ornamented with dots arranged in symmetrical pat-

terns ; animal very small ; tube sometimes lackin

9, F. algicola Hudson (Fig. 8)

e2. Larger, corona not ornamented as ie er

o, F. ambigua Hudson

d5. The five lobes forming long slender iid arms, like those

of Stephanoceros (Fig. 9 ents . I, F. millsii Kellicott

c4. Lobes seven . 12, F. regalis Hudson

a2. Setæ arranged in odi or ge io rows on the five long

pointed lobes of the corona. . . . hi cos n eichhornii

r. (Figs. 9 and to)

B2. Corona without sete and apparently without d (a minute ciliary

wreath can usually be detected on careful search)

ar. Corona with one dorsal lobe, the coronal cup edged with a delicate

festooned membrane ; body long, with a long slender tapering stalk,

by which it is attached. Living in colonies of Méegalotrocha albo-

flavicans . - . 14, Acyclus inguietus Leidy (Fig. 11)

42. Coronal op a IS po HE. sack ; body short and thick ; no

foot, — the animal being attached by a flat disk . . . . Apsilus

ér. Coronal cup not oblique, its frontal margin horizontal

15, A. vorax Leidy

62. Coronal cup and its frontal margin oblique ; ganglion or brain in

the nec

ez. Ventral margin of the coronal PY with a central convex lobe-

like projection . . 16, A. bipera Foulke (Fig. 12)

c2. Ventral margin of cup even, fh i central projecting lobe

17, A. bucinedax Forbes (Fig. 13)

B3. Corona without sete and not produced into long lobes, but with

Strong conspicuous moving cilia forming a marginal continuous curve

about the corona. In this curve there is a more or less conspicuous gap

on the dorsal side. Mouth near the ventral side of the corona. On the

body just below the corona either a single dorsal antenna, or two ventral

ones, or all three, are noticeable. Trophi malleo-ramate (Fig. 16)

ar. Individuals attached, separate, or in branching non-spherical colonies

of few specimens (1-30 or thereabouts)

^. Corona of four lobes (or in No. 21 of three lobes) ; ventral

antennz obvious ; dorsal antenna minute Melicerta

cI. Lobes of the corona when expanded wider than the tube; a

Short, blunt chin on the ventral side below the corona; tube

forme i Loe os S8, M. ripe

d of nearly spherical pellets Seb (Figs. 14, r5, and 16)

732

THE AMERICAN NATURALIST. [ VOL. XXXV.

c2. Lobes when expanded of the same width as the tube; chin

long and pointed ; tube of oP having the form of a pointed

cylinder i 19, M. conifera Hudson (Fig. 17)

£3. Lobes when DAAN more than three times the width of the

body ; ventral antennæ very long ; tube gelatinous, without pel-

lets : : . 20, M. tubicolaria Ehr.

c4. Ures (ventral) pair at tikes dias: by a large notch;

lower (dorsal) pair almost confluent, so that the corona seems to

have but three lobes ; ventral antenna short ; chin two-pointed.

Tube with ovoid fecal pellets, or floccose, without pellets

21, M. janus Hudson (Fig. 18)

62. Corona broad, of two lobes, with a wide dorsal gap; dorsal antenna

minute ; ventral antennae obvious ; tube without pellets Limnias

cz. Tube cylindrical, transparent, ringed by transverse ridges at

regular intervals ; five horny processes on the dorsal surface of

the body, below the corona 22, L. annulatus Bailey (Fig. 19)

c2. Tube nearly cylindrical, not ringed, often partly covered with

débris ; no horny processes on dorsal surface of the body ; ven-

tral antennz very short . . . 23, L. ceratophylli Schrank

¢?. Tube roughened with transverse rows of raised points; seven

horny processes on the dorsal surface of the body below the

corona; ventral antennz T equal in length to the diameter

of the tube . 24, Z. shiawasseensis Kellicott

63. Corona nearly ticdu dd x diid dorsal gap ; dorsal antenna

very large, with two projections or hooks at its sides ; ventral

antennæ small or absent . . . Cephalosiphon

cI. Tube tapering to the foot, PREE EE with extrane-

ous material ; foot very long and slender n . 25, C. D

Ehr. (Fig. 20)

c2.

Tube irregular, semitransparent, uere per

6g. Corona a wide oval (or nearly circular), indistinctly two-lobed ;

dorsal gap minute, ventral antennz obvious ; dorsal antenna incon-

pene or absent (Ecistes

One or more dorsal hacks or ‘projections ‘eee ihe corona ;

ventral antennz minute ; tube absent or small and irregular

dr. Two dorsal hooks below the corona; — n

branched into antler-like structures . . . 27

es (Fi ig. i

42. A single dorsal hook below the corona; foot very —

animal without a tube, living in the mucilaginous matrix of t

alga Gloiotricha pisum . . . 28, O. mucicola Kell

c2. Antenne (ventral) very short ; no dew hooks

T. ntennz set wide apart; tube very irregular an

often beset with extraneous matter. . . 29, O. cry sia n

Ehr. (Fig. 22)

No.4175.] MORTH-AMERICAN INVERTEBRATES. 733

d2. Dorsal gap of the corona very wide ; tube opaque, regular,

tapering slightly from top to bottom . . 30, O. intermedius

Davis (Fig. 23)

c3. Antenne long

dr. Antenne very long and recurved, tube floccose ; very small

1, O. longicornis Davis

d2. Corona large, nearly circular, crossed with thick ribs, tube

oose, very irregular, clay colored . 32, O. umbella Hudson

42. Not attached ; inhabiting a tube ; individuals separate or one adult

grouped with its youn

ór. Corona horseshoe-shaped ; two antenne on the ventral surface

of the body, united almost to their tips 33, Conochilus dossuarius

udso

43. In clusters of many individuals, forming usually a spherical colony,

appearing to the naked eye as a small yellowish or grayish ball

br. Clusters attached

cI. Body (in known American species) with two or four opaque

warts in a transverse row on the ventral side. No tube. Corona

broad, kidney-shaped, with short axis dorso-ventral; antennz

inconspicuons ig. vd sc s xd ie Megalotrocha

dr. Opaque wartstwo. . . . . . . 34, M. semibullata

Hudson (Fig. 24)

da. Opaque warts four . . .... 2 oe 35, M. alboflavicans

r. (Fig. 25)

c2. Dwelling in transparent gelatinous tubes ; body without opaque

warts or denticles ; corona heart-shaped with long axis dorso-

ventral; antennz inconspicuous. 36, Lacinularia socialis Ehr.

62. F ree-swimming colonies or clusters . . . . . . onochilus

cI. Antenne two, separate except at the base, situated on the

corona, between the mouth and the ventral gap ; colonies spheri-

cal, of many individuals . . . . 37, C. volvox Ehr. (Fig. 26)

£2. Antennz united, so as to appear single, large and conspicuous,

and situated on the corona; clusters usually unsymmetrical and

containing comparatively few individuals . . 38, C. unicornis

Rousselet (Fig. 27)

C. dossuarius — see No. 33)

42. Not fixed when adult ; not forming colonies nor living in tubes (in

a few rare cases the animal lives in a tube, but is never attached by its

foot to the bottom of the tube, as in 47

Br. Without lorica; i.e., the cuticle of the animal is flexible, not stiffened

to form an unyielding armor or Zorica

#1. Swimming with the corona and creeping like a leech ; body usually

nearly cylindrical (dorsal and ventral surfaces not being conspicuously

differentiated), and composed of rings which may be drawn one within

the other in a telescopic fashion. Foot (reckoned from the cloaca to

734 THE AMERICAN NATURALIST. [Vor. XXXV.

the tip) usually ending in three toes, and bearing two to four spurs

some distance from the tip. A dorsal proboscis behind the corona.

Trophi or jaws ramate (Fig. 34). (Ovariestwo.) Order BDELLOIDA

ér. Corona of two nearly circular retractile lobes, transversely placed

Family PHILODINIDE

cr. Eyes two

dr. Eyes on the frontal column or proboscis . . . . Rotifer

ez. Constructing and dwelling in tubes 39, Æ. mento Anderson

e2. Not living in tubes

ft. Antenna very long (one-half to one-third as long as the

body) ; foot short (one-third body length) ; spurs short and

DER. s un 40, R. macroceros Gosse (Fig. 28)

/?. Antenna not remarkably long

£I. Body very long and slender, white and transparent ;

foot extraordinarily long (one and one-half times the

length of the rest of the body) ; toes long and slender

41, R. neptunius Ehr. (Fig. 29)

£2. Foot not remarkably long

hr. Spurs not twice as long as the width of the joint to

which they are attached. Body transparent or whitish,

not colored or dark.

ir. Body whitish, opaque, passing gradually into the

foot; spurs one and one-half times as long as the

width of the joint to which they are attached, and

forming an obtuse angle with each other

42, R. vulgaris Schrank (Fig. 3°)

i2. Body thick, suddenly decreasing in size to form

the foot, which makes up half the entire length of the

animal ; spurs produced to a longish point at the tip

43, R. macrurus Schrank

h2. Spurs at least twice as long as the width of the

joint to which they are attached.

the joint to which they are attached, and with a

constriction one-third of their length from the tip

44, R. trisecatus Weber (Fig. 31)

i2. Body slender; foot long, not distinctly marked

off from the body ; spurs twice as long as the width

of the joint to which they are attached, slightly

swollen at base — 45, R. elongatus Weber (Fig: 32)

i. Body colored — dark brown, usually covered with

débris ; transverse folds very marked ; spurs almost,

or quite, three times as long as the width of the joint to

which they are attached 46, R. tardus Ehr. (Fig-33)

No. 417.] MORTH-AMERICAN INVERTEBRATES. 735

d2. Eyes in the “neck D 2 over the brain, jt above

the jaws . . Philodina

ez. Three mts in Hiec jaw (Fig. $e»

fT. Body beset with strong spines on the dorsal surface

47, P. aculeata Ehr. (Fig. 34)

f2. Body without spines, surface usually sticky, so that it is

frequently covered with débris ; spurs long and sharp, two

and one-half times as long as the width of the joint to

which they are attached . . . 48, P. macrostyla Ehr.

22. Two teeth in each jaw

Jt. Body smooth, colorless, very short, thickened in the

middle; corona very large ; foot sharply set off from the

body ; spurs shorter than the width of the joint to which

they are attached . 49, P. megalotrocha (Fig. 35)

J2. Body colorless or reddish, not short and thick ; foot not

distinctly marked off . . 50, P. roseola Ehr.

J3. Body greenish yellow, othé wis much like the last

51, P. citrina Ehr.

EK Eyes none: 2 55. s 0E Cm . . Callidina

(The genus Callidina is a very large and difficult one; while the following

species are all that have been identified in America, doubtless many more will be

er. Living ina tube . . . 52, C. eremita Bryce (Fig. 36)

e2. Not living in a tube ;

J?. Body with stout, blunt dorsal papillæ, especially on the

segment next to the foot

53, C. papillosa Thompson (Fig. 37)

J2. Body without papillæ or spines

gf. Foot ending in toes

hi. Jaws (Fig. 38) small, with numerous (8-10) fine

ridges (“ teeth ”)

i1. Corona as wide as the neck ; ridges (“teeth ”),

ten in each jaw (Fig. 38) ; spurs drawn to a slender

point c. . 54, C. elegans Ehr. (Fig. 38)

72. Corona not so vie as the neck ; ridges or teeth,

eight in each jaw (Fig. 39) ; spurs (Fig. 40) short

and broad

55, C. constricta Dujardin (Figs. 39 and 40)

A2. Jaws with but two distinct teeth (in addition to a

number of fine ridges)

tz. Teeth two in each jaw; spurs at least twice as

long as the width of the joint to which they are

attached, straight and thick ; body colorless. Para-

sitic on Asellus 56, C. socialis Kellicott (Fig. 41)

736 THE AMERICAN NATURALIST. [Vor. XXXV.

i2. Each jaw (Fig. 42) with one large and two

small teeth ; spurs (Fig. 43) scarcely as long as

the width of the joint to which they are attached,

swollen at base ; body yellowish

57, C. musculosa Milne (Figs. 42 and 43)

£2. Foot ending in a disk in place of toes ; teeth, six to

eight in each jaw; body reddish . 58, C. magna Plate

é2. Corona a flat surface covered with cilia, on the ventral side of the

axierent (Fig.4dM) ^. Ius c 2 E

cz. Skin smooth; no eyes; proboscis without bristles and with a

few cilia ; body broad . « 59, A. vaga Davis (Fig. 44)

42. Swimming with the corona ; not creeping like a leech (sometimes

creeping with the toes). Jaws of various forms, but never ramate.

A very large number of rotifers fall within this division, and the recogni-

tion of species, or in many cases even of genera, is very difficult. In order

to reduce the difficulties to a minimum, I separate out a number of species

having striking peculiarities, by means of the first five subdivisions given

below (47-45). The following is to be especially noted ; only the species

mentioned under a given characteristic possess that characteristic. For

example, any species having swimming appendages will be found under 45,

species not named under this subdivision do not have such appendages.

br. Spherical in form ; no foot ; ciliary wreath midway between the

equator and one pole of the sphere

60, Trochosphera solstitialis Thorpe

62. Foot ending in a single pointed “ toe”

cr. Corona a circle of strong cilia ; mouth in the center ; form of

the body conical; one eye 61, Microcodon clavus Ehr. (Fig. 45)

c2. Corona an oblique ciliated disk, with two auricles; body

brownish red or cherry red in color; one eye

62, Notommata monopus Jennings (Fig. 46)

c3. Corona an oblique ciliated surface with weak cilia; no eye:

parasitic in fresh-water annelids . 63, Albertia naidis Bousfield

63. Foot ending in two pointed projections, not side by side (as in

most rotifers), but one dorsal, the other ventral. Otherwise much

like No. 61

64, Microcodides chlena Gosse (orbiculodiscus Thorpe)

64. Corona extended dorsally into a large and broad proboscis, fringed

with cilia and bearing near its end two red eyes

65, Rhinops vitrea Hudson (Fig. 47)

65. Body bearing swimming or skipping appendages, in the form

movable spines, blades, or branching crustacean-like limbs ; no foot

cr. Six branching appendages, somewhat like those of a crusta-

cean ; two small stylate appendages on the posterior dorsal

surface. Eyes two . . 66, Pedalion mirum Hudson (F ig. 48)

c2. Twelve blade-shaped appendages, with serrate edges, arrang

No.417.] NORTH-AMERICAN INVERTEBRATES. 737

in four groups of three each, at about the level of the jaws. One

B c V o n 67, Polyarthra platyptera Ehr. (Fig. 49)

c3. Two very long spines (thrice the length of the body), attached

ventro-laterally ; eyes two . . . . 68, Pedetes saltator Gosse

cq. Three very long spines, two lateral, one ventral. Two eyes

69, Triarthra longiseta Ehr. (Fig. 50)

(The remainder of the group (a2) are less easily recogniza-

ble. It will facilitate the use of the rest of the key to this

subdivision to point out in a preliminary way certain striking

characteristics of a number of species.)

No eyes. — Hydatina senta (No. 87), Albertia (No. 63), Pleurotrocha

(No. 91), 7 aphrocampa saundersia (?) (No. 92), Diglena contorta (No. 93).

Eyes two.— Rhinops (No. 65), Diglena (Nos. 124-129), Distemma

(No. 130, ZafArocampa saundersie (9) (No. 92), Triphylus (No. 8 5),

Pedalion (No. 66), Pedetes (No. 68), Triarthra (No. 69).

Eyes three. — Asplanchna priodonta (No. 74), Asplanchna herrickii

(No. 73), TriophthaImus dorsualis (No. 122), Eosphora (No. 123).

Parasitic. —. External, —Pleurotrocha parasitica (No. 91). Internal, —

Albertia (No. 63), Hertwigia (No. 80), Proales wernecki (No. 110).

Viviparous (the developing embryo frequently seen within the mother). —

ier mam (No. 70), Asplanchna (Nos. 71-76), Rhinops vitrea

(No. 65).

No foot. — Trochosphera (No. 60), Pedalion (No. 66), Polyarthra

(No. 67), Pedetes (No. 68), Triarthra (No. 69), Asplanchna (Nos. 71-76),

Ascomorpha (Nos. 77-79), Hertwigia (No. 80), Anarthra (No. 8t).

46. No anus, the intestine ending blindly

cI. Foot present (viviparous) . . . .

di. Foot small, on the ventral surface

70, A. myrmeleo Ehr. (Fig. 51)

Asplanchnopus

c2. No foot :

dr. Large, clear, sac-like animals, with incudate jaws (Fig. 54);

corona with two slight conical elevations. Viviparous

Asplanchna

er. Saclike body with large projections or “ humps ”

Jt. Four humps, — one dorsal, one ventral, and two lateral

71, A. ebbesbornii Hudson (Fig. 52)

J2. Three humps, — like the last but with the ventral one

lacking (perhaps a variation of 71) — . 72, A. amphora

Hudson

«2. Sac-like body without humps

Jt. Eyes three, — one large, on the brain, two small, lateral

738 THE AMERICAN NATURALIST. | [Vor. XXXV.

gi. A bilobed glandular organ (Fig. 53, 4) attached close

to the opening of the ovary and excretory organs ; 20-25

flame cells on each excretory tube

73, A. herrickii de Guerne (Fig. 53)

£2. No glandular organ as described in 73; flame cells

but 3-5 on each excretory tube 74, A. priodonta Gosse

f2. Only one eye

£I. Jaws stout, with doubly pointed ends not serrated

(Fig. 54); flame cells 10-20 on each side

75, A. brightwellii Gosse

£2. Jaws weaker, ending in a blunt tooth and a broad,

thin plate (Fig. 55) (probably a variety of the last)

76, A. girodi de Guerne

d». Very small rotifers, sac-like ; corona rising slightly to a

single apex ; jaws not incudate ; not viviparous

Ascomorpha

er. A single large projecting process at the mid-dorsal edge

of the corona

ft. Ovate in form; hyaline except the stomach; on each

side of the body a “sub-dorsal groove” . 77, 4. hyalina

Kellicott (Fig. 56)

fe. Flattened; in one view nearly circular, with a neck-like

projection ; in the edge view oblong, half as long as wide;

skin stiffened so as almost to form a lorica

78, A. orbicularis Kellicott (?)

(This species is probably founded on dead specimens of Gas-

tropus stylifer, No. 153.)

e2. No large projecting process on the corona ; dorsal view

sac-like; lateral view unsymmetrical, with a gibbous dorsal

outline ; color usually dark green . . . . 79,4. ecaudis

Perty (A. helvetica Perty ; Sacculus viridis Gosse) (Fig. 57)

à7. Foot not present; no appendages to body ; anus present

cz. Parasitic in Volvox; having a large mid-dorsal projecting

process on the corona . . . . 80, Hertwigia parasita Ehr.

c2. Not parasitic ; body an elongated, parallel-sided, flattened sacó

corona squarely transverse, with a single marginal wreath of cilia,

and with two broad, flat setz-bearing prominences on the dorsal

side and two long styles near the ventral side . 81, Anar thra

aptera Hood (Fig. 58)

(See also Trochosphera, No. =

48. Foot present, ending in two toes placed side by side ; body with-

out swimming appendages ; anus present

cr. Form a broad, often swollen, cone, of which the foot forms the

apex. Corona large, transverse, flat or strongly convex, with four

No.417.] MWORTH-AMERICAN INVERTEBRATES. 739

long styles and a number of styligerous prominences ; at the sides

of the corona two in ciliated auricles, which may be retracted.

Asingleeye . . - + . Synchata

dr. The slender foot one-third to one-half the length of the

remainder of the body ; toes minute . . . . 82, S. stylata

Wierzejski (Fig. 59)

d2. Foot very short (less than one-sixth the length of the rest of

the body)

er. Body a swollen cone ; corona strongly convex, with two

central thick, club-shaped prominences ; auricles very long,

usually inclined backward . . . . 83, S. pectinata Ehr.

e2. Body a slender cone ; coronal surface almost flat, without

club-shaped prominences ; auricles small . 84, S. tremula

Ehr. (Fig. 60)

c2. Corona large, nearly transverse (slightly oblique), surrounded

by a wreath of cilia which is interrupted ventrally, and bearing a

number (3-7) of large prominences crowned with styles ; mouth

near the ventral side of the corona

di. Eyes two; jaws forcipate ; foot slender, of three joints ;

stomach with six long cecal projections . . . 85, ZvipAylus

lacustris Ehr. (Fig. 61)

d2. Eyes one or none

er. Eye one ; body long conical, humped dorsally, its outline

with Pus distinct curves ; corona with three styligerous

prominences. Ciliary wreath encircling the corona and a

sub-square space at right angles to the corona on the ventral

surface . . . . . . 86, Cyrtonia tuba Ehr. (Fig. 62)

e2. No eye; body long conical ; a large rotifer, 87, Hydatina

senta Ehr. (Fig. 63)

€3. Body large, sac-shaped, hyaline (like an Asplanchna), foot

very small and retractile, situated at the posterior ventral

angle of the body, almost on the ventral surface. Jaws mal-

leate ; eye single . . 88, JVotops clavulatus Ehr. (Fig. 64)

*4. Body strongly arched dorsally; nearly flat ventrally ;

cuticula thickened into a number of definite but inconspicu-

ous folds or teeth (so that the animal might almost be con-

sidered loricate); foot continuing the body axis, but nearer

the ventral side, with two small retractile toes ; one eye

. 89, Motops pelagicus uet (Fig. 65)

45. Body almost square in ventral view ; foot one-third of

total length, MS a continuation of the «hs axis; one

Notops brachionus Ehr.

4. Corona miho prominences d. styles ; consisting of an

oblique (or even ventral) disk or area covered with close-set cilia

or with numerous interrupted ciliary curves. Two lateral evertible

740 THE AMERICAN NATURALIST. [Vor. XXXV.

ciliated projections (auricles) present in many species. Jaws forci-

pate. Soft, flexible, elongated rotifers ; exceedingly numerous

usse. NOTOMMATAD

dz. Withouteyes . . Pleurotrocha

er. Externally riadit on ON water peer

, P. parasitica Jennings

22. Free-swimming, body elongated, ia with numerous

marked annulations ; head enlarged, with a decurved hood

or fleshy proboscis in front; possibly one or two eyes present

, Taphrocampa saundersie Gosse (Fig. 66)

e3. Free-swimming; elongated, convex above and gibbous

posteriorly; flat ventrally ; indications of a neck between head

and body ; a small hook-like proboscis in front; corona a

ventral ciliated area one-third the length of the body; foot

very small; toes minute; auricles present; two dorsal

antenne ; baie with dark granules at the posterior end

93, P. (Diglena) contorta Stokes

d2. One eye

ez. Eye in the neck, — that is, attached to the brain, some

distance from the front of the head

ft. Body with numerous permanent conspicuous annulations

or crenulations ; minute larva-like forms; a small pro-

jection or “ tail” just dorsad of the toes ; protrusible auri-

cles present but frequently remaining hidden

Taphrocampa

gi. Body nearly cylindrical, short, thick ; brain opaque ;

two auricles present

Ar. Toes short and thick, conical, diverging so as to

make an acute angle . . . . . 94, T. annulosa

A2. Toes longer, slender and curved ; so placed that

the two together form es outline of a crescent

i: , T. selenura Gosse (Fig. 68)

£2. Body elongated Reid crenulate; head not

enlarged ; brain containing a club-shaped granular

region; toes short; stout, conical, forming an acute

angle with each other . . 96, T. clavigera Stokes

(See ib No. 92, T. saundersi@)

f2. Body without numerous permanent conspicuous annula-

tions (though divisible into a number of joints)

(Cuticula somewhat stiffened and with a slight longitu-

dinal dorsal cleft, — see Mecum Nos. 181, 182)

£I. Very large species (.2 7 mm. in length) slow

moving, usually enlarged inet the middle ; brain three

(or five) lobed, clear or opaque ; corona extending

No. 417.]

NORTH-AMERICAN INVERTEBRATES. 741

the ventral surface as a large movable ciliated « lip

(except in Nos. 99, 100, and 101); body projecting back-

ward over the foot as a jointed or sac-like tail; auricles

usually present ^, isi 7. . 4. Copeus

Ar. The two toes alike

zz. "Tailslender, usually pointed, prominent, obscurely

two-jointed ; foot two-jointe

ji. Auricles very long (longer than the width of

the head), extending laterally, ciliated yd at the

tip and along the anterior side . 97, C. copeus,

Ehr. (or C. ehrenbergii Gosse) (Fi ig. 69)

j2. Auricles very small or absent

98, C. labiatus Gosse

(97 and 98 are probably identical, 98 being founded

on specimens of 97, with the auricles retracted)

22. Tail thick, rounded, bag-like; auricles large,

broadest at base

jr. Brain three-lobed . . . 99, C. pachyurus

Gosse (Fig. 70)

j2. Brain five-lobed . . 100, C. guinguelobatus

Stokes

(Probably a variation of 99)

73. Tail small, inconspicuous; brain three-lobed

ji. Tail a minute tubercle; corona without lip ;

auricles very small ; toes very small

1o1, C. cerberus Gosse (Fig. 71)

A2. The two toes dissimilar in form; tail slender,

rounded ; no lip 102, C. americanus Pell (Fig. 72)

42. Moderate-sized or small (except No. 106); auricles

present; toes small; brain usually partly opaque, not

three-lobed ; body ss d RR behind, above the

foot, as a large or small “ tail” . Notommata

(A difficult genus with very numerous species, — many of the

American ones not yet identifie

Ar. Tail long, almost or quite as long as the toes, so

that the animal seems to end caudally in three

prominent toes. . . 103, JV. tripus Ehr. (Fig. 73)

A2. Tail inconspicuous or absent

ir. Brain clear ; body fusiform; head narrower than

the body ; auricles small; toes minute cones ; foot

very short . 104, W. drachyota Ehr. (Fig. 74)

72. Brain more or less opaque

jz. Color orange red or brick red, with a brownish

tinge; body long, cylindrical, truncate at each

end ; cilia extending onto ventral surface almost

THE AMERICAN NATURALIST. [VOL. XXXV.

one-fourth the length of the body; toes very

small; a separate foot scarcely distinguishable.

Eye near the caudal end of the brain, almost

completely hidden by the dense opacity of the

later . . 105, JV. truncata Jennings (Fig. 75)

jz. Body San aah or sac-like, swollen behind

kr. Very large and slow ; body sac-like, with a

swollen neck, between which and the body is

a constriction ; foot short ; toes minute

106, JV. collaris Ehr. (Fig. 76)

k2. Smaller; body subcylindrical ; head wide ;

brain with a spherical opaque mass behind,

bearing the eye, and connected by a tube with

the front; foot very short, of two Re e

small, bild a" in form. Length, .

3m 07, N. aurita Ehr. re s

£3. ns N. EA but smaller and more

slender ; toes much longer and decurved ; the

eye sometimes hidden by the opaque brain.

Length about .15 mm. . . 108, W. cyrtopus

Gosse (Fig. 78)

j3. Body cylindrical, long, flexible, with a number

of transverse constrictions and longitudinal folds,

slightly tapering toward each end ; auricles two

stalked spheres, with the cilia confined to the

spheres; toes very small; brain long, cylindrical,

opaque at the caudal end ; eye just in front of

the opacity . . 109, W. Zerulosa Duj. (Fig. 79)

(V. vorax Stokes would be characterized in t he same

manner; it is probably a pics ii of N. torulosa.)

(See also No. 62, N. monopus Jennings)

No auricles, corona a ciliated ince oblique or

extending onto the ventral surface ; brain clear ; body

cylindrical or larviform, usually small Proales

(A large, ill-defined genus, many of its American repre esenta-

tives unidentified or undescribed. The following American

species have been reported)

hr. With a small decurved fleshy proboscis at the front

ir. Parasitic in Vaucheria, forming galls ; body fusi-

form ; toes small, straight, pointed

110, P. w rneckii Ehr.

i2. Free-swimming ; body cylindrical, iind fluted

longitudinally ; proboscis large ; eye very large ; foot

stout ; toes slender ; pointed . , P. felis

Ehr. (Fig. 80)

No.417.] WORTH-AMERICAN INVERTEBRATES. 743

h2. No proboscis

zz. Living in gelatinous masses of alge, or some-

times free in waters full of unicellular alge. Body

nearly cylindrical or oval, slightly arched dorsally ;

eye large, in two parts, at the anterior border of the

brain; foot and toes minute. Adults yellowish

brown in color . . . 112, P. algicola Kellicott

22. Body nearly cylindrical, thick, clumsy ; head

broad, truncate; foot very broad, with a depres-

sion in the median i ; sie conica

P. sordida Gosse (Fig. 81)

73. Body thick, iced n at the head, very flexible

and changeable, colorless ; toes minute, conical ;

eye small, inconspicuous . . 114, P.(F urcularia)

micropus Gosse (Fig. 82)

74. Body slender, soft, larva-like ; toes minute ; foot

indistinguishable. . . . . 115, P. decipiens

75. Body strongly arched dorsally, flat or convex

ventrally ; foot conical ; toes minute

P. gibba Ehr. (Fig. 84)

(The animal described under this name by Hudson and

Gosse is totally different from Ehrenberg's)

«2. Eye frontal, placed near or at the anterior end of the body.

Body nearly cylindrical, somewhat larviform, frequently en-

larged in the lumbar region ; anterior end conical ; corona

oblique ; the two toes usually rather large and conspicuous

Furcularia

$ lender

JT. Toes unequal, very long and s Lo enirere Eli

J?. Toes equal

41. Toes blade-shaped, acute, decurved, the ventral edge

of each notched with two (or sometimes three or more)

teeth... . « » 118, F. forfiicula Ehr, (Fig. 85)

£2. Toes sae and stout at base, and abruptly passing

at about a third of their length from the base into a

hair-like filament . . . . . . 119, £F. semisetifera

Glascott

£3. Body slender, compressed, the ventral line making a

prominent angle; front rounded; gut hup : toes

slender, straight, acute... . . o, F. gracili

"hr. (Fig. 86)

£4. Body oblong, slightly compressed, convex on the back,

abruptly falling off steeply to the foot; toes stylate,

straight, acute, nearly half the length of the body

121, F. gibba Ehr. (Diaschiza semiaperta Gosse?)

(Fig. 87) (F. micropus, — see No. 114)

744 THE AMERICAN NATURALIST. [VoL. XXXV.

dj. Three eyes

er. Eyes in a transverse row near the posterior end of the

brain . . . 122, Z?ZopAhthalmus dorsualis Ehr. (Fig. 88)

e2. One large eye on the brain (‘ cervical Fee two smaller ones

on the front Eosphora

fr. Body — broie tehid? convex doii; ; head

separated from the body by a neck, and bearing two

prominent auricles ; foot slender ; toes short, acute

123, E. aurita Ehr. (Fig. 89)

d4. Eyes two

ez. Eyes frontal; body usually swelling behind and tapering

toward the head; toes cesa large; jaws prominent,

forcipate . Diglena

SH A PETE fei Vibe or hook-like proboscis hanging

onto the face from the dorsal margin of the anterior end

gi. Body massive, subcylindrical, dorsum convex, swollen

ehind; corona a ventral ciliated area; foot a single

large joint ; toes parallel-sided, abruptly pointed

4, D. grandis Gosse (Fig. 90)

£2. Body cylindrical, stout, obtuse at each end, not

swollen behind ; corona a long ventral area ; toes long,

curved, somewhat enlarged at base

125, D. forcipata Ehr. (Fig. 91)

Eyes colorless ; body slender at each end, swollen in

the middle; proboscis acute ; toes long, slender, curved

strongly inward and downward . . 126, D. cércinator

Gosse (Fig. 92)

J?. No proboscis

gt. Body cylindric, long, slender ; front broadly truncate

(obliquely) ; foot short, ves toes long, straight, slender

27, D. caudata Ehr. (Fig. 93)

g2. Body short, cylindric, dicia truncate at each end ;

toes short, straight, acute, projected from the ventral

side, nearly at right angles to the body ax

128, D. catellina ‘Ebr. (Fig. 94)

£j. Body oblong, swollen posteriorly ; head larger than

neck ; toes long, slender, straight, perfectly even in

thickness pointed ; eyes very close together; jaws Pro

trusile ; alimentary canal large, always filled with green

matter . . . . . 129, D. biraphis Gosse de ig. 95)

e2. The two eyes cervical in position . . Distemma

Jr. am a slender cone ; toes stout, Vai toothed at

o, D. forficula Ehr.

B2. Cuticula stiffened, to forts a an armor or lentes Sober LORICATA

41. Foot absent

No. 417.| MORTH-AMERICAN INVERTEBRATES. 745

ór. Lorica oval, compressed, without teeth or spines, formed of two

subequal convex plates ; one tse

, Anapus ovalis Bergendal (Fig. 96)

ġ2. Lorica truncate in front pei: behind, formed of a convex dorsal

and a concave ventral plate ; no teeth nor spines

132, Anuræa hypelasma Gosse (Fi ig. 97)

43. Lorica with six teeth or spines at the anterior edge (usually box-

like in form, open at the anterior and posterior ends ; frequently

with spines at the posterior end also o)

cI. Lorica oblong, convex above, flattened beneath ; dorsal surface

marked off into polygonal areas

dr. Lorica subconical in dorsal view, prolonged at the posterior

end into a long, strong spine which is slightly narrowed at its

base ; polygonal areas of the dorsal surface divided into right

and left sets by a median longitudinal line

3, Anurea cochlearis Gosse (Fig. 98)

42. Lorica as in dz, but porci spine lacking

Anurea cochlearis, var. tecta

4j. Like Zr, but there is a median dorsal row of polygonal areas

on the lorica . . . Anuræa cochlearis, var. stipitata

44. Lorica subquadtangular with a spine at each of the two

posterior lateral angles 134, Anuræa aculeata Ehr. (Fig. 99)

eI. As in d4, but the Mor separating the polygonal areas

Strongly serrate . nurea aculeata, var. serrulata

e2. Asin ZZ, but the two posistidt omm unequal

urca aculeata, var. valga

c2. Dorsal surface of the lorica area with Do mene furrows

or striations . . . Notholca

dr. Loricaa inne te narrow, ebur ey prolonged at the

posterior end into a long spine, equal in length to the body.

Anterior spines large, the right one of the two median spines

immensely developed, so as y be as long as the bo

135, JV. longispina Kellicott (Fig. 100)

d2.. Ovoid, truncated in front, the anterior spines rather short ;

body rounded and without spines posteriorly ; longitudinal

striations of dorsal plate strongly marked; dorsal plate of

lorica much wider than the ventral plate 136, JV. striata Ehr.

dj. Much like JV. striata, but with a subquadrangular projection

from the posterior orifice . striata, var. labis

44. Cylindro-conical in form ; duni plate ixteedal posteriorly

to form a tooth or spine; ventral plate elevated posteriorly to

form a high angular projection . . . 137, N. foliacea Ehr.

42. Foot present

br. Foot transversely wrinkled or ringed (as in Figs. 101-102) ; very

retractile

746 THE AMERICAN NATURALIST. [Vor. XXXV.

cr. Foot ending in a ciliated cup ; lorica dorso-ventrally flattened,

thin; corona transverse, ciliary wreath two lateral semicircles ;

eyes two. . Pterodina

dr. Lorica very flat, pito, Mig. thin Per without teeth or

projections . . 138, P. patina Ebr: (Fig. 101)

(P. valvata is ihe: young g^ 138)

d2. Lorica oval in dorsal view, concave on the dorsal side ;

edges of the lorica very thick ; no teeth or other projections

139, P. reflexa Gosse (Fig. 102)

d3. Lorica thin, broadly ovoid or nearly circular, with a prom-

inent tooth on each side at the posterior lateral margi

140, P. bidentata Ternetz

c2. Foot ending in two toes (usually small). Lote arched dorsally,

flat or slightly convex ventrally. Spines or teeth usually (not

always) present at the anterior dorsal Pu of the lorica (fre-

quently elsewhere also) . . . Brachionus

dI. Lorica with szx teeth or spines at he’ anterior margin of the

dorsal plate (with or without posterior spines)

ez. The two middle anterior spines longest and curving out-

ward; ventro-posterior part of lorica slightly prolonged to

form a sort of sheath for the foot ; on the dorsal side of

sheath a subsquare piece cut out, so that in dorsal view

oot orifice appears to be bounded by two or three

Ed Two posterior lateral spines long, short, or absent.

(Excessively variable) . . 141, B. ġakeri Ehr. (Fig. 103)

ft. Posterior spines short B. bakeri, var. brevispinus Ehr.

fe. Sheath for foot unsymmetrical (right side less developed) ;

body covered with tubercles

B. bakeri, var. tuberculus Turner

e2. Two middle anterior spines longest and curving outward ;

a square plate projecting from the middle of the posterior

edge over the foot orifice. Usually two posterior lateral

spines on the lorica, these disappearing with age

42, B. variabilis Hempel

£3. Anterior spines straight; a deep sinus between the two

median ones. No posterior spines — 143, B. wrceolaris Ehr.

(fr. Reddish, spines Ary

rubens, a variation of B. urceolaris)

d2. Lorica with but four en or spines at the anterior dorsal

margin

er. Median anterior spines very short ; lateral ones very long.

Toes large, each ending in two minute points, so that the foot

at first view,appears to be bifurcated

fr. Two posterior lateral spines on the lorica, the left one

very small, the right one lon

144, B. (or Schizocerca) diversicornis Daday

No. 417.]

NORTH-AMERICAN INVERTEBRATES. 747

J2. The two posterior lateral spines equal

B. (or Schizocerca) diversicornis, var. homoceros Wierz

e2. Four variable (often subequal) anterior spines or teeth.

Anterior ventral margin of lorica sinuous. Posterior part of

the lorica rounded, either without teeth or with two or four

teeth or spines. (Excessively variable) . 145, B. pala Ehr.

JT. Median anterior spines very d no posterior spines

B. pata, var. dorcas Gosse

f2. Median anterior spines Pic long ; two marked posterior

spmes . . B. pala, var. spinosus Wierz.

23. Four anterior spines — about equal length. Lorica sub-

quadrate in dorsal view ; thick, so that a transverse section

would be nearly a circle. A slight invagination in the sides

of the lorica in the lumbar region, causing blunt angles on

the sides of the lorica. Entire surface covered with minute

spinules. No posterior spines

B. punctatus Hempel (Fig. 104)

140,

d3. Anterior margin of lorica without teeth or spines ; at most

c3.

merely sinuate

eZ. Anterior dorsal margin sinuate, with a slight rounded notch

in the middle. Lorica usually with irregular bluntly angled

outlines and with faceted surface. Posterior extremity with

two short blunt processes close to the foot

7, B. angularis Gosse

Jt. The two posterior aee draia into spines

B. angularis, var. bidens Plate

e2. Anterior dorsal margin straight and truncated, with a

small median sinus for the dorsal antenna; lorica thin,

smooth, flexible. Dorsal surface highly arched; ventral,

nearly flat. Toes pointed, and having the inner edges convex,

the outer concave . . . 148, B. mollis Hempel (Fig. 105)

Foot ending in two toes, and situated on the ventral side o

the body ; lorica an irregularly oblong or ellipsoidal box, marked

with grooves and sometimes with ress closed behind, opened

ventrally for the protrusion of the foo à Ploesoma

dr. Lorica firm, with two wide et benedi ies ridges

passing transversely across the middle of the dorsal surface ; a

number of deep longitudinal grooves passing forward and back-

ward from the ends of the transverse grooves ; surface covered

with fine areolations. Lorica in dorsal view about twice as long

as wide, projecting at the anterior dorsal margin in a sharp

median point, flanked by two small points or angles

149, P. lenticulare Herrick (Fig. 106)

42. Body short, little longer than wide ; lorica with grooves and

ridges much as in the last, but more thin and flexible, and not

748 THE AMERICAN NATURALIST. [Vor. XXXV.

covered with areolations. Anterior dorsal margin of the lorica

truncate, without points 150, P. ¢runcatum Levander (Fig. 107)

Lorica soft, covered with coarse areolations, partly arranged

in very irregular rows ; hinder part of the body retractile and

extensile ; when extended the body ends posteriorly in a blunt

Dant uos e eoo 151, P. molle Kellicott

dg. Lorica soft, covered with coarse irregular cuticular vesicles ;

body thick, short, rounded behind

152, P. hudsoni Imhof (Fig. 108)

c4. Foot projecting from the ventral surface, small, lightly ringed,

and ending in a single pointed toe. Lorica compressed later-

ally so as to be nearly circular in side view, with a sort of

projecting collar for the protrusion of the head; in dorsal or

ventral view oblong. Lorica rose color; internal organs blue,

green, and orange . 153, Gastropus stylifer Imhof (Fig. 109)

£5. Foot very small and ending in two small toes ; lorica thin,

compressed laterally, flask-shaped, with the foot projecting from

the ventral surface ; size about .10 mm.

154, Gastropus minor Rousselet

(See also No. 223, Cochleare turbo)

62. Foot present, not transversely wrinkled nor ringed (though often

jointed

cr. Foot distinctly jointed and ending in two small toes; lorica

with an arched dorsal plate and a nearly flat ventral one ; dorsal

surface tuberculate or faceted but not bearing spines. Spines

present at the anterior and posterior margins of the lorica

dr. Lorica with ten spines in front and four behind, its whole

form markedly unsymmetrical ; surface faceted and covered

with raised points . . . . 155, Brachionus militaris Ehr.

d2. Lorica flattened, only slightly arched dorsally ; dorsal sur-

face faceted and roughened ; two spines in front, turned

ventrally at their tips ; two posterior lateral spines. Wo ye

156, JVoteus quadricornis Ehr. (Fig. 110)

c2. Lorica flattened and bearing one or more spines on its dorsal

surface (spines at the posterior margin of the lorica may or may

not be present also). Foot distinct, jointed, ending in one or two

toes

dr. Spines on the dorsal surface of the lorica eight or twelve

(including four at the posterior margin); lorica subquadrate

rough, toothed at the edges; head covered by a chitinous

£dd Xo s cocum c PNE

ez. Eight spines on the dorsal surface of the lorica

Jt. Foot bearing two large dorsal spines or spurs

157, P. collinsii Gosse

fe. Foot without dorsal spines . . 158, P. serica Thorpe

No. 417.] WORTH-AMERICAN INVERTEBRATES. 749

e2. Twelve spines on the dorsal surface of the lorica ; foot

with two long dorsal spines |

159, P. subquadratus Perty (Fig. III)

d2. Two spines on the dorsal surface of the lorica, a very long

one rising from the middle, a short one from near the posterior

margin. Head covered with a large semicircular shiel

160, Stephanops bifurcus Bolton

43. One spine from the middle of the dorsal surface of the

lorica ; otherwise much like the last

161, Stephanops longispinatus Tatem (Fig. 112)

d4. One long spine from in front of the middle of the dorsal sur-

face of the lorica ; also four curved spines from the posterior

margin, and a short median point at the anterior dorsal margin

162, Brachionus (?) gleasonii Up de Graff (Fig. 113)

¢3. A broad, nearly circular projecting plate over the head, appear-

ing like a halo. Lorica cylindrical or pyriform, covering both

dorsal and ventral surfaces ; not faceted nor bearing spines on

the surface (though there may be spines at the posterior edge).

Foot distinct ; toes two

dr. Lorica pyriform, with a narrow neck, slightly prolonged

behind into three subparallel acute spines. Foot with a toe-

like spine above the two toes

163, Stephanops lamellaris Ehr. (Fig. 114)

` 42. Lorica cylindric, with a distinct neck ; dorsal plate prolonged

eg.

behind, over the foot, as a spoon-like shield

164, Stephanops muticus Ehr.

The very short foot ending in one, two, or more very slender,

stiff, bristle-like toes; where more than one are present, these

may be equal or unequal. Lorica cylindrical, fusiform, ovate, or

conical, closed all around but open at each end, smooth, rather

thin; frequently showing a tendency to be spirally curved or

otherwise unsymmetrical in form ; often with one or two longi-

tudinal ridges. Jaws unsymmetrical. One eye

Family RATTULID/E

(The Rattulide are so badly in need of revision that it is difficult to

give a usable key to the species. Moreover, it is certain that many

Species in addition to those hitherto recorded will be found in

America.)

dı. One long, straight, bristle-like toe (if others are present,

these are much shorter than the largest one) . Mastigocerca

er. Lorica with a single marked ridge, passing from the mid-

dorsal line of the lorica forward and to the right. No teeth

or spines at the anterior margin of the lorica. A single long

toe, with three or more minute * substyles ” at its base

750 THE AMERICAN NATURALIST. [Vor. XXXV.

ft. Body fusiform ; ridge high (one-fourth the diameter of

the body). . . . . 165, M. carinata Ehr. (Fig. 115)

fz. Body fusiform ; ridge low . . . 166, M. rattus Ehr.

J3. Body cylindrical, elongated (length about four or five

times the greatest diameter), slightly enlarged in front.

Ridge low, hardly noticeable. Toe about the length of the

brio vos ovs. 7, M. elongata Gosse (Fig. 116)

22. Lorica with two parallel longitudinal ridges close together,

with a groove between them. No teeth at the anterior

margin of the lorica

ft. The two ridges extending three-fourths the length of the

body, which is fusiform in shape ; toe very long, straight

168, M. bicristata Gosse (Fig. 117)

f2. The two parallel ridges close together and extending

only one-half the length of the lorica ; body shorter, ovoid,

or oblong in side view . 169, M. mucosa Stokes (Fig. 118)

£3. Lorica without a ridge, cylindrical, ovate, or conical. (See

also No. 167, M. elongata)

ft. One or more teeth or spines present at the anterior

margin of the lorica

£I. Two unequal teeth or spines at the anterior margin

of the lorica ; body long-fusiform

170, M. bicornis Ehr. (Fig. 119)

£2. Two sharp, slender, equal spines close together at the

anterior margin of the lorica; body a cone, largest in

front and tapering to a toe about one-fourth the length

ofthe body . 171, M. birostris Minkiewicz (Fig. 120)

£j. A single large median tooth projecting over the head

from the anterior dorsal margin of the lorica. Body

cylindrical, somewhat curved ; toe little more than half

as long as the body

172, M. capucina Wierz. & Zach. (Fig. 121)

£4. A single short broad median tooth from the anterior

dorsal edge of the lorica. Body ovate-fusiform, nearly

symmetrical, constricted anteriorly to form a cylindri

neck. Toe not so long as the body ;

173, M. multicrinis Kellicott (Fig. 122)

f2. Anterior dorsal margin of lorica smooth, without teeth

£1. Body very broad, ovate, compressed dorso-ventrally,

unsymmetrical ; toe four-fifths the length of the body

174, M. lata Jennings (Fig. 123)

g2. Body short and thick, arched dorsally, nearly flat

ventrally ; lateral antennæ, at the posterior lateral pus

of the lorica, protected by two prominent projecting

spines ; toe longer than the body

175, M. bicuspes Pell. (Fig. 124)

No.417.] WORTH-AMERICAN INVERTEBRATES. 751

42. Two equal bristle-like toes; these not more than half the

length of the body . . Rattulus

ez. Lorica loug-cylinidvieal, uentos ; anterior margin of the

lorica with a single prominent tooth on the left side, and with

a number of crenulations ; toes very nearly half the length of

the body, each with two substyles at the base

176, R. tigris Müller (Fig. 125)

e2. Body shorter, cylindrical, with two deep furrows encircling

it just in front of the middle. Toes very short (less than

half as long as the body is thick), and frequently retracted

within the lorica, so as to be concealed

177, &. sulcatus Jennings (Fig. 126)

4j. Two unequal bristle-like toes, the shorter being more than

half as long as the longer. (Additional short substyles may be

present) . Coelopus (or Diurella ?)

er. Body holt xd thick (abest adf: as thick as long); curved,

with a ridge on the right side ; one or two short teeth at the

anterior margin of the lorica; longest toe almost or quite

half the length of the body, the other little shorter ; four sub-

styles in addition to the two toes

178, C. porcellus Gosse (Fig. 127)

e2. Like the last, but ses pid about one-fourth as thick as it

islong i . + 179, C. tenuior Gosse

e3. No teeth at initio margin " lorica ; body curved, cylin-

drical or fusiform ; no ridge ; the two toes nearly equal, not

quite so long as ilis body is thick

180, C. dbrachyurus Gosse (Fig. 128)

£5. Lorica cleft down the middle of the back by a fissure, whose

sides are united by membrane ; open at both ends for the projec-

tion of the head and foot. Toes blade-shaped

Family SALPINAD/E

dr. Lorica only slightly developed, covering only the dorsal half

of the body ; median fissure not strongly marked (resembling

the species of Furcularia, Nos. 117-121). Oneeye Diaschiza

eZ. Minute, swiftly moving ; body short, cuneiform or cylin-

drical ; head broad ; lorica covering only the posterior third

of the body ; median fissure broad, inconspicuous

181, D. lacinulata O. F. M. (Fig. 129)

e2. Larger, body laterally compressed, arched dorsally ; toes

equal in length to the height of the body, curved toward the

dorsal side . 182, D. semiaperta Gosse (Fig. 130)

d2. Lorica well divelóged, an oblong box enclosing the body,

Open at both ends and distinctly cleft down the middle of the

back ; furnished with spines or teeth at the anterior or pos-

terior margins or both ; one eye . . + Salpi

752 THE AMERICAN NATURALIST. [Vor. XXXV.

ez. Spines or teeth on the lorica as follows: two anterior

dorsal two anterior ventral, one (median) posterior dorsal,

two posterior ventral. (Spines all rather short, tooth-like)

. 183, .S. mucronata Ehr. (Fig. 131)

e2. Spines or teeth as follows : no anterior dorsals, two anterior

ventral one posterior dorsal, two posterior ventral. (The

four following species are distinguished, all having the above

characters ; it is possible that they should be considered

mere variations of a single species, S. ventralis Ehr.)

fr. Anterior ventral spines short and straight ; posterior

spines all short, the ventral ones recurved

184, S. brevispina Ehr.

fe. Anterior ventral spines very short, posterior dorsal spine

short, decurved ; posterior ventral pair longer than the

posterior dorsal ; lorica with a stippled collar

185, S. ventralis Ehr.

f$. Anterior ventral spines short, straight ; posterior spines

all long and straight, the ventral ones much the longer ;

lorica surface not stippled ; dorsal cleft wide

186, .S. macracantha Gosse

f4. Anterior ventral spines short, incurved ; posterior dor-

sal conical, short, arched ; posterior ventral long, stout,

incurved ; dorsal cleft narrow ; lorica frequently stippled

187, .S. eustala Gosse (Fig. 132)

. No posterior spines; anterior dorsal pair very long ;

anterior ventral pair also long and slender, but only about

half as long as the dorsal ones. Lorica broad

188, S. macrocera Jennings (Fig. 133)

d3. Lorica as in d2, but without spines ; the eye lacking ——

Diplax

er. Body triangular in section ; lateral outline of lorica nearly

ovate, but strongly arched dorsally and nearly flat ventrally

189, D. trigona Gosse

£2. Body much compressed, long and narrow in dorsal view, in

side view nearly a parallelogram . 190, D. compressa Gosse

có. Foot very long, of several joints; the two toes very long ;

lorica entire (not cleft dorsally nor ventrally), vase-shaped Ot core

pressed, not bearing spines. Head furnished with a chitinous

covering. Eye one :

dr. Lorica vase-shaped, faceted, and with surface roughened ;

head retractile within a chitinous cap; eye one; foot bearing

two spines dorsally ; foot and toes together nearly or quie

twice the length of the body. . . . + +: : Dinocharis

ez. A short spine dorsally between the two toes

191, D. focillum Ehr. (Fig 134)

No.4175.] MWORTH-AMERICAN INVERTEBRATES. 753

e2. No spine between the two toes . 192, D. Zetractis Ehr.

d2. Lorica somewhat vase-shaped, thin, smooth, and transparent ;

head with a thin chitinous covering ; eye very close to or upon

De mastar oo 22000. Ved See pl ee ae

ez. Body nearly cylindrical or slightly compressed laterally ;

body, foot, and toes of about equal length

193, S. longicaudatum Ehr. (Fig. 135)

22. Body broad, ovate or pear-shaped ; toes about as long as

body and foot together 194, S. eudactylotum Gosse (Fig. 136)

c7. Lorica of two dissimilar plates, one dorsal and one ventral, the

former usually larger and arched (except in No. 201), the latter

flat or slightly convex. Large transparent rotifers, with a single

eye. Foot jointed, the two toes large, usually blade-s aped

Euchlanis

di. Lorica oval or ovate, gently arched above, nearly flat below ;

transverse section a low segment of a circle

ez. Ventral plate with a flange projecting laterally from its

junction with the body; anterior dorsal edge with a broad

gap having a straight bottom (Fig. 138); hind dorsal edge

notched. . . . . . . 195, E. dilatata Ebr. (Fig. 137)

e2. Ventral plate without a lateral flange; anterior dorsal

margin with a small subsquare notch | 196, Æ. deflexa Gosse

42. Lorica a long, narrow oval or an ellipse, depressed ; anterior

dorsal edge membranous ; transverse section a low circular

segment ; ventral plate elliptical and broadest at the hind end

197, EK. lyra Hudson

43. ‘Lorica broadly oval or ovoid, constricted near the middle,

the dorsal plate much wider than the ventral and turned

downward and inward . 108, Æ. pyriformis Gosse (Fig. 139)

d4. Lorica roof-shaped, with sloping sides ; not rising to a ridge;

cleft for a short distance behind. Ventral plate flat, smaller in

outline than the dorsal. A small species, 199, Æ. oropha Gosse

d5. Lorica with a high median dorsal keel; lateral edges of the

lorica extending laterally into two wide shelves, so that the

animal seems to have three keels, and is triangular in section.

Outline from above ovoid . . . . 200, E. triquetra Ehr.

dó. Dorsal plate carinate ; lorica ovate, flask-shaped ; foot four-

jointed ; toes more than half the length of the lorica

201, E. ampuliformis Herrick (E. propatula) Gosse (?) (Fig. 140)

£8. Lorica of two plates, the ventral one nearly flat, the dorsal one

slightly or considerably elevated. The two plates separated by a

deep lateral furrow covered with flexible membrane. Body ending

posteriorly in either one or two large rod-shaped toes (the foot

Proper being short and inconspicuous). One eye

Family CATHYPNADE

754 THE AMERICAN NATURALIST. [Vor. XXXV.

di. Toes two

ez. Lorica subcircular or broadly ovoid, with a wide and deep

lateral furrow. Cathypna

ft. Lorica rather ftoi, ‘dorsal and yeiai pisti nearly

equal in size, broadly oval in outline ; toes two-fifths as long

as the lorica, each with a distinct ange or shoulder at the

side, near the tip, and with a small sharp claw, which con-

tinues the inner side of the toe 202, C. Zuna Ehr. (Fig. 141)

J?. Lorica ovoid, the dorsal plate ending behind in a sub-

square plate with its posterior lateral corners extended into

distinct angles. Toes rod-shaped, very long (almost as

long as the lorica). . 203, C. /eontina Turner (Fig. 142)

J3- Very large (.31 mm. in length); lorica long ovoid or

truncate elliptical in form ; dorsal plate projecting pos-

teriorly as a semicircular plate over the foot. Toes about

half the length of the lorica . . 4, C. ungulata Gosse

£2. Lorica longer, usually a ve ris aed in front ; lateral

furrow not pronounced . Distyla

Jt. Dorsal plate lacini: £e, iid off into polygoni

are

gI. Dorsal plate ending behind in a quadrangular projec-

tion with may pita sides

205, D. ohioensis Herrick (Fig. 143)

£2. Dorsal plate ending behind in two sharp points sepa-

rated by a broad gap . 206, D. stokesii Pell (Fig. 144)

£3. Dorsal plate ending behind in a single sharp point

207, D. ludwigii Eckstein (Fig. 145)

£4. Dorsal plate not ending behind in projections, but

bearing in the posterior lateral region two short project-

ing teeth, protecting two lateral antennz. Foot conspic-

uous, three-jointed 208, D. spinifera Western (Fig. 146)

f2. Dorsal and ventral plates marked with small crescentic

elevations arranged in somewhat regular patterns. Lorica

of truncate elliptical form, without posterior projections

209, D. signifera Jennings (Fig. 147)

J3- Lorica flexible, smooth, or marked with irregular ar wrin-

kles or longitudinal folds

£I. Small, lorica parallel-sided, soft, with irregular wrin-

kles and longitudinal folds . . 210, D. flexilis Gosse

£2. Lorica long, flexible, nearly parallel-sided when

extended (Fig. 148); when completely retracted broader

in front and ending in two strong incurved points (Fig.

149). Dorsal surface with a few irregular long itudinal

folds . 211, D. gzssensis Eckstein (Figs. 148 - 149)

No. 417.] MWORTH-AMERICAN INVERTEBRATES. 755

&3. Lorica soft, much broader behind and tapering for-

ward to the narrow head when extended ; broad ovate,

truncate, when retracted. Toes short, with claw about

one-third the length of the toe . 212, D. inermis Gosse

d2. One rod-shaped toe . . Monostyla

eZ. Lorica broad ovate, flat, Fuels apta d into two long

sharp, outwardly curved spines, separated by a narrow rounded

NBNER oso 213, M. quadridentata Ehr. (Fig. 150)

€2. Lorica ovate, toe Manit fusiform

214, M. closterocerca Schmarda (Fig. 151)

e3. Lorica without spines ; toe rod-shaped or tapering to the tip

Jr. Lorica broadly ovate, with a crescentic concavity in

front when the head is retracted. Toe with parallel sides

215, M. lunaris Ehr. (Fig. 152)

J2. Lorica ovate, without a crescentic concavity in front

when the head is retracted, but nearly truncate. Toe

short, rod-like . . . 216, M. cornuta Ehr. (Fig. 153)

J3. Dorsal plate high eres ventral somewhat convex ;

anterior dorsal edge of lorica with a shallow, rounded or

quadrate notch ; ventral plate with a much deeper, broader

notch . . 217, M. bulla Gosse (Fig. 154)

du. E. ovate ; two anterior lateral points when the head

is retracted. Ventral plate flexible, with irregular longi-

tudinal folds . . . 218, M. arcuata Bryce (Fig. 155)

J5. Lorica subcircular, ae little longer than broad; two

inwardly curved sharp points at the anterior edge when the

head is retracted ; toe rod-like, ending in two short claws

projecting from between two minute spines

219, M. robusta Stokes (Fig. 156)

Jó. Lorica subovate, with a straight anterior margin when

contracted (insufficiently described)

220, M. truncata Turner

J7. Lorica broadly ovate, truncate, rounded dorsally. Dor-

sal plate strengthened by two diverging longitudinal ribs

about equidistant from each other and from the lateral

edges of the plate, jig the anterior margin slightly

angulate where they join it. . 221, M. ovata Forbes

*4. Lorica a soft, flexible, destin skin, hardly deserving

the name of lorica. Body oblong, samba se Toe rod-

Shaped, with a short claw. . . . 222, M. mollis Gosse

c9. Lorica of a single piece, like a coat, iios only the anterior

half of the dorsal surface of the — or less ; foot long, jointed,

with two minute toes . . . . . Cochleare

dr. Lorica threesided . . . 225 C. indii Gosse (Fig. 157)

756 THE AMERICAN NATURALIST. [VoL. XXXV.

cro. Head surmounted by an arched chitinous shield, appearing in

side view like a hook; lorica either arched, and compressed

laterally, or dorso-ventrally flattene

dr. Lorica arched, somewhat compressed laterally, so as to be

higher than wide; open behind and in front, and sometimes

open on the ventral side. Minute, inconspicuous rotifers

Colurus

ez. Lorica ovate as viewed from above ; produced caudally

(as seen in dorsal view) into two acute spines, separated by a

deep sinus ; ventral surface of lorica cleft

224, C. deffexus Ehr. (Fig. 158)

e2. Lorica not cleft ventrally ; excavate behind, so as to form

two teeth with a shallow notch between them

225, C. bicuspidatus Ehr. (Fig. 159)

£3. Pear-shaped, widest behind, in dorsal view ; ventral sur-

face cleft; caudal notch very shallow, between two short

terminal points ; foot and toes three-fourths the length of the

MODA Secu . . 226, C.caudatus Ehr.

e4. Ovate in ion. or dein view, the posterior end rounded,

without points ; lorica cleft ventrally. Foot very small

227, C. obtusus Gosse (Fig. 160)

£5. Elongate-ovate in dorsal view ; caudal margin rounded,

without cleft or points ; ventral cleft interrupted in front of

the middle, so that the openings for the head and foot are

not continuous. . 228, C. agilis Stokes (Fig. 161)

eó. Lorica ovate in decl view, high behind, low in front; a

long, stiff, pointed, hyaline crest extending backward and .

upward from the "re of the anterior dorsal edge of the

lorica . , C. cristatus Rousselet (Fig. 162)

d2. Height and width of Vitas about the same ; lorica forming

an ovate box with surface marked into areas ; a thin ridge on

the mid-dorsal line ; a similar ridge on the mid-ventral line from

the anterior margin to pag the middle of the length

o, Metopidia salpina Ehr. (Fig. 163)

d3. Lorica flattened, bas than high, usually turtle-like in

appearance; open only in front and behind, not along the

ventral middle line ... Metopidia

ez. Outline of lorica varying tom elliptical through oval and

ovate to orbicular, but without teeth, spines, or prominent

angles (except at the sides of the foot and head, where angles

are necessarily formed, owing to the openings in the lorica)

ft. Lorica nearly circular, much depressed ; dorsal plate

with a very low decia median ridge and with a submar-

ginal line of corrugati . . 231, M. solidus Gosse

Je. Lorca oval or ges ph depressed, evenly rounded

No. 417.] MWORTH-AMERICAN INVERTEBRATES. 757

above ; ventral plate excavate behind for the foot; eyes

two . 232, M. lepadella Ehr. (Fig. 164)

Fa. Similar to d fat but with four eyes. (Perhaps only a

variation of the last) . . 233, M. bractea Ehr.

J4. Lorica elliptical or ord nae: toes long, slender ;

,eyes two or four ; foot of three nearly equal joints

234, M. oblonga Ehr. (Fig. 165)

J5. Like the last (of which it is probably only a variation),

but with the third joint of the foot equal in length to the

other two together ; the two toes protruded between two

minute lateral spines . 235, M. dentata Turner (Fig. 166)

fo. As in M. oblonga, but with the anterior margin of the

lorica covered with points, so as to form a stippled collar ;

the angles of the lorica at the sides of the head very sharp,

so as almost to form spines (variation of M. oblonga ?)

» M. collaris Stokes (Fig. 167)

e2. Lorica rhomboid-ovate in outline ; dorsal plate roof-shaped,

lower behind and ending in an obtuse point ; ventral surface

237, M. rhomboides Gosse

£3. Loca siis viding bekindi in an acute point

238, M. acuminata Ehr.

e4. Lorica with three wide, thin wings, one dorsal and two

lateral ; the dorsal view nearly circular

239, M. triptera Ehr.

e5. Lorica broad behind, having four prominent angles, two

lateral and two posterior

240, M. ehrenbergii Perty (Fig. 168)

e6. Lorica with two great spines at the anterior dorsal margin ;

these separated by a narrow median notch

241, M. cornuta Schmarda (Fig. 169)

ANN ARBOR, MicH., Feb. 5, 1901.

SOME OF THE MORE IMPORTANT LITERATURE ON THE

ROTATORIA.

I. Most IMPORTANT GENERAL SYSTEMATIC WORKS.

EHRENBERG, C. G. Die Infusionsthierchen als vollkommene Organismen.

Leipzig. 1838. (Still one of the most important works.)

N, C. T., and Gosse, P. H. The Rotifera, or Wheel Animalcules.

2 vols., with supplement. London, Longmans, Green & Co. 1889.

(Indispensable. )

Hupsox

758 THE AMERICAN NATURALIST.

JANsoN, OTTO. Versuch einer Uebersicht über die Rotatorien-Familie der

Philodinaeen. Beilage zum XII. Bande der Adhdlg. d. Naturw.

Vereins zu Bremen. 1893. (Monograph of the Bdelloida.)

ROUSSELET, C. F. List of New Rotifers since 1893. Journ. Roy. Micr.

Soc. pp. 450-458. 1893

Second List of Rotifers since 1889. Jd¢d. pp. 10-15. 1897.

WEBER, E. F. Faune Rotatorienne du Bassin du Leman. Revue Suisse

de Zool. Tome v, pp. 263-785, 25 plates. (Beautiful figures of

many species.)

II. PAPERS ON AMERICAN ROTATORIA.

HEMPEL, A. A List of the Protozoa and Rotifera found in the Illinois

River and Adjacent Lakes at Havana, Ill. Bull. TIl. State Lab. of

Nat. Hist. Vol. v, Art. vi, pp. 301-388. 9

HERRICK, C. L. Notes on American Rotifers. Bull. Sci. Lab. Denison

Univ. Vol. i, pp. 43-62. 1885.

JENNINGS, H. S. The Rotatoria of the Great Lakes and of Some of the

Inland Lakes of Michigan. Bull. Mich. Fish Commission. No. 3,

pp. I-34. 1894.

Report on the Rotatoria. From A Biological Examination of

Lake Michigan in the Traverse Bay Region. Bull. Mich. Fish

Commission. No. 6, pp. 85-93. 18

Rotatoria of the United States. Bull. U.S. Fish Commission for

1899. pp. 67-104. 1900. (List of all American species thus far

recorded, with literature on American Rotifera.

KELLICOTT, D. S. Partial List of the Rotifera of Shiawassee River at

Corunna, Mich. Proc. Amer. Soc. Micr. Vol. x. 1888.

The Rotifera of Sandusky Bay. Zid. Vol. xviii, pp. 155-164, -

1896, and vol. xix, pp. 43-54, 1897.

TURNER, C. H. Notes upon the Cladocera, Copepoda, Ostracoda, and

Rotifera of Cincinnati, with Descriptions of New Species. Bul.

Sci. Lab, Denison Univ. Vol. vi, pt. ii, pp. 57-74. 1892.

III. SEE ALSO NUMEROUS RECENT PAPERS BY ROUSSELET (MOSTLY IN

THE JOURNAL OF THE QUEKETT MICROSCOPICAL CLUB, IMPOR-

TANT), Hoop, STENROOS, LEVANDER, BILFINGER, WIERZEJSKI,

SCORIKOW, WESTERN, DADAY, AND OTHERS.

760 THE AMERICAN. NATURALIST. [VoL. XXXV.

LIST OF FIGURES— PLATE I.

, ee a n Rousselet.

tata, a W

p dietis. sud Hudson and Gosse.

eber.

. 5 algicola, after "nei and Gosse.

Y EDA TS after Weber

IO. , jaws, after Hodséh and Gosse.

11. Acyclus inquietus, after Leidy, hon Hudson and Gosse.

12. Apsilus bipera, ventral view of head, after Foulke, from Stokes.

1i. * bucinedax, after Stokes

14. Melicerta ringens, in tube, after Hudson and Gosse.

No.41] MORTH-AMERICAN INVERTEBRATES. 761

PLATE I.

THE AMERICAN NATURALIST. [Vor. XXXV.

LIST OF FIGURES — PLATE II.

: medi ringens, removed from tube, after Weber.

, jaws, after Hudson and Gosse.

"S conifera, after Hudson and Gosse.

" janus, after Hudson and Gosse

Limnias annulatus, after Weber.

. Cephalosiphon limnias, after Weber.

. CEcistes melicerta, side view.

* crystallinus, after Hudson and Gosse.

«intermedius, after Hudson and Gosse.

s eh anis semibullata, after Weber.

. alboflavicans, Lad part, after Hudson and Gosse.

; o ugs tolres, after Hudson and Gos

3 icornis, Min dE

Rotifer macroceros, after Weber.

” rins nius, after Weber.

ris (corona retracted), after Weber.

after Weber.

; vulga

s rn of Rotifer trisecatus

dicbus after Weber.

M UM * — tardus, after Weber.

ROR

R Q.

Fos

764

THE AMERICAN NATURALIST. [Vor. XXXV.

LIST OF FIGURES— PLATE III.

. Jaws of Philodina aculeata (ramate aet after Weber.

. Philodina "ue igne after Weber

. Callidina e

« Balink after Janson.

* elegans, jaws, after Janson.

* ^ constricta, jaws, after Janson.

" P , spurs, after Janson.

* socialis, spurs, after Janson.

* musculosa, jaws, after Janson.

7 a , Spurs, after Janson.

. Adineta vaga, anterior end, after Weber.

. Microcodon clavus, after Weber.

otommata monopus

i Msi ie anterior end, after Hudson and Gosse.

Pedal m, after Weber

Palais; platyptera, after robes

. Triarthra longiseta, after Ehrenberg.

. Asp domom myrmeleo, after Weber

* Aue ebbesbornii, side view, after Hudson and Gosse.

herrickii, after Wierzejski.

" brightwellii, jaws page type), after Wierzejski.

gs girodi, jaws, after Wierzejski.

x appa ans: Hs after Kellico

caudis, after "ab ae Gosse.

No. 417.] WORTH-AMERICAN INVERTEBRATES. 765

Toast

PA EE EI N

STO PEDIS jer

ia an

fags]

PLATE III.

766

THE AMERICAN NATURALIST. [VorL. XXXV.

LIST OF FIGURES — PLATE IV.

. Anarthra aptera, after Ho

od.

vanga ges after poeta ski.

Hudson and Gosse.

. Triphylus venen dani Western.

. Cyrtonia tuba, after MN

. Hydatina senta, after

^ AME clavulatus, kid an and Gosse.

pelagicus.

i bis aegros saundersiz, after Hudson and Gosse.

annulosa, after Hudson and Gosse.

selenura, toes, after Hudson and Gosse.

“

[11

. Copeus copeus, after Ehrenberg.

pachyurus, after Weber.

cerberus, after Hudson and Gosse.

icanus, after Pell.

[11

ameri

i cime see after Stokes.

bra

"gin ota, after Weber.

"i truncata.

“ ok after Ehrenberg.

“ aurita, after Hudson and Gosse.

No. 417.]

NORTH-AMERICAN INVERTEBRATES.

HA.

3 — — Án

SS = a

E APTA

Mee ^t

E, C

e sr vip 1

PLATE IV.

767

768

THE AMERICAN NATURALIST. [Vor. XXXV.

LIST OF FIGURES— PLATE V.

: s Speen iar after Hudson and Gos

rulosa, after Cohn, from vedi and Gosse.

T Peon fais, nidi Hudson and Gosse.

ordida, after Hudson and Gosse.

: Pnn Hei after Weber.

gra

s, after Hudson and Gosse.

» gi ees after Ehre

y nberg.

. Triophthalmus J orenntin: after Ehrenberg.

9. Eosphora aurita, after Ehrenberg.

: gases grandis, after. Hudson and Gosse.

orcipata, after Ehrenber

" circinator, after Hudson Lam Gosse.

" caudata, after Hudson and Gosse.

s catellina, after Weber.

s ie a after Hudson and Gosse.

. Anapus ovali

. Anurza eed after Hudson and Gosse.

ix co Sii after Weber.

p leata, after Hudson and Gosse

Nihol iouis kik, lorica, after Hudson uid Gosse.

: diee de patina, after Hudson and Gosse

refl

exa, after Rousselet.

No.417.] MWORTH-AMERICAN INVERTEBRATES. 769

PLATE V.

THE AMERICAN NATURALIST. [Vor. XXXV.

LIST OF FIGURES — PLATE VI.

. Brachionus bakeri.

i capa spen Hempel.

ft s, after H

5- mpel.

: Agente E erue ies Wierzejski.

amp after Weber

e qaas i, after Wierzejski.

Gastropus datos after Weber

. Noteus quadricornis, after i oltes and Gosse.

. Polychattus subquadrat

. Stephanops epatis. after Weber.

Brachionus (?) gleasonii, after Vorce, from Hudson and Gosse.

Weber.

. Stephanops lamellaris, after

nata.

pun dicus cari

" bicristata.

icornis.

us birostris, after Minkiewicz.

No. 417.] WORTH-AMERICAN INVERTEBRATES 77

. I

M c)

ESO Q ok OFAN

[ OQ Oe VE }

O oe Vy M

e, Mig TQ esf

; fo q

MO à D OS

Q e $]

MON,

w g Jy

PLATE VI.

712

THE AMERICAN NATURALIST. [Vor. XXXV.

LIST OF sii o Hia ies VII.

à Maegan capucina, after Wierzejski.

multi

ticrinis, after Kellicott.

p ie

* bicuspes.

: bsc. €—

: Saa sem sob:

“ brachyurus.

— lacinulata, after Weber.

semiaperta,

after Hudson and Gosse.

i Supin mucronata, ata Hudson and Gosse.

eustala, after Hudson and Gosse.

« — macrocera

4. Dinocharis po oiim, after t Hadron and Gosse.

. Scaridium longicaudatum, af

eber.

" eudactylotum, after Hudson and Gosse.

No.4175.] MWORTH-AMERICAN INVERTEBRATES. 7731

PLATE VII.

774

137-

138.

THE AMERICAN NATURALIST. (Vor. XXXV.

LIST OF FIGURES — PLATE VIII.

re pin after Hudson and Goss

i V amps luna.

; PPA ohioensis.

€

, anterior dorsal edge of oie after Hudson and Gosse.

ikia , after Hudson and Goss

ampuliformis, after Herrick.

ntina.

stokesii.

ludwigii.

simi after Western.

signifer

gissensis, zii extended.

i ovv quadridentata

osterocerca.

No.4175.] NORTH-AMERICAN INVERTEBRATES. 775

776 THE AMERICAN NATURALIST. [VoL. XXXV.

153.

170.

171.

s "—- hic ag

LIST OF FIGURES — PLATE IX.

Monostyla cornuta.

bulla.

Ns arcuata.

u robusta, after Stokes.

. Cochleare turbo, after Hudson and Gosse.

: Meraba deflexus, after Hudson and Gosse.

TRN after Hudson and Gosse.

u obtusus, after Hudson and Goss

" agilis, ventral view of lorica, par Stokes.

u cristatus, after Rousselet.

after We

padella, after xhcdibdig.

es anm after Ehrenberg.

" dentata, after Turner.

? collaris, after Stokes.

“4 ehrenbergii. ;

" ?) cornuta, after Schmarda.

Malleate jaws, after Hudson and Gosse.

Forcipate jaws, after Hudson and Gosse.

No.417.] WORTH-AMERICAN INVERTEBRATES. 777

REVIEWS OF RECENT LITERATURE.

ZOOLOGY.

Reptilian Affinities of Primitive Mammals. — Sixta’ has inves-

tigated the osteology of the foot in monotremes with the view of

ascertaining the affinities of this primitive group of mammals. The

monotremes possess a humerus with an epicondyloid foramen like

that in reptiles, especially in the fossil Dimetrodon. Their radius,

ulna, and carpal elements are arranged on the primitive reptilian

plan, and yet their front feet are not unlike those of some mar-

supials (Dasyurus). In the hind foot the astragalus and calcaneum

resemble those of reptiles, and the foot, on the whole, is oriented

as in that group. The tarsals are as numerous as in the marsupials,

but their forms and positions are typically monotrematic. The

structure of the hind foot indicates an intermediate position for the

monotremes between reptiles and marsupials. This opinion is

further supported by a large body of evidence drawn from other

organs, such as the heart and chief blood vessels (Hochstetter), the

brain (Ziehen, Smith), the copulatory organs (Gadow), the embryonic

membranes (Semon), the teeth (Poulton, Cope, Osborn), and skull.

Anatomical Miscellanies. — The three numbers of the Hopkins

Hospital Bulletin? for April, May, and June are combined into a

single part and devoted to a collection of some twenty-three essays,

mostly anatomical The series opens with Barker’s address on

the Study of Anatomy, delivered at the Rush Medical College in

October, 1900; Bardeen describes a new carbon-dioxide freezing

microtome, Born's wax-plate reconstruction method as used in the

Anatomical Laboratory at Hopkins, and calls attention to the

Importance of ordinary dissection-room material in scientific study.

The cold-storage method of preserving bodies for dissection is dwelt

* Sixta, V. Vergleichend-osteologische Untersuchung über den Bau der Füsse

der Reptilien, Monotremen, und Marsupialier, Zool. Anzeiger, Bd. xxiv (1901),

- 321-332.

? Bulletin of the Johns Hopkins Hospital, vol. xii, Nos. 121-122, 125, April-

May-June, rgor.

779

780 THE AMERICAN NATURALIST. [VoL. XXXV.

upon by Kerr. Harrison discusses the occurrence of tails in man,

and reports on an interesting case brought to his notice by Dr. Wat-

son. Brush contributes notes on cervical ribs, and Walker gives

an account of cases of hereditary anchyloses of phalangeal joints.

The development of the pig’s intestine is described by MacCallum,

and the structure of the gall-bladder by Sudler; of a less anatomical

character are Halsted’s report on the effects of injecting bile into

the pancreas, and Opie’s account of the etiology of acute hemor-

rhagic pancreatitis. The axillary artery and its variations are

described by Hitzrot, the blood vessels of the lymphatic glands by

Calvert, and the anatomical origin of the lymphatics in the liver

by Mall, who also contributes a note on the basement membranes

of the kidney tubules. Lewis gives an account of the pectoralis

major muscle in man, The development of the human diaphragm

is fully described by Mall. Neurology is represented by Mellus’s

article on the bilateral relations of the cerebral cortex, Long’s paper

on the development of the nuclei of the pons, and F owler’s descrip-

tion of a model of the dentate nucleus of the cerebellum. The

development of the generative tracts in white ants is described by

Knower, and the modification of normal menstruation by Mosher.

The series gives substantial evidence of the great quantity and high

quality of advanced work done in the Hopkins Hospital. P.

The Position of the Centrosome in Resting Cells. — By a thor-

ough study of the segmenting eggs in Ascaris, zur Strassen" has

shown that at each resting stage the centrosome takes up 4 position

between the nucleus and the center of the exposed surface of the

blastomere, Ze, lies in the axis of the cell near its distal pole.

This position is always attained, even though it involves à consider-

able migration on the part of the centrosome. As similar oe

tions have been figured by other authors in the segmenting €g85

several animals, and as the centrosome in ordinary epithelium regu-

larly occurs in this position, zur Strassen believes that the rule

discovered by him for Ascaris may prove to be of general sies

tion for all kinds of epithelial tissue, including the blastoderms

eggs. These observations favor the view that epithelial cells gens

a true polarity, but this polarity is not one which has arisen first st

adult epithelia. As is shown by its occurrence in the early qe

mentation stages, it is a polarity that is fairly comparable with tha

1Zur Strassen, O. Ueber die Lage der Centrosomen in ruhenden ers

Archiv fiir Entwickelungsmechanik der Organismen, Bd. xii (1901), PP- IM^

No.4175.] REVIEWS OF RECENT LITERATURE. 781

of a flagellate protozoan cell, the ancestor of the colony of cells from

which the metazoan body is believed to have taken its origin. Thus

zur Strassen regards the polarity of epithelial cells not as a newly

acquired feature, but as an inherited one derived from forms as

primitive, possibly, as the protozoa. P.

The Intestine of Cetaceans. — Süssbach! has described in detail

the structure of the intestine in a number of cetacean embryos

recently collected by Kükenthal. In the toothed whales there is no

division into a large and small intestine, and, except in Platanista,

there is no ccecum present, structural conditions always observed in

the baleen whales. 'The toothed whales always possess a simple

mesentery, without any trace of the complications introduced in the

baleen whales by the folding of the intestine about parts of the mes-

entery. The toothed whales have a relatively longer intestine than

the baleen whales, but the configuration of the intestinal surface

seems to be independent for the two groups; thus some of the

toothed whales with short intestines have much the same kind of

Intestinal surface as that in the baleen whales, though a general rule

Was found to the effect that the shorter the intestine is, the more

complicated are the folds on its surface. Notwithstanding this last

circumstance, the condition of the intestine points to the complete

separateness of the two groups of living cetaceans, the baleen and

the toothed whales. — P

Notes on Recent Fish Literature. — In the Proceedings of the

California Academy of Sciences (Zool, Vol. II, Nos. 7, 8) Jordan

and Snyder describe two very remarkable new genera of Japanese

fishes, The one, Ereunias (grallator), is a cottoid, allied to Trig-

lops, but without ventrals, and with the four lowermost pectoral

Tays developed as detached feelers, as in Trigla. The other genus,

seus (sachi), is like Podothecus, but with enormously developed

orsal and anal fins. It belongs to the Agonidz.

= e same Procedings Jordan and Starks describe three new

due an Japan, Snyderina Jyamano&ami, Pomacentrus celestis, and

c Doer deani. Snyderina is a new genus of Scorpznidse, allied

la osopodasys. All these species are represented by admirable

P'ates, the work of Mrs, Chloe Lesley Starks.

1 "

Ste aen, S. Der Darm der Cetaceen, Jenaische Zeitschrift, Bd. xxxv

) PP- 495-542, Taf. XVI, XVII.

782 THE AMERICAN NATURALIST. [VOL XXXV

In the Abhandlungen der Senchenbergischen Naturforschenden Gesell-

schaft of Frankfurt (Vol. XXV, No. 2), Dr. Steindachner gives an

account of the fishes collected by Dr. Kükenthal in Molucca and

Borneo. Two hundred and eight species are enumerated, six of

them new, most of these being well figured.

In the Records of Australian Museum (Vol. IV, No. 1), Mr. E. R.

Waite discovers that the sharks of the genera Hemiscyllium and

Chiloscyllium bring forth their young alive. He therefore very

properly separates these genera from the Scylliorhinidz as a distinct

family, Hemiscylliide. The genus Orectolobus (Crossorhinus) is

also viviparous, and is recognized as a distinct family, Orectolobidz.

A figure is given of Hemiscyllium modestum, and also of the Austra-

lian dogfish, Sgualus megalops, a species which has a very close

relative in Japan.

As fishes of Lord Howe’s Island, Mr. Waite figures Upeneus pleu-

rostigma, Apogon norfolcensis, Iniistius cacatua, Chatodon tricinctus,

and Monacanthius homensis. We may note that the deep green color

of the teeth of Pseudoscarus guacamaia and related species is not the

result of staining through the food. It is inborn, unvarying, and a

result of distinct specialization. DS

An Elementary Book on Lepidoptera. — This is really an excel-

lent book, both in conception and in execution. Reversing the

order of her title, Miss Dickerson deals in* Part I with the butter-

flies. The monarch Danais archippus holds the place of honor,

and its life history and structural characters are given accurately

and with sufficient detail ; the other species, twelve in number, with

the exception of two swallowtails, Papilios, a white, Pieris, and a

sulphur, Colias, are all closely allied nymphalids.

In Part II Callosamia promethea is made the starting point, and

the twenty and more species that follow are Bombyces, sensu lat.,

and Sphinges. :

The treatment in both these parts is so happy that the omission

. of an adequate account of the life history of a blue, Lyczna, and of

a skipper, hesperid, among the butterflies, and of two or more of the

lower moths, is especially regrettable. i

Part III is divided between a chapter on relationship, showing

classification and ancestry of moths and butterflies, and practi

suggestions how to collect, keep, and study butterflies and moths.

! Dickerson, Mary C. Moths and Butterflies. Boston, Ginn & Company,

I9OI. xviii + 344 pp., 244 illustrations.

No.417.] REVIEWS OF RECENT LITERATURE. 783

The chapter on relationship is from its nature more open to criti-

cism ; it is, however, clear in statement and well balanced in pro-

portion. The practical directions are adequate, though exception

must be taken to the recommendation for mounting moths and

butterflies in glass-covered tablets; a sealed mount for any object

preserved for study — and specimens not for study should not be

collected — is undesirable.

A sbort list of books for reference, a glossary, and an index are

also given. s

Throughout the book errors of statement — such as, ‘the tus-

socks are very often included, by competent authorities it is to be

presumed, in the Noctuidz,’ and that there is but one brood of the

white-marked tussock each year — are infrequent.

The illustrations are more numerous than either the title or the

list indicates, and they show to what good advantage the camera

may be applied in nature study. S.H

Sesiidz. — Mr. William Beutenmiiller’s long-expected monograph

of the Sesiidz of America north of Mexico appears in sumptuous

form as Part VI of Vol. I of the Memoirs of the American Museum

of Natural History. After a brief introduction there are sections on

the position of the family, its characters, characters of the genera,

Synopsis of genera, historic review of generic names, habits of imago,

mimicry of imago, characters of the larva, synopsis of larva, habits

of larve, synopsis of food habits of larva, characters of pups.

descriptions of genera and species with synopses of the species,

and a bibliography.

Mr. Beutenmüller recognizes seventeen genera, 100 species, and

ten varieties ; one new species and one new variety are described.

Of the eight plates, five are devoted to the imagoes and three to

larval borings. The figures of the imagoes, though inconveniently

arranged on the plates, are admirable ; only five species and three

varieties are unfigured. Text-figures. showing structural details are

also given.

A bibliography of more than thirty-five pages and with 542 titles

Would seem adequate, but is not sufficient to include all the works

quoted in the body of the paper. siis

The date, March, 19o1, on the cover is entirely unjustifiable ;

Copies were not received until June, and were certainly not 1s$

earlier than the end of May. S. H.

784 THE AMERICAN NATURALIST. (Vor. XXXV.

Mosquitoes. — Dr. Howard's well-known interest in and practical

studies of mosquitoes take form in a volume! that will be of great

value to naturalists, physicians, and municipalities. After a brief

introduction he treats, in turn, with mosquitoes in general, malaria

and mosquitoes, the common mosquitoes of the genus Culex, the

malarial mosquitoes of the genus Anopheles, mosquitoes and yellow

fever, mosquitoes and filiariasis, other genera of North American

mosquitoes, natural enemies of mosquitoes, remedies against mos-

quitoes, how to collect and preserve mosquitoes, and the. classifica-

tion of the United States mosquitoes.

Dr. Howard's style is clear and direct, though a little inclined to

redundancy and to repetition. An occasional lack of precision of

expression and of accuracy may be noted ; mosquitoes, for instance,

being held to include species that are not Culicidz, and again to

exclude forms always comprised within that family. Chapter VI,

dealing with genera of North American mosquitoes other than

Culex and Anopheles, mentions six *genera known to occur in the

United States," but considers and refers to seven!

'The illustrations are excellent; a few are original, but the greater

number from the facile pencil of Miss Sullivan appeared in the

Author's Notes in 1900. Figure 42 does not represent Aédes fuscus,

and in the Notes was attributed to 4. sapphirinus.

S TIE

BOTANY.

Bergen's Foundations of Botany? is virtually a much-improved

and enlarged edition of the author's Elements of Botany, which has

been one of the most successful of recent elementary text-books.

The author's intimate knowledge of the needs and limitations of

high schools, gained by long experience as a teacher, is shown even

more effectively in the present volume than in its predecessor, for the

publishers have done their part much better than before, improving

the illustrations as well as adding largely to their number and in

other ways doing fuller justice to the plan of the book.

! Howard, L. O. Mosquitoes. How they live; how they carry disease p

they are classified ; how they may be destroyed. New York, McClure, Phillips

& Co., 1901. xv + 241 pp. pl., 50 figs. :

? Bergen, Joseph Y., Instructor in Biology, English High School Bostol™

Foundations of Botany. Boston, Ginn & Company, 1901. 12mo. ii + 412 +

257 pp. 12 pls., 306 figs.

No.417.] REVIEWS OF RECENT LITERATURE. 785

A first part deals with the structure, functions, and classification

of plants. Beginning with the seed and its germination, this part

treats comparatively of the various organs of seed plants as regards

their gross anatomy, histology, and physiology, then considers various

types of cryptogams, and finally outlines the evolutionary history of

the vegetable kingdom. Following this comes a part devoted to

ecology, which ends with a brief discussion of the causes of organic

evolution. "The last third of the book consists of a key and flora in

which are described nearly seven hundred species of plants available

for study in the springtime. A Handbook for the Use of Teachers is

published separately. This includes. sixty-two pages of practical

suggestions and references supplementary to the numerous direc-

tions for observations, experiments, and extra reading which are

given in the text-book itself.

The book abounds in fresh, interesting facts illustrating the prin-

ciples discussed. These, too, are presented in a way to be intel-

ligible and significant to young people. It will lead them to many

intellectual delights.

Comparatively few errors have been noticed, and these are for the

most part such as will be readily detected by teachers who use the

book. They may be misled, however, by the unqualified adoption

of the calyx-adnate theory of inferior ovaries given in Chapters XV

and XVIII. To-say the least, this complicated view of the morphology

of many flowers and fruits is open to serious question, The modern

view which regards an inferior ovary simply as one imbedded in the

torus accords much better with what is known of the development

of flowers, and avoids much perplexity to the student of flower and

fruit morphology. FREDERICK LERoy SARGENT.

Notes. — Professor Jepson’s Flora of Western Middle California

(Berkeley, Cal., 1901, iv + 625 pp.), which covers a region already

locally handled, is really an expression of the author’s own success

in a field recognition of species, and, as he is conscientious and has

drawn up his descriptions from fresh material instead of compiling

them or basing them on herbarium fragments, the book should be

helpful

Dennert’s Plant-life and Structure, translated from the German

by Clara L. Skeat, forms one of the “ Temple Primers,” of which

The Macmillan Company are the American publishers. It is easier

to say why some books see the light than others. 'This little book

belongs in the second category.

786 THE AMERICAN NATURALIST. . [Vor. XXXV.

Dr. Robinson’s latest “Contribution from the Gray Herbarium of

Harvard University,” constituting No. 26 of the current volume

of Proceedings of the American Academy of Arts and Sciences, includes

synopses of Melampodium and Nocca (or Lagascea), and a number

of additions to the Mexican flora, chiefly in the genus Eupatorium.

An enumeration of the plants known to grow on Mt. Rainier, by

Piper, is published in Mazama for April. Flowering plants seem to

disappear above 10,000 feet, at which elevation Smelowskia ovalis

still occurs, and the timber line is at 6500 feet.

A considerable part of Agora for June is given to articles on

Mt. Katahdin and its botany.

Fascicle 2 of the current volume of the Mémoires de [Institut

Egyptien is a contribution to the flora of Egypt, by E. Sicken-

berger. Though the fascicle bears date 1901, the preface is dated

January 31, 1895.

Mr. J. M. Macouns’s “ List of the Plants of the Pribilof Islands,”

with notes on their distribution, has been reprinted from Part IH

of Jordan’s Fur Seals and Fur-seal Islands of the North Pacific Ocean.

A paper on new spermatophytes from Mexico and Central

America, by M. L. Fernald, constitutes No. 27 of the current volume

of Proceedings of the American Academy of Arts and Sciences.

A thirteenth selection of new plants from Wyoming, by Aven

Nelson, is published in the Buletin of the Torrey Botanical Club

for April. .

The third of a numbered series of papers which Professor Sargent

is publishing in the Botanical Gazette under the title “ New or

Little-known North American Trees,” in the April number of that

journal, adds a considerable number of species of Crategus for the

United States, and a new Alaskan Betula, and raises the Californian

Cupressus Goveniana pygmea to specific rank.

The forms commonly referred to Ribes rubrum have been passed

in review by Hedlund in recent numbers of Botaniska Notiser.

The native plums form the subject of Bulletin No. 87 of the

Wisconsin Experiment Station, by Professor Goff.

A revision of thirty-five western and northern Antennarias of the

plantaginifolia set, by Elias Nelson, has recently been separately

distributed from the Proceedings of the United States National

Museum.

NUBE e =

No. 417.] REVIEWS OF RECENT LITERATURE. 787

Anatomical studies of Primulaceæ and Simarubez are contained

in recent numbers of the Annales des Sciences Naturelles.

The Gardener's Chronicle for June 8 contains an illustrated article

on the species of Platanus usually planted as street trees, etc.

The morphology and anatomy of Aloinez is the subject of an

illustrated paper by Hausen in the Verhandlungen des Botanischen

Vereins der Provinz Brandenburg for 1900.

The grasses of Iowa are considered. at length as to their biology,

chemical composition, etc., by Pammel, Weems, and Lamson-

Scribner, in Buletin No. 54 of the Iowa Agricultural College.

Professor Lamson-Scribner gives an account of some Arizona

grasses — several of them described as new — in Circular No. 32 of

the Division of Agrostology of the United States Department of

Agriculture.

A revised edition of Buletin No. 17 of the Division of Agrostology

of the United States Department of Agriculture (Professor Lamson-

Scribner’s * American Grasses, II ") has been issued from the

Government press.

Mr. C. L. Shear publishes some notes on Fournier's Mexican

species and varieties of Bromus in the Bulletin of the Torrey Botani-

cal Club for April.

The coralloid rootlets of Cycas revoluta and their symbiosis are

considered by A. C. Life in the Botanical Gazette for April.

Professor Shimek contributes a paper on the pteridop elvis 2

Iowa to the current volume of the Bulletin of the laboratories of

natural history of the State University of Iowa.

The eighty-two Alaskan Hepaticze noted in Professor Evans's paper

on the representatives of this group collected by the Harriman Expe-

dition are increased by the addition of three others collected by

Professor Setchell, in a list of species published in Nos. 6, 7 of Zoe.

A third series of preliminary diagnoses of new species of Laboul-

beniaceze, by Dr. Thaxter, is issued as No. 23 of the current volume

of Proceedings of the American Academy of Arts and Sciences.

A convenient pocket host index for the commoner European para-

sitic fungi, by Dr. Lindau, is published by Gebrüder Borntraeger, of

Berlin,

788 THE AMERICAN NATURALIST.

A revision of forty species of Hydnum, by H. J. Banker, appears

in the April number of the Bulletin of the Torrey Botanical Club.

A report on the agricultural resources and capabilities of Hawaii,

by Professor Stubbs, which constitutes Bulletin No. 95 of the Office

of Experiment Stations of the United States Department of Agri-

culture, contains reproductions of a number of excellent photographs

of tropical plants and fruits.

The cultivation and manufacture of tea in British India and

Ceylon is the subject of No. 2 of the current volume of Beihefte zum

Tropenpflanzer, published in Berlin.

An illustrated article on breeding new wheats, at the Minnesota

Experiment Station, is a prominent feature of Zhe World’s Work

for May.

The manufacture of sago is illustrated in Der Tropenpflanzer for

May.

“The Stock-poisoning Plants of Montana” is the title of an exten-

sive illustrated paper by Chesnut and Wilcox, published as Bulletin

No. 26 of the Division of Botany of the United States Department

of Agriculture.

The third volume of Meddelanden fraan Stockholms högskolas bota-

niska institut —an assembled series of papers by Professor Lagerheim

and his associates — is especially interesting because of its myco-

logical contents.

An illustrated catalogue of the plants of the Alpine garden “La

Linnza," of Bourg-St.-Pierre, has recently been issued by the direc-

tor, M. Correvon, of Geneva.

An account of the botanic garden of the Czernowitz University,

planted in 1877, is contained in the Festschrift of the recently

celebrated quarter-centennial of the University.

A short illustrated article on the botanic gardens of Malta is

published in the Gardener’s Chronicle for May 4.

The Youth’s Companion, than which no more potent medium could

be desired, has undertaken to aid in creating a national sentiment

which shall eventually result in the universal beautifying of the

grounds of the rural schools of the United States.

CORRESPONDENCE.

Editor of the American Naturalist :

Sir, — In the course of recent bibliographical studies the writer

has discovered a number of cases in which the names given to

recently described American hydro oids have been given to other

species by previous writers. For t

ally responsible, and takes the first opportunity to correct the errors.

The other cases will be discussed la er.

Halecium geniculatum Nutting. Hydroids from Alaska and Puget

Sound. Proceedings of the — States National Museum, Vol.

XXI, p. 744.

This name was used. by Norman in a paper published in 1866,

entitled “ On the Hydrozoa (etc.) of the Hebrides,” p. 1

or this species the following name is now proposed: Halecium

washingtoni Nutting.

Halecium robustum Nutting. Papers from the Harriman Alaska

Expedition, XXI. The Hydroids, 1901, p. 182.

Allman gave this name to a species from gore Bay. Chal-

lenger Report. The Hydroida, Part II, 1888, p. 1

For this species I propose the following name : Kalohi harri-

mani Nutting.

Thuiaria elegans Nutting. Papers from the Harriman Alaska

Expedition, XXI. The Hydroids, 1901, p. 187.

Name preoccupied by Kirchenpauer. T ocdindie Gattungun und

Arten von Sertulariden, 1884, p. 21

This species will now be known as Thuiaria kincaidi Nutting.

Named in honor of Mr. Trevor Kincaid, one of the members of the

Harriman Expedition.

Judging from the numerous cases that have "n come to my

attention of the use of preoccupied names in zoólogy, I am more

and more of the opinion that it is unsafe to give descriptive specific

names to forms belonging to long-established genera. It is safe to

Say that no one investigator has at his command all of the papers

relating to any group of considerable size. The names which occur

to a writer as particularly apt in a descriptive way are the very ones

that have been most likely to appeal to previous writers.

C. C. NUTTING.

UNIVERSITY oF Iowa, July 13, 1901.

(Wo. 416 was mailed August 22.)

789

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Ward’s Exercises in Meteorology . . . . . ae

Davis’ Elementary Meteorology - - ° ° ° Se

Williams’ Elements of Chemistry . AE TEL e

Hastings and Beachi General Pages o 0 5 c T

Wentworth and Hill's Physics - S ed , Ue ME

Gage's Principles of Physics * acu n. * eus n

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VoL. XXXV, No. 418 J

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

I. Se 10i i ae ee 5e ^ E

(Conclusion) ; Professor W. M. WHEELER

IL Notes on deg Nautilus eer . Professor BASHFORD DEAN 819

HI. Baseleveling and its Faunal Signifoanee — e f eL

from Southeastern United States . CHARLES C. ADAMS z

IV. Notes and Literature: Anthropology. The Seri. Indians—

Zoólogy, Schmeil's Zoólogy, Herrick's 2 of Wild Birds, '

The American Naturalist.

ASSOCIATE EDITORS:

J. A. ALLEN, Pu.D., American Museum of -— = = York.

E. A. ANDREWS, Pu.D., Johns Hopkins University, Baltim

WILLIAM S. BAYLEY, Pu.D., Colby Dann, Water.

CHARLES E. BEECHER, Pu.D., Yale University, New Haven.

WILLIAM M. DAVIS, M.E., Harvard University, Coie

ALES Se. M.D., New York Ci

D. S. JORDAN, LL.D., Stanford Univ oak

CHARLES a KOFOID, PH.D., University of e Urbana.

W. E. RITTER, Pu.D., University of California, Berkeley.

FRANK RUSSELL, PH.D., Harvard University, Cambridge.

ISRAEL C. RUSSELL, LL.D., University of Michigan, Ann Arbor.

ERWIN F. SMITH, S. D, U. S. Department of Agriculture, pun cua

LEONHARD STEJNEGER, Smithsonian Institution, Washington

W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, PH.D., University of Nebraska, Lincoln.

WILLIAM M. WHEELER, PH.D., University of Texas, Austin.

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Vor. XXXV. October, 190r. No. 418.

T : THE COMPOUND AND MIXED NESTS OF

E AMERICAN ANTS.

WILLIAM MORTON WHEELER.

Part III. SymMBIOGENESIS AND PSYCHOGENESIS.

“Eine Psychologie in Spencer-Darwin'schem Sinne auf Entwicklungslehre

gegründet, aber auf positiver Detailforschung fussend, verspricht reichere Resul-

tate als alle bisherigen Speculationen." — E. MACH.

ALL writers on the behavior of ants have been deeply

impressed with the cases of social symbiosis, more especially

with those of an extreme type like Polyergus and Anergates.

The deadly animosity of the members of a formicary, not only

towards ants of another species but even towards ants of the

. Same species belonging to different colonies, is so striking and

...80 nearly universal that an extraordinary explanation seems to

. be demanded to account for the cases of amicable consociation

_ of two species. In the presence of such phenomena, instinct

and consentience, or sense-experience, have been abandoned

as inadequate, and the existence in ants of some higher form

of intelligence, like understanding and ratiocination, has been

. postulated without further ado. At the same time the evo-

lutionist has been stimulated to broach a phylogeny of social

791

792 THE AMERICAN NATURALIST. [Vor. XXXV.

symbiosis. Both the stress laid on the psychical manifesta-

tions of ants and the attempts at establishing a phylogeny

of the compound and mixed nests have been clearly appre-

hended and set forth in considerable detail by Wasmann (91).

While my own observations lead me to agree with this able

investigator in many respects, I must, nevertheless, dissent

from his attitude towards the genetic method as applied to

the study of the compound and mixed nests. It is necessary,

therefore, to attempt a critical revision of this matter so far

as this is possible within the limits of the present paper. I

shall deal first with phylogeny as applied to the cases of social

symbiosis and conclude with a very brief consideration of some

of the pertinent psychical problems.

Darwin was the first to attempt an explanation of the origin

of dulosis in the European ants. In a well-known passage in

the “ Origin " ('61, p. 244) he says: “ By what steps the instinct

of Formica sanguinea originated I will not pretend to conjec-

ture. But as ants, which are not slave-makers, will, as I have

seen, carry off pupz of other species, if scattered near their

nests, it is possible that such pupz originally stored as food

might become developed; and the foreign ants thus uninten-

tionally reared would then follow their proper instincts, and do

what work they could. If their presence proved useful to the

species which had seized them — if it were more advantageous

to this species to capture workers than to procreate them —

the habit of collecting pupz originally for food might by nat-

ural selection be strengthened and rendered permanent for the

very different purpose of raising slaves. When the instinct

was once acquired, if carried out to a much less extent even

than in our British F. sanguinea, which, as we have seen, is

less aided by its slaves than the same species in Switzerland,

natural selection might increase and modify the instinct —

always supposing each modification to be of use to the species

— until an ant was formed as abjectly dependent on its slaves

as is the Formica rufescens.”

Apart from the statement that the English and Swiss sa”

guinea differ in their behavior —a statement ‘which has been

since disproved — Darwin’s views have been accepted by Forel

No. 418.] NESTS OF AMERICAN ANTS. 793

(74, p. 440), who has also called attention to the fact that the

frequent occurrence of pupae without cocoons in the nests of

F. fusca would add to the plausibility of Darwin's hypothesis,

for such free pupze would be able to hatch without the assist-

ance of the enslaving species.!

Pursuing Darwin's line of thought, Forel (74, p. 443)

called attention to the following series, which seems to have

been previously in great part suggested by von Hagens (67):

“1, Working ants pure and simple; 2. ants dwelling in

abnormal mixed colonies; 3. F. sanguinea (sometimes with-

out slaves); 4. Polyergus rufescens (here the working instinct,

which is merely diminished in F. sanguinea, disappears com-

pletely and the slave-making instinct attains its apogee) ;

5. Strongylognathus huberi (the slave-making instinct is cer-

tainly still alive); 6. S. ¢estaceus (the slave-making instinct

no longer exists except in the form of derisory vestiges, the

worker is on the road to atrophy and tends to disappear) ;

7. Anergates atratulus (the worker has disappeared ; only para-

sitism is admissible). This last ant, it seems to me, is a

remarkable example of reversion to ancestral traits (incom-

plete societies, without workers) through parasitism ; its gene-

alogy is explicable through S. ¢estaceus, the workers of which

have become so rare in comparison with the females and

males." Essentially the same series of cases was adopted by

Lubbock (94, pp. 88, 89). At the present time it could be

still further perfected, as Wasmann suggests, by the insertion

of Tomognathus sublevis between F. sanguinea and Polyergus.

The views initiated by Darwin have not been allowed to

pass unchallenged. The first to take up the cudgels was

McCook in the concluding paragraphs of his paper on Polyer-

gus lucidus (80, pp. 383, 384). After presenting Darwin's

views he writes: «Whatever credit we may give to this

ingenious hypothesis, it must be said that in the case of our

F. schaufussi, natural selection has not operated to degen-

erate the soldierly courage and faculty, and remand the duty

of defense to those associates in whom the military faculty

1 The American slaves of Z. sanguinea and Polyergus, viz., F. fusca, Vats., sub-

sericea and subenescens, and F. nitidiventris also often have free pup®.

794 THE AMERICAN NATURALIST. (Vor. XXXV.

has been specialized. In other words, if Lucidus has become

specialized as a warrior, dropping an original disposition and

ability to labor, her slave has not become specialized as a

worker, nor dropped her combative faculty, but seems to be

possessed in all respects of the riormal habits and nature of

ants of her species. At least I could trace in her no effects

of slavery, other than the strange association with and care of

her abductor. One, therefore, who accepts Dr. Darwin's sug-

gestion, must allow that natural selection has wrought toward

specialization in one section of the colony, but has been sus-

pended in its operations upon the other section. It is doubt-

ful if the anomalous conditions thus raised by Dr. Darwin's

explanation be not more difficult to explain than the original

conditions to which the hypothesis was applied."

Assuredly, if all the arguments against natural selection were

as easily refuted as the one here produced by McCook, Dar-

win's followers would have occasion for great rejoicing. The

refutation is equally easy whether we consider the colonies of

the auxiliary species with which Polyergus wars, or only the

larvae and pupz which it abducts from these colonies and rears

as its slaves. So far as the latter are concerned, McCook has

completely overlooked the very obvious fact that Polyergus

rears only the workers of the auxiliary species, and these have

never been known to reproduce in the mixed nests. But even

if we suppose, for reasons to be given below, that these workers

may often reproduce normally, the difficulty is not removed,

since they would not .only have to be able to transmit charac-

ters acquired in their imaginal stage, — and the possibility of this

has by no means been demonstrated, — but they would have to

transmit these characters to male or female offspring if there

was to be any permanent modification of the species. Now

this is impossible, because Polyergus will not permit the winged

sexes of the auxiliary species to reach maturity. Hence the

detached auxiliaries cannot impart any modifications to the

species to which they belong, no: matter what peculiarities

they may take on in the Polyergus nests.

Though the fallacy in McCook's argument is very obvious,

Wasmann still contends that it involves “ einen vorzüglichen

No. 418.] NESTS OF AMERICAN ANTS. 795

Gedanken," which he puts in the form of the following ques-

tion: “If natural selection can bring about such a peculiar

development of the instincts in the dominant species, why has

it not exercised a corresponding influence over the enslaved

species?" Wasmann is evidently bent on rescuing that por-

tion of the argument not expressly stated by McCook, viz.,

the necessity of a change of character in the ants of the

colonies attacked by Polyergus. But closer inspection shows

this attempt to be unsuccessful, for he has not considered the

question of the relative abundance of the dominant and aux-

iliary species. That this is a matter of some moment in a

discussion of this kind is seen from a general survey of para-

sitic and predaceous organisms. These must of necessity live

on that margin of surplus vitality and reproductivity so char-

acteristic of all animals and plants; for it is obvious that

organisms which depend very decidedly for their sustenance

on special hosts or prey must endanger their own existence

to the extent that they endanger the existence of the species

on which they depend. The serious injury or death of the

host species implies the death of the parasitic species in all

cases where the relations between the two are of a highly

specialized character. This argument could be adduced if the

Polyergus were more abundant or quite as abundant as the

auxiliary species. But it is quite unnecessary to make use of

it, because Polyergus is a rare ant of local occurrence, and the

various forms of F. fusca and F. pallide-fulva which it enslaves

are widely distributed and vastly more abundant both in col-

onies and individuals. In fact, no other insects are as com-

mon in our Northern States as the varieties of these two ants,

and even in Texas, near the southernmost limits of the dis-

tribution of F. fusca, a form very closely related to F. sub-

sericea, viz., F. gnava Buckley, is the most prolific of ants. Its

nests often contain upwards of thirty fertile queens, and the

number of eggs, larvae, and pupa which are reared between

the end of February and the first of June is enormous. Why,

then, should these very prolific and widely distributed species

exhibit a special development of valor or a particular defensive

form of nest architecture in adaptation to a rare predaceous

796 THE AMERICAN NATURALIST. [VoL. XXXV.

ant of sporadic occurrence? As well might we expect the

human dermis and its appendages to present hereditary modi-

fications in adaptation to occasional parasites like Pediculus

capitis. The advantages that would result from the develop-

ment of a courageous disposition and specially protective forms

of nest architecture in F. fusca and pallide-fulva are more

than outweighed by those derived from their unusual powers

of reproduction. These species run to offspring, not to valor.

That the survival and even the predominance of species does

not necessarily depend on the development of moral and

psychic endowment is demonstrated on a grand scale in the

vegetable kingdom.

This defense of McCook's argument is, ipsc only a

small portion of Wasmann's criticism of the position held by

Darwin, Forel, and Lubbock in regard to the phylogeny of the

mixed colonies ('91, pp. 214-254). Wasmann sees in these same

facts irrefutable arguments against the theory of indeterminate

variation and natural selection, and arguments equally strong

in favor of tracing the modicum of physical and psychical

development which is acceptable to him to *'innere gesetz-

mássig wirkende Entwicklungsursachen." In my opinion,

he has succeeded — if indeed he has really succeeded —

only in showing that the genetic method has been somewhat

awkwardly applied to the cases of compound and mixed nests.

On the whole, I believe that he has neither invalidated the

principle of natural selection, nor made it perfectly clear that

we must forthwith deliver ourselves up to anything so impal-

pable as “innere gesetzmüssig wirkende Entwicklungsursa-

chen." In support of these statements the following remarks

on some of Wasmann's arguments are offered.

Wasmann seems to regard it as an established fact that

worker ants do not reproduce, or do so only under unusual or

even pathological conditions. He is also inclined to emphasize

the differences between the instincts of the queens and those of

the workers. Hence the workers are debarred from transmit-

ting their peculiar characters, either congenital or acquired,

and the instinct modifications, so characteristic of different

species of ants, must be explained as arising from determinate,

No. 418.] NESTS OF AMERICAN ANTS. 797

internally regulated variations. The premises to these conclu-

sions I cannot accept, since they do not appear to me to be

indefectibly established. They are based on a rather limited

study of a few species of highly specialized European ants, and

cannot, therefore, lay claim to great generality. My own

observations, still incomplete, to be sure, on several Texan

ants representing both the most primitive and the most special-

ized subfamilies (Ponerinze and Camponotinz) convince me

that worker ants not only very frequently lay eggs in consider-

able numbers, but that these produce perfectly normal offspring.

When workers are properly fed in the artificial nests, they

seem to have no more desire to devour their own eggs than

to devour those which are deposited in their keeping by the

queens.. During February last a carefully isolated lot of

workers and soldiers of a handsome Camponotus (C. texanus

n.sp.)laid dozens of eggs in my Fielde nests. By the first

week in June many of the larvae were mature, and a few of

these had spun their cocoons before I was compelled to leave

my laboratory for the summer. These cocoons were found to

contain perfectly normal male pupz, thus adding fresh evidence

for the generally accepted belief that the parthenogenetic off-

spring of worker ants are males. Similar observations were

made on workers of Camponotus marginatus n. Var. and Pachy-

condyla, except that in these cases I did not follow the larva

quite to the pupal stage. I am, moreover, convinced that

numerous eggs are laid (probably by the soldiers) and reared

in the frequently queenless nests of a gall-inhabiting Colo-

bopsis (C. etiolata n. sp.) from Texas. In fact, for aught we

know to the contrary, every well-developed ant colony may

contain one or more fertile workers. Where the worker caste

is dimorphic the soldiers probably have the greatest tendency

to lay eggs. Judging by analogy with other Hymenoptera,

like Polistes among the wasps, it is also probable that the older

and more vigorous the ant colony, the greater the tendency for

workers to take on the reproductive powers of the gecm

That these conditions clearly imply the possibility of the inher-

itance of worker characters through the male offspring goes

Without saying. The comparatively frequent development of

798 THE AMERICAN NATURALIST. [Vor. XXXV.

the reproductive power in worker ants makes it possible to

account for the conditions presented by Leptogenys and

Tomognathus. In these genera worker forms (ergatoids) have

usurped the functions of the winged queens, which have com-

pletely disappeared.!

The statements of Wasmann and other authors concern-

ing the differences between the instincts of the queens and

workers seem to me to require some qualification. This differ-

ence is rather quantitative than qualitative, for the recently

fertilized queen, even in highly specialized ants, during the

establishment of her colony displays nearly all the worker

instincts, even to excavating and defending the nest and caring

for the first brood of young. In some species (Ponerinz ?) she

may even exhibit the foraging instinct so characteristic of the

workers, for aught we know to the contrary. It is true that

in the more highly specialized ants like Formica, these instincts

lapse into desuetude as soon as the workers make their appear-

ance in the nest, but it is equally true that they may be retained

throughout life as in the queens of the Ponerine, Leptothorax,

and probably also many other ants. While I do not wish to

lay unwonted stress on these fragmentary observations and

reflections, they are, nevertheless, quite sufficient to bid us

hesitate in the use of arguments which start from the assump-

tion that the worker ants reproduce only under pathological

conditions and present instincts essentially different from those

of the queens. :

Wasmann encounters the gravest difficulties in the genetic

explanation of dulosis. His remarks are mainly confined to

the two well-known cases, Formica sanguinea and Polyergus,

the former, in the opinion of most writers, an incipiently, the

latter a perfectly, dulotic species. He attempts to show that

dulosis could not have arisen in sanguinea by selection, since

flourishing or medium-sized colonies of this species could have

derived no advantage from the possession of a small number of

slaves, while the advantage that would accrue to a small colony

1 Silvestri ('O1), in a paper received while these paragraphs are going through

the press, expresses some very similar views concerning the fecundity of worker

Termites.

No. 418.] NESTS OF AMERICAN ANTS. 799

would be more than outweighed by the difficulty such a colony

would experience in obtaining slaves! And even if the dulotic

habit had manifested itself repeatedly as a chance variation,

and had proved useful, there would still be lacking the heredi-

tary basis for the instinct, since this is exhibited only by the

workers. Wasmann states the difficulty in the following words

(91 p. 236): ‘ Before the inclination to rear slaves had proved

itself permanently beneficial during many generations, it could

not have been established by natural selection as an hereditary

Anlage; but the possibility of inheriting the Anlage must

exist before the incipient inclination could be transmitted from

one colony to another — ergo natural selection lacks the very

point of departure for the development of an hereditary slave-

making instinct from the accidental forms of mixed colonies.

We must leave natural selection like Baron von Münchhausen

to drag itself out of the morass by its own hair."

Surely natural selection, however numerous its shortcom-

ings, deserves better treatment at our hands. A careful

perusal, however, of the above-quoted passage from the

“Origin” and a consideration of the facts brought to light

since its publication leave little cause for anxiety. Wasmann

assumes that sanguinea robs the pupæ of other ants for the sake

of rearing slaves. This is scarcely borne out by the facts.

The young of the auxiliary species are sought for and appro-

priated to serve as food, in obedience to an ancient and wide-

spread formicid instinct that emerges to view very clearly in

many often distantly related species of ants. Thus Adlerz

has shown (96a) that on exceptional occasions even Lasius

niger robs the larve and pupz of Z. flavus, and these may

hatch and function as slaves in the nest of the dark-colored

species. Wasmann, too (99a), has observed a colony of L fuli-

ginosus appropriating the larvæ and pupæ of a neighboring

colony of Myrmica levinodis. I have repeatedly observed the

same instinct in Mexican and Texan Ecitons ('01; E. crassi-

corne, schmitti, pilosum)? It is also probable that many cases

pæ in their nests for some

ut even if some of them

800 THE AMERICAN NATURALIST. (VoL. XXXV.

of synclerobiosis (especially those of /. exsectoides with sub-

sericea) are due to a sporadic outcropping of this ancient

instinct. In view of the further facts that sanguinea can get

on perfectly without auxiliaries, that it is the young colonies

which usually contain the greatest number of slaves, and that

the number of these is often highly variable in different colo-

nies even in the same localities, we are certainly justified in

demanding more stringent proof that sanguinea really robs for

the sake of rearing slaves? Laying most stress on the fact that

the booty serves as food, — and of this Forel's observations

contain sufficient evidence (74, p. 258)? — we may regard the

imaginal auxiliaries in the sanguinea nests as a mere by-product,

as it were, of the colonial activities. The sanguinea often

appropriate more food than they can devour, and the residuum

merely adds workers to the colony, which are not harmful and

may even be advantageous. This is evidently the interpreta-

tion intended by Darwin, who does not pretend to invoke the

principle of natural selection in his genetic explanation of

the sanguinea stage of dulosis, so that Wasmann has taken

unnecessary pains to refute an imaginary argument. With

should hatch, the nomadic habits of the Ecitons and their poorly developed

deportation instinct would prevent the formation of permanent mixed colonies

since the larvae and pupae which they kidnap belong to home-loving spect

1 At Colebrook, Conn., I recently found three cases of synclerobiosis in 1 addi-

tion to those enumerated in the second part of this paper: (1) acolony of F. exsec-

toides with F. subsericea, similar to the mixed colonies of these species observ:

by Forel and Schmitt; (2) F. nitidiventris with F. rufa var. obscuripes Forel ;

(3) Z. nitidiventris with F. rufa var. difficilis Emery. All of these colonies were

small, and in none of ihn could I find the queens of either of the consociating

species.

? Wasmann believes (91, p. 198) that in sanguinea the perception of the

cases in which I am inclined to believe that Wasmann has ae overestima

the power of association in ants. His statement, however, as he bim

which I gave a fabulous number of Z. pratensis cocoons during the course

summer failed to rear a single one. The same was true of several other formi-

caries to which I gave fewer cocoons."

HTC cy cay

No. 418.] NESTS OF AMERICAN ANTS. 80I

F. sanguinea, however, there is already given the generalized

condition required as a starting point for the action of natural

selection and the development under its guidance of cases like

Polyergus, as Darwin suggests. In Polyergus the predatory

instincts have been developed to a highly specialized condition,

while the domestic instincts have retrograded pari passu as a

natural result of the survival of the prey, till the presence of

slaves in the nest has become a conditio sine qud non of exist-

ence. This correlation of instincts has involved the corre-

sponding correlation of structure which we find so beautifully

exhibited in Polyergus.! In the predatory instincts every

slight variation in advance would be beneficial to the species,

while slight retrogressions would not under the circumstances

be disadvantageous. I cannot, therefore, agree with Wasmann

when he says ('91, p. 247) : “ Natural selection could only main-

tain and augment useful instinct variations: but the develop-

ment of slavery up to the Polyergus-, Strongylognathus-, and

Anergates-stage is beneficial neither to the masters nor the

slaves — ergo natural selection cannot have produced the

instincts of the slave-holding ants." As good an answer as I

can conceive to an argument of this nature is a reference to

the cases of extremely specialized parasitism like the Cestode

and Sacculina, both of which are connected by tolerably com-

plete series of intermediate forms with the more generalized,

non-parasitic members of their respective phyla.

The symbiotic sequence suggested by Forel and Lubbock

is objected to by Wasmann on fairly good grounds. It is by

no means clear that the development has passed successively

through the stages represented by these forms. Indeed, as

Wasmann shows, the problem of symbiogenesis is much more

complicated than it appeared when the above sequence was

Suggested. It now seems evident that several lines of devel-

opment have proceeded independently from cases of plesio-

biosis (and possibly also parabiosis), which constituted the

necessary initial stages of symbiogenesis. Thus it is probable

that cleptobiosis, xenobiosis, and dulosis represent at least three

"In accounting for this development of instincts and structures, it is, of course,

necessary to regard the whole mixed colony as a single evolutionary unit.

802 THE AMERICAN NATURALIST. [Vor. XXXV.

discrete lines of development, the two former starting from con-

ditions of plesiobiosis, the latter from the widespread instincts

of ants to prey on the offspring of other Formicide. Other

cases which obviously resemble true dulosis may have arisen

from xenobiosis. This appears to be true of cases like Zepto-

thorax emersoni and possibly also of the species of Strongy-

lognathus and Tomognathus. On the other hand, the cases of

colacobiosis may be conceived to have originated either from

xenobiotic conditions like that of Z. emersoni or from dulotic

conditions like that of Strongylognathus testaceus. I cannot

believe that Forel or Lubbock really intended their sequence

as anything more than a rather general attempt in concrete

language to account for the phylogenetic derivation of the

remarkable cases of social parasitism (Anergates) from the

simpler forms of mixed nests. It is therefore superfluous to

waste many words for the sake of showing that the ants of the

Forel-Lubbock series are not phylogenetically related. It is

not only easier to sketch the phylogeny of the compound and

mixed nests in bold outlines than to fill in the details, as Was-

mann somewhat reproachfully suggests, but this is the only

available method of procedure at the present time. Still even

the attempt at detailed speculation in this direction scarcely

merits our disapproval as it does Wasmann's, for free and

open speculation is necessary to the advancement of a scien-

tific subject, if only as furnishing the necessary incentives

and guides to the attainment of profounder insight. Mere

fact-culling is not and never can be science.

Another argument on which Wasmann lays some stress is

drawn from the supposed immutability of instinct! The

instincts of F. sanguinea and Polyergus are regarded as iden-

tical both in Europe and America, and these instincts must

therefore have remained unchanged for a very long period of

time (9L p. 249). “Huber’s amazons of 1804 fought and

! [t is unnecessary in this place to deal with the doctrine of the immutability

of instinct so brilliantly advocated by Fabre ('79—00). That it is quite unten-

able has been demonstrated by Dr. and Mrs. Peckham ('98), Whitman (99), an

others. It could, in fact, be demonstrated to be false from Fabre's own mag

nificent observations. The genus Leptothorax, considered below, furnishes addi-

tional evidence, if this were needed.

No. 418.] NESTS OF AMERICAN ANTS. 803

conquered exactly like Forel’s amazons of 1870, and proved

themselves to be quite as dependent on their slaves as their

modern descendants; and there can be no doubt, that if

Adam had studied and described the habits of the amazons,

his account would agree very accurately with Huber's and

Forel’s.”” Similarly, Wasmann stresses the long-existing

fixity of instinct in Formicoxenus and Leptothorax (p. 226).

No issue can be taken with him on this point — but what

phylogenist would not take it for granted? If structure

can remain stable during æons of geological time, certainly

instinct may also remain relatively unchanged. It is, how-

ever, equally true — and this point seems not to have been

considered by Wasmann — that structure may undergo little

change as compared with instinct. In support of this state-

ment I would include in this place a series of facts which

may have arrested the attention of the reader in the previ-

ous portions of this paper, vzz., the remarkable differences of

instinct exhibited by the species of the single genus Leptothorax.

Morphologically, this very large and widely distributed genus

has been justly styled * homogeneous" by Forel (74, p. 339),

since the numerous species are closely related to one another

and often separable only on rather trivial characters. Even

the subgenera :Dichothorax and Temnothorax are based on

relatively slight differences. In their habits, on the other

hand, the species of Leptothorax are singularly diverse.

Many of the forms have no tendency to consort with ants

of other species, but differ considerably in the stations which

they inhabit. Some prefer to live under stones, others in

moss, others under bark or in dead wood, and still others,

like one of the Texan species, in cynipid galls, or, like our

New England Z. longispinosus Rog. in worm-eaten hickory

nuts among the dead leaves under the trees. Many species,

however, have a pronounced penchant for entering into more

or less intimate symbiotic relations with other Formicidze, as

shown in the following conspectus : :

I. The European ZL. muscorum often lives in plesiobiosis

With Formica rufa (see pp. 519, 520).

2. A similar tendency is undoubtedly exhibited by our

804 THE AMERICAN NATURALIST. [VoL. XXXV.

American L. canadensis Provencher,! which I have had occa-

sion to observe since the second part of this paper was written.

June 21, I found at Cudahy, near Milwaukee, Wis., two nests

of this Leptothorax compounded with a large nest of Cremas-

togaster lineolata Say in an old oak stump. The galleries of

the Cremastogaster extended far into the dry rotten wood and

contained numerous worker larvae and pupa. The two Lep-

tothorax nests, which together scarcely contained more than

sixty to seventy ants, were about twenty inches apart. Each

was a simple chamber one-half to three-quarters of an inch in

diameter excavated in the thick bark, and each communicated

by means of a slender passage with the subcortical space,

which was used as a common runway by the workers of

both species. When the bark was stripped off, several of the

Leptothorax were seen running on the surface of the wood and

mingling with the Cremastogasters, which they closely resem-

bled in coloration and sculpture, though differing in size,

shape, and movement. The Leptothorax were very timid, and

when touched with the fingers or tweezers, at once curled up

and “feigned death,” after the manner of Myrmecina. One of

the nests contained several mature larvae and pseudonymphs.

These the workers and the single deálated queen were

hastily removing to a place of safety. Rain began to fall

soon after I had opened this interesting compound nest, and

I was obliged to continue my observations on some living

specimens of both the species hurriedly confined to a small

vial. When the two species met, as they often did in these

narrow quarters, the Cremastogaster stroked the Leptothorax

with its antenna. On such occasions the latter at once

crouched motionless and folded its antennz along the sides

of its head. Then sometimes the Cremastogaster would

stand over the little ant and lick its rugose head and thorax ;

at other times it would pass on without bestowing these atten-

tions on the Leptothorax, which at once sprang to its feet

and ran away. This performance was repeated so often dur-

ing the remainder of the day that there could. be no doubt

1 According to Emery (’94, p. 318) this ant is perhaps only a subspecies of

the European Z. acervorum.

No. 418.] NESTS OF AMERICAN ANTS. 805

concerning the friendly relations of the two species. By the

following morning the Leptothorax and most of the Cremas-

togasters were either dead or dying, having been suffocated

by the pungent exhalations of the latter species. Thus the

little I could observe of the relations of the two species

resembled those which I have recorded for L. emersoni and

Myrmica brevinodis — only reversed, the L. canadensis behaving

like the Myrmica, while the Cremastogaster behaved some-

what like Z. emersoni.

3. L. pergandei lives, probably as a guest, in the nests of

Monomorium minutum var. minimum (see p. 539).

4. The single colony of the Mexican Z. petiolatus which I

have seen was living in parabiosis with species of Cryptocerus

and Cremastogaster (see p. 527).

5. L. tuberum var. unifasciatus lives with the European

- Formicoxenus vavouxi, the relations between the species being,

perhaps, the same as those which obtain between Formica rufa

and Formicoxenus nitidulus (see p. 538).

6. L. muscorum, L. acervorum, and L. tuberum live as slaves

or auxiliaries with the European Tomognathus sublevis (see

PP. 70, 71).

7. L. curvispinosus probably performs the same rôle in the

nests of 7. americanus (see p. 715).

8. L. tuberum has been found associated with Strongy-

lognathus testaceus. Here, too, the Leptothorax probably acts .

as the slave of the dulotic species (see p. 710).

9. L. emersoni lives with Myrmica brevinodis as described

in the first part of this paper. The compound nest resembles

that of Z. canadensis with Cremastogaster and of Formicoxenus

nitidulus with Formica rufa, but the relations between the two

Species of ants are like those existing in mixed nests. In

one sense L. emersoni is the dominant species and the huge

Myrmicas are its auxiliaries, or slaves; in another sense the

806 THE AMERICAN NATURALIST. [VoL. XXXV.

Leptothorax is a guest or social parasite resembling Anergates,

though still retaining intact its own household and its domestic

instincts. Z. emersont may therefore be said to combine in

itself the instincts of ants belonging to several categories of

mixed and compound nests.

The range of habits clearly indicated in this brief survey of

our very fragmentary knowledge of Leptothorax species is

still further enlarged if we include the genera Tomognathus

and Formicoxenus, both of which are closely allied to Lepto-

thorax. In fact, Tomognathus is indistinguishable from Lep-

tothorax in the male sex and larval stages (Adlerz, '96). That

the three myrmicine genera under consideration must have had

a common origin is evident from their morphology. Neverthe-

less the habits of the various species are so diverse as to repre-

sent all the forms of social symbiosis except colacobiosis of the

extreme type found in Anergates. It is evident, furthermore,

that the ants of these genera must have originally possessed

certain traits which made it especially easy for them to enter

into symbiotic relations with other species of Formicide. I

believe that we may still recognize in many of the species of

Leptothorax several of these traits, such as the following:

1. The genus has a very wide geographical distribution, a

prerequisite to the establishment of such numerous and varied

relations with other ants.

2. The species are all of small size. This must undoubtedly

facilitate their association with other ants.

3. The colonies consist of a relatively small number of indi-

viduals. This, too, must greatly facilitate life as guests oF

parasites in the nests of other ants.

4. Most of the species are rather timid, or at any rate not

belligerent. They are, therefore, of a more adaptable tempera-

ment than many other ants even of the same size (eg. Tetra-

morium cespitum). Forel (74, pp. 339, 340) has shown that

L. tubero-affinis will rear pups of L. nylanderi and even of

Tetramorium cespitum and live on good terms with the

imagines when they hatch.

5. There is no very sharp differentiation in habits b

the queens and workers of Leptothorax. This, too, should

etween

inl

No. 418.] NESTS OF AMERICAN ANTS. 807

facilitate symbiosis. The queens, as I have shown in the case

of L. emersoni, may retain the excavating instinct and the

instincts which relate to the care of the larva. Wasmann

(91, p. 219, footnote) saw the queen of L. tuberum var. nigri-

ceps removing the larvae and pupz when the nest was disturbed,

and I have recorded above a similar observation on L. canaden-

sis. More recently I have seen both the winged and deálated

queens in a nest of Z. Jongispinosus carrying away the larve

quite as busily as the workers. It is as Forel says (74, p. 3 39):

“Les 9 vivent presque comme les $; elles sont seulement

moins aptes au travail."

6. The similarity in instinct between the queens and workers

of Leptothorax finds its physical expression in the frequent

occurrence of intermediate, or ergatogynous, forms. So-called

microgynic individuals, or winged queens no larger than the

workers, have been frequently observed by Forel (74, p. 341)

and Wasmann ('95, p. 618) in L. acervorum. Those observed

by the latter author also showed color transitions between the

normal queens and workers. Adlerz (86, p. 77) found micro-

gynic individuals in Z. acervorum, muscorum, and tuberum.

Still other ergatogynous forms, which may be called ergatoid

queens, are represented by the large ocelligerous workers of

L. emersoni described and figured in the first part of this

paper (pp. 434, 436). These individuals bear a striking and

suggestive resemblance to the ergatoid queens of Fi omognathus

sublevis described and figured by Alderz Wasmann (95

P. 619) also records the occurrence of what he calls ''erga-

togyne Mischformen," or individuals completely transitional

between the queens and workers, in colonies of L.acervorum

and Formicoxenus nitidulus.

It is possible to draw still further inferences from the hetero-

geneous instincts exhibited by the genus Leptothorax and its

allies. Viewed as a whole, these different symbiotic relations

` cannot be said to bear the ear-marks of internal developmental

Causes operating in a perfectly determinate manner. Indeed,

appearances are quite otherwise and seem rather to pomt 1

indeterminate variations which have been and are still in

process of being seized on and fixed by natural selection. 1t

808 THE AMERICAN NATURALIST. [Vor. XXXV.

must also be admitted that the same appearance is presented

by the whole complex of conditions in compound and mixed

nests, but the demonstration is more cogent when it can be

shown that we have relations as different as those of dominant

species (Z. emersont) and slaves (L. acervorum) not only in

the same genus but among closely allied forms. This fact also

suggests that the instincts of the same species may be so general-

ised as to enable it to function like man, either as a slave or

master, according to the circumstances.

Although these considerations may seem to lack precision

they certainly show that we cannot dogmatize on the inade-

quacy of natural selection from a study of a few highly special-

ized ants like FÆ sanguinea and Polyergus rufescens. The

complicated phylogeny of the mixed nests can only be estab-

lished after a patient study of genera like Leptothorax and

Tomognathus. Another even more neglected group of small

ants which promises to throw some light on this subject

comprises the species of Monomorium with the allied genus

Xenomyrmex. At present so very little is known concerning

the habits of these genera that it must suffice merely to call

attention to them in this connection.

It is necessary in conclusion to consider very briefly the psycho-

logical problems suggested by the phenomena of social symbio-

sis, since, as above stated, the cases of dulosis have led authors

to postulate unusual mental powers in ants. The accounts of

sane and critical workers like Forel have been distorted by the

* popularizer," till one almost believes that the ante-bellum

Southerner might have learned many things in the management

of his slaves from a conscientious observance of Proverbs vi. 6.

‘Wasmann in his numerous writings ('91, '97, '99, '99b, etc.)

has undoubtedly done much, at least in Germany, towards the

exposure of this pseudo-psychology and a more rational con-

ception of ant behavior. His long familiarity with these ARE

mals and their guests has given him a singularly lucid insight

into their activities. My own more limited observations On

our North American species lead me to agree with him so ud

as the facts are concerned and many of the inferences which

he has drawn from them. I am constrained to say, however,

No. 418.] NESTS OF AMERICAN ANTS. 809

that I cannot adopt either his psychological definitions or his

psychogenetic reservations.

Wasmann seems to me unduly to expand the conception of

instinct in one direction, while circumscribing it rather too

narrowly in another. It is true that he distinguishes instinct

sensu stricto, the equivalent of the term as employed by many

comparative psychologists to designate the purposeful, auto-

matic, or stereotyped hereditary activities which are performed

prior to all experience and without awareness of their object,

and instinct sensu lato, which embraces also the activities

depending on the sense-experience of the individual, and all

that this implies, — adaptation and choice, associative memory,

etc., — activities which have come to be very generally desig-

nated as “intelligent.” But he does not appear to regard

these differences as sufficiently important to merit sharp dis-

tinction. Indeed, he even attempts (99, p. 12 et seg.) to show

that the presence of an element of experience in the associative

process of an animal is not of sufficient moment to merit dis-

tinction from purely hereditary associations. This, I believe,

few psychologists will admit. The detection of such a differ-

ence, however difficult it may be in practice, is surely not

beyond the possibility of carefully devised experimentation and

induction. And theoretically the two kinds of activity should

certainly be distinguished and separately designated. Was-

mann traces the non-stereotyped activities depending on choice

to a “sinnliches Erkenntniss- und Begehrungsvermógen," which

he regards as being the distinguishing trait of instinct. Thus

he comes to include under instinct both the instinct and intel-

ligence of other authors. I believe with Emery (93, '98) and

Bethe (98) that Wasmann has overshot the mark and attempted

to include too much in his conception of instinct. I should

continue, therefore, to emphasize the difference between activi-

ties which are compelled by inherited mechanism and those

Which imply choice on the part of the individual organism.

For the latter the term “ intelligence ” has been so very gener-

ally used that it seems both hopeless and idle to try to restrict

it, as Wasmann so emphatically desires, to the ratiocinative

Process in its clearest manifestations.

810 THE AMERICAN NATURALIST. [VoL XXXV.

That instinct activities and activities implying choice should

not be included under the same name is also evident from the

difficulties which we experience when we attempt to show how

the former could pass over into the latter; although the

ingenious hypothesis of Spencer, James, Morgan, and Whit-

man ('99, p. 333 e? seg.) may indicate where we are to seek for

this transition, which these authors find in the progressive

complication and mutual interference of instincts. Such con-

ditions, it is claimed, must lead to a diminution in the automa-

ticity of instinct and the supervention of a state of hesitancy

and choice on the part of the organism.

While on the one hand, as above stated, Wasmann improperly

expands the conception of instinct by including in it also the

simpler manifestations of intelligence, he narrows it in another

direction when he attempts to distinguish rather too sharply

between reflex action and instinct. His criterion that reflex

action depends essentially on the function of subordinate

ganglia, whereas instinct depends on the activity of a brain,

or sensorium, can only be maintained if the conception of

instinct is restricted to the Metazoa and understood as includ-

ing intelligence (sensu auctorum). But with the rejection of

this definition of instinct we must also reject such a distinc-

tion between reflex action and instinct.

It may be said in this connection that the attempts of others

to distinguish between instinct and reflex action are almost

equally unsatisfactory. This is true, e.g., of the distinction

emphasized by Romanes, when he says (95, p. 12) : “ I endeavor

to draw as sharply as possible the line which zz theory should

be taken to separate instinctive from reflex action; and this

line, as I have already said, is constituted by the boundary of

non-mental or unconscious adjustment, with adjustment 1n

which there is concerned consciousness or mind." It is well

that Romanes has stamped his distinction as a theoretical one,

for its application in comparative psychology is obviously

impracticable, since it must fluctuate with our opinions COD

cerning the presence or absence of consciousness in different

animals. It is not at all certain that consciousness is present

in the cases of pure instinct; or, if present, it may exist as à

No. 418.] NESTS OF AMERICAN ANTS. SII

mere epiphenomenon as Morgan explains ('00, p. 208): “ An

organism — if such exists —in which all the activities are

throughout life purely automatic and purely instinctive, might

indeed be conscious, but its consciousness would be of no

practical value ; for all the activities being, ex hypothest, auto-

matic, there would be no conscious guidance or control.

Consciousness might be present as a spectator of the activi-

ties, but it would be a mere spectator without power of

guidance, since, in so far as guided by intelligence, activities

cease to be instinctive. It should be clearly grasped that, in

so far as an activity is guided by individual control towards

more complete accuracy, just so far does it cease to be instinc-

tive, as the word is here used, and become intelligent. And

when an instinct is, as so often is the case, modified and

adapted to meet new circumstances, the modification and

adaptation is no part of the instinct as such, but is due to

intelligent control.

“I repeat, then, that in instinct as such consciousness is an

epiphenomenon or adjunct. But this does not, of course,

imply that it is absent. Only in so far as consciousness

accompanies the performance of instinctive activities can

intelligence get a hold on them for the purpose of control and

guidance. The performance of automatic activities affords to

consciousness data, which form a foundation upon which the

psychical structure reared by intelligence is based."

Finally, the distinction noted by Spencer, Morgan, and others

that reflex action is “localized response involving a particular

Organ or a definite group of muscles initiated by a more or less

specialized external stimulus," whereas “ instinctive activity is

a response of the organism as a whole, involving the coópera-

tion of several organs and many groups of muscles," implies

only a difference in degree as Marshall (98, p. 100 £7 seq.) and

Loeb (00, p. 77) have pointed out; for when we extend our

view to simple as well as complex organisms, and forget for the

moment the staple experiments of neuro-muscular physiology,

we must agree with James when he says (90. p. 384) that the

"actions we call instinctive all conform to the general reflex

type," and with Marshall (98, p. 100) when he says: “All

812 THE AMERICAN NATURALIST. [Vor. XXXV.

instincts appear as modes of that simplest of all forms of

activity, the reaction of a living cell to the stimulus received

from its environment. And we are led to conclude with

Loeb ('00, p. 177) that “the discrimination between reflex and

instinctive actions is chiefly conventional.” The work of this

last author, above all others, is of great value in freeing us

from some of the traditional misconceptions of instinct. He

has succeeded in tracing a number of instincts to simple trop-

isms (or taxes) and has shown good reasons for maintaining

that many of the more complicated instincts are only catenary

reflexes (Kettenreflexe). These, however, have not yet been

sufficiently analyzed.

Loeb’s conception is also fruitful in another direction, for

as Mach says (00, p. 64) it throws light on the relations

between development and instinct. Numerous “ instincts " of

ants, such as their reactions to moisture, heat, light, and con.

tact, are evidently simple reflexes and may, I believe, be

treated as cases of hygro-, thermo-, helio-, and thigmotaxis.

The first and last of these reactions are especially striking.

Similarly the olfactory reactions, which are such an extremely

important factor in the lives of ants, are probably not essentially

different from the chemotactic reactions of simpler organisms.

The consociation of ants in mixed and compound nests is

undoubtedly dependent to a very considerable extent on

olfactory reactions. The young auxiliaries that hatch from

stolen pupz are at once adopted in the mixed nests because

they have acquired the nest odor of the dominant species.

Moreover, the fact that these intimate relations are established

only between ants of rather close taxonomic affinities is prob-

ably due to their having very similar odoriferous secretions to

begin with. The animosity of ants seems to be at once excited

by species which emit peculiar or unfamiliar odors. While I

accept Loeb’s conception as simplifying to a considerable

extent the problem of instinct, I cannot suppose with Bethe

('98,'00) that the behavior of ants and bees is entirely of ,

reflex nature; I can only indorse Wasmann's (99b) and Forel's

('00—01) comments on this author's extreme views. :

Wasmann (91, p. 179 e£ seg.) has shown in detail why ts

No. 418.] NESTS OF AMERICAN ANTS. 813

quite unnecessary to assume the existence of anything beyond

instinct and simple intelligence in the ants which form com-

pound and mixed nests. I should even be inclined to place

a more moderate estimate than Wasmann on the psychical

endowments of these animals. The manifestations of intelli-

gence are very feeble, as any observer who tries to free himself

from anthropomorphism will surely find. There are distinct

traces of associations with indications of some permanence of

these associations, or what might be called animal memory as

restricted in its meaning by Thorndike ('98, pp. 98, 99). Imita-

tion is clearly manifested, but in a form which does not neces-

sarily imply the existence of consciousness. There is a certain

ability to profit by experience, and considerable power of

adaptation to new circumstances, both remarkably developed

as compared with these powers in other insects. There is

evidence of choice and of that which it necessarily presupposes,

viz., will, but there are no evidences of anything resembling

abstract thought, cognition, or ratiocination as manifested in

man. Nor are there the slightest grounds for postulating the

existence of these powers, which would be a hindrance rather

than a help in the activities of ants under existing conditions.

Having arrived at the same conclusion as Wasmann that

there are no evidences of ratiocination in ants we have reached

the limits of our brief inquiry. This conclusion, however,

even if it be extended so as to exclude all animals except man

from a participation in this faculty, does not imply the admis-

sion of a qualitative difference between the human and animal

psyche, as understood by Wasmann. Surely the sciences of

comparative physiology, anatomy, and embryology, not to men-

tion paleontology, distribution, and taxonomy, must have vm

cultivated to little purpose during the nineteenth century if we

are to rest satisfed with the scholastic definition of ratiocina-

tion as an adequate and final verity. And surely no one who

is conversant with modern biological science will accept the

views that the power of abstract, ratiocinative thought, which

is absent in infants and young children, scarcely developed in

Savages and highly developed and generally manifested only in

the minority of civilized men, has miraculously sprung into

814 THE AMERICAN NATURALIST. [Vor. XXXV.

existence in full panoply like the daughter of Jove. Such con-

ceptions recall the rhetorical figure which Houzeau (72, Vol. II,

p. 264) aptly uses in his discussion of this same matter: “The

spark," he says, “which we draw from a rod of wax is analo-

gous to that from the Leyden jar and we attribute it to elec-

tricity. Had we from the very first sought to liken it to the

thunderbolt, the difference of proportions, the quantitative ine-

quality might have been such that we should have been scan-

.dalized by the comparison." But we need not dwell on mere

opinions respecting the status of ratiocination in the animal

kingdom. That the task of tracing reason to more generalized

and primitive psychic processes is not impossible is shown by

Binet's recent investigations (01), the gist of which is included

in the following quotation (p. 159): “ There is no decided differ-

ence between perception and logical reasoning; the two opera-

tions are both reasonings, transitions from the known to the

unknown. The analogy is so close that we were able to compare

perception with formal reasoning, and to show that perception

contains all the essential elements of a peripatetic syllogism

(see p. 88). In short, perception and logical reasoning are only

the two extremes of a long series of phenomena, and when we

place ourselves in the middle of the series we find inferences

which belong to both at the same time (see p. 70). Further,

we have shown that a kind of filial relationship exists between

perception and the reasonings of conscious logic. Thus when

we make systematized anzesthesia, which has been developed in

a patient relatively to a cerfain person, gradually disappear the

thing which appears first of all is the perception of the person

as species; and it is only afterwards, by a kind of ascending

evolution, that the recognition of the person as individual

takes place; now, we know that recognition is a complex oper

ation which touches closely upon reasoning properly so called.

All these reasons lead to the belief that perceptive reasoning

and logical reasoning imply the same mechanism (see p. 77)

A somewhat similar conclusion respecting the derivation of

ratiocination is reached by Wundt (01, pp. 342; 395):

However doubtful we may be of the complete. succes

attempts like that of Binet, we may be confident, nevertheless,

s of

No. 418.] NESTS OF AMERICAN ANTS. 815

that they contain the germs of promise, for the consensus of the

biological sciences leads us at last to one point of view: “ We

are prepared," as Mach says (98, p. 235), “ to regard ourselves

and every one of our ideas as a product and a subject of uni-

versal evolution ; and in this way we shall advance sturdily and

unimpeded along the paths which the future will throw open

to us."

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'00b

NOTES ON LIVING NAUTILUS.

BASHFORD DEAN.

DuRiNG a recent visit to Manila I learned from Commis-

sioner Dean C. Worcester that Nautilus, a form which one

usually associates with remote and cannibal islands, could be

collected quite readily in the straits between the islands of

Negros and Cebu. This locality was accessible, and a short

stay there, I was told, would afford one an opportunity of

examining this, the only living picture of orthoceratids and

ammonites, to say nothing of its greater interest as the

probable key to the puzzles of cephalopod descent. So it

came about that I visited Negros. Thanks to the kindness

of Professor Worcester, the trip itself proved a zoólogical

excursion de luxe, for he secured for me the services of his

long-time guide, the taxidermist Mateo Francisco, and he put

us in charge of his good friend, Don José Bocanegra, whose

large sugar estate is near the town of Bais. It was during this

trip that the following notes and sketches were outlined. They

have been allowed to stand practically as jotted down, as first

impressions, — with my apologies, and a dedication, if they are

worth one, to the student of Nautilus, my good friend and

former colleague in Columbia University, Dr. Arthur Willey.

Nautilus is doubtless common throughout the waters of the

southern Philippines.! It can, however, be obtained so readily

in the region of southern Negros that such a station, if only

because of its accessibility, deserves to become a classic one.

The reason why a deep-water form like Nautilus can be secured

with little difficulty here is a simple one: it is a regular

by-product, so to speak, of the traps of the fishermen. Its

* I note, in passing, that dead shells of Nautilus occur not uncommonly as fat

north as Japan. One was picked up a few years ago near the laboratory at

Misaki (lat. 35? 10’ N.).

819

THE

j /

AMERICAN NATURALIST.

MI Rs

| XR

ari M - “4 J

À ER ;

WNI

» KY if [ ul X /

MU Um

" T

| ACE ]

[Vor.

In the background the Philippine fish-trap in which they are taken.

_Fic. 1. — Sketch illustrating living Nautilus.

XXXV.

No. 418.] NOTES ON LIVING NAUTILUS. 821

flesh is, indeed, eaten, but it is little esteemed. The shells have

hitherto found little market ; locally they are used as drinking

cups, or flower vases, and are sometimes cut into roughly

shaped spoons. But they are now bought extensively by

Chinese shopkeepers, and at such good prices that there will

probably be a goodly Nautilus fishery before long. The

Chinese, I was told, export the shells to China, where they

are used as material for button-making.

The best catches of Nautilus are made in deep water, that is,

according to the estimates of the fishermen, at a depth of from

250 to 400 bragas (— about 450-700 meters). I notice, how-

ever, that in the Spanish charts the maximum depth recorded

is 200 meters, and it is probable accordingly that either the

fishermen make extraordinary allowance for the sag of their

line, or that the measurement given on the chart represents

the maximum sounding limit of the Coast Survey steamer. As

the water shallows, Nautilus becomes less abundant : in water

of 100 meters it is still taken, although in small numbers. A.

single specimen, as Señor Romero informed me while at Bais,

was collected in that bay by the native divers while gathering

pearl oysters. There can, I believe, be little doubt of this

instance, for my informant is well acquainted with Nautilus,

and states that the animal was brought to him living. It was

obtained in water less than 4 m. in depth.

The most favorable station thus far discovered is, as Professor

Worcester stated, near the small fishing village Manjuyod,

about six miles distant from Bais. This and other localities

where I learned that -Nautilus is taken are indicated in the

accompanying outline map (Fig. 2). That it is taken off the

villages of Cebu, Malaboyoc, Ginatilan, and Sambuan, I know

only from hearsay. I was told that a Chinese shopkeeper made

a tour of these places and collected 3000 shells ; and he

reported that the fishing was done not far from shore. In

the locality indicated, near the pueblo of San juan on the

Southern coast. of Siquijor, Nautilus is taken in relatively

Shallow water, 100 m. and less; here, however, its abundance

is inconstant. In this neighborhood I picked up a shell gem

in diameter.

822 THE AMERICAN NATURALIST. [VoL. XXXV.

Season of the year has an important bearing upon collecting.

Spring and early summer are the most favorable, probably

because the water is quiet in the absence of the northern

monsoon. June is generally stated to be the month in which

MALABOYOC

+ CEBU

GINATILAN

SAMBUAN

N. 9.30.

TANON

DuMAGUETE

SANDUGAN

NEGROS

PASIGAJON

SIQUIJOR

E. 139.30| Merio.

Sanj FERNANDO

Fic. 2.— = map of the neighborhood of Bais, southern Negros, showing in stippled areas

ome of the fishing grounds for Nautilus. (After Spanish coast chart.)

the largest catches are made, as many as twenty shells. having

been taken in a single basket. The least favorable season is

winter, for the wind sweeps down the straits between Negros

and Cebu, raising a sea which soon swamps the small prahus of

No. 418.] NOTES ON LIVING NAUTILUS. 823

the fishermen. It happened, therefore, that at the time of my

visit, in January and February, I was able to get but few

specimens.

The fishing, as has already been stated, is carried on by

means of a fish-trap, a sketch of which is given in Fig. 3. It

is a cage-like, flattened affair, about six feet in length, con-

structed of strips of bamboo woven in coarse mesh. At one

end there is a funnel-shaped insinking. through which the

animals gain entrance. The funnel operates somewhat on the

plan of the mouth of a rat live-trap, the animal pushing through

it, not indeed at the pointed end, but on one side (the median

ventral line, to borrow a phrase). Intercrossing strips then

LAY Ss

< AAN EE

rh eS jr 1

TAR DS ame

PONS Es

Fic. 3. — The Negros fish-cage, or “ bo-bo," in which Nautilus is taken.

close elastically and prevent egress. The cage stands upon

stout strips of bamboo, to which weights are usually attached.

Bait is hung from the roof supports at various points. This

consists of whole fowls, viscera or joints of kid, or the remains

of dogs or cats when the latter attractions can be found.

Freshness of bait is not regarded as a sine qua mon. The

cage “bo-bo,” as it is called in Viscaya, is attached, in the

Way shown in the figure, by a strip of rattan, to the main line,

also of rattan, which connects it with the float. The latter

consists usually of several joints of bamboo. As to the iat

of fishing, it appears that the bo-bo is put down at a favorable

Opportunity, ánd is allowed to remain for several days, often

for a week or longer. At San Juan I found that it was

Customary to examine the cages on stated days, weekly or

824 THE AMERICAN NATURALIST. [Vor. XXXV.

bi-weekly ; at Manjuyod, on the other hand, visits to the

cages are made at irregular intervals. The bait is then

renewed, the captured fish removed, — this through a trap-

door in the bottom, — and the apparatus sunk again to the

bottom. |

In the accompanying cut (Fig. 1) I have attempted to repre-

sent a bo-bo in position, with Nautilus in and around it. This,

I believe, gives a moderately accurate idea of some of the posi-

tions assumed by the animals. Several are represented, partly

A B

Fic. 4, A and B.— Diagrams of shells showing in stippled areas the oral apertures of

ale (A) and female (B) Nautilus

retracted, in a position of rest, and one of them, attracted by

the bait, is shown swimming rapidly near the bottom some-

what in the direction of the cage. The degree of shadow is, of

course, hypothetical, and it is doubtful whether the bottom 1$

as free from other living forms as I have indicated.

The Shell. — In examining fresh specimens I notic

there appeared to be sexual differences in the shells. That of

the female was wider at the sides of the oval aperture and here

showed a somewhat angular contour. This peculiarity Lhar

expressed in the accompanying Fig. 4, B, and it may be con-

trasted with the condition of the shell of a male shown in the

ed that

No. 418.] NOTES ON LIVING NAUTILUS. 825

same figure, 4. This difference was clear in the case of the six

specimens I examined ; but I later found that there must be

considerable variation, for in looking over the shells obtained

at Manjuyod I was unable to distinguish those of the females

from those of the males in as many instances as three out

of ten.!

Another character of the shell is the great number of irreg-

ular growth lines which they often present (Fig. 5). These

lines show frequently, as in the figure at *, an undulation of

a somewhat regular pattern, reminding one of the markings

Fic. 5. — Shell of Nautilus. Outline sketch showing undulatory lines of growth.

formed at the septal rims in Ceratite or Goniatite. Doubtless

this peculiarity in recent shells has already been commented

upon, and the point made that the irregular growth lines can-

not be directly related to those of the fossil forms, since they

are not connected with the formation of septa, several being

Sometimes found on the wall of a single chamber. They are

interesting, however, I suggest, as representing à tendency

uring special periods of shell-forming activity, perhaps after

3 ‘I find that Willey gives (1895, Wat. Sci., p. 412) a similar note, stating that

"in most adults the shell can readily be identified as belonging to male or feine,

but often this identification is very difficult, and in young shells impossible.

figures extremes of difference.

826 THE AMERICAN NATURALIST. | [Vor. XXXV.

some unusual shock has been given to the free rim of the

shell, for the mantle to contract in crenulate lines, a tendency

which during the descent of the tetrabranchs may well have

been seized upon by selection and made of use in the formation

of the specialized margins of the septa. And from this stand-

point the recent markings may be regarded as related to

the curiously expressed lines on the ancient shells. They

seem of entirely too regular a character, as close examination

shows, to be interpreted merely as lines marking the repair of

the free lip of the shell. Such lines of repair do, indeed, occur

and are common, for the shell-lip is delicate and often apt to

be injured in an animal with the evident habits of nautilus. I

have in mind one instance when the shell had been repaired,

after the lip had been crushed badly ; in this case the main

fracture passed backward from the rim of the lip to a distance

of 4.5 cm. f

General Appearance of the Living Animal. — The first glimpse

at a living specimen showed it at the bottom of the vessel in

which it had been brought, its position upright somewhat as

shown in the present Fig. 6. If the vessel is sufficiently large -

the animal is usually found with its back (Z.e., its aboral pole,

so to speak) pushed as far as possible into a corner. And

if several specimens are thus confined they will often be found

to have backed away from one another as far as possible.

They remained thus almost motionless sometimes for hours ;

at other times they will exhibit active movements and subside

quite suddenly. The color of the animal one gets little idea of

from preserved specimens. The general color of the exposed

parts, hood excepted, is white, clean, opaque, almost the same

tint, in fact, as the body of the shell. The latter was in every

case I observed brilliantly clean. The face of Nautilus, then,

is white, as is also the dorsal fold of the mantle, which rises

into sight within the concave hinder rim of the hood and covers

when extended the jet-black portion of the coil of the shell.

The dorsal surface of the hood, as one would infer from pre

served specimens, is of a brownish color. Looking at this

surface for the first time in a living specimen, one is given the

impression that it has been spattered with burnt umber, raw

No. 418.] NOTES ON LIVING. NAUTILUS, 827

sienna, and yellow ochre, and that the pigment has then been

carelessly wiped away, leaving the warty prominences in white

relief. Nowhere are observed the characteristic chromato-

phores of the dibranch. The only pigmentation which may

further be noted is on and around the eye, and in the creases

between the thickened proximal portions of the oral tentacles,

also on the edge of the mantle, margining the oral aperture ;

from this the shell itself receives an inky border, which is

continued from the blackened portion of the coil around the

entire aperture. This line measures about 2mm. in width

and can be detected from the outer surface of the shell; it

does not appear, however, in shells which have been carelessly

cleaned. ^

Nautilus, as far as my experience goes, lives but a short time

in confinement, twenty hours being the longest time I was able

to keep one alive. And even from the beginning the animal

was evidently not at its best. It is true that the conditions

under which I observed them were unfavorable, for in no case

were specimens brought me which had been confined less than

three hours in a water bucket. And the best aquarium I could

improvise was a tank which held only about two hundred liters

of water. Changing the water at frequent intervals had little

effect in keeping the animals alive. I soon found that it was

difficult to know just when the animal did die, for it retains a

very lifelike position. This I discovered as follows : with

a view of finding whether at any time movements became

more active I caused my helper to watch a specimen through

out the night, the man to call me in case it showed change of

Position or any peculiar behavior. In the early morning I was

told that the animal had shown no movements, a statement I

soon had little reason to doubt, for I found my Nautilus dead.

On returning it to the water I observed that it floated, retain-

ing, however, its customary position ; and so it stayed 1n spite

of my efforts to cause it to sink. This is the only case 1M

which I observed a specimen remain at the surface ; and the

thought suggested itself that it was from such floating speci-

mens that the idea originated that Nautilus was more OF less

à surface form.

828 THE AMERICAN NATURALIST. | [Vor. XXXV.

The usual position and appearance of nautilus in confinement

are sketched in Fig. 6.1 It will be noted that the lower rim of

the hood lies in approximately a horizontal position, the dorsal

NNN i

Fas. 6, 7, 8. — Diagrams showing the position of Nautilus at different stages of closing.

fold of the mantle protruding slightly above the concave hinder

rim, entirely concealing the black portion of the shell. The

1 In the photograph which Willey has given (Q. 7. M. S. 1896, p. 179), the a

ing specimen appears to be somewhat droopy, or perhaps it is just recovering

from its alarm at being confined in so small a vessel, evidently a m

Thus the base of the hood has shrunken away from the shell, es ially at the

sides (but then this is not’ unusual) and the dorsal fold of the mantl

normally rises to the upper margin of the black area of the shell can hardly be

seen; the tentacles, too, sag down in a way which I think rather uncustomary,

cramped, perhaps, by the smallness of the glass vessel. y impression, t00, -*

that they have drawn down the ventral lip of the shell somewhat lower than 1$

usual. w easy it is, though, to criticise a zoólogical result, even in the form

of an admirable photograph, and to forget that it may have been my own speci-

mens that were sickly!

No. 418.] NOTES ON LIVING NAUTILUS. 829

eye protrudes conspicuously, but not, however, beyond the

plane of the surface of the shell; its lower rim lies more or

less within the lateral indentation of the shell's aperture.

This indentation, as will later be seen, is apparently a provision

to enable the animal to see, even when largely retracted. The

tip of the siphon is also seen to protrude (S), and between it

and the hood lie the closely arranged oral tentacles. When the

animal is in this position

the proximal ends of the

tentacles appear long,

narrow, and moderately

well rounded ; the distal

portions taper out deli-

cately, the tips extended

and weaving together,

somewhat as shown in

the figure. Examining

a similar specimen from

in front (Fig. 10), we

observe a more or less

regular arrangement of

these tips of the tenta-

cles; they cluster to-

gether, lying in the main

in front of and almost Fico. Vae patio ret 1a position.

concealing the opening

of the mouth. In this position the shape of the dorsal surface

of the hood is well shown ; there is a median flattened tract

from which the sides shelve away, moulding the hood to ue

whorl of the shell and to the region of the eyes and tentacles.

Looking at a specimen from above, we obtain the appearance

shown in Fig. 12. ` This, however, pictures a more retracted

Stage. It will be seen that the hood is neatly adjusted to the

outline of the aperture of the shell. Anteriorly the distal ends

of the tentacles protrude in a tuft and are arranged quite

symmetrically.1

‘Willey (Q. 7. M. S. 1898, Pl. X) figures them in complete extension, diverg:

ing widely at the sides of the head, and still symmetrical in position.

830 THE AMERICAN NATURALIST. [VoL. XXXV.

Appearance when retracting. — When a specimen is picked

out of water (and it bears handling with considerable compo-

sure) it slowly retracts, presenting the appearance shown in

side view in Figs. 7 and 8, in three-quarter view in Fig. 9, in

dorsal view Fig. 12, and in oral view Fig. 11. The first symp-

toms of retraction are seen in the shortening of the tips of

the tentacles and in the drooping of the hood. These processes

an early stage of retraction.

Nautilus resting and partly

Figs. 10, 11, 12. — 10, Nautilus, front view, showing oral region, at

I , Oral region at a lat t: f retraction. 12, D 1 t of

g g , Dor p

retracted, showing tentacles.

are more or less rapid, depending upon the condition of the

animal ; and I am led to believe that under normal circum-

stances retraction can take place very quickly. In the short-

ening of the tips of the tentacles the fleshy proximal portions

become. notably stouter, at the same time more angular in

section, dovetailing one beside the other so tightly that their

common outer surface is flattened where it is later to be oppos

to the inside of the shell. Observe also that the hood becomes

notably depressed, at the same time exposing the dorsal rim of

No. 418.] NOTES ON LIVING NAUTILUS. 831

the mantle; this then recedes, exposing the blackened portion

of the shell. The eye is still seen peering over the notch at

the lip of the shell. In this stage of retraction the animal will

often remain for many minutes; if handled somewhat roughly

it will retract still further, presenting a stage of closure shown

in Fig. 8. Here the hood has closed over the aperture of the

shell like a lid, or rather like an operculum; it has shrunken

and flattened somewhat at the same time; behind it the :

mantle has receded, leaving the. blackened area of the shell

much in evidence. At the anterior end (Fig. 11) can still -

be seen the turgid ends of some of the tentacles. The final

closure of the aperture of the shell appears to take place by

the bending down of the anterior flap-like portion of the hood,

a process, however, which does not readily occur. Only by

sharply stimulating this region have I seen total closure, and

even then but for a few moments. If undisturbed, even if

kept out of water, the animal slowly reopens. The hood first

rises sluggishly, often separating itself behind from the shell,

and the tentacles begin to protrude. More slowly does the

mantle expand again over the blackened area. If still kept

out of water the tentacles protrude and hang down over the

edge of the shell in a flaccid mass.

If an animal be taken out of water and held in an inverted

position it will at first close just as has been described. After

some time, hours (Professor Worcester tells me) in the case

of specimens just out of water, minutes in the case of those

Which have been kept in aquarium, the heavy fleshy “ head ”

'of the animal will begin to droop downward, and will finally

appear as shown in side view in Fig. 14 and in ventral view

in Fig. rs. Whether under such circumstances the animal

will ultimately fall out of its shell, I neglected to observe.

I found, however, that by this means one can readily remove

the animal from the shell and without breaking the delicate

Siphuncular tube. The muscles attaching the body to the

shell occupy very definite areas within the latero-dorsal region

of the aperture and at the sides of the coil of the shell 2 and

these areas are landmarked quite clearly in the inverted animal.

The muscles may accordingly be separated from the shell, one s

832 THE AMERICAN NATURALIST. [Vor. XXXV.

finger-tips answering adequately for the purpose. Before the

operation is completed, however, the siphuncular tube should

be loosened ; this is effected by thrusting the hand within the

shell below the siphon. At such treatment the animal retracts,

but after several attempts the finger-tips find their way to the

base of the siphuncle. In the inverted position the animal

presents several peculiar features. The hood is bent and mis-

shapen, and evidently possesses considerable plasticity. Adja-

cent to it the black portion of the shell is largely exposed,

owing to the retraction of the mantle. The tentacles are

tumid, retracted, and are closely pressed together. The eyes

stand out prominently, like mushrooms. The siphon shows

its broad separation from the tentacular portion of the head,

and hangs downward as a stout, muscular mass; at its base

on either side droops a thin fold of the mantle (M A). |. Ot

this the inky black edge corresponds to the black line already

referred to as lying within the aperture of the shell. The

entire mantle is now greatly retracted: its posterior ventral

rim (M P) exposes a portion of the extensive mantle cavity, in

which, although not shown in the figure, can be seen the gills.

The animal while in this position is observed to give an occa-

sional gasping movement which involves the adjacent mantle

rim as well as the siphon folds; these are expanded and

contracted quickly.

The Tentacles and their. Movements, Extension and Retrac-

tion. — Extension and retraction play an important and con-

stant part in the economy of these organs. The maximum

extension observed is about as shown in Fig. 8; shortly after

death it is possible, however, to draw out the tentacles con-

siderably further, to the degree which I have indicated in the

swimming Nautilus of Fig. 1, or in the diagram, Fig. 13, C

. Indeed, I suspect that the tentacles can be everted even fur-

ther than this, should the animal be so disposed. It is of

course doubtful if this protraction can be carried to the degree

which would cause the obliteration of the distinct boundary

line which separates the distal from the proximal portion of

the tentacles. Judged from their varied movements, these

organs have come to be highly specialized in the matter of

NO. 418.] NOTES ON LIVING NAUTILUS. 833

lengthening and shortening, if for no other purpose than that

of their protection within the shell How these processes

came to be evolved there is no good evidence for determining ;

an aftractive hypothesis would connect them with the trans-

verse foldings which permit the tentacles to serve as adhesive

organs. And one can readily conceive that a

process of shortening based upon the principle

of transverse folds would result in the production

of one highly effective fold, to the loss or detri-

ment of the rest. This highly serviceable fold

would of course be represented in the present

boundary between the proximal and distal portions

of the tentacle. According to such a view the

proximal portion has now lost all traces of trans-

verse segmentation, becoming, in fact, but a kind

of sheath for the rest. The distal portion, on the

other hand, on account of its varied move- B

ments, has still retained them. It will

accordingly be seen that the slender distal

portion of the tentacle is not to be regarded

as the homologue of a single highly A

specialized sucking organ, as some

writers have maintained. There is,

I believe, no evidence that such

sucking organs are to be looked for

in the tetrabranch division of the

cephalopods. On the contrary, this

division, possessing tentacles many

Pit pans cont soscd any Ce

; y have

required the specialized sucking rings which characterize the

few-armed cephalopods. à

: The Movements of Nautilus. — In captivity Nautilus remains

in one position, sometimes for hours. This sluggishness, how:

ever, may reasonably be due to the shock it suffers in being

suddenly brought to live in surface water. Slight movements

of the ends of the tentacles are sometimes the only signs that

the animal is still alive. Spasmodically, however, a strength

834 THE AMERICAN NATURALIST. (Vor. XXXV.

of movement is exhibited which convinces the observer that

under natural conditions Nautilus must be looked upon as an

active form.! It will suddenly sweep from the bottom and

bump its shell into a side or corner of the aquarium, — this,

too, with considerable energy. At the shock the animal sinks to

the bottom, partially retracts, and again remains almost motion-

less. In these spasmodic movements the animal rarely rises

more than three or four inches from the bottom, although on

one occasion I observed it swim nearly to the surface, a ver-

tical distance, however, of not more than forty-five cm.

During the night-time the movements appear to be if any-

thing less active than during the day. A curious rocking

movement is often observed, the animal swinging forward or

backward, or from side to side, but never to a degree suggest-

ing a change in the resting position. Rotation — that is, turn-

ing the shell to right or left — is very readily accomplished,

and without, as far as I could see, any special movement on

the part of the siphon. A very slight change in the direction

of the water current exhaled from the siphon, aided by a slight

leaning of the shell, is evidently enough to account for this

ready movement, for the animal is delicately poised and little

mechanical impulse is needed. Occasional distinct move-

ments of the hood begin, and tentacles are observed. The

hood. will partly close, then open again, somewhat abruptly.

The tentacles show a peculiar weaving movement ; sometimes

they separate quite widely from one another, and then

converge. In their function as adhesive organs they will

attach themselves somewhat delicately to objects presented to

1 Professor Moseley, in his interesting Votes of a Naturalist on H. M.S. €

lenger, observed that Nautilus swims with its tentacles “extended radially Hom

the head, somewhat like the tentacles in a sea-anemone; but each pair has its

definite and different direction, which is constantly maintained.” As far ves my

own observations go, the exact position of the tentacles during swimming is not

easy to determine, for the specimens were closely confined and their movements

sudden and short; I am led to conclude, however, that the tentacles are drawn

together during sustained movement, like those of other cephalopods. And

this position I represented them, somewhat stiffly, perhaps, in Fig. 1.

above volume, together with other references to Nautilus, I am indebted to. ud

fessor Ijima, who kindly secured them for me in the University of Tokyo

forwarded them to Misaki.

No. 418.] NOTES ON LIVING NAUTILUS. 835

them.! When roughly touched, however, they will merely retract.

On one occasion only did I observe the separation of the tentacles

around the mouth and the protrusion of the jaws, to the degree

shown in Fig. t. I was not able to induce the animals to feed.

I noticed, however, that in specimens which had recently died

little effort is needed to draw the jaws forward to a surprising

degree, so that they would stand well outside of the proximal

portions of the tentacles. And I think that there is little

doubt that the jaws can be used in a position which one would

: tof its shell. Th

Fic. 14. — Side view of Nautilus shown hanging is positi

after the animal has been held for some time hood downward.

hardly be led to expect from an examination of preserved speci-

mens. Accordingly I infer that the tentacles are of less impor-

tance in the mechanical operation of feeding than is popularly

believed. When the jaws are drawn out, as above noted, the

tongue is found to lie far forward, curving outward between

the tips of the jaws. Nautilus is already well known to have

a similar diet to that of other cephalopods. Its fondness for

animal food in a decomposing condition I note from hear-

Say. I found, however, that with its strong jaws and rasping

1 Willey in his notes convinces me that I have not examined the tentacles

When at their best, for he has seen them “adhere so firmly to à foreign quet

to e torn away from the animal when the foreign body is forcibly cca

(Q. J. M. S., 1898, p- 207.) His figure indicates that the tentacles when firmly

attached become partially retracted, at least as far as their tips are concemed.

836 THE AMERICAN NATURALIST. [VoL. XXXV.

tongue it has little difficulty in feeding upon animals which are

enclosed in a stout leathery skin. Reference has already been

made to the spasmodic movement of the siphonal flaps and

of the ventral rim of the mantle; under normal conditions the

movement of exhalation is practically confined to the siphonal

flaps, the ventral rim of the mantle being closely apposed to

the aperture of the shell, just as the portion of the mantle

behind the hood is apposed to the black region of the coil.

It is not difficult to convince one's self that under natural

conditions Nautilus is well able to

protrude its * head ” quite widely

from the aperture of the shell in

a way suggested by the position

of Fig. 14.

Breeding Season, and the Ques-

tion whether Eggs and Young have

been collected by Fishermen.— From

the condition of the reproductive

organs in five specimens examined

I am led to infer that the breeding

T de of Nautilus in the region of south-

Fic. 15. — Individual of preceding figure Ern Negros extends over a con:

Hn M eet one eee siderable season ; this evidence,

however, is too meager to be of particular value.! There is

certainly, it seems to me, a fair chance of securing the embryos

of Nautilus in this region if what I learned from the fishermen is

to be relied upon. One of these men in particular gave such a

satisfactory account that I cannot very well doubt that he had

at some time seen deposited eggs. And it may be well to state

parenthetically that no data were given to the fishermen which

they could make use of in their reminiscences. The fisherman

in question assured me that on one occasion, as nearly as he

could remember it was during the early part of the summer of

! I find that Willey at first believed that in the case of Nautilus in New Guinea

“reproduction takes place all the year round” (Q. J. M. S., 1896, p- 176). aan

his observations in Lifu, however, he abandoned this conclusion, stating that "1t

now seems probable that the breeding of Nautilus . . . is subject to a definite law

of periodicity” (Mature, 1897, p. 403).

No. 418.] NOTES ON LIVING NAUTILUS. 837

1900, he noticed eggs attached to a fish-cage in which Nautilus

had been taken : there were several eggs, he did not remember

how many, but they were attached not far from one another,

and stood up on end, each as big as two joints of his finger,

and with a leathery skin. Their color was white, and—this

was the curious part of his story —they had brown markings

across them! which reminded him of those on the shell of the

adult animal. It was from the latter feature that he was sure

the eggs belonged to Nautilus. His account seemed to me

worth repeating, since it is hardly probable that he could have

invented offhand so tolerably accurate a story, or that he would

have been apt to describe an object which has nothing to

do with eggs of Nautilus and yet resembles them so clearly.

Moreover, there is a greater probability of accuracy when one

considers that such a man is apt to be a skillful observer in mat-

ters relating to his work, —a thing which a stay-at-home zoól-

ogist often finds hard to realize. I was assured on every hand

that small specimens of Nautilus are relatively common during

the fishing season ; that specimens whose shell is the size of

à silver dollar are often thrown away, and that still smaller

specimens are occasionally taken, in spite of the large mesh

of the fish-cage.

! Could these be the rows of foldings and “fenestrations " of the outer egg

capsule which Willey described (ature, 1897, p. 402)?

BASELEVELING AND ITS FAUNAL SIGNIFI-

CANCE, WITH ILLUSTRATIONS FROM

SOUTHEASTERN UNITED STATES.

CHARLES C. ADAMS.

In considering some of the peculiarities of the fauna of the

southeastern United States I wish to call attention to certain

principles that are fundamental to an understanding of the

present distribution of many forms of life, especially those of

land and fresh-water. Especial stress is laid on the relation of

physiographic changes, and baseleveling in particular, to faunal

changes and to the differentiation of land and fresh-water

animals.!

The necessity of correlating the results of baseleveling, and

river histories in particular, with the distribution of habitats

and their fauna, was very emphatically impressed upon me in

connection with some studies upon the Pleuroceridz of south-

eastern United States. This relation was clearly demonstrated

to me when I learned that the former course of the Tennessee

River from Chattanooga was direct to the Gulf via the Coosa-

Alabama system. Having thus become fully convinced of the

great biological significance of the baseleveling factors, it was of

considerable interest that I later learned that Woodworth (94)

had already appreciated this significance.

/ Ina general way, this relation of physiography to topographic

: and geographic distribution has been well understood, but such

\ Studies need to be carried out in more detail. The geologists,

from their familiarity with the physiographic processes, seem

to have had a better appreciation of this relation than zoólo-

gists. Woodworth (94) has fully realized in a detailed way the

Influence of the baseleveling processes upon faunal changes.

1I am under obligations to Messrs. Salisbury, Hayes, and — aS per-

T ion to use their figures and to Prof. E. A. Birge of the Wisconsin . and

atural History Survey for the loan of the cuts for Figs. b ? and 3.

840 THE AMERICAN NATURALIST. [Vor. XXXV.

His paper is of great value to the general student as well as

to the special student of faunal problems. I find that he

had anticipated many of the conclusions to which I had come,

but, on account of their imperfect recognition, they will be

well worth repeating. Geologists have been primarily con-

cerned with the submarine aspect of physiographic changes.

Chamberlin has discussed this in a very suggestive paper ('98).

Since there is a very evident correlation between the erosion

of the land and the deposition of the eroded material in the

sea, we are able to see that under some circumstances both

the submarine and subaerial factors of physiographic change

must be taken into consideration in explaining habitats and

the struggle for existence of aquatic organisms. This may be -

illustrated if we suppose that as a continental shelf becomes

reduced in area the crowding of its fauna might force some

of its members into fresh water, a habitat with a relatively

poor fauna, which consequently is open territory.

It appears that many biologists are unfamiliar with the idea of

baseleveling. For this reason the following brief account is

given, showing how rivers cut down land and tend to reduce it

to a baselevel. If a comparatively level country be elevated

out of the sea a few hundred feet, and rains begin to work

upon it, the slight inequalities of the surface will cause the

waters to collect in the hollows and then run off, carrying

detritus and cutting a trench or gully. This trench becomes

with succeeding rains deeper, wider, and longer as it cuts its

way into the uplands, and thus valleys may be formed. With

this growth or increase of the area of valleys there is an

increase of lowland and cliff habitat and a decrease in the

upland habitat. These progressive changes are beautifully

shown in Figs. 1, 2, and 3, after Salisbury. It is important

to notice here the changes in the relative proportions of the

habitats, with a premium placed upon those forms which can

1 Dr. H. C. Cowlethas recently applied these principles to the study of plants:

The Physiographic Ecology of Chicago and Vicinity; A Study of the Origin,

Development, and Classification of Plant Societies, Bot. Gazette,

(1901), pp. 73-108, 145-182.

The Plant Societies of Chicago and Vicinity, Bull. of the Geographic Society f

Chicago, No. 2, 1901.

No. 418.] BASELEVELING. 841

occupy the habitat that is increasing in area. These changes

wil produce tension lines between the inhabitants of the

various habitats.

With regard to the river fauna, at first when the fall of the

river is great, rapid-water forms have the advantage, but with

the development of sluggish, meandering streams upon bottom

land, quiet-water forms find a home. Since the lowland or

swampy conditions begin at the mouth of the stream, quiet-

Fic. 1.— Young valleys. (After Salisbury.)

Water forms will be introduced there, and with the noe

"p stream of lowland conditions this fauna is led in inn —

tion ; similarly the head of the valley leads its rapid-water

fauna in the same direction. Thus there is a definite succes-

"lon of forms working up the valley, and, in a oe

by passing down stream one gets a recapitulation ien gun

of fauna which inhabit a given point of the stream during Its

Ideal history, Dendritic tributary streams help to carry vois

same conditions in diverging directions, thus leading the fauna

n Various directions toward the divides.

842 THE AMERICAN NATURALIST. [Vou. XXXV.

Later on, with the wearing away of the inter-stream uplands,

which at first were flat-topped, but which become more and

more reduced by the dendritic etchings of the smaller streams,

the divides gradually become more sharp, and the country

eae -— n -

*3 d « é

"1 A» P Sar tus.

Fic. 2. — Valleys of later stage. (After Salisbury.)

reaches its maximum degree of roughness and cliff habitat,

and consequently the greatest isolation of its streams and their

faunas. By degrees these sharp divides are lowered, and head-

water streams favorably located begin to capture fellow streams

with their faunas, and thus there is a mixing of rapid-water

faunas.

The time of maximum roughness in the topography coin-

cides with that of maximum isolation, but with the fusion and

concentration of drainage lines a period begins which favors

wide distribution by the removal of barriers ; thus in a country

approaching baselevel a wide distribution of the fauna will be

greatly facilitated with a premium placed upon the lowland

fauna. This stage of wide distribution favors interbreeding,

No. 418.] BASELEVELING. 843

and thus a period of swamping by intercrossing may be pro-

duced. The cliff fauna would have its maximum distribution

and area of habitat when the country was at its maximum

roughness.

It will not be necessary here to go further into the details

of these processes, as the preceding remarks, with an exami-

nation of the diagrams, will suffice to show that these princi-

ples may be applied almost indefinitely, but the details vary

greatly in different localities. One point perhaps needs special

emphasis, and that is, we must think of these processes as

active and taking place before our eyes. -

The bearing of these factors upon the large amount of

endemism seen in southeastern United States will be illus-

trated by a few examples. The physiographic changes of this

Fic, 4. — Valleys of further development than shown in Fig. 2. (After Salisbury.)

region have been very carefully studied, but comparatively little

3 been done to correlate these changes with the geographic

'stribution of the fauna of this region, and especially with its

endemism. I hope, by calling attention to a few of these

844 THE AMERICAN NATURALIST. [VoL. XXXV.

correlations and to some of the principles involved, that others,

more familiar with special groups of animals, may be led to

make similar comparisons and correlations, and also to empha-

size the fact that causal and genetic faunal studies must in the

future take into account these factors, if work of permanent

value is to be done. Biological surveys will, carried out in

this way, become decidedly more than faunal lists, with miscel- .

laneous biological information. In order to emphasize certain

aspects of the baseleveling processes special phases will be

discussed under separate heads.

Migration of Divides.

The migration of divides has long been recognized as an

important factor in the mixing of river faunas, but without a

distinct recognition of it as a phase of the baseleveling process.

The following examples will show the influence of the migra-

tion of divides upon faunas. But first, attention will be called

to some of the facts of the distribution of the family Pleu-

roceride, or Strepomatidze of the older authors. This family

is distinctly North American, living forms occurring here only.

Tryon’s monograph ('73) on this family, from which I take my

data, recognizes four hundred and sixty-four species. Excepting

those species whose exact locality is not definitely known and

certain Pacific coast forms, most of these are limited to the

Tennessee River and its tributaries above Florence, Alabama,

and to the Coosa River. Two-thirds, or about three hundred,

of these species are confined to these two river systems.

The region in which this family reaches its maximum

development in both individuals and species is remarkably

limited and is bounded by Tryon as follows. On the “North, by

the Tennessee River and tributaries. The Cumberland Moun-

tains prevent the dispersion of the species of this river to the

northward until its course is directed into Alabama. Here the

character of its species (which we will again allude to further

on) changes, and they become gradually less numerous and of

greater geographical dispersion as the river runs toward the

west. Eas/,the mountain range of the Blue Ridge, running

49V) "wprooodno[q A Jo uonvpuo49gip umurxeur jo voae ayy sr yoya *&opea uenpejeddy jo dey — + "514

No. 418.] BASELEVELING. 845

southwestwardly into the interior of northern Georgia ; thence,

the Chattahoochee River and tributaries, to within a hundred

miles of the Gulf. South, the species are restrained from

: 7 [PP PH i.

; T c 777 2

i: ; viaa

772 2o:

Spreading by the influence of the Gulf of Mexico. West, the

id labama, Cahawba, and Black Warrior Rivers and their tribu-

ries, those of the latter reaching almost to Florence, on the

8 46 THE AMERICAN NATURALIST. [VoL. XXXV.

Tennessee River, which may represent the northwestern point

of our boundary.

* These limits are : necessarily imperfect, but, nevertheless,

include at least three-fourths of our species within an area of

three hundred miles extent, either north and south or east and

west." By reference to Fig. 4 (after Hayes and Campbell), one

will see that these limits are practically those of the Appala-

chian valley, and thus enclose a natural physiographic region.

Again, Tryon remarks: * Assuming the Ohio River as a

dividing line, we find Vat ninety-five per cent of all the species

originate south of it.’

So much for these general facts of the distribution of this

family which are necessary for an understanding of the influ-

ence of the migration of the divides upon them.

With the above facts of the distribution of these shells fresh

in mind, about two years ago I learned from Hayes's (95) paper

of the view that the upper Tennessee River formerly flowed by

way of the Coosa-Alabama Rivers into the Gulf. I saw at once

in this an adequate explanation for the peculiar distribution of

many of these shells in these two river systems. Since the

data for this paper have been collected Mr. C. T. Simpson ('00)

has published similar conclusions, drawn from the Unionidz.

According to Hayes's and Campbell's ('94), and also Hayes's (99),

opinion, the Tennessee River above Chattanooga was captured

by a westward-flowing stream, and thus led away from the

Gulf by the Coosa-Alabama to the Gulf again by way of the

Ohio and Mississippi. Thus we see that the region in which

the maximum development of this family is reached was broken

into two parts (Fig. 5). The bearing of this upon the shell

fauna is very marked.

This family is divided into two sections, and as to their dis-

tribution Tryon (/c. cit., p. xl) says: “ While the Trypanosto-

moid forms attain their maximum development in size and

number in the Tennessee River, they are to a very great

extent replaced by the Goniobasic forms in the Coosa River,

which is undoubtedly the metropolis of the latter. The most

striking genus of each of these groups is absolutely confined

to the respective streams in which the groups had their origin.

No. 418.] BASELEVELING. 847

Thus, Io and Schizostoma are inhabitants, the first of the

Tennessee and branches, the second of the Coosa, and neither

of them are elsewhere found." This is an excellent illustra-

tion of how the migration of a divide may favor differentiation.

Fic. 5.—

Map showing ancient (broken line) and present (continuous line) drainage in the vicinity

of Chattanooga. (Adapted from Hayes, '99.)

The family has thus been broken up into two parts, both

geographically and biologically. |

The genus Pleurocera is credited with eighty-four species,

twenty-three of Which are found in Tennessee, and twenty-one

ie apparently peculiar to it. This same genus is represented in

Alabama system by fourteen species, and three of them are

Peculiar to it. «These species,” says Tryon, “are generally

848 THE AMERICAN NATURALIST. [Vor. XXXV.

confined, however, to those portions of the Coosa and branches

that approach to east Tennessee." Lithasia and Pleurocera

are other genera divided in their distribution between the two

systems. In the genus Strephobasis one species (Bitzniata)

is found in the Black Warrior of Alabama drainage and in the

Tennessee River at Chattanooga.

In the Goniobasic section there are two types of shells: one

group is characteristic of the Tennessee and the other of the

Coosa. In the genus Goniobasis, about ten of the elevated

smooth species are found in the Tennessee, and about the same

number in the Alabama system. Tryon (/oc. cit., p. xlviii) says

of this genus : * There are over sixty species in the group which

I have designated as ‘compact and ponderous’ for want of a

better name. They are essentially a distinct group from the

other Goniobases, and all the species except three are peculiar

to the branches of the Alabama River." |

Consequently, we see that this family reaches its maximum

development in two distinct river systems, the upper Tennessee

and the Coosa-Alabama, which formerly formed one river sys-

tem (Fig. 5), and that by the diversion westward of the upper

Tennessee, by the migration of its divide, this fauna has been

split into two parts, each part having several genera, as well as

species, in common. This process has led to an isolation of

the Coosa-Alabama fauna and a mixing of the waters of the

upper Tennessee with those of the Mississippi valley. In

faunal studies some of the most striking peculiarities of the

Tennessee River are very apt to be overlooked if we consider

it a part of the Mississippi system only. - %

That these two systems have been separated for a consider-

able period of time, or that these forms have evolved very

rapidly, is shown by the fact that it is the genera or groups of

species which are found common to both streams rather than

the individual species.

The family Pleuroceridz is not the only group which has

divided by the diversion of the waters of the Tennessee. The

family Viviparidae has a geographical distribution very similar

to.that of the Pleuroceridz, its greatest variety being found In

southeastern United States. The genus Tulotoma is confin

No. 418.] BASELEVELING. ' 849

to the upper part of the Coosa River, and Lioplax has two

species, one from the Coosa and the other from the Mississippi

valley drainage. The species Campeloma ponderosa is found in

both river systems.

An interesting illustration comes from crayfish. Cambarus

spinosus, extraneous, and erichsonianus are found in both the

Tennessee River and in the head-water parts of the Coosa-

Alabama system.

A head-water fauna is very different from a lowland fauna;

thus a knowledge of the habitats of animals will furnish criteria

by which we are able to recognize, to some degree, the condi-

tions under which divides have migrated. Since valleys con-

tinually lead their rapid-water fauna to the divides, these forms

are the first which become transferred when they are shifted.

What evidence is there to support this view? This is well

shown if we consider those forms which are the firs? to sur-

mount mountain barriers.

The Rocky Mountains have been a very formidable barrier

to our fauna, there being but few passes or gaps through them.

The fish fauna on each side of these mountains in the United

States are remarkably distinct, but two species, according to

Jordan (96, p. 118), are found on both sides of this great bar-

rier, and these species are very naturally rapid-water species,

and not lowland forms. These two fish are the Rocky Moun-

tain trout (Salmo mykiss Walb. and the Rocky Mountain

whitefish (Coregonus williamsoni Gir.). In the Appalachian

Mountains the upland living brook trout (Sa/velinus fontinalis

Mitch.) illustrates the same point, it being found on both sides

of the range.

Extension of Lowland Faunas into Uplands by Baseleveling.

River valleys are well known as highways for the dispersal

of animals. These valleys, of. course, are not always due to

the process of baseleveling, but in the southeastern Appala-

chians we have good illustrations of the extension of a valley

due to this process. The valley of the Tennessee River above

Chattanooga is an excellent illustration of this. By means of

850 THE AMERICAN NATURALIST. [VoL. XXXV.

a narrow neck of lowland being carried back into the uplands

an isolated, elongated land habitat is developed which reminds

one of river isolation. Merriam’s Life Zone Map (98, plate)

shows that there is an extension of the Upper Austral Zone

into the Transition Zone due to the Tennessee valley. With

the baseleveling process the isotherms gradually migrate up

the valley and thus help to make it possible for the fauna to

follow. Of course valleys, of whatever origin, may act as

highways of dispersal.

It is important in this connection to recall the more or less

definite succession of forms up the valley, because the problem

of dispersal among some animals is largely a problem of the

continuous distribution of the habitat.

Segregation of the Upland Faunas by Valleys.

Correlated with the extension of the valleys into uplands is

the segregation of the uplands by the growth of the valley.

This influence is beautifully shown by the land shell fauna

on each side of the Tennessee valley. Since the Tennessee

valley divides the uplands of the Cumberland plateau from

those of the Smoky Mountains, I was led to anticipate that

this isolation, which has existed for a considerable period of

time, would show upon the land shell fauna, because these

animals are very sensitive indicators of such influences. Later

I was interested to learn that Pilsbry (00) had very recently

divided the «Cumberland sub-region” of Binney into an east-

_ern and western division, corresponding to the uplands segre

gated by the Tennessee valley. He gives about twenty

species and varieties characteristic of the Cumberland plateau,

and about twenty-five to the Smoky Mountains. The French

Broad River has for a long time separated the Great Smokies

from Roan Mountain, and corresponding with these facts it has

been found that six forms are peculiar to Roan Mountain and

about a dozen to the Smoky Mountains.

No. 418.] BASELEVELING. 851

Influence of Baseleveling upon Tension Lines.

The relation of the faunas of the different levels or base-

levels to each other is also important from the standpoint of

tension zones. On account of the large area which they may

occupy at certain stages of topographic development these

zones have great importance, for in them there is a large

population subject to a peculiar environment and hence liable

to important evolutionary changes. A stage of equilibrium is

not reached here, as erosion makes this area relatively unstable

and subject to continual changes. It is important to determine

which is the more powerful, the physiographic or organic

influences, in the production of these lines. The. powerful

influence of the physiographic processes in general leads one

to expect that at first the physiographic factors will dominate,

but later, in a fairly uniform habitat, the struggle will become

more organic. In the study of dispersal and distribution of

animals, it is important to see that the physical conditions

lead, and that in a more or less definite succession the flora

and fauna follow ; thus the fauna comes to fit the habitat as a

flexible material does a mold.

The time is passed when faunal lists should be the aim of

faunal studies. The study must not only be comparative, but

genetic, and much stress must be laid on the study of the habi-

tat, not in a static, rigid sense, but as a fluctuating or periodi-

cal medium. The bearing of faunal studies upon the problem

of differentiation and the origin of species, is very close, and

in our search for the factors we must not lose the perspective,

and overlook those factors which are fundamental and work

through long periods of time.

HULL ZoéLocicaL LABORATORY,

UNIVERSITY OF CHICAGO.

THE AMERICAN NATURALIST.

REFERENCES.

CHAMBERLIN, T. C. A Systematic Source of Evolution of Provin-

cial Faunas. /ourn. of Geol. Vol. vi, pp. 597-608.

Hayes, C. W. The Southern Appalachians. Nat. Geograph.

Monogr. Vol. i, No. to.

Hayes, C. W. Physiography of the Chattanooga District in Ten-

nessee, Georgia, and Alabama. Nineteenth Ann. Rep. U. S. Geol.

Survey, 1897-98. Pt. ii, pp. 1-58, Pls. I-V.

Hayes, C. W., and CAMPBELL, M. R. Geomorphology of the South-

ern Appalachians. Nat. Geograph. Mag. Vol. vi, pp. 63-126,

Pls. IV-VI.

JorpAN, D. S. Science Sketches. New edition. Chicago.

MERRIAM, C. H. Life Zones and Crop Zones of the United States.

Bull. No. 10, Division of Biol. Survey, U. S. Dept. of Agriculture.

Pitspry, H. A. Mollusca of the Great Smoky Mountains. Proc.

Acad. Nat. Sci. of Philadelphia (1900), pp. 110-150.

Simpson, C. T. On the evidence of the Unionide regarding the

Former Courses of the Tennessee and the Other Southern Rivers.

Science. Vol. xii, N.S., pp. 133-136.

Tryon, G. W. Land and Fresh-Water Shells of North America.

Pt. iv, Strepomatide. Smithsonian Miscellaneous Collections,

0. 253. Washington.

WOODWORTH, J. B. The Relation between Baseleveling and Organic

Evolution. Amer. Geologist. Vol. xiv, pp. 209-235.

REFERENCES ON THE BASELEVELING PROCESSES.

GILBERT, G. K. Report on the Geology of the Henry Mountains.

U. S. Geog. and Geol. Survey, Rocky Mountain Region. Chap-

ter V. Land Sculpture.

SALISBURY, R. D., and Atwoop, W. W. The Geography of the

Region about Devil’s Lake and the Dalles of the Wisconsin, with

Some Notes on its Surface Geology. Wis. Geol. and Nat. Hist.

Survey, Bull. No. V. Educational Series, No. 1. Chapter III.

General Outline of Rain and River Erosion.

NOTES AND LITERATURE.

ANTHROPOLOGY.

The Seri Indians.|— It is especially fortunate that Seriland

should have been explored by an ethnologist eminently fitted to

describe the physiographic features of that little-known region.

Professor McGee's memoir upon * The Seri Indians" is furnished

with a new topographic map, a detailed description of the country

and of its fauna and flora, as well as a succinct account of the arts

and social institutions of the natives.

After presenting a comprehensive summary of Seri history, the

author deals with the somatic characters of the people. The Seri

differ from other Amerinds in their physiological and physical char-

acters as strikingly as in their demotic. They are remarkable espe-

cially for their fleetness of foot to an extent that we should regard as.

quite incredible were it made known to us merely by travelers' tales.

The description of Seri demotic characters more nearly approaches

completeness in the chapters devoted to Symbolism and Decoration,

Industries and Industrial Products, and Social Organization. Per-

sonal decoration is confined almost entirely to facial painting, and

Pen 1S * feminine prerogative. In discussing the “ significance of

M " Professor McGee outlines a scheme of progressive

ibn et from What he terms automacy to autonomy that is

"i ut interesting. Industrial development is surprisingly

among the Seri. The “industrial use of stone is fortuitous

Cnr mii The author offers a new classification of primitive

tolithic ased upon his observations among the Seri. The pro-

Pu EA that in which the stones are not shaped to conform

fractured eis vem pauer; the technolithic, wherein the stones are

"c o ke be obigen shaped. Seri marriage customs are instruc-

BRA ui e of social organization ; they exhibit an almost

Dance of the tr; E and show a deep-seated interest in the mainte-

because ofthe a autonomy. The Seri are polygenous, apparently

reduction in the number of warriors in recent years.

1 Sevente

enth Annual Report of th ican Ethnology, 1 :

Washington, Dc ‘port of the Bureau of American E Y, 895-96.

» 1898 (distributed 1901). 344 pp. map and plates.

853.

854 THE AMERICAN NATURALIST. [ VoL. XXXV.

In addition to the store of geographic knowledge this memoir is

a positive contribution to ethnologic science of the highest value.

It establishes a new linguistic stock (Serian), it affords illustra-

tions of priscan phases of culture of extreme rarity, and the author

advances many suggestions of theoretic interest.

FRANK RUSSELL.

ZOOLOGY.

Schmeil’s Zoélogy.!— This work was originally prepared for stu-

dents in German Gymnasia and “Realschulen,” and has as its

primary object a disciplinary use of the facts of zoology. It tells

certain facts, it leads the student to infer other facts and reasons by

numerous and carefully worded questions. It differs from the old-

time zoólogies, which merely gave a description of selected animals,

by some very important features. It takes representative forms and

describes them from an cecological standpoint; showing how they

are adapted to their environment, and how they are fitted to a

certain kind of life. Then follow shorter descriptions of allied

forms. There is a minimum of structural details throughout, but

the biological side, so interesting to young students, is everywhere

emphasized. We would advise that all secondary schools teaching

zoology have a copy of this work in the reference library ; its price

and its unnecessarily large size forbid its use with us as a class-room

text. : E

Herrick's Home Life of Wild Birds. — The subtitle of Professor

Herrick's book? if it is understood that only the home life of birds

is to be studied and photographed by the new method, will serve as

a guide to the nature of this very valuable addition to the list of

books dealing with bird life. Professor Herrick has, by the help of

the strong parental instinct in birds, overcome the difficulties with

which the photography of birds in the wild state has hitherto been

attended. Instead of trusting to mechanical devices for arranging

ASchmeil, Otto. Zext-Book of Zoilogy, treated from a biological standpoint,

translated from the German by Rudolf Rosenstock, M.A., edited by J. F.

Cunningham, M.A. London, Adam and Charles Black, 1901. xvi + 493 PP-

2 Herrick, Francis Hobart. The Home Life of Wild Birds. A New Method

of the Study and Photography of Birds. With 141 original illustrations from

are by the author. New York and. London, G. P. Putnam's Sons, kii

149 pp.

No. 418.] NOTES AND LITERATURE. 855

the camera in the branches of trees, and taking a chance shot or

two when the bird shyly returns to the nest, he cuts off the limb or

twigs on which a nest containing young birds is fixed, and removes

it to a favorable situation near by. Then, concealing himself and

his camera in a small green tent within a few feet of the nest, he

waits for the parents to adjust themselves to the new conditions.

This they do in a surprisingly short time. In a few hours the old

site is forgotten, and the birds are as firmly attached to the new one

as if they had themselves chosen it. The operations of feeding,

cleaning, brooding, and other incidental actions of old and young

may now be observed and registered with a completeness and

clearness impossible under the old method.

Professor Herrick's book contains introductory chapters dealing

with the instincts and habits of birds and the psychological princi-

ples involved in the successful practice of the new method, followed

by other chapters explaining and illustrating the method as employed

by Professor Herrick. Seven of these are devoted to a detailed

account of the home life of a number of our common birds as

observed at close range. The text is supplemented by a series of

photographs, which surpass for clearness and scientific value, as well

as for popular interest, anything of the kind heretofore published.

The birds thus treated belong for the most part to the passerine

order; the night hawk and the kingfisher are the only exceptions.

The very fact that in the study of such familiar birds as the robin,

catbird, and cedar bird, we are offered so much interesting observa-

tion, much of it original in its accuracy of detail, gives a good idea

of the value of the method, and promises rich results when it is

applied to less familiar birds |

The concluding chapters of the book deal with general questions,

Pig as the force of habit, fear, etc. There is an excellent index;

3t E typogr aphy, and the reproduction of the photographs are

lie ste quality. The excellence of the book in such matters is.

rdc nce that it is intended not as a scientific treatise, but as

eiii r a iai and if it is reviewed as such, it should be

AE iin y the highest praise. There is, however, much reference

er iMm and the three last chapters to the questions of instinct

abit, and they are treated in several paragraphs in a purely

oig qas spirit. The author, moreover, puts in at the outset a

Ar against what he justly terms the “gross anthropo-

tin rus characterizes much of what is now written op

T first reading of Professor Herrick's book, however, will

856 THE AMERICAN NATURALLIST. [Vor. XXXV.

leave the ordinary reader with no very clear notion of the author's

ideas on the subjects of instinct, habit, fear, etc. His conclusions,

though in accord with such an excellent authority as Principal

Morgan, are by no means clearly presented. The reader follows on

one page a rather discursive account of an encounter with a black

snake, and, turning the page, finds himself confronted with a difficult

paragraph on the instinct of fear. The anecdotes which are to elu-

cidate the principles laid down, or to be laid down, overweigh the

reasoning, and often have no especial bearing. More systematic

arrangement of ideas, an occasional paragraph recapitulating con-

clusions, and an introductory line here and there to show the bearing

of the coming anecdote would greatly increase the value of the semi-

scientific chapters. It might have been better, moreover, to avoid

the dogmatic attitude of the paragraphs on the nest-building instinct

on page xvi, and make reference at least to the possibility of tradi-

tion playing a part in handing on the art of nest building. The

nests built by chaffinches turned loose in New Zealand (A. R. Wal-

lace, Darwinism, p. 76) may be adduced as evidence that a bird like

the robin perhaps does not make mud nests **as instinctively as it

lays blue eggs."

Of undoubted scientific value are Professor Herrick's observations

on the food offered to the young, on the way the food is carried to

them, and on the sanitation of the nest. The sensitiveness of the

gullet of young birds and the parent's habit of taking out food not

immediately swallowed and transferring it to another gullet, throws

light on what has been hitherto an obscure process. It is doubtful

whether the home life of other passerine birds will vary much from

the types observed by Professor Herrick, but there is, as he suggests,

a great unexplored field in the orders.

It is as a popular but unusually accurate and thoughtful presem

tation of a subject at present much in vogue that the book has -

special claim to recognition. Professor Herrick's patience, ingenuity,

and quick power of observation deserve the success with whi the

book is sure to meet. The wealth of extraordinarily good illustra"

tions, the intimate relations with birds into which the reader is

brought, will fascinate any one who has any interest in the study

of living animals. If the author's style lacks the inspiration which

a more imaginative temperament might give to it, his scientific habit

of mind, joined to his evident enthusiasm, make ‘him an excellent

guide and model for those who may begin observation by his me"

The method is admirably suited for instruction in summer schools,

No. 418.] NOTES AND LITERATURE. 857

and has been already adopted in at least one outdoor school of

natural history.

The objections to the method arising from the possibility of

danger to the young thus removed from the site selected by the

parent are carefully discussed by Professor Herrick. It is well,

however, to repeat his warning against interfering lightly in the home

life of wild birds. In the opinion of the present writer, none but

trained naturalists should use the method, for they, if animated by

Professor Herrick's genuine love for the individual bird, will be on

their guard against the dangers likely to be incurred. RA

The Fishes of Ohio is the title of a paper by Professor Raymond

C. Osburn, published as a Bulletin of the Ohio State University

(Ser. 5, No. 20), The paper is a descriptive faunal list of the fishes

known to occur within the borders of the state. In the Introduction

is ES an historical sketch of ichthyological investigation of Ohio

shes.

Each species is briefly, though sufficiently, described ; following

each description is a list of localities in the state where the species

is known to occur; there is also given a few notes concerning its

habits, etc. Keys to facilitate identifications are also given. The

paper is neatly and carefully gotten up and indicates very careful

and thorough work.

The publication of descriptive faunal lists like the present one, by

institutions which have the facilities for such work, is to be highly

commended.

In s footnote Notropis fretensis (Cope) is regarded by Mr. Osburn

m being allied to Notropis heterodon Cope and Notropis cayuga Meek,

Pons shared by Drs. Jordan and Evermann. It is, however, a

i vetet itg (Cope), differing from the typical rubrifrons in

Gu eight anal rays. In his original description Professor

[UR s E to the fact that this species resembled Minnilus.

ele rifrons usually has ten anal rays. It is not, however,

The ty EM some specimens with nine or even eight anal rays.

Stein. pe of Notropis rubrifrons is in the Philadelphia Academy of

5, where I had the pleasure of examining it a few years ago.

S. E. MEEK.

__ The Otocysts of Decapod Crustaceans. — An exhaustive study of

Peat |

Structure, development, and function of the otocysts of decapods

858 THE AMERICAN NATURALIST. (Vor. XXXV.

has been made by C. W. Prentiss.' In Palzemonetes, as in most

other macrurans, the otocyst is a sac lodged in the basal segment

of the antennule and opening dorsally by a constricted aperture

partly covered by a scale-like fold. The sac is lined with cuticula

which at the aperture is continuous with the animal's external shell.

On an elevation rising from the floor of the sac is a horseshoe-

shaped double row of from forty-five to fifty-eight hairs. The hairs

are plumed and instead of being straight, as the tactile hairs of the

outer surface are, they are bent so that the distal part of each shaft

makes an angle of 120 degrees with its shorter base. Each hair is

attached to the sac by a thin-walled chitinous bulb, thus allowing the

hair as a whole to move freely. In the tangle formed by the cross-

ing of the hairs are lodged fine grains of sand and organic detritus

constituting an otolith. Every hair has at its base a group of matrix

cells by which its chitinous wall was secreted. A single nerve fibre

leads from the base of the hair inward to the brain, where it termi-

nates in many fine branches. Each fibre has on its course a single

cell body, so that each “auditory” hair is innervated by a single

neurone. The same is true of the tactile hairs of the general surface,

but the olfactory hairs, on the contrary, are innervated each by as

many as a hundred neurones. Not only do olfactory and tactile

hairs differ in this respect, but they can also be distinguished by the

fact that in the olfactory hair the nerve fibres pass far out throug

the axis of the hair towards its tip, but in the tactile hair the single

fibre ends at the base of the hair. An otocyst essentially similar to

that in Palamonetes was found in Crangon and in Cambarus.

In the common green crab, Carcinus, the otocyst is closed, contains

no otolith, and its hairs are arranged in three groups instead of one.

The innervation of these hairs is the same as in the macrurans

studied.

Every time a shrimp or crab casts its shell, the cuticular lining of

the otocyst, the attached hairs, and the otolith, if such be present,

are discharged. As a preparatory step to this change, the matrix

cells form a new hair under each old one, the new hair being half

inverted in that the tip is pushed back into the base as the end of

a finger of a glove might be infolded into the rest of the finger.

When the skin is shed the new hairs are in part drawn out by the

retreating skin to which they are slightly attached and in part

1 Prentiss, C. W. The Otocyst of Decapod Crustacea: its Structure, Devel

opment, and Function, Bull. Mus. Comp. Zoil., vol. xxxvi (1901), pp. 167-75"

No. 418.] NOTES AND LITERATURE. 859

expanded by blood pressure. The discharge of the contents of the

otocyst is through the natural aperture of the cyst, which remains

open in macrurans but rapidly closes in brachyurans. Where an

otolith is discharged, as in the shrimp and other macrurans, the ani-

mal immediately after ecdysis gathers in its claws small sand grains

and puts them in the opening of the otocyst. These become attached

to one another and to the auditory hairs by secretions from the walls

of the otocyst and thus form a new otolith.

_ The otocyst was originally described as an organ of hearing.

When sounds are produced under the water of an aquarium in which

shrimps are contained, the animals respond by a darting movement

if near the source of sound. The vibrations which stimulate the

shrimps, however, can be /z// by the submerged hand at a distance

of ten to twenty centimeters greater than that at which the shrimps

react. Moreover, shrimps respond to these vibrations even after the

otocysts are removed. ‘The reactions are inhibited, however, by the

removal of the antenne and antennules with their tactile hairs. It

follows from these observations that whether we call the reactions

auditory or tactile, the otocysts take little or no part in producing

em.

If, then, the otocysts are not stimulated by sound, what is their

function? When the otocysts are removed, shrimps swim with a

more or less rolling motion and may even turn ventral side up.

Their equilibration is thus shown to be seriously interfered with.

When their eyes are covered with opaque materials so as to blind

ves they swim with little or no rolling motion; but when both

poo deprived of otocysts, they move with the greatest irregu-

c uos sometimes on their backs and sometimes in irregular

em ii Pe eir capacity for orientation has disappeared completely,

? experiments show, though the eye plays some part in

keeping the animals upright, the otocyst is the chief sense organ in

tation reflexes. The otocyst is stimulated through

Mi Ri the otolith on the sensory hairs, as is shown by the

cgi So OR of Kreidl, who induced shrimp to form otoliths

€s instead of sand grains and then found that the

ani .

n became oriented to the lines of force of a magnet as they

fo st y did to gravity. These results are confirmed by Prentiss.

in the um ment of the otocyst is also dealt with. It is not present

the second rval stage of the lobster nor is it more than indicated in

almost the ie third, but at the fourth stage it suddenly appears with

860 THE AMERICAN NATURALIST. (VoL. XXXV.

the animals shows a corresponding change ; young lobsters swim with

great irregularity until the otocyst is developed, after which they

regularly assume the upright position. It is also interesting to

observe that immature lobsters newly molted and not allowed to

form a new otolith reassume the rolling movements of their earlier

stages and continue in these until opportunity is given them to form

new otoliths. Thus the results obtained by Prentiss confirm com-

pletely the view first advanced by Delage, namely, that the otocyst

has as its chief function that of originating orientation reflexes.

That it is not an organ of hearing cannot with so much certainty be

maintained, although its importance in this respect has assuredly

been shown to be very slight. P.

Recent Work on Anopheles. — In a recent study on the geograph-

ical distribution of Anopheles in relation to the former distribution

of ague in England, G. H. F. Nuttall records with L. Cobbett and

G. Strangeways (Journal of. Hygiene, Vol. I, January, 1901) a series

of observations of general interest in addition to much that is purely

medical. He finds that three species, Anopheles maculipennis, A. bifur-

catus, and A. nigripes, occur in Great Britain in all districts formerly

malarious, but extend into regions in which no ague is known to

have been prevalent at any time. To-day Anopheles is most numer-

ous in low-lying land containing stagnant or slow-flowing water and

corresponding to the districts where ague was formerly prevalent. As

the disappearance of this disease does not depend upon the extinc-

tion of Anopheles, possible causes, therefore, are: (2) a reduction in

the number of these insects consequent upon drainage of the land;

(6) a reduction of the population in infected districts by emigration ;

intermediary host besides man capable of harboring the parasite.

The coincidence of the geographical distribution of ague and Ano

pheles is certainly not as precise as claimed by Grassi, and probably

the numerical distribution will prove of equal importance. The

presence of Anopheles in non-malarious districts explains the occa-

sional occurrence of ague if a malarious subject comes in from other

parts.

In another paper on the structure and biology of Anopheles,

Nuttall and Shipley (Journal of Hygiene, Vol. I, January, 1901) 8°

a full summary of our knowledge, together with personal observe

tions. Noteworthy is the fact that the larva of Anopheles, as

that of Dixa, browse upon matter adhering to the surface film, like

No. 418.] NOTES AND LITERATURE. 861

certain fresh-water snails and many Turbellaria. Palmate hairs on

the third to the seventh abdominal segments, with branches in the form

of a cup, constitute the means by which the larva cling to the surface

flm. The studies are to be continued, and from their precise and

detailed character bid fair to become a most valuable source of

information regarding the genus. H. B. W.

The Insect Book. — The complete title! indicates the effort of

the author and of the publishers. The former is widely known for

many valuable monographs, both systematic and biological, and for

energetic and successful practical work ; the publishers are equally.

well known for their enterprise in numerous undertakings tending

towards the promotion of nature study. Such a combination should,

it would seem, be eminently satisfactory. That it is not so is disap-

pointing ; that the title should be modified is evident. The Insect

Book should not exclude the groups to which the “ majority of col-

lectors of insects confine their attention”; moreover, the standard

must be low that considers the life histories “full”; the tables are

confined to families or higher groups, and the bibliography, pp. 405-

416, copied with but few additions from Banks’s List (Bulletin 24,

Division of Entomology, U. S. Department of Agriculture, 1900),

however useful from a taxonomic standpoint, is not of primary impor-

tance in a book planned “to encourage the study of life histories of

insects"; moreover, the very evident deficiencies of Banks’s work

are unnoticed and the mistakes not only are uncorrected but are

augmented.

The Text is readable, though not altogether well balanced in pro-

portion; it shows evidence of haste, of a lack of continuity, and

also of the most important essentials for a popular book, namely

dau v conciseness. Dr. Howard evidently does not believe

we k ie should be left for the tombstones, but his extravagant

adi several contemporary authors can but jar many of his

ak errors of statement are amusing, others are serious :

oral) rimi may be placed the classification (p. 295) of the

w as one of the “older books " relating to insects ;

"^ai dre £n 1 " popular account of the bees, wasps, ants, grassh pers;

and beetles, with £u SAEPE: Insects, exclusive ot the butterflies and moths

Howard, "t uo an uu py tables, and Bibliographies. By Leland O.

Bibicilture. vision of Entomology, U. S. Department of

265 text EL Doubleday, Page & Co., 1901. xxvii + 429 pp., 47 pls.,

862 THE AMERICAN NATURALIST. [Vor. XXXV.

among those of a more serious nature may be mentioned the repro-

duction (pp. 343, 344) of Scudder's original notation attributing a

day song and a night song to Oecanthus niveus. ‘These supposed

“day” and * night" songs have long been known to be the songs

of distinct species, and so far back as 1893 Scudder recognized his

error and stated that a revision of his score was desirable.

The plates are all original; most of the text-figures have been

previously used and the source is acknowledged, though not in all

cases satisfactorily.

Very few of the illustrations, either in the plates or in the text, can

be commended; this can be clearly seen by comparing the colored

plates with those of Holland's Butterfly Book, and the text-figures

with those from the same blocks as used in various publications of

the United States Department of Agriculture and elsewhere; in the

make-up of the book the plates are poorly placed.

Inconsistencies in typography and lack of ordinary care in proof-

reading are so apparent that even an inexpert corrector of the press

could not turn many pages without finding glaring errors. — S, H.

Michaelsen's Oligocheta.!— When Dr. Michaelsen began his

work he had before him no easy task, and it is safe to say that no

student of this class of animals could have discharged his duty in 4

more satisfactory manner. The museum in Hamburg is in possession

of the largest, and as regards species most complete, collection of

Oligochzta, and when we remember that it has been brought

together entirely by Dr. Michaelsen, it may be readily understood

that the author has been eminently fitted for his work. The litera-

ture of this class of animals is already very large, and it is also so

scattered that it is impossible for any one living outside of the large

literary and scientific centers of Europe to procure or have access to

all. Dr. Michaelsen has had access to all this literature, and has

personally examined almost every species described by different

_ authors, as far as they yet exist, and the result has been a work

the very highest value, and one which no student of the group Ca^

do without.

The first part of the book contains a list of abbreviations for

reference to literature. It seems to the reviewer that it would have

been better if such abbreviations had been adopted as are used T

the publications abbreviated. This suggestion refers equally t9 .

| Michaelsen, W. Das. ZYerreich. 10. Lieferung, Vermes, Oligocheta. + "i

Friedlaender, 1900. xxix + 575 pp» 13 figs. x

No. 418.] NOTES AND LITERATURE. 863

almost every publication issued, and is a subject worthy of interna-

tional consideration. Next after the abbreviations we come to a

systematic index. When we consider that this index of families

and species occupies not less than sixteen pages in double columns,

we may begin to comprehend the enormous increase in described

species during the last few years. All in all, there are probably

1100 species of Oligocheta now sufficiently well described to be

readily identified. As is well known, the majority of species of this

class can only be identified by anatomical characters. This makes

identification difficult without the aid of illustrations, and it is much

to be regretted that so few are found in this otherwise so valuable

monograph. The morphological part is confined to ten pages, and

it is here that the want of illustrations is especially felt. The author

introduces a number of new words to indicate structures, such as

tanilobic, zygolobic, prolobic, etc., according to the various and

different encroachments of the prostomium on the anterior somite,

etc. Many of these names have been derived from the Greek and

Latin and are most happily found, and we can only regret that the

author has not seen fit to revise also the general nomenclature to

such an extent that students of other languages could readily under-

stand what parts and organs are referred to. Why not use, for

instance, * spermatheca" instead of “ samentasche," etc.? The

larger part of the book is devoted to a systematic description of the

species, their genera and families. These descriptions are models

of conciseness, including all the principal exterior and interior

Characters necessary to define the species. A large number of

names have been changed, priority being given to the oldest ones,

according to the rules adopted by the German society of naturalists.

In this department the author has made very thoroug

as a consequence we meet with many novelties which will be referred

to in their respective places. Every genus is preceded by a key

for identifying the species. Many will object to the style of key

opted. In our opinion the most desirable key in all: systeme

works is the one which, besides facilitating the finding Mite

Species, at the same time gives us an idea of the systematic arrange

ment and relationships. The system adopted by the author gives E

idea of the affinity of the species, but simply facilitates finding tn

In adopting the present style of key the yste a" né

genera and species certainly suffers. This is especially the aa

large genera, as, for instance, Dichogaster (Benham, pun

the species number over 150. oi n

864 THE AMERICAN NATURALIST. [ VoL. XXXV.

The treatment of the limicolide groups should prove especially.

interesting to the American students, as the scattered literature of

these groups is especially difficult of access, as much of it is in Slavic

languages. We are glad to see that the genus Ilyodrilus has been

restricted to its three California species and separated from Bran-

chiura, with which genus it has really no affinity. The treatment of

Enchytrzidz is especially good, the author having previously studied

this family, and being the first one to bring order out of the chaos

which existed previous to his monograph on that family. Among

innovations we remark the genus Lumbricillus substituted for

Pachydrilus. It is to be regretted that this change should have

been necessary, as we have already too many names with a similar

sound. The family which probably interests us most is the large

one of Megascolecide. This family is made to contain the majority

of North American terrestrial earthworms. The most interesting

fact connected with the family as defined by the author is that it is

made to contain Ocnerodrilus as well as Diplocardia, Trigaster,

Plutellus, Chilota, etc. The genus Achantodrilus, which a year ago

contained more species than almost any other genus, is now restricted

to a single species. The Old-World species of the once large genus

have now been separated from the New Zealand ones, while the

majority of the species have been referred to the genus Notiodrilus,

etc. It is most interesting to note that as a result of the author's

investigations we now find Achantodrilus placed next to Microscolex

and its allied genera, while a few years ago these were referred te

different families. The genus Plutellus has been resurrected, as 1$

quite proper, but what will astonish the general student more is that

Plutellus and Pheretima have been referred to the same family, —

the reason being that we now place very little importance on the

number of seta in each somite, as long as the interior organs resem-

ble each other. ‘The author has had special opportunity to examine

Kinberg’s types in the Stockholm Museum. As a result of this

investigation it has been possible to identify nearly all of Kinberg's

genera and most of his species, and consequently many later names

have to give way to the old Kinberg names, — Pheretima instead of

Perichzta, etc. The subfamily Diplocardinz has been well treated,

and so has its near relative Trigastrine. The Zapotecia has been

raised to a full genus, it previously having been considered a sub-

genus only. Dichogaster is made to contain both Benhamia and

Dichogaster, which certainly is an easy way out of the difficulty in

defining the relationship of these two genera. But even the author's

No. 418.] NOTES AND LITERATURE. 865

lately established genus Balanta has had to give way to the all-

engulfing Dichogaster. A division into subgenera retaining such

names as Benhamia, Balanta, and Dichogaster would probably have

been more in accordance to the ideas of several other investigators,

and would have served to give a better review of this genus — we

confess one of the most puzzling in the class. Under Ocnerodrilinz

we now meet with the genus Kerria, the structure of which is the

one that gave the clue to the relationship of these genera with

one and two pairs of prostates. Several of the subgenera under

Ocnerodrilus have been raised to independent genera. We have

thus Nematogenia, Pymeodrilus, and Ocnerodrilus side by side.

Nematogenia panamensis has been raised to an independent species.

Among the subgenera Enicmodrilus has been fused with Ilyogenia.

It would have been an improvement if the species followed each

other in the general text in the same manner as in the key, or vice

versa: this is rarely the case. The name Geoscolecide has been

changed to Glossoscolecidz, the latter being an older name.

The last family, Lumbricide, has more of a negative interest to

students in this country, as comparatively few indigenous species are

found here. While it is true that several species of this family have

been described as new from North America, it is doubtful if they

really are indigenous, and we may find that the species so considered

are mere importations. This fact is certain, that no new Lumbricidz

have been found west of the Rocky Mountains. All the species on

the Pacific coast are undoubtedly importations from Europe, and a

very careful search by the reviewer has failed to reveal a single new

species of this family between Alaska and Central America. This

is the more interesting as the Pacific coast is in many respects

related to Japan and eastern Asia. Thus Pillsbury has shown that

the dart-bearing Helices are only found on the Pacific coast of

North America, while they-are widely distributed through the conti-

nents of Europe and Asia. Similarly among the Crustacea um

genus Astacus is found all along the Pacific coast, and in Asia and

Europe, while the genus Cambarus is found only in central sg

eastern North America. Among plants also there is a similarity

between the Pacific coast and Japanese forms. But when we come

to the earthworms the conditions are different. As far as —

not a single species is found common to Japan or Asia and io

Pacific coast. The indigenous California species seem er d

from south of the

equator and to genera which with good reason may pe conem m

866 THE AMERICAN NATURALIST. (VoL, XXXV.

having emanated from a former antarctic continent. The indigenous

terrestrial species of Lumbricidz found in North America should for

the present be looked upon with suspicion in connection with their

true habitat. The family of Lumbricide has received model treat-

ment in Michaelsen's monograph. The book ends with a very per-

fect register, — a desideratum not always found in transatlantic

scientific works. Taking it all in all, this splendid monograph

cannot be too highly praised. It is not only indispensable to all

students of the class, but it is so complete that for the identifica-

tion of the species the older literature is made almost superfluous.

As regards nomenclature, Dr. Michaelsen's ideas are almost final ;

while as regards classification they are sure to remain unchanged

for many years. A perusal of the volume shows one thing which

should be encouraging to all students of earthworms; that is, com-

pared to the terrestrial Oligochzta the limicolide forms are almost

unknown, and it is evident that it is among the latter that the

reatest novelti vi i :

g s ties will be found in the future Gustav EISE.

BOTANY.

A Botany for Children. — Professor Atkinson in an attractive

little book entitled Hirst Studies im Plant Life! follows the current

practice of presenting vital phenomena to the young beginner, rather

than details of form. The first part gives a brief account of the

growth and parts of plants, the second and third discuss their work

and behavior, the fourth gives the life story of a sweet pea, an 02

a-férn,a moss, and a mushroom, and the fifth treats of the battles of

plants in.the world. The illustrations are of exceptional excellence.

Many ingenious and simple experiments are introduced. Much 0

the text seems likely to interest. young peóple, and any teacher may

gain valuable suggestions from the book: e

_ A prévailing-fault, however, isxa more or less obvious “writing

down ” to the young reader, becoming at times mere sentimentality

such as intelligent children resent. - It is hard to see how the follow:

ing passage, for example, can serve any useful purpose: “ ique

1 Atkinson, George Francis, Ph.B., Professor of Botany, Cornell University.

First Studies of Plant Life. Boston, Ginn & Company, 1901. 12mo. xii + 266 it

308 figs. : i

No. 418.] NOTES AND LITERATURE. 867

tell you that such interesting plants as the ferns, mosses, mush-

rooms, and puffballs are cryplogams, and that therefore you should

not try to read the stories they have to tell. But why call them

cryptogams? ‘That is a terrible word, that ought to be blotted out

of the English language. Why not call them plants, as they are?

They are just as much God's creatures as the dandelion and thistle

and smartweed are. They are just as interesting too, and mean as

much in our lives as they do."

Furthermore, it may be questioned whether some of the topics to

which considerable space is given—as, for example, turgidity,

plasmolysis, and various microscopic details — are really within the

comprehension of young beginners. Every one who has studied

children knows what confused and perverted ideas they will often

get regarding matters of much greater simplicity: than belongs to

some of the physiological topics here presented. It is hard enough

to give college students clear ideas of microscopic mechanisms and

life processes.

Only a few errors of statement have been noticed, but there is one

which is sure to bewilder the pupil On page 14 it is said of the

bean that the embryo plant (meaning the plumule) is attached to

the cotyledons. Then on page r9 it is asked if this small object

which looks like a tiny plant is the embryo, and the reader is left to

suppose that it is. Finally, on page 22 the reader is told most

impressively that all inside the seed coat and its lining is the

embryo, and that the embryo thus includes the cotyledons.

F. L. S.

A New Publication on Woody Plants. — Houghton, Mifflin & Co.

announce that they will issue next autumn the first part of a new

publication, Trees and Shrubs, consisting of text edited by Professor

Sargent, and plates from drawing by Mr. Faxon, pertaining to woody

plants, particularly those adapted to the gardens of Europe and the

United States, or of commercial or economic importance. The

sample pages and plates that have been distributed with the pro-

spectus show, as would have been expected, excellence in drawing

and publication, and the happy mean between technicality and pop-

ular writing which mark the Si/va is likely to be maintained by

Professor Sargent in this new publication, which in size and general

appearance will bear considerable resemblance to the Si/va. Two

parts, each consisting of twenty-five plates and costing $5.00, are

expected to appear yearly. —

868 THE AMERICAN NATURALIST. |. [Vor. XXXV.

The prospectus of this proffered work, curiously enough, raises a

question that may bother the botanical nomenclaturists, for Pl. II

and the accompanying text depict and describe what is called a new

species in that much-vexed genus Cretzgus, so that when the initial

number of the publication appears it will doubtless be found neces-

sary to refer in it to this prior distribution of the species referred to.

Flowers and Ferns in their Haunts.' — Where to draw the line

between pleasant summer reading and didactic literature is often

hard to decide, and yet when we go to the country there is a limit

to what our trunks will carry, so that some kind of discrimination

becomes necessary. This pretty little book will hardly come amiss,

from whichever point of view selected, though it may not hold the

attention of the reader like a novelette nor suit the needs of a class

in botany; but through its pages runs a chatty narrative that is pleas-

ing, and the illustrations show much that can be done by aid of the

camera when intelligently used. T.

The Dictionary of Gardening.? — As was stated in the Naturalist

for November, 1900, the excellent Dictionary of Gardening of Mr.

George Nicholson, curator of the famous Kew Gardens, which for

years has been the reference book for gardeners wherever English is

read, has had planned for it a supplement, bringing it up to the end

of the century. The first volume of this supplement appeared in

June, 1900, and a second volume, completing it, was distributed in

July of the present year.

No more favorable place than Kew could be found for the elabo-

ration of a compendious work.on cultivated plants and the most "e

cessful ways of growing them. Not far from 25,000 species are said

to be cultivated there. Kew is probably freer than any other estab-

lishment in the world from the common fault of botanic gardens, that

the collections are grown uncritically under whatever names are

attached to them when they are procured; and the very common, if

often necessary, defect of botanical gardening, that a great variety of

plants requiring dissimilar treatment are huddled together into à

! Wright, Mabel Osgood. Flowers and Ferns im their Haunts, with illustra-

tions from photographs by the author and J. Horace McFarland. New York,

The Macmillan Company, 1901. xix + 358 pp

? Nicholson, George. The Century Supplement to the Dictionary of Garden

ing, a practical encyclopedia of horticulture for gardeners and botanists. George

T. King; Hyde Park, Mass.

No. 418.] NOTES AND LITERATURE. 869

single house, where they collectively get the handling that best suits

their average needs while it is not precisely adapted to any one spe-

cies, is so far overcome at Kew that representatives of the great Eng-

lish plant houses sometimes go there for training. Mr. Nicholson is

at once a good gardener and a lover and student of plants, and he

has had the assistance of the best specialists at Kew and elsewhere,

so that the Dictionary, as now completed, is a work alike valuable to

the student of plants and their amateur and professional grower, and

it cannot be spared from the shelves wherever plants are grown in

variety. Ti

Bailey's Botany.!— Carrying out his well-known ideas that botany

in the secondary schools should begin with the commoner and grosser

plants rather than by the use of those demanding the aid of the micro-

scope, Professor Bailey has added another to the series of text-books

already well and favorably known. Observation, experiment, and

thought are thrust at the pupil throughout it, and the author very

neatly acknowledges his obligation to hundreds of young people in

many places for instruction “in the point of view,” for the book, he

tells us, is made for the pupil and, therefore, most appropriately views

things as he sees them, even though it may enlarge his view of them

before they are dropped.

Notes. — Under the guidance of Professors F lahault and Geddes,

the late Robert Smith had begun the preparation of a series of botanical

maps of Scotland, and since his death two sheets, respectively of

Edinburgh and vicinity and northern Perthshire, have been issued in.

convenient pocket form, with a brief descriptive pamphlet, by John

Bartholomew & Co. of Edinburgh.

W. N. Suksdorf has recently distributed excerpts from the Deutsche

botanische Monatsschrift, extending over the period between Novem-

ber, 1898, and June, rgor, in which are published descriptions of a

Considerable species of Washington plants believed by him to be

new to science.

Several new species and varieties of Californian plants are described

by H. M. Hall in the Botanical Gazette for June.

Professor Dudley contributes an interesting and well-illustrated

Paper on the Big Basin Redwood Park to the Forester for July.

y Bailey, L. H. p An elementary text for schools. New York, The

n Company, 19oo. xii + 356 pp., 500 figs.

870 THE AMERICAN NATURALIST. [Vor XXXV.

A second part of Professor Nelson’s “Contributions from the

Rocky Mountain Herbarium," published in the Botanical Gazette for

June, deals with various Colorado species and new western Arnicas

and Eupatoriez.

Four papers on the botany of the Yukon territory, as exemplified

in the collections of R. S. Williams and J. B. Tarleton, are published

in No. 6 of the current second volume of the Buletin of the New York

Botanical Garden. Á

The Botanical Seminar of the University of Nebraska has pub-

lished the results of recent studies on the vegetation of that state.

A catalogue of the flora of Montreal Island, Canada, is being pub-

lished in current numbers of the Bulletin de 1’ Académie Internationale

de Géographie Botanique.

A systematic list of the plants collected by Schmidt on the

Danish expedition to Siam in 1889 and 1900 is being published in

the Botanisk Tidsskrift.

Pilger’s “ Beitrag zur Flora von Mattogrosso ” is concluded in the

second Heft of Bd. XXX of Engler’s Botanische Jahrbücher, which,

like other recent numbers, is also devoted in large part to studies of

African plants.

Stanleya pinnatifida is figured in the Gardeners’ Chronicle of June 15.

Dr. Small reviews the Mimosacez of the southeastern United States

in the Bulletin of the New York Botanical Garden of May 27.

Rosa Engelmanni is figured in the April fascicle of /cones Selecta

Horti Thenensis.

Two species of Epilobium are characterized by Suksdorf in the

West American Scientist for May.

An interesting “Study of the Papaw,” Carica Papaya, by F : B.

Kilmer, has run through several recent numbers of the American

Journal of Pharmacy.

The Canada thistle is the subject of a revised issue of Circular 27

of the Division of Botany of the United States Department of Agricul-

ture, by Mr. Dewey.

Several new Canadian gentians are described by Holm in the

Ottawa Naturalist for July.

In the Bulletin of the Torrey Botanical Club for June, Dr. Small

proposes the generic name Brayodendron for what has been known

as Diospyros Texana, and describes several species in the genera

Quercus, /Esculus, Hypericum, Azalea, and Dendrium.

No. 418.] . NOTES AND LITERATURE. 871

Dr. Rydberg discusses the oaks of the continental divide north of

"Mexico in No. 6 of the current volume of the Bulletin of the New

Vork Botanical Garden.

Vol. I of Kraenzlin's OrcAidacearum Genera et Species, devoted to

the groups Apostasiex, Cypripediez, and Ophrydez, is completed

with the 16th fascicle, the preface of which bears the date May,

1901. The volume, though its titlepage bears the date 19or, has

been in publication since 1897.

‘An important article on the Texan home of Pinus Cubensis, by

Professor Bray, appears in the Forester for June.

The report of the society /s7s of Dresden for 1900 contains Parts I

and II of a paper by Menzel on the gymnosperms of the North-Bohe-

mian lignite formations, illustrated by a number of plates.

To the rather numerous check lists of ferns and fern allies is added

another by B. D. Gilbert, bearing the imprint of L. C. Childs & Son,

‘Utica, N. Y., 1901. Unlike most such lists, this one, which includes

438 numbers (species and varieties) which occur in North America

above the Mexican line, contains a considerable number of critical

notes and descriptions of new species.

A study of the nectar glands of Pteridium aquilinum, by Professor

Lloyd, is published in Science for June 7.

A monograph of North American Sordariacez, by David Griffiths,

constitutes the opening number of Vol. XI of the Memoirs of the

Torrey Botanical Club. It is illustrated by 19 plates and several

figures in the text, and contains a bibliography and index.

Part III of Arthur and Holway’s “ Descriptions of American Uredi-

nez," in No. 2 of the current volume of the Bulletin from the Labora-

tories of Natural History of the State University of Lowa, deals with

‘Tusts of several groups of grasses.

From a short paper by Professor Burt, published in the Bulletin of

the Torrey Botanical Club for May, Tremella mycetophila Pk. appears

teferable to the genus Exobasidium.

Professor Atkinson, in Bulletin 193 of the Cornell University Exper i-

ment Station, gives an account óf several of the fungi which attack

the wood of shade and timber trees, — a subject heretofore investi-

gated in this country by Dudley and von Schrenk.

The Manchester Literary and Philosophical Society, in a recent

number of its Memoirs and Proceedings, prints an interesting lecture

9n the flora of the human body by Dr. Metchnikoff.

872 THE AMERICAN NATURALIST.

A doctor’s thesis by S. L. Schouten, on the methods of securing

pure cultures of micro-organisms from a single cell isolated under the

microscope, has been issued from the University of Utrecht.

The present knowledge of the causation of certain plant diseases

by bacteria is well brought out in a series of controversial articles

by Professor Fischer and Dr. E. F. Smith, in the Centralblatt für

Bakteriologie und Parasitenkunde, extending over the last two or

three years, separates of which have recently been distributed by

Dr. Smith.

Ule shows, in Heft 2 of the current volume of Engler's Botanische

Jahrbücher, that a number of Amazonian Epiphytes seem to owe their

distribution in part to certain ants which take to their nests in trees

seeds of the other plants, some of which vegetate there.

` The initial number of what promises to be an interesting seriés of

articles on the pollination of Chilian flowers is published by Dr.

Jokow in Heft 3-4, Bd. IV, of the Verhandlungen des deutschen

wissenschaftlichen Vereins zu Santiago de Chile. `

Prometheus, of June 12, contains the first of a series. of articles by

.Sajó on plums and other fruits of American origin.

Mr. Pinchot has issued as. Bulletin No, 30 of the Division of fun e

try of the United States Department of Agriculture a very practical

forest-working plan for. a portion of the New York forest preserve,

prepared by Messrs. Hosmer, Bruce, and : Newell

The Imperial Botanic Garden of St. Petersburg. feels the angi

experienced by most active scientific establishments of a medium of

publication under its own control as to time.and other important

conditions, and the. director. of.the garden, Professor A. Fischer de

Waldheim, announces the early appearance of the. first. number of a

Bulletin which the. Garden i is to issue.

Phe. Plant World for: June: contain rtis intanet ehan arti-

cles, among them one on the Cuban uses ofi Oreodoxa, one on Ber-

muda, and some suggestions . as to. the points. to be observed in

collecting . specimens. of Cratzegus. mcn

A portrait and appreciative: dintie sited i Forbes Meehan

appear in the — oe of inn ise kd de

i odor Geyer ew 4

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CHAMBERLAIN, C. J. Methods in Plant Histology. Chicago, University

Chicago Press, 1901. viii, 159 pp., 8vo, 73 figs. — Gasser, H. The Circulation

in the Nervous System, Plüttévills Wis., Journal Publishing Co., 1901. 156 pp.

Siler L. O. The Insect Book: A Popular Account of the

j Wisi Ants, Grasshoppers, Flies, and Other North American Insects

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and Bibliographies. New York, Doubleday, Page & Co. 1901. xxvii, 429 pp»

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1901. xi, 347 pp. 8vo, 14 ills. $3.00. — Verhandlungen der Anato-

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ALLEN, J. A. "The Generic Names Myrmicophaga and Tamandua, and the

Specific Names of the Opossums of the Genus Didelphis. Proc. Biol. Soc. Wash.

Vol. xiv, pp. 91-93. — ATwoop, H., and STEWART, J. H. Poultry Experiments :

Loss of Weight in Eggs pit age ee n. W.Va. Agr. Exp. Sta., Bull. No. 73-

— Baker, F. C. e Digitations of the Mantle of Physa. Bull.

Chicago WA Sc. Vol. ii, a 4, pp- 225-228, 2 pls. — BAKER, F. C. Descrip-

tion of a New Species of Limnea. Bull. Chicago Acad. Sci. Vol. ii, No. 4,

PP. 229, 230, 1 fig. — BECKER, Gro. F. Report on the mrs a the Philippine

Islands. From Twenty-First Ann. Rept. U.S. Geol. Surv. Pt. iii, 139 pp, m

and ills. — BENEDICT, J. E. Four New Symmetrical Hermit Chabe (Pagruids)

from the West Indian Region. Proc. U.S. Nat. Mus. Vol. xxiii, pp. 771-776.

— CERTES, A. Colorabilité élective, “intra vitam" des filamens sporiféres du

Spir ohacillus gigas (Cert.) et de divers microorganismes d'eau douce et d'eau de

mer par certaines couleurs d'analine. Compt. Rend. Assoc. Trans. Avani. Sci.

i New

ls. V i

Known Coccidz. I. Ripersiella and Ceroputo. Proc. Biol. Soc. Wash. Vol. xiv,

PP- 165-167. — CocKERELL, T. D. A. On a Slug of the Genus Veronicella from

Tahiti. Proc. U.S. Nat. Mus. Vol. xxiii, pp. 835, 836. — COCKERELL, T. : À-

Notes on the Food of Birds. N. Mex. Agr. Exp. Sta, Bull. No. 37. Pp. 35-57.

— CoRBETT. Apple Districts of West Virginia. W. Va. Agr. Exp. Sta., Bull.

No. 75. Pp. 83-178, 11 figs. —CorBEtT, L. C. Cold Storage. W. Va. Agr.

Exp. Sta., Bull. No. 74. Pp. 51-80. — COVILLE, F. V. Juncus Columbianus, an

Undescribed Rush from the €— Plains. Proc. Biol. Soc. Wash. Vol. xiv,

PP. 87-89. — Curriz, R. P. Papers from the Harriman Alaska Expedition.

XXII. pu qas Results 1 14): The Odonata. Proc. Wash. Acad. Sci.

Vol. iii, pp. 217-223. — Dorsey, G. A. Archzological Investigations on the

Island of ta Plata, Ecuador. ye Columbian Museum, Anthrop. Ser. Vol. ii,

No. 5 pp. 247-280, Pls. XL-CII. — Dorsey, G. A. and VorH, H. R. The

874 THE AMERICAN NATURALIST. [Vor. XXXV.

Oraibi Loyal Ceremony. Field Columbian Museum, Anthrop. Ser. Vol. iii,

No. 1, pp. 1-59, Pls. I-XXXVII.—ELtiot, D. G. A List of the Land and Sea

Mammals of North America North of Mexico. Supplement to the Synopsis,

Field Columbian Museum, Zool. Ser. Nol. ii, No. 2, pp. 477-522, Pls. L-LVI.—

ELLIOT, D. G. A List of Mammals obtained by Thaddeus Surber in North and

South y een Georgia, and Florida. Field pagos Museum, Zool. Ser.

Vol. iii, No. p. 31-57, Pls. V-X. — ELLIoT, D. G. e Caribou of the Kenai

Peninsula, oaks. Field Columbian Museum, Zool. Sea Vol. iii, No. 5, pp. 59-

62, Pls. XI-XIII.— FISCHER, A. Die Bakterien der Pflanzen; Antwort an

Herrn Dr. Erwin F. Smith. Centralli. Bat. u. Parasitol. Bd. v, pp. 279-287.—

GarciA, F. Orchard Notes. N. Mex. Agr. Exp. Sta., vni No.

gate with a € Fi idm Ann. Rept. N.Y. Zool. Soc. Pp. 77-119,

ls, map.—I1jima, I. Studies om the Hexactinellida. Contribution I:

Soho: e, at is Imp. Univ. Tokyo. Vol. xv, pp. 1-299

Pls. I-XIII, map. — JoHNsTON, J. B. The Brain of per A Contribution

to the Morphology of n Vertebrate Brain. Zool. Jah Bd. xv, Heft 1, 2,

pp. 1-204, Pls. I-XII.— JORDAN, D. and SNYDER, J. a A Review of the

Lancelets, Hagfishes, and eae ave 2 dudes with a Description of Two New

Species. Proc. U.S. Nat. Mus. Vol. xxiii, pp. 725-734, Pl. XXX. — JORDAN,

n d SNYDER, J. O. List of Fishes C Hoc in 1883 and 1885 by Pierre

boas ro and Preserved in the United States National Museum, with Desc

tions of Six New Species. Proc. U.S. Nat. Mus. Vol. xxiii, pp- ath

Pls. XXXI-XXXVIII. — JoRDAN, D. S. idi SNYDER, J. O. Description of

Two New Genera of Fishes (Ereunias and Draciscus) from Japan. Proc. Cal.

Acad. Sci. Ser. 3, Zoöl., vol. ii, pp. 377- ape Pls. XIII, XIX.— Jorpay, D. S.

and Snyper, J. O. "o of Nine New Species of Fishes contained in

Museums in Japan. Journ. College Sci. pel Univ. Tokyo. Vol. xv, pt. !b

1-311, Pls. XVI, con —Jorpan, D. S., and Starks, E. C. On the

PP.

ET S of the Lutianoid Fish Aphareus E Proc. U.S. Nat. Mus.

Vol. xxiii, pp. 719-723, Pls. XXVIII, XXIX. — Jorpan, D. S., and STARKS, E-

C. Description of Three New Species of Fishes from Japan. roc. Cal. Acad.

Sei. Ser. 3, Zoól, vol. ii, pp. 381—386, Pls. XX, XXI. — KUWANA, S. S. The

San José Scale in Japan. Contrib. Hopkins Seaside Lab. No. xxv, 13 pp ! 1 fig.

— Mearns, E. A. A New Pocketmouse from Southern California. oc. Biol.

oc. Wash. Vol. xiv, pp. 135, decree on E. A. The American Jaguars.

Proc. Biol. Soc. Wash. Vol. xiv, pp. 137-143, — MEARNS, E. A. Description of

a New Ocelot xia Texas and Northeastern Mexico. Proc. Biol. Soc. Wash.

Vol. xiv, pp. 145-148. — MEaRNs, E. A. Two New Cats of the Eyra Group from

North America. Peay Biol. Soc. Wash, Vol. xiv, pp. 149-151. — MEARNS, E. A.

T the Mainland Forms of the Eastern Deermouse, Peromyscus leucopus (Rafi).

- Biol. Soc. Wash. Vol. iv, pp. 153-155, — MiRIHA We G: ls Syno

cad. Sci. Vol. iii, pp. 273-295.— MERRIAM, C. H. Six New Mammals pw

Cozumel Island, Yucatan. Proc. Biol. Soc. Wash. Vol. xiv, pP- 99-104 7

denm CN New Brocket from Yucatan. Proc. Biol. Soc. Wash.

P- 105, 106. — MERRIAM, C. H. Descriptions of Twenty-Three ET

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PP. 107-117. — MERRIAM, C. H. Descriptions of Four New P :

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Mexico. Proc. Biol. Soc. Wash. Vol. xiv, pp. 119-124. — MERRIAM, C. H. Two

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pp. 125, 126. — MERRIAM, C. H. Descriptions of Three New Kangaroo Mice of

the Genus Microdipodops. Proc. Biol. Soc. Wash. Vol. xiv, pp. 127, 128. —

MILLER, G: S., Jr. The Alpine Meis Hare. Proc. Biol. Soc. Wash. Vol. xiv,

pp: 97, 98. — MILLER, G. S., Jr. New Shrew from Switzerland. Proc. Biol.

Soc. Wash. Vol. xiv, pp. 95, 96. pee G. S, Jr. Descriptions of Three

New Asiatic Shrews. Proc. Biol. Soc. Wash. Vol. xiv, pp. 157-159. — NELSON,

E. W. A New Species of Galictis from Mexico. Proc. Biol. Soc. Wash. Vol. xiv,

pp. 129, 130. — NELSON, E. W. Descriptions of Two New Squirrels from Mexico.

Proc. Biol. Soc. Wash. Vol. xiv, pp. 131, 132. — NiPHER, F. E. The Relation

of Direct to Reversed Photographic Pictures. Trans. Acad. Sci. St. Louis.

Vol. xi, Nò. 4, pp. 51-61, Pls. II-X. — Nipuer, F. E. The Specific Heat of

Gaseous Nebulze in Gravitational Contraction. Zvans. Acad. Sci. St. Louis.

Vol. xi, No. 4, pp. 63-70. — NuTTING, C. C. The Sea Bottom: Its Physical

Conditions iod its Fauna. Science. Vol. xiii, pp. 841-852. — NUTTING, C. C.

Papers from the Harriman Alaska Expedition. ed The Hydroids. Proc.

Wash. Acad. Sci. Vol. iii, pp. 157-216, Pls. XIV-XXVI. — NUTTING, C. C.

The Laboratory Equipment of the * Bahama dei " from the University of

Iowa. Journ. Appl. Microscopy. Vol. iv, No. 4, pp. 1229-1234, 7 figs. —

Com. for 1899. Pp. 325-386.— OsBuRN, R. C. The Fishes of Ohio. Special

Papers Ohio Acad. Sci. No. 4, 104 pp. — PALMER, T. S. The Earliest Generic

Name of the Northern Fur Seal. Proc. Biol. Soc. Wash. Vol. xiv, pP. 15$ 1 134-

(a:

[t]

Tam

XON S

of West Virginia. pue Bi iol. Soc. Wash. Vol. xiv, pp- 161-163. — RANKIN.

oo hice off the West Coast of Greenland by the Princeton Arctic

Expedition of 1899. Proc. Acad. Sci. Phil. 1901. Pp. 169-181.— REPP, J. J.

Parturient Padi and the Schmidt Treatment. Iowa Agr. College, Bull.

No. 58. Pp. 17-30. — Rirrer, W. E. Papers from the Harriman Alaska Expe-

dition. XXIII. The Ascidians. Proc. Wash. Acad. Sci. Vol. iii, pp. 225-266,

Pls. XVII-XXX.— SHUFELDT, R. W. The Osteology of the Cuckoos. den

Amer. Phil. S l „ 2 pa—an, E F rud

‘oc. Vol. Ix, No. 165, 51 PP: 2 pls in Dr. Alfred

Bacterial Diseases ‘of Plants? A Consideration of Som e Statements in

Fischer’s “ Vorlesungen über Bakterien." Centralbl. Bakt. u. Parasitol. Ba. v,

PP. 271-278.— SwrrH, E. F. Dr. Alfred Fischer in the Róle of Pathologist.

Centralbl. Bakt. u. Parasitol. Bd. v, pp. 810-817. — SMITH, E. F. iio.

auf Alfred Fischer’s “Antwort” in Betreff der Existenz von durch Bakterien

verursachten Pflanzenkrankheiten. Cen£ra/M. Bakt. u. Parasitol. . vii, pp. 88-

199, Pls. I-XI.— SregLE, E. S. Sixth List of Additions to the Flora of r

ington, D.C., and Vicinity. Proc. Biol. Soc. Wash. Vol. xiv, pP- "s aea

Stevens, N. M. Studies on Ciliate Infusoria. Proc. Cal Acad. S DP. 3i

Zoól., vol. iii, 42 pp. 6 pls.— bees vs D, dre egt tun Bull.

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876 THE AMERICAN NATURALLIST.

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CHARLES A. KOFOID, PH.D., rue of Tllinois, Urbana.

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THE

AMERICAN NATURALIST

VoL. XXXV. November, IQOI. No. 419.

THE PARASITIC ORIGIN OF MACROÉRGATES

AMONG ANTS.

WILLIAM MORTON WHEELER.

. THE genus Pheidole among ants is in several respects

noteworthy. The unusually large number of species which

it comprises afford valuable materials for the taxonomist,

While their wide distribution and geographical variation can-

in habits of the species of this extensive genus. And,

finally, a fascinating congeries of morphological and physio-

of the

E = all the known species of Pheidole the worker phase

is represented by two very different forms : small-bodied,

EU big-headed,

ag prions from the Zoilogical Laboratory of the University x bese

877

878 THE AMERICAN NATURALIST. [Vor. XXXV.

Until very recently the genus has been characterized as

presenting no forms intermediate between the workers and

soldiers, in contradistinction to the Old World genus Pheido-

logeton, the species of which exhibit even in the same col-

ony numerous intermediates between the gigantic, big-headed

soldiers and the minute workers. During the past year I

found that two of our Texan and Mexican species of Pheidole

(Ph. kingii André, ver. instabilis Emery, and PA. vaslitit Per-

gande) resemble Pheidologeton in presenting, in the very same

nest, complete series of intermediates.! My attention was

directed to this singular condition by Professor Emery, who

has described the Texan ZZ. instabilis from specimens col-

lected at Austin. He has also rectified the synonomy of the

Mexican Ph. tepicana Pergande, which presents a like poly-

morphism and has therefore led to the description of several

species from soldiers of different dimensions? Recently Pro-

fessor Forel, utilizing some observations which I made at

Queretaro, Mexico, has been able to rectify a similar error

in the synonymy of PA. vaslitit Pergande.?

In the present paper I desire to call attention to another

peculiar modification of the workers of Pheidole, traceable to

a perfectly definite, though obviously very different cause from

that which must bring about the above-mentioned di- and poly-

morphism. June 3, 1901, I found at New Braunfels, Texas,

on a shady hill that slopes to the lovely sources of the Comal

River, two medium-sized nests of P/. commutata Mayr. They

were under stones about sixty feet apart. One contained

ants of the typical dark variety of the species, while in the

other nest both workers and soldiers were decidedly paler.

In either case on lifting the stone my attention was attracted

by several very large and conspicuous workers, with huge

abdomens, moving about among the workers and soldiers of

normal dimensions. I had been collecting and observing the

1 Wheeler, W. M. Notices Biologiques sur les Fourmis Mexicaines, Ann.

Soc. Entomol. de Belgique, tome xlv (1901), pp. 199-205. us

? Emery, C. Remarques sur un Petit Groupe de Pheidole de la Région

Sonorienne, Bull. Soc. Entomol. de France (année 1901), No. 5, pp- 119-127

? Forel, A. Fourmis Mexicaines récoltées par M. le prof. W. M- Wheeler,

Ann, Soc. Entomol. de Belgique, tome xlv (1901), pp. 123-141-

No. 419.] MACROERGATES AMONG ANTS. 879

little fungus-growing ants, Cyphomyrmex rimosus Spinola, which

abounded on the same hill-slope, and all my bottles and bags

used for living colonies were filled with these remarkable ants.

I was therefore compelled to preserve in some small vials of

alcohol as many of the Pheidole workers as could be captured.

At the time I supposed that the huge individuals might repre-

sent some hitherto unknown guest-ant which had taken up its

abode in the nests of the Pheidole.

On returning from my collecting trip I found that the con-

spicuous individuals were nothing more nor less than gigantic

workers of P4. commutata. One of the nests had yielded six,

the other three, of these creatures. There were besides from

each nest two or three somewhat smaller individuals clearly

intermediate in size between the typical and the gigantic

workers. All of these large individuals are evidently to be

regarded as belonging to Wasmann’s category of macroérgates,!

since they are certainly “ individuals which approach the females

in an abnormal manner only in the size of the body, but in

other respects (even in the development of the abdomen) are

normal workers." Although the abdomen is enormously dis-

tended in the macroérgates of PA. commutata, it is nevertheless

clearly of the worker type.

The length of the normal workers of the Texan PA. commu-

‘ata is not greater than 3 mm. Many of them are scarcely

more than 2.5 to 2.8 mm., which was the length of Mayr's

type specimens from Florida? The largest macroérgates, how-

Ever, measure 5 mm., while the smaller ones are fully 4 to 4.5 mm.

long. Thus the volumes of the normal workers and the

extreme macroérgates would be in the ratio of 27 to 225 if

they had the same form. But the abdomens of the latter are

So €normously distended that the ratio must be 27 to at least

200. In other words, the large macroérgates are nearly eight

times as large as the normal workers. They are even larger

than the soldiers, which measure about 4 mm., though in this

A Pepa E. Die ergatogynen Formen bei den Ameisen und ihre Erklä-

ge bes Centralbl., Bd. xv (1895), Nr. 16 u. 17, pP- ee D ud

d. Zoo] cde Die Formiciden der Vereinigten Staaten von Nordamerik

` - Ges., Bd. xxxvi (Wien, 1886), pp. 419-464.

880 THE AMERICAN NATURALIST. [VoL. XXXV.

case the difference in size is not so striking on account of

the enormous heads of the latter. The size relations are

shown in the figures, which represent the soldier, normal and

macroérgatic workers, drawn with the camera lucida under the

same magnification.

Examination even with a good pocket lens reveals the cause

of the great abdominal development of the macroérgates. One

sees distinctly the white coils of a parasitic worm distending

the abdomen till its dorsal and ventral sclerites are widely

separated by the tense intersegmental membranes. Thus the

abdomen of the Pheidole comes to resemble externally that of

replete individuals of the honey ant (Myrmecocystus melliger)

or our common northern Prenolepis imparis. In some of the

alcoholic specimens the tense abdominal wall has burst and

allowed a few of the coils of the parasite to protrude. Such

specimens may perhaps suggest the way in which the parasite

ultimately effects its escape from the ant, if indeed it ever

leaves its host.

My friend, Dr. T. H. Montgomery, who has kindly exam-

ined a few of the Pheidoles, writes me that the parasite is a

species of Mermis. Its exact location in the ant's body is

not easy to determine, Ze, whether it occupies the lumen

of the enormously distended crop, or ingluvies, or lies in

the body cavity outside of the alimentary tract. From

careful dissection of a single large macroérgate — the one

represented in the figure — I conclude that the Mermis lies

within the ingluvies. In this case the head of the parasite

extended forward through the postpetiolar and into the

petiolar segment, thus occupying the attenuated neck of

the ingluvies. The fat-body in the parasitized ants is almost

or completely absent and the walls of the enormously dis-

tended crop are practically in contact with the walls of the

abdomen. The large macroérgate figured contained only 4

single closely convoluted Mermis, which was ful 50, Me

long, or ten times the length of the ant. One individual

dissected by Dr. Montgomery contained two somewhat smaller

parasites, together with many of their eggs. According to

Dr. Montgomery, the parasites are “either fully mature Or

No. 419.] MACROERGATES AMONG ANTS. 881

in what von Linstow calls the ‘second larval stage,’ which is,

however, really the immature stage.” 1

While it is certainly somewhat singular that a species of

Mermis should occur in ants, even greater interest attaches

to the case under discussion on account of the manifest effects

of the parasite on its host. The fact that all the infested

individuals are of huge size as compared with the normal

Soh

TENA

7 PLA PST

ESTANY

(/ N TAT LEAN ANN

Ki Were Ny

Lar ae

Cann yg

; Pheidole commutata Mayr. a, normal soldier; 5, normal worker} c, parasitized macroérgate.

: (Drawn under the same magnification.)

Workers is remarkable, for, on first thought, one would cer-

tainly expect an animal infested with such a large parasite to

be stunted or, at any rate, below the average stature of the

Species. This paradoxical condition of the macroérgatic Pheil-

doles is easily understood, however, when we make due allow-

ance for certain peculiarities in the behavior of ants. In the

first place it is obvious that the parasites must enter the body

"Von Linstow, O. Das Genus Mermis, Archiv. f. mikr. Anat. Bd. liii (1898).

882 THE AMERICAN NATURALIST. [Vor. XXXV.

of the worker ant while she is still a larva. This is proved by

the fact that two of the large macroérgates are callows, one of

them still very soft and pale yellow, the other — again the one

represented in the figure — with harder integument, but with-

out the deep coloration of the mature workers. Such huge

parasites could scarcely have made their appearance in ants so

recently escaped from their pupa. But even if there had been

no callows among the macroérgates, the truth of the above

statement would still be patent, both because the macroérgates

were all infested while none of the normal workers were found

fo contain parasites, and because the stature of an ant is, of

course, fixed in the pupal stage and cannot be subsequently

increased to the dimensions exhibited in the cases under

consideration.

It is evident, furthermore, that the macroérgatic stature,

which is very apparent not only in the distention of the abdo-

men but also in the greater dimensions of the head, thorax,

petiole, antennze, and legs, can have its origin only in an unusu-

ally large amount of food consumed during the growth period

of larval life! Now, as I have shown in former papers? differ-

ent species of ants employ very different methods of feeding

their larva. Species of Camponotus, Formica, Lasius, and

Myrmica feed their larvae with liquid food regurgitated from

their crops, and possibly also with the secretion of the salivary

glands. Other species, however, like the Ponerinz and some

Myrmicinze (Aphzenogaster, Pogonomyrmex, Tomognathus, and

some species of Pheidole), feed their larvae with comminuted

insects. Unfortunately I have not been able to observe the

method of feeding in Ph. commutata, but it is safe to say that

it must conform to one or both of these methods. If the larva

are fed by regurgitation, we must suppose that the parasitized

1 The opposite condition, że., a small amount of food consumed during larval

life, results in what may be called microérgatic forms. Such are the firstborn

workers of all incipient ant colonies. These forms are, of course, perfectly

normal products of underfeeding, whereas the macroérgates of Pheidole are pre

ducts of overfeeding induced by a pathological condition. a

2 Wheeler, W. M. A Study of Some Texan Ponerinæ, Biol. Bull., vol. a

(1900), No. 1, pp. 1-31, Figs. r-10; and The Habits of Ponera and 5

tomma, Biol. Bull., vol. ii (1900), No. 2, pp. 43-69, Figs. 1-5-

No. 419.] MACROERGATES AMONG ANTS. 883

individuals have some means of informing their nurses that

their appetite is unusually keen — like that of a human being

infested with a tapeworm. If, on the other hand, the larva

are fed with comminuted insects, they could simply of their

own accord eat much more food than is consumed by the

larvae of normal workers. In either case, however, the stimu-

lus to the increased feeding that finally results in the macroér-

gatic stature must, of course, reside in the larva aud not in the

worker ants which supply the food.

The ability of a small animal like the worker of Ph. commu-

tata to nourish a parasite larger than the normal individuals

of the host species is accounted for by the fact that the larvæ

and adults of these social insects are so readily fed by other

members of the colony. The infested ant therefore suffers rela-

tively little inconvenience when compared with an animal which

must rely entirely on its own efforts in securing food. Both

during the larval and adult stages the macroërgate must be fed

by the other ants, for it is extremely doubtful whether these

heavy-bodied individuals ever leave the nest for the purpose of

foraging. They probably remain at home: like the heavy-

headed soldiers.

It is not difficult to understand how the Pheidole larva

become infested with the Mermis, since the parasite extrudes

its eggs within the crop of the adult worker. Such eggs or the

embryos arising from them could easily find their way into

the gullet and mouth of the ant and be transferred thence to

the larvæ while the latter are being licked and cleansed; or,

in case the workers of Ph. commutata feed their larvze by

regurgitation, the transferring of the parasite would be still

easier and more direct. ;

Other interesting conclusions follow from a consideration of

the fact that all the macroérgates are structurally of the pure

Worker type. Except in the excessive size and peculiar hyper-

trophy of the abdomen, I can detect no morphological differ

ences between the parasitized individuals and their diminutive

‘ister ants. There is certainly no appreciable tenaa m

Approach the soldier or female type of structure. From tus

we may conclude either that the larvae must become infested with

884 THE AMERICAN NATURALIST. [Vor. XXXV.

the Mermis after they have developed as workers so far that

their structure can no longer be affected except in volume, or

that the still undifferentiated larvae are infested but neverthe-

less develop into workers because so much of the food which

they devour is appropriated by their parasites. A decision

between these alternatives would require more precise study

than was possible under the circumstances.

While there can be no doubt that macroérgatism in Ph. com-

mutata is due to the presence of the Mermis, we cannot with

certainty exclude the possibility of an atavistic tendency

towards macroérgatism in the workers of this genus; for, as

Emery has shown in a very suggestive paper,! in those ants

which have the sterile females represented by huge soldiers

and diminutive workers, the latter have without doubt under-

gone a reduction in size during phylogenetic development.

It would be possible, therefore, to explain macroérgatism as

an attempt to regain the ancestral worker stature which was,

of course, that of the queen. This is probably the explana-

tion of macroérgatism in many ants, e.g., in Solenopsis, and

possibly also in the small group of Pheidole mentioned in

the introduction to this paper. While a similar reversional

tendency may also be present in PA. commutata, it is perhaps

unnecessary to lay much stress upon it, since the presence of

the Mermis is of itself quite sufficient to account for the

stature of the macroérgates.

It is interesting in conclusion to compare the production

of macroérgates in the nests of Ph. commutata with certain

phenomena observed by Wasmann? in mixed nests of Polyer-

gus rufescens and Formica fusca. He finds that such nests

are peculiarly liable to contain ergatoid females of the former

species, Z.e.,, “individuals which in size and in the development

of the abdomen (even of the ovaries) belong to the true female |

type, but have the thoracic structure of the workers and are

therefore wingless." From a biological point of view these

are, as Wasmann claims, really secondary queens. He believes

that they are produced by the slave ants (F. fusca) through

! Die Entstehung und Ausbildung des TTEA bei den Ameisen, Biol.

Centralll., Bd. xiv (1894), pp. 53-59. 2 Loc

No. 419.] MACROÉRGATES AMONG ANTS. 885

excessive care and feeding of certain larvae which had previ-

ously been permitted to develop as workers beyond the stage

in which the wing rudiments would make their appearance in

queen larvae. In other words, the fusca workers attempt to

change worker larvae of Polyergus into queens but succeed

only in producing the wingless ergatoids. In explanation of

such conduct, Wasmann suggests that the F. fusca usually

have several queens even in very small nests and may perhaps

retain the instinct, when enslaved by Polyergus, to educate

numerous female ants. If, after the nuptial flight of the

Polyergus, they find no fertile queens of their own species in

the nest they may endeavor to transform the young worker

larve into queens with the above-mentioned result. Was-

mann's hypothesis is of interest, as it points to the existence

of a peculiar instinct in ants which regulates the number and

character of the personnel in the colony. We know that such

an instinct is well developed in termites, and it is more than

probable that it exists also among ants. It offers an interesting

field for future investigation.

Both Wasmann's hypothesis to account for the ergatoid

females of Polyergus by excessive feeding of the worker larve,

and his interesting “ Lomechusa-Hemmungs-Hypothese," in

Which he accounts for the pseudogynes of Formica through

an attempt on the part of the ants to transform queen larvae

into workers, seem to start from the assumption that the

larvae are quite passive and that the worker ants feed them

entirely in obedience to certain instinctive promptings of their

own. This accords with Emery's view! that the sexual poly-

morphism of the ant colony is the result of the development of

an instinct in the workers to feed the larvze in different ways.

Hence, “the characters in which the worker differs from the

Corresponding sexual form are not congenital, or blastogenous,

but acquired, z.e., somatogenous. Nor are these characters

transmitted by heredity, except as a peculiarity of the germ-

Plasma to enter on different paths of ontogenetic development

according to the different circumstances of existence." While

I5 view is undoubtedly supported by many facts, and while

1 Loc. cif.

886 THE AMERICAN NATURALIST.

considerable importance may indeed be attributed to the ini-

tiative of the workers in determining the character of the

adult ants which they rear, the macroérgates of Ph. commu-

tata prove, nevertheless, that we must also attribute a certain

amount of initiative to the larvae themselves. If this be

granted, it is but a short step to the admission that the initia-

tive of the larva, even under normal circumstances, — 7.e., when

not infested with internal parasites, — may be considerable.

It is not altogether improbable that further investigation with

this possibility in mind may lead to some alteration or emen-

dation of the various hypotheses that have been framed for the

purpose of explaining the complicated phenomena of sexual

polymorphism. Thus we may find eventually that the tend-

ency to develop abortive ovaries is really inherited (through

the fertile queens of course), and that differences in the chem-

ical nature of the internal secretions, perhaps analogous to

those which are supposed to obtain between castrated and

non-castrated animals, may furnish the different stimuli that

induce the larvae to demand of their own accord more or less

food, or food of a different quality, and to develop accordingly

into queens or workers.

COLEBROOK, CONN., August Io, rgor.

ON SOME POINTS IN THE ANATOMY OF A

COLLECTION OF AXOLOTLS FROM

COLORADO, AND A SPECIMEN

FROM NORTH DAKOTA.

HENRY LESLIE OSBORN.

I. THE COLLECTION FROM COLORADO.

IN a previous article in this journal by the writer ('00)

reference was made to a collection of axolotls which were

kindly loaned by Dr. Lee for comparison with the specimen

. from Amenia, North Dakota, therein described. By an unfor-

tunate mistake these specimens were located from Montana, but

in fact they are from Colorado. The material was collected by

Dr. Thomas G. Lee, of the University of Minnesota, during a

brief stay in the mining town of Crede, in southern Colorado, -

during the latter part of August and the beginning of Septem-

ber, 1896. Having heard mention among the people there of

“dogfish ” and «fish with legs on 'em," he succeeded in locat-

ing the wonders in a lake twenty-five miles distant and paid

the placea visit. He found the lake occupying a narrow valley

hemmed in between two mountain ridges and dammed back by

a natural formation, apparently of glacial origin, giving it much

the appearance of an artificial lake. The elevation of the place

is about eight thousand feet above the level of the sea. The

Tuer is very cold. The lake appeared to be very deep. Ata

distance of about a hundred feet from the shore a line one

hundred feet in length would not reach the bottom. There

were water plants growing abundantly in the lake, and among

them there were areas that were free and open, in which the -

axolotls were seen coming up from time to time apparently to

“ cathe. They were then captured with a dip-net. The larger

Sized ones were found farther out, while smaller ones Were

reached from the shore. Dr. Lee informs me further that he

887

888

*OPLAO[OD Ur01] Upas JO MIIA 9pIG — `I ‘DIA

THE AMERICAN NATURALIST. [Vor. XXXV.

found the ordinary larvae of Ambly-

stoma tigrinum ranging in length

from one to four inches in the irri-

gation ditches that were common in

the San Luis valley in the vicinity

of Garrison.

The material was preserved in 5%

formalin. It contains in all twenty-

six specimens, with the following

lengths in millimeters, vzz.: 89, 92,

98, 100, 103, 105, 105, 105, 106,

100, 110, £12, 113) 113, 185, 10

118, 120, 121, 160, 190, 215, 220,

250, 250, 262.

Most of the specimens only cor-

roborate the descriptions of previous

writers as to the external character-

istics of these interesting forms, and

are mentioned here chiefly for the

sake of making a record of the facts

and the locality. One of them is in

a an advanced stage of metamorphosis,

and has nearly reached the terrestrial

form of Amblystoma tigrinum. This

specimen is of especial interest, be-

cause it is in the act of undergoing

its development in the midst of its

natural surroundings. Marsh and

Tegetmeier, as well as others, have

described the metamorphosis of sire-

dons under artificial conditions, but

I do not know of an account of a

siredon transforming in its natural

environment. A number of

measurements of these specimens

were made, and are given in 4

table of measurements at the end

of this article.

No. 419.] THE ANATOMY OF AXOLOTLS. 889

The external characteristics are indicated in Fig. 1, which

is drawn from specimen No.2 of the following lists. In the

account which now follows, the siredon described as the type

of the collection is No. 2. It will be compared with the other

siredons, with No. 10, a metamorphosing specimen, and with

adult specimens of Amélystoma tigrinum from the collection in

the museum of Hamline University, from St. Paul, Minn.

The coloration (of the formalin material) is uniform and

not mottled or spotted. The head and trunk are dark above

and light beneath, the division line running from the jaw along

the side of the head and on the body on the level of the

ventral borders of the limbs. The post-abdomen, or “ tail,” is

dark throughout, as are also the dorsal and ventral folds of

skin, “fins.” One of the larger specimens (No. 8) differs

from the rest in being distinctly spotted. The spots are very

numerous and small, about 1 mm. across, and are found in all

parts of the dark area of the animal. Both of the Dakota

specimens are spotted, as can be seen by a reference to Fig. 7

of this article and Fig. 1 of the preceding. In both of these,

however, the spots are fewer and larger and more distinct.

Baird's S. gracilis (59 Vol. X, Pl. XLIV) is also spotted, and

the Colorado specimen is more like to it, according to the illus-

tration, than to the Dakota forms. In the smaller specimens

of the Colorado collection the dorsal darker area is not uniform

in tone, but is mottled with dark irregular patches scattered

irregularly upon it. These patches are not distinctly bounded,

but shade into the general color tone at the edges, and are

very irregular in shape. They also extend out upon the dorsal

and ventral fins.

The head in the Colorado specimens differs markedly from

that of terrestrial amblystomas in several particulars. - In

order to afford the opportunity to test some of these points,

I give a number of measurements made on specimens of

Amblystoma found at Hamline, Minnesota. I may say that

these are fully metamorphosed and strictly terrestrial. |

By dividing the length of the head, from the snout to the

fold Crossing between the bases of the hinder gills, by the

length from the chin to the posterior boundary of the cloacal

890 THE AMERICAN NATURALIST. [VoL. XXXV.

opening, we obtain the ratio of head length to head and body

length, and can then make direct comparisons, irrespective of

difference of length. By measuring to the cloaca we avoid

the influence of the variation of the post-abdomen. These

measurements and the ratio are shown in this table.

Ratio or HEAD LENGTH TO LENGTH OF HEAD AND Bopy.

COLORADO SPECIMENS dx hui AMENIA SPECIMENS

AMBLYSTOMA M

No. | Sizes Ratiol No. | Sizes Ratio| No. | Sizes Ratio|| No. | Sizes| Ratio| No. | Sizes | Ratio

12 | 48 |37%|| 1 | v |32%|| 10 |f 12776|| 12 | 33 |32% X | fx | 31%

$ |36 6 2 Serer a) aes

3; ü 3 % Yos 31% 4 "n 8% KENMARE SPECIMENS

5 | 1$ 335 2 | wus |29% 8 | fus | 28% || —— m

3| 8 |342|| 7 | 3s |30% 5 | e's | 26%|| x | ates | 29%

8 | $8 |37%|| 9 | Ahr | 31% 10 | 49s |27%

13 | fs |32% | 7 |2 | 24%

Average 36% || Average 31% | Average 27.5%

These figures show great variation in the head length in

general among these forms, so that inferences from single

cases will need to be carefully guarded. Thus in the first

group we find ratios of 33% to 37%, and in the St. Paul forms

we find a range of from 24% to 32%. The size variation thus

shown to exist would be an interesting topic for study, and has

not as yet been studied. Referring now to the ratios, we see

clearly that on the whole the Colorado forms have a larger

head than the St. Paul forms. In the latter the ratio is 27.5%,

while in the former it is 31% for the larger sizes and 36% for

the smaller ones. These figures are in reality an underesti-

mate, for they are reduced by the fact that in the comparison

no allowance has been made for the tendency of smaller speci-

mens to have larger heads. This tendency is very well shown

by comparing specimens of different sizes in the foregoing

statement. Thus in the St. Paul specimens it is very clearly

seen, and in the Colorado forms the smaller ones have a ratio

of 36%, while the larger ones have a ratio of only 31%. If the

series of specimens were large enough to enable us to eliminate

No. 419.] THE ANATOMY OF AXOLOTLS. 891

individual variation, a more accurate statement of the amount

of difference could be made. But if we compare a Colorado

specimen with a St. Paul specimen of the same size, we find

in No. 1 of the Colorado forms a ratio of 32%, while Ambly-

stoma, No. 8, of the same size has a ratio of 28%, and the

larger St. Paul specimens have a still smaller ratio than even

larger specimens of the Colorado series. Thus No. 7 from

St. Paul has a ratio of 24%, while the largest of the Colorado

series, a much larger animal, has a ratio of 32%. One should

not, however, place much reliance on the comparison of indi-

viduals, but I believe that on large series of individuals of the

same size it would be found that the head ratio in the siredons

. is much larger.

Not only is the head as a whole larger in the aquatic forms

than in the terrestrial ones, but the proportion of its parts is

also different. A reduction of the head in the throat region

would be expected in connection with the degeneration of the

branchial apparatus, and does occur. This is seen by reference

to the following statement showing the lengths of these regions

of the head in a number of different Colorado and other sire-

dons and in six St. Paul amblystomas, 775. :

COLORADO SERIES | St, Papi Sen

No, Shes Snout to T y No Sizes E uu

12 f 27% 73% - u 41% 59%

II É 26% 74% 5 1 41% 59%

5| 4 26% 74% 8 36 40% n

ES A 26% | 74% | 1 | H "T ln :

Ado 35% | 65% || 12 | d 35% 7

; Hn 33% 67% 4 ho dove pov

D 27% 73% . 60.3%

. Hou oz €—

E xx a

0 o

9 35 23% 77%

13 m 28%, 72% x | H 22% 7 y

Average . ee EAE ee

R eee: r KENMARE SPECIMEN

Meram ae eer ee ug

- m SPECIMEN r 4 | 29% | 73%

Ex i | 284, | 72% i RM

892 THE AMERICAN NATURALIST. [Vor. XXXV.

These figures show that the average ratio of the length of

the hinder part of the head to the front part is 72% in the

siredons and only 60 in the terrestrial amblystomas.

The form of the outline of the head anteriorly is another

point in which the aquatic and terrestrial forms differ con-

siderably. The difference is shown in Fig. 2. In the former

the outline of the head in front is very blunt and broadly

convex; in the latter the outline bends suddenly after the

Fic. 2. — Dorsal view of head of Colorado siredons, No. 2 and No. 1o.

nostril is reached, producing a sharply convex curve. This

is due to the presence in the aquatic form of a cheek fold of

the skin bordering the jaws, not found in the terrestrial state.

In the terrestrial forms the skin is drawn and tight-fitting over

the whole of the head. This is obviously correlated with the

need of providing against the desiccating influences of the

atmosphere. In the aquatic forms, on the other hand, the skin

is very loose and ample. This is noticeable generally in the

head, not only in the general skin, but in the presence of gills,

opercula, and such special modifications of the outer surface.

No. 419.] THE ANATOMY OF AXOLOTLS. 893

The folds at the side of the mouth reduce the diameter of its

opening. This difference in the size of the mouth is another

marked difference between the axolotls and the terrestrial

forms. It is indicated in Fig. 3, where the angle of the jaws

is seen in 3 a to be anterior to the eyes, while in 3 c it is

considerably posterior. This

smaller size of mouth is per-

haps correlated with the proc-

esses involved in taking water

into the mouth chamber for the

purpose of forcing it back and

out of the sides of the throat

during aquatic respiration.

The gills (in No. 2) do not

differ essentially from the ac-

counts and figures of Baird and

others. They are shorter than

the head. They are flattened

outgrowths of the sides of the

throat. Proximally they are

continued in the floor of the

throat to the hyoid bone in

thecenter. On the underside

Me the opercula, thin flaps of Eme

skin, which merge distally into :

the under margin of the gill.

The gill plates are borne on

the posterior side of the free T

Portion of the gill, which by . pre. 3.—Side views of bed. a siredon Xo, 3

1G. 3

from Colorado ; 4, siredon No. 10

m E

fans of a muscle running rado ; c, salamander from St. Paul.

ns length can be drawn back

against the side of the body, thus protecting the plates. The

latter are thin and placed in rows crossing the long axis of the

gill, They are so arranged that two longer plates next the edge

of the gill are followed by two smaller plates, set nearer to each

Other in the middle, as shown in Fig. 4 4. Many of the sp ü id

Mens in the collection show the gills in a state of degeneration,

the gills being much shorter and the plates reduced in number.

894 THE AMERICAN NATURALIST. [Vow. XXXV.

The trunk is essentially the same in the siredon and terres-

trial states, excepting as to three points, vzz., coloration, the

dorsal fin, and the ** webbing" of the toes. All of these changes

have been commented on by previous writers, and may be

passed without further notice. ,

The post-abdomen is acutely tapering from its origin to

the tip. Its outline as a whole is very acute, unlike either of

the Dakota specimens. It bears a very thin and membra-

neous “fin,” which becomes noticeably wider posteriorly.

Fic. 4. — End view of gill, showing the gill plates. a, in Colorado forms ; 4, in Amenia specimen.

This broadening of the fin is greater than anything indicated

in the figures of axolotls generally, and is quite different

from the outlines in the two Dakota specimens.

The internal anatomy of the siredons, excepting of the mouth,

is strictly the same as that of the mature terrestrial forms. A

thorough dissection of one and a careful examination of several

others of the collection demonstrated an anatomical condition

of the viscera very similar to that indicated by Wiedersheim (79)

in his Fig. i. The reproductive system in both the males

and the females is fully matured, and in the females there are

accumulations of eggs ready for discharge, and the albumen

No.49.] | THE ANATOMY OF AXOLOTLS. 895

glands are swollen and active. The heart and pulmonary

organs have reached their final form, the former having two

auricles and a connection with the lungs, and the latter con-

sisting of the two elongate lungs communicating with the

throat by means of a glottis. The siredon is thus morpho-

logically a pulmonate or “air-breathing ” animal, and so it is

physiologically, for, notwithstanding its aquatic environment

and branchial equipment, it uses its lungs for breathing.

A single one of the members of the Colorado series, the one

numbered 10 in the list, is especially interesting, because it was

taken in the act of transformation, the process nearly completed.

The head of this specimen is shown in Figs. 2 4, 3 4, and 5.

The general appearance of the animal is very similar to the one

illustrated by Tegetmeier (70). The head is intermediate in

form between the aquatic and the terrestrial states. Its total

length is 27% of that of the head and trunk. This is the ratio

found in land salamanders, while 31-36 is that found in aquatic

forms. The posterior part of the head, however, is here found

not to be relatively shortened, as might at first be expected. It

I$ 72% of that of the whole head, as in siredons, as against 60/,

the ratio for land forms. As to outline in front, the head is as

In the land forms, being strongly curved (see Fig. 2 b). The

skin of the head is tightly drawn and not loose and abundant.

This tension of the skin produces the bulging of the eyes noted

by Marsh (68) and Weismann (75) among the changes of the

transformation, Dorsally the head is arched, as in the land

forms, and not any longer so flat as in the aquatic stage. The

mouth opening is not yet as wide as it is in the definitive form.

The fold at the angle of the jaws has lessened but not entirely

disappeared, and the angle of the mouth is on the level of the

“Yes and not behind them (compare Figs. 3 and 3c). Ven-

‘rally the gular fold has become confluent with the floor of the

mouth and throat in the center, and some distance from the

Middle line on each side the vestiges of the gill apparatus

E^ The three clefts are still present on each side, but

Er been pushed back and out from the throat, es ah

. “ed into the small space left behind the gular fold (se

Ig. 5). Structurally the apparatus is almost completely

896 THE AMERICAN NATURALIST. [Vou XXXV.

aborted. Besides loss of length, the opercula are gone and

the external parts of the gill are reduced to mere warty lumps.

On the floor of the mouth the tongue had not yet developed.

The trunk region is also intermediate and presents both sire-

don and salamander features. The coloration is still of the

aquatic kind, showing no hint of the future dense black general

tone spotted with irregular blotches of brilliant lemon yellow

so characteristic of the land forms of the species. On the other

hand, the dorsal fin is wholly absorbed,

and the toes are no longer “ webbed";

instead, the skin fits closely and the toe

is cylindrical and tapering. With this

change of the toes and the fins the

power of locomotion by swimming in

water must be very considerably dimin-

ished. The post-abdomen has also

reached the final form, the tail fin being

nearly completely absorbed.

Fe ee cin Ne’ -legetmeier reports in 1870 his obser-

showing the transition to the vations upon a transforming axolotl. His

stem account is very brief. He says: “The

specimens were hatched in the summer of 1868, and kept

under similar conditions without any change having taken place

beyond steady increase of growth during the succeeding winter

and the summer of 1869. In the autumn one began to change;

the external gills disappeared, the jaws became more pointed,

and the skin assumed a singularly mottled appearance. The

animal did not leave the water, but when the temperature was

warm usually breathed by standing erect against the side of the

aquarium and elevating the nostrils above the surface; during

the cold weather it usually remained submerged, rising at inter

vals to breathe.” This case of Tegetmeier’s seems to be like

the Colorado specimen in that the metamorphosis took place

while the animal still maintained its habits of aquatic life.

Apparently the same may be said of the transformations reported

on by Marsh (68), Dumeril (70), and Chauvin (75).

We do not know the cause of the transformation

siredon. In the St. Paul form of Amblystoma the animal first

of the

No. 419.] THE ANATOMY OF AXOLOTLS. 897

becomes terrestrial and afterward matures. We have come to

regard the land forms as mature, when as a matter of fact the

aquatic form may be just as truly mature, since its organization,

excepting in a few minor points, is identical with that of the

land forms. There are two kinds of transformation distinguish-

able in the metamorphosis of an Amblystoma. One of these is

in the development of the limbs, the alteration of the circula-

tory system, the development of the lungs, and the matur-

ing of the organs of reproduction; these may be considered as

the primary adult characteristics. In addition to these are a

number of secondary changes, which do not involve radical mor-

phological processes, but are largely confined to points in the

external anatomy. These latter are secondary adult character-

istics. The whole axolotl. question turns on these secondary

characteristics ; for, as to the primary changes, they take place

alike in axolotls, siredons, and salamanders, and one is as truly

adult as the other. But for some reason an interval has arisen

in axolotls and siredons between the primary and the secondary

changes, so that instead of all of them progressing par? passu,

the secondary changes are postponed, and take place either

much later or not at all In the neighborhood of St. Paul,

amblystomas are fully adult when only 100 mm. in length,

perhaps even less. In the Mexican axolotl, when the animal

reaches this length, the secondary transformations do not take

place, though the primary ones do, and the animal thus remains

aquatic, though adult in all other respects. In the siredons we

find that the condition is intermediate; instead of secondary

transformation being indefinitely postponed and not taking

Place at all, it takes place much later. In specimen No. 10 It

ms taking place after the animal has reached a length of 235 mm.

instead of having occurred at only 100 mm. In No. 2, of the

Same size and apparently from the same environment, it has

not taken place, and no signs of its approach are visible.

898

"Ejoxe(T (HON 'S1euruo'w uro4j uauiroads ayy jo Mata IPIS — ‘9 IA

THE AMERICAN NATURALIST. [VoL. XXXV.

II. THE SPECIMEN FROM KENMARE,

Nortu DAKOTA.

The axolotl referred to in the

former article (American Naturalist,

July, 1900, p. 544) as having been

found in Kenmare, North Dakota,

has been kindly loaned to me for

examination by its owner, Mr. W.

H. Makee of Kenmare. It was

somewhat injured by cutting in the

throat, possibly when it was cap-

tured, and has been somewhat dis-

torted by preservation in too small

a bottle; still it is adequate for

study and comparison with the

Amenia specimen. As it had been

preserved in alcohol, while the

Amenia specimen was preserved in

formalin, and as the Kenmare speci-

men was somewhat distorted in

fitting it into its bottle, the meas-

urements of the latter will not stand

a rigid comparison with those of the

other siredons. |

Mr. Makee writes me as to the

locality of the specimen, that it was

caught among a number of fish in

a seine during June, 1898, in Des

Lacs Lake. The lake is thirty miles

long and half a mile wide, and has

an estimated depth of twenty-five

feet. It is located in Ward County,

North Dakota, and discharges

through a chain of lakes into the

Mouse River, a tributary of the

Red River of the North, and finally

of Hudson Bay. Kenmare is at

No. 419.] THE ANATOMY OF AXOLOTLS. 899

the opposite end of the state from Amenia, and is the most

northern locality from which siredons have been reported. The

coloration resembles the Amenia specimens and is unlike that

of the forms from Colorado. It is dark above, excepting in a

narrow tract on each side of the dorsal skin fold. There are

numerous dark rounded distinct spots, the largest of which

have a diameter of 4 mm. The spots are scattered over the

body and post-abdomen and on the upper surface of the head.

They are more numerous and larger than in the Amenia form.

The undersides of the trunk and head are several shades lighter

in color than is the.case in the Amenia specimen. The skin is

smooth, as in the Colorado specimens. The “coarse and warty -

roughness" of the Amenia form is not found in the Kenmare

specimen.

The appearance of the Kenmare specimen is shown

in Fig. 6. With the exception of its gills, it is more

like the Amenia form than it is like the Colorado speci-

mens. Its dimensions are shown in the table at the end

of this article.

The head in the Kenmare specimen is shorter relatively to

the length of the head and trunk than it is in the Colorado

forms. This can be seen best by means of a table, thus:

Ratio of Head to Head and Trunk in

(2)the Kenmare specimen . . . +» + + * 7 * 29%

(@) the Amenia specimen |... 0 * otov ? 31%

(4) the St. Paul specimens |. roseo ACT 27%

The injury of the head made it difficult to be sure of the

length of head within 1% or 2% in the Kenmare specimen, but

€ven so, the head is much shorter in both of the Dakota

forms than in the Colorado specimens, and more nearly the

length that is characteristic of the terrestrial forms. Too much

stress should not, however, be laid on this fact, as the measure-

ment is based on a single specimen in each case, and some of

the single ones of the Colorado series have the head as short as

29%, as in this specimen.

900 THE AMERICAN NATURALIST. [Vor. XXXV,

The length of the head behind the eyes is also interesting

here. Tabulating this measurement in the four cases, we find

that the posterior length is

(ayin the Kenmare specimen . . . . + + + + 73%

(6) in the Amenia specimen. . . . + + + = - 78%

GM the SE Paul series o e 0, o 16r ee

of the total length of the head. In this respect the Amenia

specimen is very extreme, and the Kenmare specimen agrees

with the rule for siredons, as determinetl by the Colorado

series.

The head is also blunt anteriorly, as in general. The gular

fold in the Colorado forms, as in siredons generally (see Baird,

52, Fig. 3), is emarginate, and this is the case in the Kenmare

specimen, unlike the Amenia one.

The gills are three on each side. They are vertical out-

growths of the side of the throat, flattened so as to present a

dorsal and a ventral margin; the latter, at the base of the gill, is

continued inward under the branchial bone to form a thin flap,

the operculum, which covers the gill slit in the floor of the throat

(see Fig. 6). The gill presents two surfaces — one anterior,

the other posterior. The former is naked, the latter bears the

thin triangular plates in which the capillaries are placed which

expose the blood to the aérating action of the water. The gills

can be bent by the contraction of a muscle located within

them, so that the posterior side is parallel to the body, and the

gill plates are thus covered and protected.

There are four rows of gill plates. Two of the plates are

larger and more external. They are on the same level, and are

followed by two smaller and more internal plates. Each one

of the plates is a tall triangular and flattened structure. At

its base a blood vessel can be seen entering on one side.

Capillaries filled with blood corpuscles can be seen in the

interior, and running through the gill in the center and near

the bases of the plate is a large vessel. The blood vessels are

deeply pigmented, and the surface of the plates is minutely

spotted with black branching chromatophores.

NO. 419.] THE ANATOMY OF AXOLOTLS. 9OI

The gills differ from both the Colorado and the Amenia speci-

mens. The lengths of the filament-bearing areas of three gills

in succession, beginning with the anterior one, are respectively

76%, 90%, and 125% of the length of the head in the Kenmare

specimen. The corresponding figures for the Amenia specimen

are 76%, 90%, and 117%. For the Colorado forms they are, in

case No. 21, 33%, 50%, and 72%. In both the Dakota speci-

mens the gills are thus noticeably much longer than is the rule

among siredons, as hitherto reported, and among the Colorado

specimens which are in the main in accord with the cases

reported by Baird, Marsh, and others. The gill in the Kenmare

specimen does not broaden distally, but is tapering from base

to tip. This is in contrast with the Amenia specimen, where

the gills are spatulate in form, which point may, however, prove

to be an abnormality. The form of the gill plates in the Ken-

mare specimen is different from either those from Colorado or

the Amenia specimen. They are not long and filamentous as

in the latter, and they are not short and blunt as in the former ;

but they are wide and plate-like, tapering to the tip, the outer

row being longer than the inner; they are arranged as in the

Colorado forms.

The body offers nothing for special comment. The colora-

tion has already been mentioned, and the feet show the same

broad foot margin of skin on the sides of the toes as is found

in other siredons. The post-abdomen and the fin are much as

in the Amenia form and decidedly unlike the siredons generally

and the Colorado series. The body part is heavy and broad

from base to tip; it is broadly convex at the tip, in marked

contrast with the acute taper found in all the Colorado series.

The fin is thick and strong, contrasting with the thin mem-

branous texture of the Colorado forms, and it is not so wide

asin them. In all of these points the post-abdomen is like

that of the Amenia specimen. :

An examination of the internal anatomy was made only dl

far as it could be done by means of a short incision in the side

Wall of the body, Through this it was possible to see the lungs

fully developed, as in a strictly terrestrial form, and the various

viscera were similar to those found in the St. Paul material.

902 THE AMERICAN NATURALIST. [VoL. XXXV.

The sexual organs were not in an active state; a large corpus

adiposum is present, and behind it a small elongate but not dis-

tended genital organ, which upon examination proved to be an

ovary containing numerous slightly developed but recognizable

eggs. I did not find any evidence to show whether the animal

had ever laid eggs or was still immature, but the latter suppo-

sition seemed more likely.

BIOLOGICAL LABORATORY, HAMLINE UNIVERSITY,

St. PAUL, MINN., April 18, 1901.

LITERATURE REFERRED TO.

'52 BAIRD, S. F., and GIRARD. C. App. C. Reptiles, Stansbury's Expe-

dition, p. 335, Pl. I, Fig. i.

'59 BaiRD, S. F. Rep. U. P. R. R. Survey. Vol. x, Pl. XLIV.

"76 CHAUVIN, MARIE VON. Zeitschr. f. wiss. Zool. Bd. xxv.

'68 Marsa, C. C. American Journ. of Sci. Series 2, vol. xliv.

‘00 OsBOoRN,H. L. American Naturalist. Vol. xxxiv.

"OQ TEGETMEIER. Proc. Zodl. Soc. London.

'75 WEISMANN, AUGUST. Zeitschr f. wiss. Zool. Bd. xxv.

'79 WIEDERSHEIM, R. Zeitschr. f. wiss. Zool. Bd. xxxii.

903

THE ANATOMY OF AXOLOTLS.

NO. 419.]

Catalogue Number.

Chin to tip of tail... . +» -

Chin to margin of gular fold .

Chin to post. edge of cloaca .

Snout to level of bases of in-

ner gills :

Greatest diameter of head . 1

Width of mouth . os

Distance between nostrils. .

Distance between centers of

eyes.

Width of head at eyes .

Snout to line joining centers

of eyes .

Length of filamentous border

of gill I

Length of filamentous border

of gill 2

Length of filamentous border

Length of arm . uw ed

Greatest height of dorsal fin :

dà

A PARASITIC OR COMMENSAL OLIGOCHA:TE

IN NEW ENGLAND.

M. A. WILLCOX.

Chetogaster limnei von Baer appears to be the only oligo-

chæte which is generally recognized as a parasite ; at least it

is the only one to which Michaelsen in his recent work on the

subclass (Das Tierreich, Oligocheta, 1900) ascribes such habits.

The species has been reported only from Europe, and it is there-

fore with pleasure that I am able to announce the discovery in

the neighborhood of Boston of a very similar form, regarding

which I can at present give only the following incomplete notes.

The worms were first found about the end of May, 1901,

infesting a species of Physa, probably P. heterostropha, which

was obtained in a small stream in Wellesley. They were also

afterward observed upon an undetermined species of Planorbis.

They were abundant about the head and in the respiratory cav-

ity. Ina few instances the hinder part of the body seemed to

be imbedded in the tissues of the snail, but of this I cannot be

Sure. Most of them certainly were entirely free, and, waving

about like tiny white threads, had much the appearance of a

fungus, Occasionally I have noticed them, like the European

form, crawling over the outside of the shell ; but they ordinarily

restrict themselves to the body of the snail.

This is undoubtedly the animal mentioned by Gould (Report

on the Invertebrate Animals of Massachusetts, 1841, P. 213)

under the name of Gordius inguilinus Müller, and supposed

by him to be a parasite of P. heterostropha. In none of

specimens taken under ordinary conditions, boweven n s

alimentary tract give any indications of a parasitic ha idit

the contrary, I have seen in it no other food tha n :

It will be remembered that Lankester in his €— :

the European form («A Contribution to the €: e VI

the Lower Annelids,” Trans. Linn. Soc., Lond., Vol. $

905

n diatoms.

906 THE AMERICAN NATURALIST. | [Vor. XXXV.

pp. 631—646, 1870) speaks of finding the worms in winter within

Limnzea, and gorged with the kidney cells, while he seems to

have found no such evidence of parasitism at other seasons.

I should add that in one instance, in which the worms had

been left for some time in a watch glass with a torn Physa,

the alimentary canal of almost every one was found stuffed

with the blackish pigment characteristic of the snail, which

was floating in the water.

The worm is a small, transparent animal, about 2 mm. in

length and 1-5 mm. in breadth at its widest point. Both these

measurements are taken in extension, but as the creature repro-

duces by fission, and as colonies of at least three persons are

common, the first impression is of a much longer organism than

I have described. On the ventral side, near the anterior end,

are two clusters of cephalic setze ; each seta is sigmoid, about

1-14 mm. long, and ends in a fork whose prongs are equal and

are bent nearly at right angles to the shaft; each cluster ordi-

narily contains seven setze; though there may be six or, very

rarely, five. In one case, one side bore the full number of seven,

while there were but two on the other. It would seem probable

that the remaining ones had been torn away. At some little

distance behind these clusters are the abdominal ones, which

differ from the former only in that the setze are smaller (about

1-23 mm. in length), more numerous (being, as a rule, eleven,

rarely ten, in a cluster), and borne-on a slight projection of the

body which apparently corresponds to a rudimentary parapo-

dium, This last peculiarity, combined with a habit of holding

the hind end fixed while extending and waving about the ante-

rior one, gives the worm, as Lankester has already remarked, a

curious likeness to a geometrid larva. There are sometimes as

many as thirteen pairs of seta clusters behind the cephalic one ;

it is difficult to determine how many of these should be reck-

oned as belonging to a single individual, but for reasons which

will be given in discussing the budding, I am inclined to con-

sider with Lankester that the adult has four pairs of abdominal

clusters.

The large mouth lies at the anterior end, but the dorsal edge

projects rather over the ventral one, so that the tip of the body

No. 419.] A PARASITIC OLIGOCH.ETE. 907

in side view appears obliquely truncate. The mouth leads

directly into the pharynx, which occupies rather more than one-

third the length of the animal, is extremely distensible, and is

connected by muscle bundles to the body wall. The pharynx

is succeeded by an cesophagus so short and narrow as to be,

when the other parts are greatly dilated, quite indistinguishable.

Next comes the stomach, which as seen in optical section is

nearly square and is separated by a constriction from the

intestine, This division is somewhat longer than broad, and

its walls are usually characterized by a yellowish brown hue,

sometimes to be observed also on the stomach and suggest-

ing the chloragogue cells of Lumbricus. Opposite the third

abdominal setze is a constriction, at which point this yellow

Color suddenly disappears. The remaining part of the intes-

tine varies in shape according to the degree of development of

the buds. The hind end of the worm is slightly notched.

The stomach is often found filled with diatoms, with which

are likely to be intermingled a few pebbles. The other parts

of the alimentary tract, if not entirely empty, are usually

nearly SO.

There are five septa: one opposite the anterior and one

Opposite the posterior end of the cesophagus; one at the

constriction between stomach and intestine, one opposite the

middle of the intestine, and one at the point of its posterior

constriction, Behind the fourth abdominal seta is a zone in

Which the coelom may be absent, and behind this region may

appear the ccelom of the bud.

The large blood trunks are a dorsal pulsatile and a ventral

nonpulsatile one. They are connected by a pair of pulsatile

Vessels which encircle the cesophagus. Lankester has figured

a pair óf anterior ones extending obliquely down and back

around the pharynx; I have seen such vessels, but am not

Sure of the way in which they connect (as they undoubtedly do)

With the longitudinal ones. The blood is colorless.

Of Segmental organs I have observed only three pairs ; they

Appear as coiled granular organs in the last three segments. I

have not found them in the segment which corresponds to the

anterior abdominal setze.

908 THE AMERICAN NATURALIST. [Vor. XXXV.

The nerve cord is a fibrous-looking band encircling the

cesophagus and running back along the ventral wall. In the

region underlying the pharynx the ventral cords, although close

together and connected by several cross branches, are perfectly

distinct ; this portion sometimes shows clearly in living speci-

mens, and I have remarked in it no ganglionic swellings. Ina

stained and mounted specimen, however, when seen in profile,

two ganglia are clear and there are traces of a third.

Upon the reproductive system I have made no observations.

The worms at the season at which I examined them appeared

to have these organs undeveloped.

Every animal examined was in process of active budding. A

comparison of different stages indicates that this process takes

place in the following way. The first part to be formed is

what I may call the abdominal portion; that part, namely,

which bears the abdominal seta clusters and which consists in

the adult of four segments. These segments arise by terminal

budding from the parent ; when two of them have been formed

so that there is a series of six in all, there arises between the

third and fourth a zone in the anterior part of which is differ-

entiated a terminal segment, the fourth segment of the parent,

while its posterior part is converted into the anterior part of

the bud. During this process of differentiation another seg-

ment has arisen at the terminal end of the chain, so that when

its anterior end is complete, the bud has four abdominal seg-

ments. Before it breaks away from the parent, however, a new

individual has begun to develop between the two.

The series upon which these conclusions are based consists

of the following stages į a, a terminal bud just ready to break

away, and possessing four abdominal segments ; b, an older

individual with five abdominal segments, the fifth one the

youngest, as shown by its size and by the number and develop-

ment of its setze ; c, one with six abdominal segments present,

but a zone as yet undifferentiated into segments between seg-

ments 3 and 4 ; d, a similar specimen, except that a seventh

segment has been added at the hind end of the chain; ^ *

specimen like Z, except that two pairs of seta clusters are devel-

oping in the hitherto undifferentiated space between segments

No. 419.] A PARASITIC OLIGOCHETE. 909

3 and 4; f, one like e, but with a deep groove formed between

the two sets of developing setae.

Before the bud breaks away from the parent, a new individ-

ual has begun to develop between the two. In my experience,

which has extended over only a single month, it has been rare

to find a chain consisting of more than three persons, and as in

such chains the older bud is often so far developed as to be

readily detached from the rest by the pressure of the cover, I

am inclined to doubt whether colonies of as many as sixteen

zoóids (cf. Claus, Warzburger naturwissenschaftliche Zeitschrift,

Bd. I, pp. 37-40, 1860) are ever found in our form.

Upon comparing this description with that given by Lan-

kester for the European form, it will be seen that the only

important difference which has been shown to exist relates

to the number and arrangement of the setze in the different

bundles. I am uncertain how much importance should be

attached to this difference. Lankester describes his species

as having twelve setæ in the cephalic bundles and eight in the

abdominal ones, and after having for more than three years

“taken every opportunity of examining the little worms "

states that this number is ** almost invariable” (Quart. Journ.

Micr. Sci, N.S., Vol. IX, pp. 272, 279). Vejdovsky, on the

other hand (System und Morphol. d. Oligochaeten, p. 36), in

describing the same species, says : “ Bezüglich der Anzahl der

Borsten variieren die Bündel bedeutend, indem man auf den

hinteren Segmenten gewöhnlich eine gróssere Menge derselben

vorfindet als in den vorderen Borstenbündeln. Im allgemeinen

trifft man in einzelnen Bündeln, 8, 9, 10, 10-12 Borsten." I

have not, however, in an incomplete but somewhat careful

review of the literature, been able to find published authority

for this statement of Vejdovsky. -

In view of this uncertainty and of the fact that edi tip s

paper, which contains the only full anatomical description of the

European species, is somewhat difficult of access, it has seemed

worth while to give a fairly full account of our own torm. -

nd so admirable that it has not

kester’s plates are so complete a hat

ions to this description-

seemed desirable to add illustrat

ZOÖLOGICAL LABORATORY, WELLESLEY COLLEGE.

OBSERVATIONS ON THE BREEDING HABITS OF

AMEIURUS NEBULOSUS.

ALBERT C. EYCLESHYMER.

ALTHOUGH the bullhead, or horned pout, is one of the most

common of our fishes, but little was known of its breeding

habits until Professor Birge, several years since, discovered the

nests and eggs and made some interesting observations on the

behavior of the fishes during the spawning period. The earlier

naturalists had given us hints as to the breeding time, but these

suggestions were little more than indefinite surmises, partaking

of the character of the following remarks by Thoreau: “The

horned pout are dull and blundering fellows, fond of the mud,

and growing best in weedy ponds and rivers without current.

They stay near the bottom, moving slowly about with their bar-

bels widely spread, watching for anything eatable. They will

take any kind of bait, from an angleworm to a piece of tin

tomato-can, without coquetry, and they seldom fail to swallow

the hook. They are very tenacious of life, ‘opening and shut-

ting their mouths for half an hour after their heads have been

cut off? They spawn in spring, and the old fishes lead the

young in great schools near the shore, seemingly caring for

them as the hen for hér chickens DRM see ninos

set of rangers, with ever a lance in rest, and ready to do battle

with their nearest neighbor."

While the observations made by Profess

been published, some of the facts have been given me in à private

letter, from which I have permission to quote. Professor Birge

writes : * The bullheads on which I made my observations made

' their nests in a shallow bay with sandy bottom, in water not

more than two feet in depth. Some of the nests were In —

not six inches in depth; hardly deep enough to cover cem :

While sitting on the eggs. erflowed land an

This bay was ov din

contained numerous stumps, which had become hollow in the

911

or Birge have never

912 THE AMERICAN NATURALIST. [VoL. XXXV.

course of time, and perhaps three of the nests were in these

stumps, to which access was easily gained by the spaces between

the roots. The others were among weeds and differed consid-

erably. One or two.bullheads kept the weeds away from the

eggs, so that it was not difficult to see the nest from the shore,

while others were concealed so that it was almost impossible to

see the eggs or fish without removing the weeds from above

them.

* There was a surprising difference in the disposition of the

fishes on various nests. One of them was extremely tame. If

approached cautiously he would not swim off, and it was quite

easy for me to put my hand under him and lift him off the nest.

He seemed to enjoy being scratched gently, and when lifted off

the eggs would remain where placed or would swim off a short

distance, and, in general, was very little disturbed by handling.

Others were exceedingly shy, so that as soon as one had

approached within a few yards of them they would dart off,

throwing the eggs out of the nest as they went, with a jerk of

the tail. Of course they always came back and brought the

eggs together again. But this violent treatment of the bunch

of eggs was apt to break it up, and I observed that a consider-

able portion of the eggs was lost in such cases. One of the

catfish, whose eggs were in a stump, was particularly ferocious,

and this was the only one which I found that had a violent dis-

position. I found the nest and put my hand down into the

stump to take some of the eggs, when the fish seized it and

worried it with all his force. After that, I found it necessary

to remove the eggs from this nest with a pair of long forceps;

which the fish would bite in spite of being rapped on the nose

with them rather vigorously. It was this difference in disposi-

tion that especially attracted my attention in studying the

catfish.”

In June, 1892, I secured a number of bullheads from the

small ponds in the vicinity of Worcester, Mass. Some of these

contained ripe ova and sperm, and an attempt was made to arti-

ficially fertilize the eggs, but with ungratifying results. Many

eggs passed through the cleavage stages, but all perished before

the embryos were discernible. Although repeated efforts were

No.419] HABITS OF AMEIURUS NEBULOSUS. 913

made to find the nests, they were unsuccessful until June 8,

1896, when three nests were found in Fowler Lake, Wis. Two

of these were in pieces of stovepipe, the third in an old pail.

The nests were in clear water, near a bold, rocky shore, and at

a depth of four or five feet; all contained embryos, and each

was guarded by a parent fish, — which one I did not ascertain.

On the following day, in searching for other nests, I raised a

small piece of tin pipe and was surprised to find a pair within.

Through the raising of the pipe they became so wedged that it

was impossible for either to get free. They had not yet begun

spawning, although the eggs were so ripe that they were easily

extruded by slight pressure. Artificial fertilization was again

tried, but was only partially successful. A small percentage of

the eggs segmented, most of which died before the embryos

appeared.

During the month of. June, 1898, I chanced to camp near

Mud Lake, Mich., and learned from the fishermen that the lake

abounded in large bullheads. An extended search was made

on June 9, 10, and 11, and we had almost given up the search

when one of my companions found a nest in a small bay with

shallow, sandy shoals. Soon a dozen or more were found along

this sandy shoal, and in a depth of water not exceeding three

or four inches. They were usually concealed beneath logs,

stumps, or boards, which lay against the bank. One would

often observe a slight depression, and upon turning the shelter-

Ing object would find the pair engaged in spawning or watching

over the freshly laid eggs. In two nests which were found

beneath logs on June 11, the parent fishes were moving about

in the small sheltered excavation. The eggs were removed in

each case ; those of one lot were in early cleavage, while those

in the other were in late gastrula. Both nests were visited

on the following day, but the fishes were no longer present.

Another nest, in which the eggs were in late gastrulation

Stages, was uncovered and left exposed. When visited on the

next morning neither fish nor eggs were found. I suspect

that the eggs are devoured by the parent fish when the nests

are too much disturbed, as is known to be the case with some

of the Amphibia. I have, on several occasions, found eggs 1n

9I4 THE AMERICAN NATURALIST. [VoL. XXXV.

the stomach of Necturus. In one instance I partially removed

the eggs from a nest, and upon returning the next day found the

old Necturus in the nest as usual, but the remaining eggs

could not be found. The parent was taken, and an examination

revealed the fact that a number of eggs had been swallowed.

It was interesting to watch the actions of the fish when the

sheltering object was removed. A fence rail covering a nest-

‘ing pair was carefully turned, when the fishes immediately

sought its shelter. As it was turned farther and farther from

the nest they followed, keeping as well secluded as possible, the

while moving restlessly about in search of the nest. When the

rail was finally lifted from the water the male lingered for a

few moments, then darted for deeper water. The female

approached the shore and began searching here and there for

her lost nest. This she passed several times without recogni-

tion, although she seemed to know the surrounding landmarks,

since she would go but a short distance in either direction,

then turning would pass back to the locality of the nest,

which she found in a short time, and despite the fact that it

was unsheltered, she remained. On the following morning the

nest was visited, but again neither fish nor eggs were to be

found.

It is not difficult to allure the fish to.an improvised shelter.

A number of boards were placed on the shore with one end

projecting into the shallow water. The fishes sought these

places and made their nests beneath the boards. It is worthy

of note that in no case did I observe more than a single nest

beneath the same cover, and this quite agrees with the pugna-

cious character of the fish.

Two nests which were occupied by both parent fishes were

left undisturbed and when visited two days later only one fish

was present. It was found by examination that in each case

it was the male. When the female leaves the nest could not

be definitely determined; as nearly as could be ascertained It

would seem to be about the time the embryos begin to move.

The eggs are laid in masses, quite unlike those of most fishes,

but similar in general form to the egg masses of the common

frog. They àre free from pigment and present when beneat

No. 419.] HABITS OF AMEIURUS NEBULOSUS. 915 `

water a rich creamy color. The period intervening between

deposition and the beginning of cleavage is not precisely known.

After cleavage has begun it continues rhythmically as long as

it can be followed, the intervals between successive cleavages

being about thirty minutes. In forty to fifty hours after the

beginning of cleavage the embryo is plainly visible, measuring

2 mm. to 2.5 mm. The larvz at the time of hatching are

about a week old and measure 7 mm. to 8 mm. When they

break through their surrounding envelopes they are quite

unable to support the load of food yolk, but lie on their sides,

now and then making a few vigorous movements. In the

course of a few days they are able to swim about and soon

leave the nest accompanied by the male fish.

During the early summer of 1899 I was able to make some

further observations on the habits of the larvae. In one of

the drainage ditches on Cottage Grove Ave., Chicago, they

were so numerous that in walking a distance of a hundred

yards one would see from seventy to eighty schools. The

larva are usually huddled so closely together that they form a

dark mass, which at a distance appears as a shadow moving to

and fro. They rarely move in straight paths, but are ever

circling, apparently in quest of food. When they come in con-

tact with aquatic plants they pause, carefully search over the

leaves, and again join in a common movement for other

grounds. Frequently one departs from the company and darts

here or there after an insect which may be passing along the

Surface of the water. One never observes them at rest, as 1S

common for the adult. During the night they seem " be

especially active, since it is during this time that they jump

over the edges of the hatching dishes. E

The broods vary widely in numbers; One occasionally

observes a group of forty or fifty and again one oan sms

Several hundred. It was at first thought that either the num-

ber of eggs deposited by the different fishes varied accordingly,

or that a much greater percentage survived in some ice

While both these factors probably modify the numbers, is

Chief cause of these wide variations was discovered pct y

When I chanced to observe four good-sized groups of heo.

916 THE AMERICAN NATURALIST. [Vor. XXXV.

approaching a common point. I awaited with interest their

movements and was surprised to see them unite to form a

single school, which, however, remained intact but a short time.

The larvae soon separated into three groups, each of which

pursued a different course. This procedure was repeatedly

. witnessed, sometimes the union of small groups to form a large

one, again the subdivision of a large one. The fact that the

larvae in some of the broods vary widely in size is thus easily

explained. .A slight disturbance of the water is sufficient to

disperse them, after which they again join in a closely aggre-

gated group. A shadow cast upon the water and quickly

removed sends them scurrying here and there. I was quite

surprised to discover that a low guttural sound caused them

to disperse, while a shrill whistle caused no commotion.

During the summer of 1900 I was able to make some obser-

vations on the behavior of the fish preceding the spawning.

While observing the habits of the black bass in the artificial

ponds of Oakwoods Cemetery, Chicago, I saw a number of

large bullheads swimming about singly and close to the shore.

I thought at first that they, like many of the other fishes in

the lagoons, were exceedingly tame and had sought the shallow

water for the purpose of obtaining food. After several inef-

fectual attempts to induce them to take worms, bits of meat,

bread crumbs, etc., I concluded that they were not feeding,

and since from previous observations I knew this to be their

spawning time I surmised that they might be searching the

shore to locate suitable nesting places. They would frequently

swim so far into the shallow water that the dorsal fin an

upper portion of the body were above the surface of the water.

Here they would wriggle about, and if an indentation or slight

excavation were found they would pass in, move rapidly about,

swim out and on, only to repeat the procedure when another

suitable locality was found. The fishes were always single ;

in no case were two observed even in close proximity.”

On May 15 and 18 increasing numbers of these wanderers

were observed. On May 20 I was gratified to witness what

seemed to me a natural sequence. A cloud of muddy water

attracted my attention, and walking cautiously to within a few

No 419] HABITS OF AMEIURUS NEBULOSUS. 917

feet of the spot I waited until the turbidity cleared, when I

saw a large bullhead lying motionless on the bottom and at a

depth of eight to ten inches. After a period of some five

minutes she swam close to the shore, placed her head in a

slight excavation, and with a violent action of the entire body

threw up another cloud of mud. When the water again cleared

she was observed a short distance from the excavation. At

short intervals the process was repeated. The excavation was

being made in soft clean sand and beneath the sod bank which

formed the shore of the lagoon. I had watched the movements

of the fish for a half-hour or more when a second fish appeared

and at once began to excavate in a manner similar to that

noted for the former, the first fish meanwhile lying motionless

a few feet away. Although both fishes were pretty well

covered with sand, a number of differences were observed.

The latter was much darker than the former, the abdomen a

brighter yellow with less protruding abdominal walls, and

although about the same size, I concluded that the latter was

the male and that they had mated. While the excavating was

now done by one, now by the other, it seemed that the female

took the more active part. After watching the actions ot the

pair for an hour or more I retired and did not again visit the

nest until the morning of the following day, when both fishes

were again observed at work in the same manner. as on the

preceding day. The excavation was now deep enough -

almost entirely hide the fishes, the tips of their tails barely

showing. The nest had been greatly enlarged, as was evidenced

by the quantity of sand which had been thrown out and which

now covered an area of about three or four square feet, with x

maximum depth of four or five inches. The top of m little

mound was hollowed out into a deep, saucerlike depression

from which a broad groove led to the nest.

i her indications that the

still in progress. The female was at this time wane ET

in the saucer-like depression, from which she d i Eos

After considerable jarring over the nest the male was trig

Out. I was not permitted to disturb th

918 THE AMERICAN NATURALIST.

and consequently was not able to make observations on the

interior of the nest; by reaching back in the hole a distance of

sixteen to eighteen inches I could feel the egg mass, and upon

the removal of a portion of the eggs found them to be in late

cleavage stages, and from previous observations inferred that

they could not be more than twelve hours old.

The above fragmentary notes embody, so far as I am aware,

the first published observations on the behavior of the bullhead

during the breeding period. They are recorded with the feel-

ing that they may be of service to those who have so long

sought the embryological material of this primitive teleost. I

may further remark that a study of the breeding habits under

natural conditions shows that Ameiurus rarely burrows in nest-

ing, but instead seeks concealed places beneath logs, stumps,

boards, or even pails or other receptacles which are easy of

access. If, however, a modified environment replaces the

natural, and the places for concealment be no longer present,

as in the artificial ponds, we find the fish adapting themselves

to this changed condition and constructing nests which often

require two or even three days of unceasing labor to prepare.

In closing I cannot refrain from suggesting that a more

extended series of observations would doubtless show that

other fishes, like Ameiurus, manifest an entirely different

behavior during the spawning period as a direct adaptation to

the changed environmental conditions. Indeed, my observa-

tions on the dogfish (Amia) and the black bass living in the

ártificial ponds indicate that this is true of these forms as well.

UNIVERSITY OF CHICAGO.

VO ee

THE SPERMATOGENESIS OF ONISCUS ASELLUS

LINN, WITH ESPECIAL REFERENCE TO

THE HISTORY OF THE CHROMATIN.

M. LOUISE NICHOLS.

Tuis study was begun in February, 1899, and finished in

January, 1901. Its more important results are described in

this paper. A more complete and detailed account of the

spermatogenesis will appear in

a later publication. X

EA.

LS A

. STRUCTURE OF THE TESTIS. Py SI

Each one of the two testes leo

consists of three narrow lobes, F CT eese

isti V J COS

distinct from one another and goo Oc

ois

{os

opening successively into the

anterior expanded portion of

the vas deferens. The interior

t each lobe or follicle is occu-

pied by the germ cells, which :

are in differing stages of devel-

opment in the three follicles of

one side. | Each follicle may

be divided into two principal

regions of growth, composed of

cells of different generations and

of differing degrees of develop-

ment. The illustration (Fig. Dm

will make this clear.1 Itshows, |

in a typical case, the compara-

hai degrees of development to whi

follicles have attained. Thus, in the most posteri

1. — Testis of Oniscus asellus = * b,

ollicles of testis; vas terior part vas de-

f c

? .

erens; f.c., follicle cells; s42, spermatogonia.

ch the cells of the three

or of the

! This and the following figures are slightly schematic.

: 919

920 THE AMERICAN NATURALIST. [Vor. XXXV.

follicles (2) the apical third is occupied exclusively by sperma-

togonia, some of which can be seen in mitosis ; the basal region,

on the other hand, by spermatids in an immature state. Fol-

licle cells occur on the outside of the follicle, being especially

abundant in the basal region. In the adjacent follicle (4) the

apical two-thirds is occupied by cells in the synapsis stage,

the remaining portion by spermatids in a stage of development

later than that of follicle (2). Along the margin of the follicle

are found scattered small groups of spermatogonia (sg). The

third and most anterior follicle (c) contains chiefly spermato-

cytes in a late prophase. Groups of spermatogonia similar to

those of follicle (^) are here also found scattered along the

margin and nearly filling the extreme apical portion. The

follicle cells in the basal region are undergoing not only active

amitotic division, but to a certain extent degeneration. Their

active multiplication or fragmentation causes them to crowd in

towards the axis of the follicle.

From a comparison of the extent of these growth regions in

the three follicles, the developmental cycle may be conceived

somewhat as follows. The spermatozoa, when fully formed,

are forced into the vas. Since they have no motion of their

own, this is probably caused by the contraction of the muscle

layer of the follicle, perhaps assisted by the pressure of the

growing cells in the apical region. During this process the

spermatogonia in the apical portion of the follicle divide and

come to fill up the space left vacant by the discharged sperm.

The majority of the spermatogonia thus filling up the follicle

proceed in their development, while the remainder form the

groups of cells along the margin of the follicle already described

in follicles (4) and (c), and which are destined to supply a new

generation of cells. The spermatids also proceed in develop-

ment and are forced into the vas.

A condition like that represented in follicle (4) thus arises,

the basal region filled with spermatozoa in a late stage about

to pass into the vas deferens, and the apical region with cells

which have progressed as far as the synapsis stage. Later, the

spermatozoa having been completely discharged, the cells of

the apical region come to occupy the basal part of the follicle,

No. j

0. 419.] ONISCUS ASELLUS LINN.

NN. 92I

being now

E. 8s compactly pressed together (cg. Their devel-

progresses until, having become mature spermatozo

they pass i

y pass into the vas deferens ; the spermatogonia again fill

the apical region and the cycle is repeated

II. SPERMATOGENESIS.

I. Maturation.

E i

E. 2 crate aros x m in the history of the chromatin

gent ipie 2 e bis maturation division separates

eae an deo therefore reducing.

EE ae rate by following the changes in the

synapsis stage through the first matura-

“4 E In the anaphase of the last

i E division the chromatin threads

E ogether near the center of the cell

other a almost indisti guishable from each

boss ^ Lo sess spread apart and are

E. r the most part V-shaped (Fig. 2a).

Eu gg an elongation of the threads,

bitrate g this preces the granules of which

n D divide into two, so that

"on osome becomes longitudinally split

) Of the entire number of chromo-

somes r EUM

£e be . *

present it is difficult to be certain, owing to

number, however, is

rmatogonia and not

the number of

therefore, at this stage

o the approximation

to form a bivalent one.

codi each other so closely. The

E uu than that present in the spe

D. an sixteen. The reduction in

icm au apparently takes place,

ioo of | ape so prevalent is due t

The Coins chromosomes t¢

Mo $ ecome more and more à

COR sis are transformed into the nucl

med gma The fact that the ¢

TRR tients just before the forma

M on : toa maintenance of their in

meshes of n preparing for the

the nuclear network

tion of the

dividuality in the rest-

first maturation

become coarser, the

Fic. 2.— a, cell in the

synapsis stage. å, bi-

of the chromatin gran-

ules; nel, nucleolus.

the fact that

the

ttenuated, and finally by

ear reticulum

hromosomes remain

of the

nuclear mem-

division, the

he granules

922 THE AMERICAN NATURALIST. [Vor. XXXV,

more distinct and aggregated into separate threads joined

together by linin. The manner of their origin again lends

support to the view concerning their individuality in the resting

cell. By a gradual process of condensation sixteen compact

masses of chromatin are produced. These

sixteen masses are of various forms. Some

---f--8 are dumb-bell-shaped, two spheres of chro-

matin joined together by linin ; some are

hu crescent-shaped, and stil others are more

oe or less complete rings (Fig. 3). The differ-

oS ent forms may occur in the same nucleus,

but apparently without constancy in the

3 ratio of relative frequency of occurrence.

Two main types may be distinguished

nid Ps sual dor among the chromosomes according to their

hens m (diag structure and mode of origin; z.e., (1) those

in which the bivalent chromosome consists

of two univalent chromosomes lying end to end, as in those

having the dumb-bell shape; and (2) those in which the

univalent chromosomes lie side by side, as in those arising

through a ring or narrow V-shape. A form intermediate

between these is represented by those having a crescent shape.

The different types and their probable mode of origin are shown

in the diagram (Fig. 3).

In the equatorial plate of the first maturation division the

two forms may again be seen. In sections stained with iron

haematoxylin and strongly decolorized a longitudinal split is

distinguishable in the chromosomes. In the

dumb-bell-shaped chromosomes this lies par-

allel to the long axis of the spindle, but in the

others more or less oblique to it, or at right

angles to it (Fig. 4). In the spermatogonic p. 4. — Equatorial plate

divisions, on the contrary, as is shown in prep- of tha bea

arations similarly treated, the longitudinal

split lies transversely to the long axis of the spindle (Fig. 5).

Karyokinesis separates originally distinct chromosomes, and

the first maturation division is therefore reducing. In many

cases the chromosomes can be seen to be composed of a

No. 419.] ONISCUS ASELLUS LINN. 923

double row of four granules. After division the halves con-

sist of a double row of two granules, or four in all, and thus

simulate tetrads.

The true nature of the second maturation division, whether

equational or reductional, is difficult to decide, because the

length and the breadth of the chromosomes are

approximately equal. Since most writers on the

€» maturation of the germ cells in the arthropods

agree in ascribing to them both methods of

Fic.s.—Equatorial division (equation and reduction), it is probable

owl cane that since the first division is reducing, the second

is equational in Oniscus.

These results do not agree with those obtained by Rückert !

and vom Rath? in the ovogenesis of the copepods. The case

of Cyclops.as described by Riickert is partic- ne

ularly clear. According to his observations

the first maturation division is equational,

the second reducing. If Rückert's inter-

pretation of the method of reduction in

Cyclops be correct, and my own concerning

reduction in Oniscus be equally so, it be-

comes clear that the cell generation in which

the true reduction takes place need not be

the same for all members of a given class of

animals.

2. Metamorphosis of the Spermatids.

The transformation of the spermatids of

the isopods has already been described by

Gilson? in his comprehensive work on the

lRückert, J. Die Chromatinreduktion der C

hromo- Fic. 6 — Anterior portion

somenzahl im Entwicklungsgang der Organismen, o

Merk. of mature sperm colony

y magnified).

u. Bon. Erg., Bd. iii, 1893. Jf. flagellum j jar Pre"

Id. Zur Eireifung bei Copepoden, Merk. u. Bon. ETS. in me spa: n

A $ SSe N

nat. Heft, 1894. Chroma- of cytoplasmic fibres.

2Vom Rath, O. Neue Beiträge zur Frage der j

tinreduktion in der Samen- und Eireife, 4. M. A., 46, 1895.

3 Gilson G. Spermatogénèse chez les podes,

1886, tomes i and ii.

La Cellule, 1884 and

924 THE AMERICAN NATURALIST. [Vor. XXXV.

spermatogenesis of the arthropods. In some respects my

observations agree with his; in others they differ.

The nuclei of the spermatids in Oniscus undergo a gradual

elongation and condensation. During this process the cell

; walls between adjacent spermatids dis-

appear and groups of nuclei are formed

lying in a common plasma. Within the

latter arise bundles of fibres of great length.

Gilson in his Fig. 320 shows a direct con-

tinuity of these cytoplasmic fibrils with

the elongated nuclei. That such a con-

nection actually exists I have been unable

to convince myself. In addition to the

bundle of fibres there are single fibres of

. greater delicacy which are continuous with

the nuclei. The appearance of the mature

sperm colony may be seen from Fig. 6.

Figs. 7 and 8 represent longitudinal and

transverse sections of immature colonies. With Wilcox’s double

stain of saffranin and malachite green the cytoplasmic fibres

stain green and are thus sharply differentiated from the nuclei,

which stain red. In cross-section the nuclei are seen as a circle

of red dots surrounding a group of green

dots, the cytoplasmic fibrils. In examining a y

a series of sections from the nuclear region

towards the flagellum, the group of green ,^—c i gi

dots in the center eventually ceases to be (38+ ;

visible, and the surrounding red dots grad-

ually merge into converging green fibres tS pec ener

of great delicacy (Figs. 7 and 8). "The prenuclear region; 4 and <,

evidence at my disposal admits of two pede oo

interpretations, — either the long bundle

of cytoplasmic fibres stops abruptly before the anterior end

of the bundle is reached, and they thus have no connection

with the nuclei, or the connection is of such a character as

to escape observation. Since the structures concerned are

extremely minute and delicate, the latter might easily be

the case.

andit à fibre res

No. 419.] ONISCUS ASELLUS LINN. 925

III. SUMMARY.

The main results of this study may be briefly summarized

as follows :

1. The spermatogonic chromosomes are joined together in

pairs in the synapsis to form sixteen bivalent chromosomes.

2. A longitudinal splitting of the thread takes place at t

stage.

3. The distinctness maintained by the chromosomes up to

the formation of the nuclear network of the resting spermat-

ocyte and the manner of origin of the spermatocytic chromo-

somes from it lends support to the theory of their individuality

in the resting nucleus.

4. In the structure and mode of orig

mosomes two main types may be distinguished : (a) the com-

ponent chromosomes lie end to end, or (b) they lie side by side.

5. Inasmuch as univalent chromosomes are separated, the

first maturation division is reducing.

6. Sphere substance (idiozome) is not observable, except for

a short time during the prophases of the first spermatocyte.

7. The nucleolus of the spermatogonia disappears 7

after dissolution of the nuclear membrane, while that of €

spermatocytes, first discovered in the synapsts persists through-

out the divisions.

his

in of the bivalent chro-

8. The spermatids become associated in groups to form

colonies of nuclei lying in a common plasma.

9. Within the latter arise bundles of fibres of great length,

Whose connection with the nuclei could not be demonstrated,

as well as single fibres of greater delicacy which are continu-

ous with the nuclei. : -

10. The mature sperm colony consists of a variable st a

of filamentous nuclei contained, together with the er :.

Cytoplasmic fibres, in a tenuous sheath which is flage!a

its anterior extremity.

urs very late.

pue occ

1 If such a connection is actual, it 15 slight and

926 THE AMERICAN NATURALIST.

I wish to express here sincere gratitude to my instructors,

Dr. E. G. Conklin and Dr. Thomas H. Montgomery, Jr., for

the valuable advice which has aided me in bringing this work

to a successful completion.

ZOGLOGICAL LABORATORY,

UNIVERSITY OF PENNSYLVANIA,

ON THE FORMATION OF SPECIFIC ANTI-BODIES

IN THE BLOOD, FOLLOWING UPON

TREATMENT WITH THE SERA

OF DIFFERENT ANIMALS.

GEORGE H. F. NUTTALL.

E o si years which have elapsed since I

blood, iraia erigere: er bactericidal properties 1n the

discovered relati pei fluids, a great deal has been

experiments ref pae other propere of the blood. The

Which” led to visae to directly stimulated the researches

agglutinative, h : wai pone: a ipe: shes ot dnt

serum. We k æmolytic, and cellulicidal properties in blood

ber of bodi Av a. learnt of the existence of a num-

ies which neutralize the action of these various

substances.

coil or specific bodies or pre

certain Tm I isis of animals treate

dde acteria, with various kinds of mil

time and different kinds of blood.

Nein. T d the first to demonstrate the exisrence of

cic precipitins in the blood serum of animals immunified

against cholera, typhoid, and plague. His results were subse

E onfirmed by Nicolle and Marmorek. i

dt hé e various anti-sera were added to clear culture fitr?

in th coin: bacterium, and a precipitum was seen to occur

Ms Itrates of those cultures only to which the homologous

“serum was added. Scr dn

cma first demonstrated the existence of specific vieron

se ` He treated animals with MEE After a ie E

Es that their serum when added to 8 milk dilution brought

k ki is precipitation. A precipitum was formed only sien

anti-serum was added to the particular milk against w "

927

cipitins have been seen

d with the products of

k, with peptone, egg

928 THE AMERICAN NATURALIST. | [Vor. XXXV.

the animal had been immunified. These results have been

confirmed by Wassermann and Schütze, and demonstrate that

there are essential differences in the composition of the albu-

minous molecule in the milks of man, cattle, and goat.

Myers demonstrated the existence of precipitins in the blood

of animals treated with peptone, the precipitins acting only

upon peptone.

The credit of having discovered the existence of specific

precipitins in the bodies of animals treated with blood belongs

to Tchistovitch. He inoculated animals with eel serum, which

is toxic, and noticed that an anti-toxin made its appearance in

the serum of the treated animals, but in addition their serum

acquired the property of producing a precipitation when added

to eel serum, whereas it did not act on other sera. A specific

anti-body was also produced in rabbits treated with horse

serum. Bordet demonstrated the existence of specific pre-

cipitins in rabbits which had been treated with fowl's blood.

The anti-serum of these rabbits also produced a slight reaction

in pigeon blood, showing that the bloods of the fowl and.

pigeon are of a somewhat similar composition. Nolf separated

the corpuscles from the serum and treated two sets of animals

with the separated blood ingredients. He found that only

the serum-treated animals yielded the specific precipitin. He

treated the serum with magnesium sulphate, thus removing the

globulin. Animals were treated with globulin and albumin

solutions derived from the serum, and it was found that pre-

cipitins were formed only in the serum of animals treated with

globulin solutions.

Myers treated rabbits with fowl's egg albumin, as also with

serum globulin of the sheep and bullock. He also observed

the formation of specific precipitins in the serum of the treated

animals, although a slight reaction took place on adding the

anti-serum for sheep globulin to that for ox globulin and vice

versa. Uhlenhuth also treated rabbits with egg albumin, and

found that the anti-cserum gave a reaction with I: 100,000

dilutions of egg albumin, whereas the most delicate chemical

tests only gave a reaction with dilutions of I : 1000. He made

the interesting observations that the precipitin appeared in the

No.419.] SPECIFIC ANTI-BODIES IN THE BLOOD. 929

serum of a rabbit which had been fed for twenty-four days

with white of egg.

Leclainche and Vallée treated animals with albuminous

urine and found that the serum of the treated animals con-

tained a precipitin which acted upon the albuminous human

urine with which they had been treated but not upon albumi-

nous urine from the cow and horse.

Uhlenhuth treated rabbits with human blood and that of the

ox, and observed the formation of specific precipitins in their

serum which was tested on nineteen bloods derived from dif-

ferent animals. He obtained a reaction with a solution made

from human blood which had been dried one month. Wasser-

mann and Schütze made similar observations, testing twenty-

three kinds of blood. Stern, who also experimented along

these lines, found that the blood of three species of monkey

gave a slight reaction with the serum of rabbits treated with

human blood. In addition to other observations of a con-

firmatory character Mertens has found that the blood of a

young rabbit born of a human serum-treated mother also

contained the specific precipitin in its blood. The last obser-

vations which I shall mention are those of Dieudonné, and of

Zuelzer, whose results are merely confirmatory in character.

The majority of the publications referred to have appeared

since I began my researches in January.

I have injected rabbits intraperitoneally with the serum of

man, the ox, sheep, horse, dog, and cat, and have been able

to observe the formation of anti-bodies in the sera of all the

rabbits excepting those treated with cat serum. The anti-sera

have been tried on forty-five kinds of blood. ?

The serum of rabbits treated with dog serum, added we

these bloods, gave a negative reaction throughout, excepting

in the case of the dog. The tested dog blood was ades sp

dissolved in salt solution or used in the form of diluted fluid

1 Full details of these experiments will appear in the forthcoming number of

the Journal of Hygiene, vol. i (July 1), No.

2Since the above was written, over one

Fully realizing the interes

the study is

hundred and forty bloods have been

tested — with uniform results. to a i

the point of view of zoólogical classification, being pu

extensive scale.

930 THE AMERICAN NATURALIST. [Vor. XXXV.

serum. Whereas a marked and almost immediate precipi-

tation occurred on the addition of the specific anti-serum to

dog's blood, all the other blood solutions remained perfectly

clear.

The serum of rabbits treated with sheep serum only pro-

duced a marked precipitum with sheep serum or blood solution.

All the other sera and bloods remained perfectly clear, except-

ing those of the axis deer, gazelle and ox, in which a slight

reaction took place, and those of the squirrel and swan, in

which there was very slight clouding.

The serum of rabbits treated with ox serum only produced

a marked precipitation in ox-serum dilutions, or dried ox-blood

solutions. All the other bloods gave a negative reaction, a

slight clouding only being produced in blood solutions of the

sheep, gnu, axis deer, and gazelle, a slight opalescence appear-

ing with that of the squirrel and swan.

The serum of rabbits treated with horse serum only produced

precipitation in dilutions of horse’s blood or serum, not even a

clouding in any of the other bloods noted.

The serum of the rabbits treated with human blood, serum,

and pleuritic exudation, only produced a marked precipitation

in human blood solutions, etc. The blood of the four monkeys

gave a slight but distinct reaction. A very faint clouding at

times appeared in the solutions of the bloods of the horse, 0X;

and sheep, whereas all the other bloods remained perfectly clear.

The test gave positive results when made with diluted human

serum, pleuritic exudation, both fresh and putrid, blood and

serum which had been dried on filter-paper and on glass plates,

with blood which had undergone putrefaction for two months,

with the blood of several persons who had cut themselves

(blood collected on filter-paper), and with the serum from à

blister following a burn on the hand and pressure on the foot.

Both nasal and lachrymal secretion gave a slight but decided

reaction. A faint clouding was produced in normal urine.

That the precipitum formed in putrid blood dilution was Spe

cific was proved by adding the anti-sera of rabbits treated

with ox, sheep, and dog serum to the blood dilution, no

reaction resulting.

No. 419-] SPECIFIC ANTI-BODIES IN THE BLOOD. 931

The tests made with dried blood, whether dried on glass

or filter-paper, gave perfect reactions, as did also 1:100

dilutions kept for two weeks in test-tubes in the laboratory.

Although chloroform had been dropped into the bottom of

these tubes, molds occasionally developed upon the surface of

the serum; but this seemed in no way to interfere with the

specific reaction. Strips of filter-paper upon which both sheep

and ox blood had been allowed to dry were placed under dif-

ferent conditions. Some were kept for two months at 37° C.

in the dark; others at room temperature in the dark, and in

diffused light for the same period ; others again were exposed

for éight days to the action of sunlight in a window. All of

these samples gave apparently just as good reactions as fresh

bloods, though of course our method cannot as yet be strictly

considered to be quantitative. The body in the serum which

is acted upon by the anti-serum, and the specific body in

anti-serum, seem to be about equally resistant. Anti-serum

dried for forty-two days on filter-paper and then dissolved

in salt solution was found to give a perfectly characteristic

reaction when added to its homologous (ox) serum, the

latter diluted (1:100) as usual; it did not, however, produce

a reaction in dilutions of other bloods. Dried normal sera

exposed for half an hour to a temperature of 100? C. still gave

à clear reaction, as did also 1:100 dilutions exposed for half

an hour to'55?. As I first showed, the bactericidal properties

of blood are destroyed at the latter temperature. Dilutions of

blood exposed to a temperature of 100? gave no reaction. ;

The first rabbit in the series treated by horse-serum injections

received old anti-toxic serum which had been kept at room tem-

perature in the laboratory for two years and seven months, We

are indebted to Dr. Louis Cobbett for this serum. The serum,

to which trikresol had been added, had been kept in a corked

bottle, exposed to diffused light, the temperature of the room

being very high during the summer months. The first and ,

second rabbits of the series treated with human pleuritic €xu-

dation, etc., received only one and two injections respectively

of fresh serum, being treated for the rest of the time with

pleuritic effusion which had been kept at room temperature for

932 THE AMERICAN NATURALIST.

five to six months. The pleuritic fluid had been preserved in

a corked bottle with chloroform. These observations seem to

me to possess a particular interest.

It seemed of interest, from a medico-legal standpoint, to

determine whether or no a mzxture of several kinds of blood

would prevent the detection of one of the bloods in the mix-

ture; the presence of another blood might inhibit the action

of the anti-serum. To determine this question 1: 100 dilutions

of two to six kinds of blood were mixed together in equal pro-

portions and tested with positive results.

CONCLUSIONS.

1. The investigations we have made confirm and extend the

observations of others with regard to the formation of specific

precipitins in the blood serum of animals treated with various

sera.

2. These precipitins are specific, although they may produce

a slight reaction with the sera of allied animals.

3. The substance in serum which brings about the forma-

tion of a precipitin, as also the precipitin itself, are remarkably

stable.

4. The new test can be successfully applied to a blood which

has been mixed with that of several animals.

5. We have in this test the most delicate means hitherto

discovered of detecting and differentiating bloods, and conse-

quently we may hope that it will be put to forensic use.

UNIVERSITY OF CAMBRIDGE,

CAMBRIDGE, ENGLAND.

THE COLORS OF THE CRAYFISH.

W. J. KENT.

Some time has been devoted during the past year to a study

of the influence of environment upon the colors of the crayfish.

The work has been confined in a large part to the species

Cambarus immunis.

It was first noticed, while studying the habits of crayfish by

observations in field work, that the color of immunis in nearly

all cases closely resembled the color of the environment. In

one small pond of water, where the soil at the bottom was a

blue clay, the crayfish were all blue in color. In another pond

with a black, muddy bottom they were all black, and in still

other places of different colors. But in nearly all cases they

were of the same color as the environment.

One exception to this was found with those which w

These were confined entirely to the shallow water in the small

streams, and the color was not always similar to the color of

the environment. The crayfish in all colors except red were

found almost entirely in the ponds with deeper water and

muddy bottoms. But it was discovered later that this red

color in crayfish may be caused by exposure to sunlight. This

Was first observed while making a microscopic study of the

pigment, to which most of the color of the crayfish is due.

This pigment when removed from the crayfish and exposed o

the sunlight turned red in a short time. The same result

occurred when all effects of heat were excluded by using

diffused light and an alum bath. a fish

The influence of sunlight upon the color of living po "

Was studied further by experimenting in the laboratory. ded :

fish of various colors were kept in an aquarium exposed to s x

light. This is a difficult experiment to carry out. Uny G

which live in deeper water become strongly ERA E

can endure only very weak light. It was necessary p

933

ere red.

934 THE AMERICAN NATURALIST. [VOL. XXXV.

place the aquarium with the crayfish in diffused light and

to allow them to become accustomed by degrees to changed

conditions.

Some very good results were obtained in these experiments.

Crayfish in black, blue, and green colors were changed to brown

and red under the influence of light. These color changes,

however, were very slow. Several months were required to

produce distinct changes.

Other experiments were made in the laboratory upon the

influence of environment on the color of the crayfish. A num-

ber were taken from different places where they were of

various colors and kept for some time in an aquarium. This

aquarium tank was made of zinc and was partly covered to.

exclude the light. In some cases very distinct color changes

occurred. This was especially true of those which were red

and black. Out of a large number of crayfish of different

colors which were kept in this way for several months, nearly

all were changed to the same gray color of the zinc aquarium.

An.excellent illustration of color changes is furnished by

another species, Cambarus diogenes. These may be found in

any of the smaller streams. In the early spring they vary

much in color, but later in the year nearly all are red. The

explanation here lies in the habits of this species. They

burrow during winter and come out in the spring with more

or less of the color of the soil. These colors are gradually

turned to red in the open sunlight. There has been no oppor-

tunity for observations upon other burrowing crayfish, but

what is true of diogenes is no doubt true of other species

which burrow in winter.

Some little time was also spent in studying color changes in

the young crayfish. This study was made during the months

of April and May of the present year, and was confined to one

species, immunis. The young of immunis are at first red.

This red color, however, is not apparent to the unaided eye.

The pigment layer of a young crayfish consists of a number of

large chromatophores which lie directly below the chitinous

integument. These color bodies are somewhat scattered, and

the little color they give is scarcely noticeable in comparison

NO. 419.] THE COLORS OF THE CRAYFISH. 935

with the colors of some of the internal organs, which can also

be seen. The color of the young crayfish should be studied

with the microscope. The integument of the young crayfish

is perfectly transparent, and with a microscope any changes in

color in the chromatophores is easily observed.

A series of observations was made upon the young crayfish

in one pond where the old were black and in another pond

where the old were blue. In this pond where the old crayfish

were black the young were at first red and gradually changed

to black. This change of color required about two months'

time. In crayfish one or two weeks old only a very few of

the chromatophores showed any change in color, while in cray-

fish one month old possibly one-half of them would be changed

in color, and in those from two to three months old the process

of color change was nearly always complete. The integument

remains transparent up to this age, so that all stages in color

changes were readily seen.

In the second pond where the old crayfish were blue the

young, which were red at first, changed to blue, this change

requiring about the same length of time. In other places

where the old crayfish were red the young crayfish underwent

no change in color. They were red at first and remained red in

color. In this work the young crayfish of different ages were

brought into the laboratory and examined. The color changes

were such as occur among crayfish in their natural environment

and under ordinary conditions. No attempt was made to keep

them in the laboratory for the purpose of experiments.

From these observations it will be séen that the colors of

crayfish are due to two causes. The sunlight produces the red

color, though this same color may be caused by the rarer:

ment. All colors excepting red are due to the influence of

environment. In all cases these colors serve as a protection

against enemies.

The greater part of this work has been confined to one

species, Cambarns imi although some work has been done

on three other species, propinquus, bartonii, and diogenes.

But there seems to be no reason why similar changes in color

should not occur among all other species.

936 THE AMERICAN NATURALIST.

A word should be said here against too hasty conclusions on

this subject of colors. Many species of crayfish are migra-

tory in their habits. This is especially true of those living in

running water. The changes in color require some weeks or

possibly months of time. For these reasons the color of cray-

fish seen in the small streams may not be at all like the color

of the environment. The best results will be obtained from a

study of the crayfish in the small ponds of water where migra-

tions cannot occur.

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Arnold's Sea Beach at Ebb Tide. — This is the type of book

which one wishes had been written years ago. Every one will recall

his struggles when he first began seashore collecting; the vain

attempts to identify the specimens found and in some cases even

to ascertain the relationship of certain forms. Such a volume as

this would have lightened the labors and have given additional

pleasure to a trip to the shore. Although late in coming, it is none

the less welcome and will doubtless be of great value to many begin-

ners. The tendency of the past dozen years has been to stand in

morphological and physiological lines, and the systematic side of

biology has been too much neglected.

This volume simplifies many of the collector’s difficulties, for with

simple descriptions and characteristic figures it enables one to recog-

nize a large number of the animals and plants found on the American

coasts from Mt. Desert to Florida ; from Puget Sound to San Diego.

It begins with hints on collecting and preserving specimens and

then takes up, in systematic order, the alga, sponges, coelenterates,

worms, molluscoids, echinoderms, arthropods, mollusks, and tuni-

cates, describing and usually figuring the more common species.

Each of these groups has a section OF chapter devoted to it, and the

descriptions are prefaced by an outline of the morphology, life-his-

tory, and economic relations of the group. : :

In looking more closely at the text one notices certain blemishes

which are almost unavoidable in à work of such large scope and

which will doubtless disappear !n for future editions

there will certainly be. Thus in the chapter on collecting there

should be directions for killing the animals properly, and there cer-

tainly should be some hints as to the of formal as a peor

fluid. Again, there is noticeable a lack of symmetry in Mr aedi

Thus among the decapods we find that Gelasimus has given way

! Arnold, Augusta Foote. The Sea Beach at Ebb Tide, à guide to the engin

the seaweeds and the lower forms of an life found between tide marks, wit :

more than 6oo Jlustratone. New AP The Century Company, 1901- X +

490 pp

937

938 THE AMERICAN NATURALIST. | [Vor. XXXV.

Uca, while Homarus remains ; in the mollusk the latest vagaries of

the systematist are introduced, while in the worms a more conserva-

tive course has been adopted. Here and there errors occur. Thus

the nemertines are regarded as a class of Plathelminthes, and (p. 164)

the flatworms are stated to lack an anus. Under the Gephyrza,

Sipunculus nudus is included as an American form, while Echiurus

and Thalassema are ignored. Yet these are minor blemishes, and

the work will prove most useful not only to the casual visitor to the

shore but to the more experienced naturalist as well. K.

PHYSIOLOGY.

that Hydra viridis, when attached by the foot and placed in the path

of a constant current of weak intensity, brings the long axis of the

body in line with the current, the oral end being toward the anode.

This orientation is accomplished by a contraction on the anode side

of the body. When the animal is not attached by the foot, the anode

side still remains the side of contraction, even though the oral end

may be turned toward the cathode. In addition to orientation, the

current may call forth general contractions. Separate pieces of the

hydra react in much the same way as whole animals. Buds and

parent animals are independent in their reactions, the buds showing

essentially the same reactions as adults. P.

Evolution of Pigment. — The interest which biologists have

shown in the chemical activities of protoplasm has evinced itself in

the study of pigment as a protoplasmic product. Bohn’s ? contribu-

tion to this subject is a timely résumé of some of the more important

recent results. The pigments are classified as hydrocarbons, deriva-

tives of chromatin, and derivatives of the aromatic series. The

vegetable pigments are described under the heads of chromogenic

bacteria and chloroleucites. The occurrence, migration, and trans-

formation of animal pigments occupies much of the volume. The

author believes that in a given cell there may be a struggle between

arl, R. The Reactions of E to the Constant Current, Amer. Journ.

has vol. v (1901), pp. 301-

2 Bohn, G. L' Evolution du Pami Paris, Carré et Naud, 1901. 96 pP-

No. 419.] NOTES AND LITERATURE. 939

the various classes of pigment granules, resulting in a selection of

the more favored kind. In consequence of this a harmony of color

would prevail, first locally and finally throughout the organism.

Excretion in Annelids. — The elimination of waste products from

the bodies of annelids, particularly earthworms, has been fully studied

by Willem and Minne.! In Lumbricus reserve products in the form

of fat and of glycogen occur, the former in the ciliated cells of the

intestinal epithelium, the latter in the peritoneal cells. True waste

products are found in the same animal as guanine in the chloragogic

cells and nephridial tubules, as uric acid in the peritoneal cells and

similar elements found between the fibres of the body musculature,

and as cholesterine probably in all tissues. The chloragogic cells

produce guanine with more or less regularity. This is periodically

discharged from these cells into the coclomic fluid, where in common

with other particles it is engulfed by the free coelomic cells. The

cclomic cells when charged with the products of excretion make

their way through the intestinal epithelium and are finally discharged

into the digestive cavity The nephridial walls excrete soluble mate-

rials exclusively. Only a small amount of coelomic fluid passes

through the nephridial canal. This fluid is kept in motion by the

cilia of the canal and the waste products are thus discharged.

Similar studies were made on Nereis, Nephelis, and Clepsine, and

the following general conclusion drawn. In all the annelids studied

the cells that line those parts of the caelom particularly connected

with the circulatory system are of service in purifying the blood.

They accumulate in their protoplasm various excretory products, in

some annelids one, in others another. Thus far the following sub-

Stances have been identified : uric acid, guanine, sodic urate, and a

substance like chitine. Many annelids show a tendency toward the

obliteration of the nephrostomes, and this is accompanied by * change

in the way in which the solid excreta are discharged. Cast out

freely in those worms with large nephridial funnels, these products

in worms with restricted nephridia are accumulated and disintegrated

in the phagocytic organs and thus prepared for discharge. P.

! Wi inne, ches sur l'Excrétion chez quelques Anné-

Willem, V., et Minne, A. Recher iengipmgh ges

ech

lides, Mém. "Acad. roy. de Belgique, tome lviii (1900),

940 THE AMERICAN. NATURALIST. [VoL. XXXV.

ZOOLOGY.

Coccide Americana. — The work bearing this title consists of a

series of specimens neatly arranged in a portfolio, with sufficient

printed matter to explain where descriptions of them will be found,

and what plants they infest. The part just issued by Messrs. Quaint-

ance and Scott of Georgia is the second of the series, the first having

been prepared in Florida by Messrs. Rolfs and Quaintance. Each

part contains twenty species, most of them of economic importance,

and all correctly named; so it will readily appear that the work is a

very useful one for experiment-station workers and others who have

to determine Coccide.

The only adverse criticism one can pass upon the part under review

is that the synonymy given for the species is in many cases wrong ;

in several instances the alleged synonyms are not even congeneric.

This results from the uncritical acceptance of the work of other

authors, and the present writer has erred too often in a similar way

to be very severe on the subject. TPA €

Coccide Stanfordiane.

It is a pleasure to receive from Stan-

ford University four excellent papers on Coccida, written by the |

students of that institution. These papers are bound together as

a contribution from the Hopkins Seaside Laboratory, and are as

follows: (1) “Notes on Cerococcus,” by Rose W. Patterson;

(2) “ New and Little-Known California Coccidze," by S. I. Kuwana;

(3) “The Redwood Mealy-Bug;" by George A. Coleman ; (4) “ The

San José Scale in Japan," by S. I. Kuwana. These papers are full

of valuable information, and are accompanied by admirable plates.

Miss Patterson describes and figures the three species of Cerococcus

which occur on the Pacific slope (C. guercus Comst., C. ehrhorni

Ckll., and C. corticis Twns. and Ckll. ), giving many new facts. She

is evida unaware that a fourth species (C. ficoides Green) occurs

in India. Mr. Kuwana gives for the first time an account of the

transformations of Pseudolecanium tokionis, including the description

of the adult male, which will be greatly appreciated by coccidolo-

gists. He also describes three new species in the genera Eriococcus,

Ripersia, and Lecanium. The Ripersia festucæ is a peculiar creature,

having the female elongated, much like the male larva of ordinary

Ripersia. It might be referred to Pergandiella were the antenna

8-jointed ; possibly the discovery of the male will indicate its closer

affinity with Fonscolombia.

No. 419.] NOTES AND LITERATURE. 941

‘Mr. Coleman describes the redwood mealy-bug as Dactylopius

seguoie, but, as Mr. Ehrhorn remarked to me, it is rather a Phena-

Boceus, notwithstanding the 8-jointed antenne. It has certainly no

affinity with the two species of Dactylopius it is said to most resemble.

The description is very full, and includes all stages. T, D. A. C

Fishes of Japan. — In the Proceedings of the United States National

Museum (Vol. XXIII, pp. 739-769) Jordan and Snyder record the

species of fishes collected in Japan by Mr. Pierre L. Jouy. These

are eighty-three in number, six of them being new to science. These

are Leuciscus jouyi, Apogon unicolor, Pomacentrus rathbuni, Aboma

tsushime, Chasmias misakius, and Watasea sivicola. Chasmias is a

new genus of gobies near Gillichthys, and Watasea a new brotulid

near Neobythites. The new species are figured. I may here note

that the name Chasmias is preoccupied by Chasmias Ashmead, a

genus of Ichneumons, published a little earlier in the same proceed-

ings. For the genus of fishes, Chasmichthys Jordan and Snyder

may be substituted.

In the same paper is given an identifi

Japanese fishes collected by Dr. Thunberg and loosely described by

Houttuyn in 1782. The adoption of Houttuyn's names necessitates

several changes in nomenclature, among others the use of the name

Scomber japonicus in place of Scomber colias, for the common chub

mackerel.

Jordan and Snyder have begun a series of monographic reviews of

families of Japanese fishes. The first now published (Proceedings of

the United States National Museum, Vol. XXII, pp. 725-734)

includes the lancelets and lampreys, the second, the eels. In the

ies are Branchiostoma nakagawe,

cation of the species of

script name of Dr. Hatta.

In the review of the eels, fifty species are described, of which

nineteen are new, all of these and some of the others being figured.

The new genera are Xyrias, near Cirrhimurena, but without cirri,

and /Emasia near Gymnothorax, but with the mouth bristling with

large canines. ns51

Fowler on Fishes in the Philadelphia Academy. — In the Pro-

Philadelphia (Vol. LIII)

ceedings of the Academy of Natural Sciences at

Mr. Henry W. Fowler gives a number of interesting notes on fishes.

The types of new species of selachians in the academy museum are

942 THE AMERICAN NATURALIST. — [Vor. XXXV.

redescribed with special reference to their anatomy. Among these

are numerous types of species named in Bonaparte's Fauna Stalica,

purchased for the academy by Mr. T. B. Wilson.

The fishes recently noted by Mr. Fowler as from the Caroline

Islands came from a coral reef called Caroline, or Thornton, Island,

remote from the Carolines, and near Samoa.

From Mazatlan, Mexico, Mr. Fowler describes a new genus of

Hemiramphide with the form of Fodiator and the long beak of

Hemiramphus. The species is called Hemiexocetus caudimaculatus.

It is midway between the halfbeaks and the flying fishes.

The generic name Odontostomus, applied by Cocco to a deep-

sea fish, is preoccupied in mollusks. In place of Odontostomus,

Mr. Fowler proposes the new name Evermannella. The family, now

composed of the two genera, Evermannella and Omosudis, becomes

Evermannellide. p; S.J

Jordan and Snyder on New Fishes in the Museums of Japan. —

In the Journal of the College of Science of the Imperial University of

Tokyo(Vol. XV, Part II), Messrs. Jordan and Snyder have descrip-

tions of nine new species of fishes in Japanese Museums of which

no specimens are yet available except the original types.

These species are :

Acipenser &ikuchii, from Sagami Bay.

Lepidopus aomori, from Aomori.

Tetrapturus mitsukurii, from Misaki and numerous other localities.

Tetrapturus mazara, from Misaki.

Bentenia esticola, from Kashima.

Ebisus sagamius, from Misaki.

Reinhardtius matsuure, from Misaki.

Trachypterus ishikawe, from off Tokyo.

Trachypterus ijime, from off Tokyo.

Of the two new genera, Bentenia is nearest Velifer and Pteraclis,

and is distinguished by the anterior insertion of its dorsal and anal,

each of which has one spine greatly enlarged. :

Ebisus is a huge bass or jewfish, allied to Stereolepis but with

the head unarmed and the teeth larger. It is locally known as

Aburabozu, the “fat priest."

Dr. Mitsukuri has illustrated the. paper by photographs of the

type specimens, in so far as these are available. WSF

No. 419. | NOTES AND LITERATURE. 943

' Check List of the Fishes of Japan. — In the series called dvno-

tationes Zoologica Japonenses (Vol. II), published by the Zoólogical

Society of Tokyo under the auspices of the Imperial University of

Tokyo, Jordan and Snyder give a “ Preliminary Check List of the

Fishes of Japan." This catalogue is a neatly printed memoir of

159 pages.

It includes 686 species, exclusive of species from the Kuriles

and Riu Kiu, mentioned in footnotes. The known localities in

Japan are enumerated for each species, and the Japanese vernacular

name of each is given. These vernacular names, corrected and

largely compiled by Dr. C. Ishikawa, give the paper à special value

to Japanese students.

The paper was in press before the authors planned their expedi-

tion to Japan in 19oo. In this expedition upwards of 250 species

were added to the list and a number of changes and corrections will

be rendered necessary. The total number of species of fishes now

known from the entire empire is between 1000 and 1100.

. We may note that the new genus Corusculus (berycoides) is identical

with the earlier Eteliscus, having the same type The appearance

in the text of this needless name was the result of accident.

The paper is *dedicated to Professor Kakichi Mitsukuri of the

University of Tokyo in recognition of his work as a naturalist and

of his character as a man." It is to be hoped that this paper,

Written in America and published in Japan, may help EE the pur-

pose of bringing Japanese science and that of the nations called

Occidental into closer touch with each other. D. S.J.

Notes. — The spouting of the finback whale has been successfully

observed by Henking (Zool. Anzeiger, Bd. XXIV, pP- moo m

While off the Norwegian coast. 'The spout is not in any sense a :

of water, but is a cloud of water particles, a5 maintained first y

Scoresby, and is driven out of the animal's nostrils by an -i

not unlike a sneeze. The cloud usually has the outline "s ond

comma and Professor Henking's paper is noteworthy as con E

the fi : i ine.

e first seca drawings of this outline gap e

been

the entoderm i uth cavity of vertebrates has :

rm in the mo y layer the hypophysis is

Obstacle in determining from which germ A Me no

derived. Zeleny (Biological Bulletin, Vol- TI, pp. 267-281) Ms

i i med before the mout

found that in turtles this organ 15 for

through and that it is derived from the ectoderm.

944 THE AMERICAN NATURALIST. [VorL. XXXV.

In a synopsis of the rice rats (Oryzomys) of the United States

and Mexico, Merriam (Proceedings of the Washington Academy of

Sciences, Vol. III, pp. 273-295) adds twenty-two new species and

subspecies to the thirteen already known.

Phoronis pacifica, a new species described by H. B. Torrey (ie

logical Bulletin, Vol. II, pp. 283-288), is interesting as the first of

this almost cosmopolitan genus to be taken on our west coast.

BOTANY.

Plant Life of Alabama.'— For nearly forty years the late

Dr. Mohr gave close attention to the flora of his adopted home,

Alabama; and the later years of his life were devoted to the prepara-

tion of a book embodying the results of his long study. He lived to

see the last proof sheets corrected, but the date of its publication is

a fortnight later than that of his death. A historical account of the

botanical work done in Alabama, a detailed study of its physiography,

geology, and meteorology, and a discussion of the general principles

of plant distribution, lead to an analysis of the features marking the

Alabama flora, which, more than any other North American work,

exemplifies the trend of modern ecological study in a varied region ;

and the book closes with an annotated systematic catalogue of the

spontaneous plants, including descriptions of new or critical species,

a discussion of the relations of the flora to agriculture, and a list of

plants known to be cultivated in the state. Considering Dr. Mohr's

advanced age, it is remarkable that he should have adapted himself

to the current trend of American botanists in regard to nomencla-

ture, but in this, as in his grasp of the newer field problems in

botany, he shows a plasticity far out of the ordinary run, and his

work, which is of lasting value, is likely to produce nowhere the

impression that a younger man, if possessed of his knowledge, could

have done it better. Ww. T.

Notes. — The varied and far-reaching character of the botanical

work being done under the geological and natural history survey of

Minnesota is shown by part s of the current series of Minnesota

1 Mohr, Charles. Plant Life of Alabama, Contr. U. S. Nat. Herb, vol vi.

921 pp., with several plates. Washington, Government Printing Office, 1901-

NO. 419.] NOTES AND LITERATURE. 945

Botanical Studies, bearing date July 20, which contains: A prelimi-

nary list of Minnesota Uredinez, by E. M. Freeman, A new species

of Alaria, by De Alton Saunders, A preliminary list of Minnesota

Xylariacee, by F. K. Butters, A contribution to the knowledge of

the flora of the Red River valley in Minnesota, by W. A. Wheeler,

Observations on Gigartina exasperata, by H. B. Humphrey, Obser-

vations on the algz of the St. Paul city water, by M. G. Fanning,

Notes on some plants of Isle Royale, by W. A. Wheeler, Revegetation

of Trestle Island, by D. Lange, Violet rusts of North America, by

J. C. Arthur and E. W. D. Holway, and Observations on the embry-

ogeny of Nelumbo, by H. L. Lyon. The number is unusually well

illustrated.

Of interest to botanists, as well as ornithologists, is a pleasantly

written account, by Dr. O. Widmann, of a visit to Audubon's birth-

place, separately printed from Zhe Auk of April.

Mr. Massee continues his redescriptions of Berkeley's types of

fungi in No. 243 of the Journal of the Linnean Society.

Science of July 26 reprints from the London 7Zimes an article on

the recent report of a committee appointed for the investigation of

the botanical work carried on at Kew and South Kensington.

A biographic sketch of the late Prof. T. C. Porter, with portrait,

appears in the July number of the Bulletin of the Torrey Botanical

Club.

The flora of the Palouse region of Washington is the subject of a

Paper by Piper and Beattie, published in’ May by the Agricultural

College of that state. It contains descriptions of all of the Sperma-

tophytes and Pteridophytes known to grow wild within 35 kilo-

meters of the town of Pullman, and is provided with keys to the

families, genera, and species. In nomenclature, the Kew and

Berlin rules have been followed.

A recent number of the Gardeners’ Chronicle contains a figure,

Copied in the Revue Horticole of July 16, showing an Abyssinian land-

Scape characterized by arboreous species of lobelia, which i "

striking as the yucca, cactus, and other bizarre landscapes of arid

regions on the American continent.

Mr. J. Medley Wood's Natal Plants, figuring and describing caes

of the more interesting constituents of a most interesting flora, has

reached part 2 of the third volume.

946 THE AMERICAN NATURALIST. [VOL. XXXV.

The first part of Vol. V of the Flora Capensis, edited * by

Sir W. T. Thiselton-Dyer, extending from the Acanthacez to Clero-

dendron in the Verbenacez, has recently appeared from the press

of Lovell Reeve & Co., of London, — unfortunately without indica-

tion of the date of issue.

An interesting essay on palms and their value to residents of

tropical countries, by Professor Schróter of Zürich, constitutes the

last /VeujaArsb/att of the Naturforschende Gesellschaft of that city.

In connection with a monograph of garden beans recently pub-

lished from the Missouri Botanical Garden, a paper by Professor

Halsted on bean diseases and their remedies, constituting Bulletin 257

of the New Jersey Agricultural Experiment Station, becomes of double

interest. Like all of Professor Halsted's papers, it is well done.

PETROGRAPHY.

Origin of Corundum. — The interesting problem relative to the

origin of corundum in basic rocks has been attacked by Pratt'

through the study of slags. He finds that the separation of corun-

dum from magmas is dependent upon the composition of the magma,

upon the character of the oxides dissolved in it, and upon the

quantity of alumina present.

When the magma is a calcium-sodium silicate, corundum separates

only when the ratio of Al O; to the other bases exceeds 1: 1, and the

ratio of SiO, is less than 6. If Mg and Fe are present, no corundum

will form unless there is more than enough ALO, present to unite

with these bases.

When the magma is a magnesium silicate without excess of Mg,

all the AlO; will separate. If Mg is in excess, some of the Al;O;

will unite with this, forming spinel, and the remainder will separate.

When Cr;O, is present and only a little AlO; and MgO, these unite

with the Cr,O;, yielding chromite. No corundum is formed.

When alkalies or alkaline earths are present, the Al,0; tends to

unite with these in the formation of feldspars. There is, however,

little tendency to the formation of Mg-Al silicates when the magma

is a magnesium silicate.

1 Amer. Journ. Sci. (1899), p. 277.

No. 419.] NOTES AND LITERATURE. 947

The Calculation of Rock Analyses. — Now that so much interest

centers about the chemical composition of rock magmas and the rep-

resentation of their composition in terms of molecular ratios, a recent

paper by Kemp! on the methods of calculating rock analyses in these

terms will be of great use to all students of rocks. In this paper

the author shows how to transform percentages into molecular pro-

portions, and from these how to calculate the mineral composition of

any given rock. The most valuable portion of the article is a series

of tables in which the * molecular proportions " of each of the rock-

forming oxides is indicated for its corresponding “ percentage "M

rock analyses.

Weathering of Granites. — The conclusions drawn by Watson °

from the results of an interesting study of the weathering of a num-

ber of granitic rocks of Georgia are as follows : Assuming that Fe:0;

has remained constant, (1) the amount of water in the weathered

rocks increases rapidly as decomposition advances. At the same

time there is a loss of SiO, and of all the metallic oxides except

ALO; which in some cases shows a relative increase. (2) The loss

of SiO, is not caused by solution of the quartz of the original rocks,

but is the result of the decomposition of silicates. (3) CaO and

Na,O disappeared in larger quantities than MgO and K,O. (4) The

total loss of constituents varies between 7.68 % in weathered phases

to 71.82 % in thoroughly decomposed forms.

Origin of Phenocrysts in Granites. — The same author 4 has also

investigated these granites with respect to the origin of their pheno-

crysts. He describes in detail a large number of occurrences and

concludes his study in these words : * The absence of (a) definite

arrangement or orientation among the phenocrysts;

crysts from the border zones of the massif — grada .

interior porphyritic facies peripherally into an even granular granite

of coarse texture and the same mineral an

(€) the further absence of evidence of magmatic resorption oe

rosion of the phenocrysts ; and (7) the presence or abundant "e

sions of all the ground-mass constituents characterizing the pepe

tabular phenocrysts of the Georgia porphyritic granites, fully yustuy

the conclusion that the phenocrysts in these rocks were formed 77

Plac, and are not intratelluric in origin." -

tion from an

1 School of Mines Quart., vol. xxii, p- 75:

2 Bull. Geol. Soc. Amer., vol. xii (1901), P- 93-

3 Journal of Geology, vol. ix (1901), P- 97-

9498 THE AMERICAN NATURALIST.

The granites are found in the Piedmont plateau region. They : :

comprise even-grained and porphyritic varieties, and granite gneisses, E

all of which are plainly intrusive. Each variety is clearly described E

by the author, who also gives analyses of many types. Averages of i

these analyses give the following figures : E

SiO, Al,O3 FeO; MgO CaO NaO K,0 P

Normal grani 69.67 16.63 1.28 AS 2.16 4.73 471

Porphyritic xai 69.28 16.73 1.25 72 2.13 4:33 4:59 ;

Granite-gneisses 73.76 14.52 1.03 a 1.14 4.16 4.63 Ro

The normal and porphyritic phases possess the same composition. a

The gneisses, however, are more acid than these, while their percent- 2

ages of Al,O3, CaO, and MgO decrease.*

Gneisses of the Adirondacks. — The gneisses of a portion of the

Adirondacks are briefly described by Cushing? in a report on the

geology of Franklin County, New York. They comprise granite-

gneisses with the composition of hornblende-granites, and pyroxene-

gneisses. The latter consist of pyroxene (augite and hypersthene),

plagioclase, orthoclase, some hornblende and quartz. Intermediate

gneisses composed of hornblende and andesine, with augite and

hypersthene as common accessory constituents, are also present in

some localities. These are identical in their features with certain

hornblende-gneisses derived from gabbros, but the author is inclined

to separate them from the latter as of different age. Intrusive in -

these gneisses and in the Grenville series of sediments which are so -

well represented in the district are great dykes and masses of anor- -

thosite, gabbro, granite, diabase, and syenite porphyries. The vari-

ous types of most of these rocks have been described many times.

The author adds new descriptions which serve to show that the types -

are quite uniform over large areas. The syenite grades into granite,

both rocks being regarded as differentiates of one magma. T |

nites are composed essentially of orthoclase and albite or oligoclase

in microperthitic intergrowths, augite, hypersthene or bronzite nd

quartz. Hornblende is nearly always present to some extent. With

the increase in this component the hypersthene diminishes. — The

rock varies rapidly in composition and structure. All the intru

except the diabases and porphyries have been subjected to |

pressure and have yielded gneisses.

1 Amer, Geologist, xxvii (1901), p. 199.

? 18th Report State Geologist, Albany, N. Y., 1900.

(No. E was mailed October 14)

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AMERICAN. 3

NATURALIST -@

aS A MONTHLY JOURNAL

E: à DEVOTED TO THE NATURAL SCIENCES

i IN THEIR WIDEST SENSE

CONTENTS

L Regeneration in the Egg, Embryo, and Adult 2

3 Denon T. 101 tte

Stichostemma » Lx dé ae

meme WO EE 1007

II. The Habits and Natural History of

v. te i tD 2 1023

oS a |

Professor 1

“BOSTON, USA.

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: EI the ge penal scientific

The American Naturalist.

ASSOCIATE EDITORS:

J. A. ALLEN, PH.D., American Museum of Natural History, New York.

E. A. ANDREWS, PH.D., Johns Hopkins University, Baltimore.

rsity

DOUGLAS H. CAMPBELL, Pu. D., Stanford University.

J. H. COMSTOCK, S.B., Cornell University, Ithaca.

WILLIAM M. DAVIS, M. E., Harvard eie Camóridge.

ALES HRDLICKA, M.D., Mew Yor£ Ci

D. S. JORDAN, LL.D., Stanford Univer.

CHARLES A. KOFOID, Pu.D., Usieeraty. of Zllinois, Urbana.

ISRAEL C. RUSSELL, LL.D., University of Michigan, Ann Arbor.

ERWIN F. SMITH, S.D., U..S. Department of Agriculture, Washington.

LEONHARD STEJNEGER, Smithsonian Institution, Washington.

W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis.

HENRY B. WARD, Pu. D., University of Nebraska, Lincoln.

WILLIAM M. WHEELER, Pu.D., University of Texas, Austin.

THE AMERICAN NATURALIST is an illustrated monthly magazine

of desees History, and will aim to present to its readers the leading

facts and discoveries in Anthropology, General Biology,

Botiniy. Paleontology, Geology and Physical Geography, and Miner-

and Petrography. The contents each month will consist of

. leading original articles containing accounts and discussions of new

discoveries, reports of scientific expeditions, biographical notices of

inguished naturalists, or critical summaries of progress in some

— -line; and in addition to these there will be briefer articles on various

. points of interest, editorial comments on scientific questions of me

day, critical reviews of recent literature, and a final department fo

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December, 190r. No. 420.

REGENERATION IN THE EGG, EMBRYO,

i AND ADULT.

T. H. MORGAN.

term *regeneration" has come to mean not only the

"cement of a lost part, but also the production of a whole

nism from a piece of an animal or of a plant. There are

Other phenomena that must be included under the same

» Such as, the production of a part different in kind from the

t, and also, as I shall try to show, those cases in which

of an egg or of an embryo produces a whole or a part of

ism. It appears that there are two principal modes by

I regeneration takes place. The first and more familiar pro-

followed by the development of new organs out of this

by means of the formation of new tissue at the exposed

950 THE AMERICAN NATURALIST. [Vor. XXXV.

It has been only within the last few years that zoólogists

have discovered that parts of the embryo, or even of the unseg-

mented egg, have also the power of producing new, whole

organisms, and while this process has been here and there

brought into relation with the process of regeneration, — espe-

cially by Roux and by Barfurth, — yet up to the present time

no systematic analysis has been carried out in order to see

how far the regeneration of pieces of an adult organism and

of pieces of the egg are similar or identical processes. It is

the special object of this paper to examine further into this

question.

In order to make clear our subsequent comparison between

the development of pieces of the adult and of the egg and

embryo it will be necessary in the first place to review briefly

a few well-known facts. It has been stated that there are, in

general, two modes by which a piece of an adult may regener-

ate: (4) either by the development of new tissue at the

exposed regions, — epimorphosis; or (2) by transformation of

the old part into a new form, — morphallaxis.

(4) a. Furthermore, we find under the first category that

when a small piece of the organism is removed (1) the organ-

ism regenerates as much as is lost ; but (2), on the other hand,

the small piece does not generally make the whole organism,

if in fact it regenerates at all. As examples of these different

processes I may cite the following cases. If the leg is cut

from a newt, the larger piece — the newt — makes a new leg,

but the leg does not make a new newt. If the head is cut

from a snail, the snail makes a new head, — if the cut has

not been made too far back, — but the head does not make a

new snail. If it be objected that in both cases the smaller

piece dies before regeneration can begin, I may cite another

experiment made on the earthworm. If a few anterior seg-

ments are cut off, they do not make a new worm, although the

piece may remain alive for a long time without showing any

signs of regeneration, and during this time the larger piece,

from which the anterior end had been cut off, may have

No. 420.] REGENERATION IN THE EGG. 951

completely replaced the lost smaller part. It may also be stated

that the result is not dependent upon a lack of material in the

small piece, for the process of regeneration does not even begin,

and, as other experiments have shown, there is certainly enough

material in the small piece for regeneration to have been carried

out to some considerable extent.

b. The other case in this category is that in which the large

piece produces only the distal end of the part removed and,

therefore, less than the part cut off. For instance, if more

than five segments are taken from the anterior end of the

earthworm, still only five are regenerated, and the intermediate

region is never regained. In the planarian, if more than the

head is cut off, only the head is aż first regenerated, but in this

case the intermediate, or missing region, is made good by sub-

sequent changes that take place in the region behind the new

head and in front of the old part. A similar thing happens

when the posterior end is cut from an earthworm. At first

only a small part comes back, consisting of the end-piece — the

telson — and a few segments, but by subsequent growth td

segments are intercalated near the posterior end just in front

of the telson. Again, if the tail of a fish is cut off near its

base, there appears at first only a small new par t that assumes

the characteristic form of the tail, even before the new part has

the tail was cut off. Subse-

grown out to the level at which ud

quently, by a process of continuous growth, the entire

replaced. ; eid.

(B) a. The second mode in which the process o ge m

tion takes place is, as I have said, by the transformation

entire piece into a new organism.

The most familiar case, and the one longest known, serit

hydra. Trembley saw that small pieces Se ved viis

whole animals, and subsequent investigators have mms cifically

same thing. They have not, however cune quisque

that new tissue is not added at the cut-ends, but from pe

Observation I can state that there is

new tissue in these regions. The

no special development of

entire piece changes over

952 THE AMERICAN NATURALIST. [Vor. XXXV.

into the new form. Another example of the same process,

first made known by Bickford, is that of Tubularia. In this

form it is even more evident that a piece of the stem may be

transformed into a new hydranth without new tissue forming

beyond the level of the cut-end.

In planarians we find not only new tissue developing at the

cut-end, but also a transformation taking place in the old parts.

The Protozoa also regenerate by changing over the old part

into a new form. From the figures given by Gruber and

others there can be little question that the piece is transformed

directly into a new whole, although this important fact has not

been emphasized by those who have written on the subject.

A piece of Stentor, for example, if it contains one or more

nodes of the macronucleus, produces a new Stentor having the

characteristic form, but on a smaller scale. I have repeated

this experiment on Stentor, and have found that small pieces

produce new organs that are of proportionate size.

In these cases of morphallaxis, just described, the small

pieces change over into the characteristic form of the species,

ie, they are new wholes of smaller size. They may subse-

quently grow to the full normal size, and even produce sexual

organs. On the other hand, there is known at least one case

in which very small pieces produce not a whole form, but only

a part of a whole. It has been shown that very short pieces

of the stem of Tubularia may produce only the distal end of the

hydranth, vzz.,a proboscis. Other experiments show that, never-

theless, the material out of which the proboscis develops is

totipotent. This problem will be more fully discussed later in

another connection. The result also suggests direct compari-

son with those cases in which only a part of what has been

removed is regenerated out of the new tissue that appears at

the cut-end, as when only a few anterior segments regenerate

in the earthworm after the removal of a much greater number.

In fact, I think it may be shown that there is something more

than a mere similarity between the two cases, and that they

both result from the action of the same factors.

We may next examine some of the cases in which a part of

the egg, or of the embryo, produces an entire organism. The

No. 420.] REGENERATION IN THE EGG. 953

earliest observation of this sort was made by Haeckel in 1870.

Pieces of the embryo of a jellyfish were found to produce each

a new smaller embryo which gave rise to an entire organism.

This discovery attracted little or no attention at the time,

because the important theoretical questions that are involved

were not appreciated. Without treating the facts in the order

of their historical sequence I shall describe a few other results

that have a direct bearing on the questions before us.

The development of isolated blastomeres is one of the sim-

plest cases of the kind. If the first two blastomeres or cells of

the egg of the sea-urchin be separated from each other, each con-

tinues to segment, in most cases as though it were still a part

ofa whole. There is produced a half blastula open at the side

that corresponds to the region at which the other blastomere lay

at the two-cellstage. The half sphere next closes in to become

a whole sphere of half size, and from this time onwards the

embryo continues to develop as a symmetrical whole, passing

through the later stages as does the normal, whole embryo.

. These results show that after separation of the blastomeres

each continues to develop for a time in the same way that it

would have done had it remained in contact with its fellow;

but the process of half development is only carried through the

cleavage stages, and then a new change takes place, initiated

by the closing of the open side, and a new whole of half size is

‘established. It is this process of closing of the opening and

the subsequent whole development of the half piece that resem:

ble the changes that take place in the regeneration of pieces

of adult animals.

In other species of se

behave in a somewhat different way.

cleavage is not so obviously a half cleavage,

the first divisions may be more or less like those of a hilasto-

mere that remains in contact with its fellow, yet there x not

formed an open half sphere, but the blastomeres close in as

they are formed. A whole embryo of half size is also pro-

duced in this case.

In another sea-urchin, Toxopneustes,

after separation of the blastomeres the c

a-urchins the isolated blastomeres

In Spheerechinus the

or at least while

it has been found that

leavage may proceed

954 THE AMERICAN NATURALIST. [VoL. XXXV.

at once as a whole, z.e, the isolated blastomere divides in the

same way that the whole egg divides, and not as it would have

divided had it remained in contact with its fellow. The blas-

tula is closed from the beginning. Later, as in the other cases,

a whole embryo develops.

These and other experiments seem to show that the form of

cleavage of the egg is the result of an arrangement or structure

of the protoplasm, and that in some cases this same arrange-

ment is retained after the separation of the blastomeres; while

in other cases the protoplasm may rearrange itself at once

into a new whole, in which the arrangement of the parts is

symmetrical and like that of the whole egg. The blastomere

divides in consequence in the same way as does the egg.

The closing in of the half blastula of Echinus presents cer-

tain obvious resemblances to the closing in of pieces of adult

animals. Several writers have supposed that in both cases

this change is due to a simple physical rearrangement of the

material, but it is, I think, not altogether improbable that at

least one other factor is also present. I base my opinion on the

study that I have made of the closing in of pieces of Tubularia,

of hydra, and of the closing of large wounds made in the side

of very young tadpoles. While the closing in seems to be the

result, in part, of the physical property of the semifluid sub-

stance to become round, or more symmetrical, it seems also

to involve certain contractile phenomena of a different sort.

The cells that are in contact draw towards each other as à

result of their own movement or contractility. There may be

also certain tactic phenomena present. At least, if we cannot

show positively that some such factors may be at work we have

at least no right to exclude them in the present state of our

knowledge as possible factors in the result.

After the closing of the half blastula has taken place we find

that the piece no longer behaves as a part, but as a new whole.

We must assume that it has become a symmetrical or whole

structure. Let us look somewhat further into this question.

We have seen that in the case of the egg the change that

takes place in a piece, making the piece symmetrical, may be

brought about by a simple physical rearrangement of the

No. 420.] REGENERATION IN THE EGG. 955

substances of the protoplasm; but after the piece has been

divided into parts or cells the contents of each cell is separated

from that of its neighbor by the cell-walls which we have come

to look upon as barriers preventing free interchange between the

cells. In many cases, however, an actual continuity of the pro-

toplasm from cell to cell has been demonstrated, and it has been

recently shown that connections may be established during the

cleavage period between neighboring cells. We shall not go

far wrong, I think, if we assume that the protoplasm through-

out the embryo is a continuous structure, and that it is not

shut up in protoplasmic-tight compartments. How, then, from

this point of view can we look upon the changes that take

place in an unsymmetrical piece so that it becomes symmet-

rical? Can we regard the first step to be the same as in the

piece of an egg, Ze, to be the outcome of a symmetrical rear-

rangement of the material or substances contained in the

egg? We may think of this as possible, but the change in itself

would not be sufficient to account for the subsequent events,

unless we assume that it leads to the symmetrical organization

of the living substance itself, for on this rather than on the

gross contents of the protoplasm the subsequent changes seem

to depend. That this must be so is shown, I think, by the

following considerations.

Numerous experiments have demonstrated that any part of

the egg may produce a whole embryo — provided, of course,

that it contains the nucleus and is large enough. The sub-

stances contained in the different parts of the protoplasm are

very similar throughout the egg, although there may be mov

orless of one or of another sort in each region. Since any

part of the egg can produce an embryo we cannot regard

the presence or absence of any of these substances as of fun-

damental importance; hence it is probable, I think, that the

result depends on the protoplasm rather than on the substances

it contains. i *

The structure of the egg is à symmetrical ory d

development shows; a piece of the egg retains at Bre T "e

Structure in most cases as its cleavage demonstrates, spe

may later become also symmetrical, as the later changés

956 THE AMERICAN NATURALIST. |. [Vor. XXXV.

prove, and we are led to ask, What is the nature of this sym-

metry! or organization that exists in the protoplasm of the

egg, and which may appear in the protoplasm of a part of the

egg? The problem is all the more difficult to understand

when we find that an egg that has divided on a radially sym-

metrical plan may produce a bilateral embryo. It is probable,

however, in such cases that a bilateral structure is really pres-

ent from the beginning of the cleavage, but that the factors

that are at work during the cleavage are not necessarily those

that determine the bilateral organization of the embryo. In

some cases, as in the frog's egg, a careful examination even of

the early cleavage stages shows that a bilateral organization is

present in the protoplasm, although the form of the cleavage

may appear in many cases to be radially symmetrical.

In order to answer our question in regard to the nature of

the organization we should have to know more of the nature of

development itself. All we can do at present is to examine

some of the implications that are involved in the assump-

tion, and at least attempt to make clear our position. The

new axial relations that are established in the piece present

certain interesting relations. If the original organization was

a bilateral one that corresponded, let us assume, more or less

with the plane of the first division, then an isolated blastomere

has at first only the organization of a half, or rather contains

the factors that lead to a half development. In other words,

it has no median plane of symmetry, yet later such a median

plane is established. It is this change, taking place in the

isolated part, that we are forced to assume, that gives us one of

the most interesting and also important problems with which the

student of experimental embryology has to deal. We know of

nothing similar taking place in inorganic nature. The most obvi-

ous change that may seem to approach this is in the formation

of an entire crystal from a piece, but in this case there is never

any change in the position of the original axes that the piece

has inherited from the old crystal; there is no rearrangement

1 The term “symmetry " does not, perhaps, express the idea entirely, since an

egg with an asymmetrical cleavage must be regarded as having also a definite

organization.

No. 420.] REGENERATION IN THE EGG. 957

and, hence, the essential condition is absent. Our conception

of the structure of a magnet must be also fundamentally dif-

ferent from the ideas involved in the organization of the egg.

It is true that any piece of a magnet at once becomes a whole

magnet of smaller size, and this occurs in the smallest pieces

that it is possible to obtain. On the other hand, the change

in the organization of a piece of the egg is a relatively slow

one, and it can take place only in a relatively large piece of the

substance. If it be objected that these differences are only

trivial ones, and not essential, still it can be shown, I think,

that we must hold entirely different views of the nature of the

polarity of a magnet and the organization of the egg. Our

conception of the polarity of the magnet rests on the idea that

itis the sum total of the polarities, or, perhaps, of the orien-

tation of the minutest elements, the molecules, of which the

magnet is made up, while our conception of the organization

of the egg is exactly the reverse (or at least I shall try to

show that we must really believe this to be the case), and we

must think of the entire egg as a whole and not the sum total

of an infinite number of smaller wholes. We may claim, I

think, that this property of the egg substance of forming itself

into a new whole is peculiar to the living protoplasm and is a

property that we do not find, or have not found as yet, 1n inor-

ganic, or perhaps we may go further and say in dead, matter.

If we choose to call this property of living matter à vital factor

in the sense that it is not found in matter that is dead there

can be, I think, little objection to so doing. If the statement

seems to be arguing in a circle, we may state more simply that

those properties of living things that are not shown by non-

living things we shall call vital properties. We may add that

we cannot be sure, at present, whether these vital factors will

conflict with our present ideas of causality or not ; they seem

rather to be, however, new causal phenomena peculiar to dé

tain organic substances or compounds, but it would be Qut, o

place here to examine further into these difficult questions.

I have dwelt somewhat at length on this topic because,

as I shall try to show, an analysis of the phenomenon m

takes place when a piece of an adult animal (hydra, tor

958 THE AMERICAN NATURALIST. |. [Vor. XXXV.

example) changes over into a new, whole organism, leads us

directly to the same conclusion.

A ring cut at any level from the body of hydra closes its

open ends quickly, and in the course of a few days it elongates

and assumes the typical proportions of this form. At the

time of the operation the piece was a part of a definite organi-

zation, yet it afterwards becomes itself a whole structure. The

material of which the piece is composed must be totipotent, since

any piece may make a whole structure. The results demon-

strate that although the piece must be regarded as a part of a

whole organization before it was removed, yet after its removal

it becomes itself a whole organism. This fact shows that there

is no contradiction in our regarding the entire egg as also hav-

ing a definite structure, or as being an organized whole, and yet

any piece of it may become a new whole. This conclusion is

of some importance because, at first, students of experimental

embryology were inclined to go too far and assume that, since

any piece of the egg could become a new whole, therefore, the

egg itself must be regarded as a very simple structure. The

experiments with hydra show that the egg may de as highly

organized as is an adult animal, and yet a piece produce a new

whole. It is scarcely necessary to add that the results do not

show that the egg zs so highly organized but only that it may

be thought to be so without contradicting the results of the

experiments.

I pointed out in my Woods Hole lecture of last year (1899)

that from whatever part of the body a piece may be cut it will

still be different in its different parts, in'so far as one part was

nearer the anterior end and another nearer the posterior end of

the animal. And similarly for the sides of pieces of a bilateral

animal, one part will always have been nearer than another to

the median plane, etc. These differences suffice, I think, for

us still to form a causal conception of how the new axial rela-

tions are attained, since the differences always present will be

the basis on which the subsequent rearrangements take place,

and the results of observations show, in fact, that the anterior

end of the new organism comes from the anterior end of the

pieces, etc. We may infer that a similar change takes place

No. 420.] REGENERATION IN THE EGG. 959

in the reorganization of the egg; and we can form here also

a causal conception of the change, although we may admit that

it is a change different from anything that we know of in inor-

ganic matter. .

We may next èxamine some other results connected with

the development of pieces of the adult and of the egg. It

has been stated that a small piece of the stem of Tubularia

sometimes produces only a partial form, — a proboscis, or a pro-

boscis with reproductive organs, or a hydranth without a stem.

The development of these partial structures occurs only in

small pieces. The production of a proboscis, for instance,

takes place only in pieces that are much smaller than the

average size of the hydranth-forming region of a long piece of

the stem. An entire hydranth without a stem may, however,

develop from pieces that are much longer than the minimal

size of the hydranth-forming region of short pieces. The point

of special interest is that the development of the proboscis

takes place in pieces whose material is totipotent, t.e., the

material can produce any part of a hydranth or of a stem.

We may restate the result by saying that there is a stronger

tendency, especially in the more distal parts of the stem, to

form the full-sized organs than to form the whole structure In

a reduced form.

These results with small pieces of Tubularia suggest at once

a comparison with those cases in which a part of an egg

develops into a part of an embryo, and not into a whole one of

smaller size. This is most strikingly shown in the ctenophore

egg and also in the egg of a snail, Ilyanassa. -It has been found

that, if the first two or first four blastomeres of the ctenophore

egg are isolated, each not only divides as à part, but each

develops into an incomplete embryo that lacks, in the case

of the one-half blastomere, a half or nearly half of the entire

Structure, and in the isolated one-quarter blastomere lacks about

three-fourths of the entire structure, etc. If Mss i

from the fertilized but unsegmented egg of the same eS

the embryos that develop from the nucleated pieces are a : :

often incomplete. The results indicate that the phenomena us

due to a lack of regulative power, or power of rearrangem

960 THE AMERICAN NATURALIST. | [Vor. XXXV.

in the egg; perhaps to a lack of lability in the protoplasm,

so that it fails to reorganize itself into a new whole. The

result is not due to the smallness of the piece, for under

certain conditions, not yet understood, a smaller piece than

a half or a fourth of the egg may make a whole structure.

We must, therefore, look upon the egg of the ctenophore as toti-

potent in all its parts, in the same way that the sea-urchin's

egg is totipotent, but an incomplete embryo develops from a

piece, since the egg after fertilization has not the power, under

ordinary circumstances, of rearrangement or regulation. It is

conceivable that at a different temperature or in a slightly dif-

ferent medium the protoplasm of the piece might rearrange

itself so that an entire embryo would develop.

If we think of the embryo as organized in the protoplasm of

the egg in the sense of there being present a definite structural

arrangement of the protoplasm, then when the blastomere of

the ctenophore egg is isolated there is simply a lack of change.

If, on the other hand, we think of the egg at the beginning of

cleavage as being organized only in so far as the cleavage is

concerned, the later organizations appearing afterwards, then

we must think of the half organization of the embryo as being

induced by the half form of the cleavage, or the half form of

the isolated part. The half development would, in this case, not

be due to a lack of change, but to the appearance after cleavage

of a half organization. If we adopt the latter alternative we

can compare the development of the one-half ctenophore embryo

with the production of a part of a hydranth in a small piece of

Tubularia. In both cases a part of a structure is formed on

account of the form or size of the piece in material that is toti-

potent. If, on the other hand, we adopt the first alternative we

find the two processes different, unless it could be shown that

towards the distal end of the piece of Tubularia a new hydranth

may be organizing before the old hydranth is thrown off. The

removal of a piece taking place at this time, each piece con-

tinues to develop in the determined direction. Since, however,

only distal organs develop from such pieces at several levels

we see that this view is insufficient by itself to give à satis-

factory solution.

No. 420.] REGENERATION IN THE EGG. 961

In regard to this latter point it seems that the development

of an incomplete structure in Tubularia is not such an iso-

lated phenomenon as it appears to be at first sight, but can be

brought into harmony with certain other known results. For

instance, when more than five segments are cut from the

anterior end of the earthworm only five, as a rule, come back

at whatever level of the anterior end the cut may be made.

The amount of material that is formed at the cut-end, before

differentiation begins, is about the same at all levels, and this

gives, I believe, an insight into the phenomenon. There is for

the earthworm, also, a lower limit of organization for the forma-

tion of the head, or of a ring of the body, and the same new part

is formed at each level, because the amount of material that is

at first formed over the exposed end is the same. We must

look upon this material as totipotent, but the number of rings

that it produces is limited, on account of the amount of material

present and of the necessary connection that exists between

the lower limit of organization and the volume of matter! The

analysis leads us in both cases to the supposition that the

results are somehow dependent on the ultimate structure of

the protoplasm from which arises the structure that we can

see. The development of a proboscis from a small piece of

the stem of Tubularia indicates that there is a quantitative ele-

ment that comes into the problem, and this conclusion is directly

in line with the conclusion that we reach from a study of pieces

of minimal size of both adult organisms and of eggs. Below a

certain point the organization of the protoplasm cannot take

place, and it is this result that shows, I think, that we cannot

look upon the whole as simply the sum total of similar parts,

but as itself having a single plan of organization or structure.

If we next proceed to an examination of the method of

development of pieces of older embryos we find some further

facts that bear on our problem. It has been found that if the

hollow blastula of the sea-urchin be cut in two, the opening in

for the same

less rings.

1 another

1 The form of the new part must also be a factor in the problem, à

amount of material, if of a different shape, might give rise to eos x

The part of the body from which the new substance arises may be st

factor,

962 THE AMERICAN NATURALIST. [Vor. XXXV.

each piece soon closes, and a new sphere of smaller size forms

from each piece. These spheres then proceed to pass through

the later stages of development characteristic of the sea-urchin

development. Driesch has found that if the embryo is cut in

two at a later stage, when the process of gastrulation has

begun, not only the piece that contains the part about to

invaginate will continue to develop and produce a whole embryo,

but the piece from the opposite side will also form a gastrula

and embryo. If, however, the same experiment is carried out

after the gastrulation has been finished, these pieces of the

outer wall fail to produce a new gastrula. A similar result

was obtained in still later stages in regard to the formation of

the two pouches that pinch off from the inner end of the

archenteron of the starfish embryo. If the inner end of the

archenteron is cut off before the two pouches have formed

there, they will be formed again from any part of the more

proximal portion of the archenteron; but if we wait until the

pouches have once been completely formed by the archenteron,

and then cut off the inner end of the latter, the two pouches

are not made anew at the cut-end. In other words, after

the archenteron has once produced the two pouches it seems

to lose ¢shroughout its entire length the power of repeating the

process, although at an earlier stage all the parts possessed

this power.

Another somewhat similar result has been recently obtairied

by Spemann. If a thread be tied around the middle of the

embryo of the frog at the time when the medullary folds are

appearing, the egg being partially constricted by the thread

into a more anterior and a more posterior part, the latter will

produce at its anterior end (where the string constricts the

nerve plate) a new head, and there is produced in this way

a double embryo with two heads, one at the anterior end, and

one behind this at the middle where the constriction is present.

The result shows that the material of the dorsal nerve plate is

totipotent in so far as the formation of all the structures of the

brain, eyes, and nerve cord are concerned, and that the position

of any part in relation to the rest determines its differentiation.

The brain forms at the anterior end of the nerve-plate, but it

No. 420.] REGENERATION IN THE EGG. 963

may also appear in the middle of the plate if the middle comes

to represent the anterior end, as is the case in the experiment

just described.

A most important theoretical question is involved in these

results, although it has not been as yet sufficiently considered

by those who have carried out the experiments, vig., that a stage

once passed through by the embryo cannot be repeated again

by the other parts of the body that may not have been directly

concerned in the operation, although these parts possessed

this power at an earlier stage. A thorough discussion of this

important question would lead us too far from our present sub-

ject, but I may point out that the result indicates that at each

stage in the development the ultimate structure seems to change

its organization, so that it no longer repeats a stage once com-

pleted. The organism has not, however, lost the power of form-

ing a new individual, or at least certain parts have not, since

eges are produced by the adult animal, each of which may make

an entire organism, and the adult animal has also the power in

many cases of replacing lost parts, although it does not repeat

the embryonic processes in doing 50. i

When we examine the method of development of pieces of

still later stages of the embryo we find the process more and

more like that which takes place in the adult. In the higher

forms (echinoderms, mollusks, annelids, and vertebrates) the

regeneration of the late stages of the

the adult takes place by the developmen

free end, while in the earlier stages in certain embryos in these

groups a new whole is established by the entire structure

changing aver into the new form. In other words, the method

of development of pieces of the early embryo 15 Ee that to-

lowed by the lower animals, j.e., by a process of morphallaxis,

he method of regen-

While in the later stages of development t

eration is more like that in the higher animals. scs

seems to be connected with what we call the spec! agen

the tissues in the later stages of development and in Ud T ¥

of the higher forms, but what is meant by this specta ing "e x

a question that needs to be further examined, although it wi

not be possible to do so at the prac en’

964 THE AMERICAN NATURALIST. [Vor. XXXV.

We have seen that there is both for pieces of the adult

animal and for pieces of the egg, or of the isolated blastomeres,

a lower limit of size below which a piece does not produce the

typical form. Even in a form like Tubularia, in which a very

small piece may produce an incomplete structure, there is still

a lower limit below which the pieces do not produce any struc-

ture. We find this lack of power to develop in small pieces in

which we know from other experiments that the material is

totipotent, and the only clue we have to account for the facts

is that the result is in some way directly connected with the

amount of material present. We find for each animal a lower

limit of size —a limit that may vary at different stages of

development —for producing a miniature copy of the type form ;

or, in other words, the organization must have a certain amount

of material in order to develop. We find this same law to hold

for pieces of eggs, of embryos, and of adult Metazoa, and it is

equally true for adult unicellular Protozoa. It has been found

that very small pieces of the egg may continue to divide,

although they may be much too small to produce an embryo;

but as several other results have also indicated, the factors that

determine the cleavage are not necessarily connected with

those that relate to the organization of the embryo. Isolated

blastomeres that are below a certain size also fail to produce

an embryo. The one-eighth blastomeres and even the one-

sixteenth blastomeres produce in some few cases the early

stages of development, but many of them fail to do so, and

beyond this point it is doubtful if a blastomere can develop

past the gastrula stage. The results show that the lower limit

is soon reached in these eggs. It is certain that for adult ani-

mals pieces much smaller than one-eighth or one-sixteenth of

the body can produce new animals of half size, but we have at

present no data for comparison between the absolute minimal

size of a piece of the egg and of the adult of the same species

that can produce a whole animal. It has been found that

pieces of planarians that contain less than ijs of the volume

of the adult may produce the typical form. Pieces of hydra

zoo of the adult produce a new hydra; and pieces of the stem

of Tubularia about 41, of the length of the stem can also make

No. 420.] REGENERATION IN THE EGG. 965

a new hydranth and stem.! The eggs produced by these forms

are much smaller — hydra perhaps is to be excepted — than

these minimal-sized pieces, and as the egg produces the typical

form we cannot expect to find any agreement between the ulti-

mate size of an isolated blastomere and of a piece of the adult

that produces a whole structure.

Peebles found that a smaller piece of a young bud than of the

wall of hydra would produce a whole organism. The result is

interesting inasmuch as the region where the bud appears is

also a part of the wall. We find in a growing region of this

sort that the protoplasm is denser than elsewhere, and may

contain at first less water than other parts of the body, so that

the absolute amount of living substance may be the same ina

small piece of such a bud as is contained only in a much larger

If this is true, and if the egg can

piece of the wall elsewhere.

r in its structure

also be shown to contain relatively less wate

than do other parts of the adult animal, it may still be possible

that the amount of living substance necessary to give rise to

the typical form may be found to be the same in the piece

of the egg and of the adult. At present, however, we lack the

data necessary to make the comparison. It is certain that in

many forms — as for instance in the starfish — a piece of the

adult that can regenerate the entire structure must be at least

several thousand, perhaps million, times greater than the egg

and must contain a much vaster amount of living substance.

Many other factors than that of mere size must come into play

in such cases, and it seems to me that the comparison 1S P

after all, of much theoretical importance, since the method 3

development of a piece of an €88 and of an adult may be

. entirely different.

Tubolars that a comparison is diffi-

be 35 mm., then a piece of the stem

hole; and pieces of this size pro-

1 The length of the stem is so variable in

Cult. If we assume the length of the stem to

measuring 1 mm. in length is about zs of the w

duced a new hydranth and stem.

966 THE AMERICAN NATURALIST. [Vor. XXXV.

II.

A year ago! I reviewed the different hypotheses that have

been advanced to account for the phenomena of regeneration

and I pointed out that if the development of a part of an egg,

or of an embryo, is also a process of regeneration, as I believe

to be the case, the same hypotheses ought to apply also to the

development of these parts. It may be, therefore, worth while

to see how far these hypotheses may account for the develop-

ment of pieces of the egg or embryo.

Bonnet extended his theory of praformed germs, primarily

invented to explain the development of the embryo from the

egg, to include the phenomena of regeneration. At that time

the development of a part of an egg was not known, but there

is no obstacle at present to applying the same interpretation to

the development or regeneration of parts of the egg and embryo.

In fact, Weismann, who also believes in a theory of praeforma-

tion both for egg-development and for development of pieces

of adult organisms, has applied the same view to the develop-

ment of parts of an egg. The theory of praformed germs as

held during the seventeenth and eighteenth centuries need no

longer be seriously discussed in the form in which it was then

expressed, since we have abundant evidence to disprove the

view; but the same conception has appeared again in a more

insidious form in our own time, and has been applied not only

to development of the egg, but also to the regeneration of both

the adult and embryo. I shall not attempt to repeat here the

arguments in favor of or against this view, but shall confine

what I have to say to those points that bear on our present

examination. The modern form of the doctrine of prefor-

mation as held by Weismann and his school is as follows.

There exists in the nucleus of the egg praformed germs

that correspond part for part to the later structures that

develop from the egg. The process of development is a

process of sorting out of the nuclear germs to different parts

of the embryo, and later, by the action of the germs con-

tained in each nucleus on the protoplasm of each cell, they

1 Woods Hole Lectures, 1899.

No. 420.] REGENERATION IN THE EGG. 967

determine the differentiation of the cell. In other words,

the protoplasm is looked upon as a kind of indifferent medium

on which the nuclear germs feed and multiply, and, replacing

the protoplasm of each cell, produce there their particular

product.

In order to account for the regeneration of parts of the adult

itis assumed that there has taken place during development

not only an unraveling of the qualities of one germ, but at the

same time at each or many of the divisions of the nucleus a

quantitative division of another original whole germ, or later of

a part of that germ, into like parts. This second germ is a sort

of reserve stuff for future contingencies, and during regeneration

this reserve stuff is supposed to become active and to do what

is required of it. Pieces of the egg or of the embryo are also

supposed to be able to regenerate, owing to the presence of

this reserve idioplasm or germ in each nucleus, that is awakened

into activity by the injury to the whole and then proceeds by

qualitative division of exactly the right sort to set free the

necessary germs to complete the embryo.

I need not repeat here the many objections that have been

brought forward in opposition to Weismann's view, or stop to

point out how the upholders of this view have been obliged

again and again to add subsidiary hypotheses to meet the

objections that have appeared, but I may indicate at least some

of the difficulties that this view meets with from the point of

view of regeneration in the egg and embryo.

The hypothesis assumes that at each division of the egg a

sorting out of the qualities of the embryo takes place. Obser-

vation shows, however, that at each division there is an equal

division of the chromatin, and not an unequal division, as would

necessarily take place in some of the divisions, if the hypothesis

is correct. The experiments with the ctenophore egg indicate

that the protoplasm is an important factor in the early divi-

sions. The experiment of compressing the frog's egg during

its period of division, so that a distribution of the nuclei in the

protoplasm, different from that in the normal egg, is brought

about, indicates that the early nuclei are equivalent ; and the

development into whole embryos of pieces of the egg 1n which

968 THE AMERICAN NATURALIST. [Vor. XXXV.

the segmentation nucleus passes through fewer divisions than

in the whole egg also points in the same direction.

The regeneration of pieces of Hydra, Tubularia, and Planaria

shows that at every level of the body the cells have the power

of producing any part of the organism,! and we must regard

them as totipotent, or nearly so. The same conclusion is

arrived at from a study of the development of isolated blasto-

meres. These facts, and others that might be given, furnish,

I think, a strong argument against Weismann's hypothesis,

and I believe we must look in a different direction for a solu-

tion of the problem of development and of regeneration.

We should be careful to distinguish between the idea of pre-

determination and that of pra formation. The former includes

the latter, but also much more, and is, in fact, an almost neces-

sary idea in connection with the phenomenon of development,

at least in one sense. Every egg of the sea-urchin is prede-

termined to give rise to a sea-urchin so long as the conditions

remain as we find them at present, but it by no means follows

that the hypothesis of preformation is a necessary consequence

of this view. It is true that since each egg passes through the

same stages to reach the same goal we must regard the series

of changes as predetermined in the structure or composition of

the egg, and in the external conditions surrounding the egg, but

the way in which the development takes place may be in no

way connected with the presence of a praformed germ of the

embryo or with the presence of praeformed germs of any sort

in the nucleus. In fact, a process of pure epigenetic develop-

ment, as generally understood nowadays, may also be prede-

termined in the egg. It is well, I think, to be clear on this

point, since otherwise we grant too readily the necessity of

assuming some sort of praeformation hypothesis.

In my Woods Hole lecture of last year I considered the

hypothesis of formative stuffs and reached the conclusion that

it is insufficient to explain many of the phenomena of regener-

ation. Let us see if this view will apply to the development of

parts of the egg and embryo. In the first place, it should be

noticed that the hypothesis was primarily invented to account

! At least so far as the different layers are concerned.

No. 420.] REGENERATION IN THE EGG. 969

for those kinds of regeneration in which a new part is added at

the end of the old part, and, as I have pointed out, it is not

well suited to explain those cases in which a piece changes

over entirely into a new whole. Since the development of

parts of the egg and of isolated blastomeres and pieces of the

embryo takes place by a process of morphallaxis, it is clear that

the hypothesis is equally inapplicable to such cases. The kind

of rearrangement or regulation that takes place in a piece of an

egg, or of an isolated blastomere, or of a piece of a protozoón, so

that a symmetrical whole is formed, cannot be made any clearer,

I think, by the assumption of specific fluids or stuffs in the

different regions that determine the later differentiation.

The idea that there is a similarity between the process by

which a broken crystal completes itself and that by which an

animal ora plant may make good a lost part has often been

suggested. The comparison rests, I think, on a superficial

resemblance, and a careful examination of the nature of the

two processes shows them to be the outcome of different fac-

tors. It may, however, be asked in what respects is the mod-

ern view of crystallization different from the view which I have

advocated in regard to the reorganization of the protoplasm ina

part of an egg or in a piece of hydra. Thetwo conceptions are

in reality entirely different. A piece of a crystal does not rear-

range its parts, much less its axes, to form a new whole of smaller

size, but has deposited on its surface, from the saturated solu-

tion in which it lies, new material that conforms in every respect

to the original axes and planes. In a piece of an egg or of an

animal, on the contrary, the entire old structure changes over

into a new whole involving in some cases à change of axes.

Pflüger's conception of the process Ub regenera aa gred

points of resemblance to the idea of the recompletion of a

crystal. He applies the conception, however, pre

cases in which the regeneration is by a process of addition -:

the old part (epimorphosis). Pflüger's hypothesis 15 that a t :

cut-end the old material attracts from the blood new ma erial,

ens : i rface and is there organized

which is deposited over the cut-su d

: ; : old part. He speaks of the

at its region of contact with the ! pt UM hf with the

new molecules being organized at the region 9 Pan

970 THE AMERICAN NATURALIST. [Vor. XXXV.

new ones, and those that have thus been organized are sup-

posed to attract others which are then deposited on the outer

end and in turn become living structures. This idea is not

fundamentally different from that held at present as to the

growth of a crystal, except in so far as the molecules are sup-

posed to be organized by those in the old part or by those that

have previously been altered, etc. But on this view we have no

explanation of the process of morphallaxis, for, in this case, the

change takes place only in the old part and throughout the old

part. If we attempt to extend Pflüger's idea so that it may

include these phenomena, we must suppose that a sort of

recrystallization takes place in the old tissue or isolated piece;

but this is really a very different conception from the former,

for the process is not reorganization by apposition, but by com-

plete working over of the entire piece into a new whole. This

idea is entirely foreign to our conception of crystal growth, and

also to the conception entertained by Pflüger. I have also

pointed out that even in some cases in which a new part is

added to the old one there are grave difficulties in the way of

accepting Pflüger's view, especially in the case of the planarian,

in which the new head does not appear in the middle of an

anterior oblique exposed edge, but to one side of the middle

line, where the influence of the old part would produce, on

Pflüger's view, a very different result.

Finally, I wish to discuss a problem that is at present one

of the most difficult questions in connection with the process

of regeneration. I mean our idea of polarity or polarization of

the living material of the animal and plant. ‘The term was

first used by Allman in 1864 to mean that a new anterior end

appears on that part of a piece that lay nearest to the old

anterior end, and a posterior end appears on that part that

lay nearest the old posterior end. This rule is followed in a

great many cases, but it is not invariable. In certain forms an

organ different in kind from the one removed has been found

to develop. The first case of the kind was described by

Bonnet, who found when certain fresh-water. worms were cut

in two that the posterior piece produced at its anterior end, |

not a new head, but another new tail. Loeb also has described

No. 420.] REGENERATION IN THE EGG. 971

several other instances of.the same phenomenon in hydroids,

and has made the additional and important discovery that in

these forms the external conditions may determine the kind

of regeneration that takes place. However, even in these

forms differences are present in the two ends- of a piece, for,

even in the most favorable forms it is found that, as a rule, a

new head or hydranth forms sooner on the anterior end of the

piece than on the posterior, and from the posterior end of the

piece a posterior end is more easily developed than from

the anterior end. In other and higher forms the factors that

determine the kind of regeneration seem to come from within

the piece itself. In some of these forms we also find occasion-

- ally that the new part may be different from the part removed

(heteromorphosis), and in some cases, as in Bonnet’s fresh-

water worm, a posterior end develops on an anterior surface.

Other cases of the same sort are also known; for instance, in

the earthworm, Allolobophora, a short posterior piece (less

than half the length of the worm) generally produces at its

anterior end another tail; a very short piece of the head-end

may produce a new head on its posterior surface, and the same

holds for short pieces of the head of a planarian.!

Sachs has vigorously combated this idea of polarity, that he

refers to as a metaphysical notion of the morphologists, and

offers as a substitute his hypotheses of formative stuffs.

have attempted to show that the idea of formative stuffs is

insufficient to explain the phenomena of regeneration, and at

present I see no escape from the idea of a polarity of some sort

existing in the organism. On the other hand, we cannot be too

careful in clearly defining what we intend to include in this

idea of polarity, for I think we can see even at present that

we must form a conception of the polarity different from

that in which the term is used by the physicist.

The term “polarity” when used in connection with phenomena

connected with the magnet means that there is a difference

in the two ends of the magnet. The idea also involves the

further one that any piece of the magnet also shows this same

1 We may call that kind of heteromorphosis in which the new part is more or

less a mirror figure of the old part “ amphimorphosis.”

972 THE AMERICAN NATURALIST. [Vor. XXXV.

difference at the ends, and since this is found to be true for

every smallest piece of the magnet, it is assumed to be a

property of each molecule or atom of which the magnet is

made up—the magnet being the sum total of the action of all

its molecules. We speak also of a crystal showing polarity in

the sense that it has a definite form that can be referred

to definite poles and axes, and it is supposed that this same

property is possessed by the smallest * crystal molecules " of

which the crystal is made.

On the other hand, in an animal or in a plant in which we find

the two ends of a piece behaving differently, yet always show-

ing the same constant difference, although we have a result

resembling in many respects the polarity of the magnet, we find -

also in several essential respects differences and, I think, fun-

damental differences (in the sense that they are the outcome of

different factors) between this phenomenon and that in the

magnet. For instance, we find in the pieces of Tubularia that

external conditions may reverse entirely, after a time, the

polarity of the piece, so that a head may appear on the poste-

rior end even while another head is present on the other end.

We know of nothing similar in the case of the magnet, unless

we consider the change that can be brought about in it by

re-magnetization; but there are quite obvious differences in

the two results. The most fundamental difference, however,

between the two conceptions is, I believe, connected with the

kind of structure that we must suppose to be present in the

two cases. The polarization of the magnet is the sum total of

a vast number of smaller units, each unit being itself a magnet.

On the other hand, we have at present clear evidence to show

that while the polarity of the organism is the outcome of its

structure or organization, this is not the effect of the presence

of a large number of units of which the organization is only

the sum total, but, on the contrary, the organism is itself the

ultimate unit that makes the polarity, and while we must

believe that this same organization may develop in a piece of

the original whole, the evidence goes to show that the reor

ganization is of a different nature from that by which a

piece of a magnet becomes af once a new magnet.

No. 420.] REGENERATION IN THE EGG. 973

In the first part of this paper I have given my reasons for

looking upon the organization of living things as a phenome-

non sui generis, and I have wished to bring here into connec-

tion with this consideration the conception of polarity as

applied to the organism. It is a phenomenon, I think, that

is different from the one bearing the same name applied to

inorganic substances. We are therefore, I think, justified in

looking upon the reorganization of pieces of the egg, embryo,

and adult as a phenomenon peculiar to the living structure of

'the egg, and without a parallel in inorganic nature. It seems

to be one of the peculiar physical properties connected with

the matter that we call living substance, and in the last analysis

we find it to be not simply the outcome of a complex of known

physical principles. We are therefore, I believe, also justified

in calling the organization of living things a vital property in

the sense, to repeat what I have just said, that it is peculiar to

this kind of substance or structure, and not the result of a

complex of known physical principles ; or, in other words, it is a

physical phenomenon as fundamental as the polarity shown by

crystals or the magnetism of the magnet, and just as the latter

are associated with certain kinds of matter, so is the organiza-

tion associated with the substance protoplasm.

THE HABITS AND NATURAL HISTORY OF

STICHOSTEMMA.

C.2M>CGHILD.

A SMALL fresh-water nemertean, undoubtedly .Szzchostemma

asensoriatum Montgomery (96), is found very abundantly in

one of the park lagoons in Chicago.

The animal has proved to be a favorable form for experi-

mental study, and in the course of such work during the past

two years a number of observations upon its habits and natural

history have been made. These observations are given here

in the belief that a connected account of the natural history of

a single form is of value because it brings together a series of

facts which, though important, often receive only incidental

attention in connection with other work, or are ignored alto-

gether, and because it affords a basis for further study.

Montgomery (96) has given a brief description of a new

American species of fresh-water nemertean which he called

Stichostemma asensoriatum. He regards Silliman's Tetra-

stemma aquarum dulcium (85) as insufficiently characterized,

but believes that it may be closely allied to S. asensoriatum.

The species which forms the subject of this paper agrees with

Montgomery's description of S. asensoriatum in all respects

except that it is not protandric, while Montgomery believes

S. asensoriatum to be protandric like S. eilhardi. Bohmig

(98) has accepted Montgomery's genus Stichostemma for his

species S. grecense, but as S. gr@cense is not protandric,

Bóhmig holds that the existence of protandry should not be

regarded as a characteristic of the genus. Ina personal com-

munication very kindly made in answer to an inquiry of my

own, Dr. Montgomery expresses himself as of the opinion

that the presence or absence of protandric hermaphroditism

is a point of minor importance. The revised genus diagnosis

of Stichostemma as given by Bóhmig (98) is as follows :

975

976 THE AMERICAN NATURALIST. [VorL. XXXV.

* Augenzahl veránderlich, gewöhnlich mehr als 4; 9 oder 10

Rüsselnerven; das Rhyncocoelom erstreckt sich nicht bis an

das Kórperende; die Nephridien dehnen sich von der Gegend

des Gehirns bis zur hinteren Korperspitze aus; zwitterig,

ovipar." The species found in the vicinity of Chicago agrees

with S. græcense in not being protandric, but since I cannot

find that it differs in other respects from the description given

by Montgomery of .S. asensoriatum, I am inclined for the pres-

ent to regard the two as identical. It appears probable, more-

over, that S. asensoriatum Montgomery is identical with Tetra-

stemma aquarum dulcium Silliman. In any case the two forms

are closely allied, as Montgomery admits.

The species under consideration is a slender worm, attaining

often a length of 20 mm. (Montgomery gives the maximum

length as 18 mm.). It varies in color from a light yellowish,

almost cream-color, in the smaller specimens, to a deep orange-

red in many of the large individuals. Even in the most bril-

liantly colored specimens, however, the terminal regions of the

body are more yellowish than other parts. The reddish colora-

tion is especially distinct in the nervous system. In many of

the larger specimens the region of the brain can be distin-

guished by the naked eye on account of its deeper coloring.

Individuals in which the eggs are nearly ripe show scarcely a

trace of the reddish tint, being almost pure yellow in color.

Animals kept in the laboratory for a long time without food

usually become more or less brownish, owing to the appear-

ance of black pigment in the integument. This pigment is

occasionally found in newly captured specimens. Iam inclined

to believe that its appearance is connected in some manner

with the lack of food.

The usual number of eyes is six, the anterior pair being the

largest and the posterior pair the smallest. Variation in the

number of eyes, or at least in the number of pigment spots,

is of frequent occurrence. Specimens with only four eyes

sometimes occur, but are rare. In case eight symmetrically

placed eyes are present, the anterior pair is usually small and

situated a short distance in front of the largest pair. The

eyes are very often unsymmetrically placed or differ in number

No. 420.] HISTORY OF STICHOSTEMMA. 977

on the two sides of the head. Variation in the number of

pigment spots can often be observed in a single individual.

One or more of the spots may divide, or in some cases a num-

ber of small masses of pigment appear in place of a single

pigment spot, in consequence of a kind of fragmentation.

One of these smaller masses may increase to the size of the

whole, while the others disappear. Migration of these masses

is often observed. A more complete account of these obser-

vations is reserved for a future time.

The body of the worm is very transparent. The alimentary

canal, blood vessels, terminal nephridial organs, nervous sys-

tem, gonads, the rhyncoccel, and the proboscis, with principal

and accessory stilets, can be clearly seen in the living animal

under slight pressure. For some time past I have employed

the form for class work on account of the ease with which

its internal anatomy can be studied.

The points of difference between this form and S. et/hardi

as given by Montgomery (96) are as follows: (1) The absence

of the supraoral sense pit, which suggested the specific name

“asensoriatum.” (2) The cephalic gland is not as voluminous

as in S. ei/hardi, and has not one main efferent duct, but the

ducts of the individual gland cells penetrate the body epithe-

lium independently of one another, in the manner of subcu-

ticular gland cells; the external openings of these cells lie at

the anterior end of the head. The want of a main efferent

duct, formed by the fusion of the individual cell ducts, might

be explained by the absence of the supraoral sense pit, since

in most metanemerteans the main duct discharges at the base

of this pit. (3) The pigment of the body epithelium occurs

exclusively in the supporting cells and is not found at all in

the interstitial tissue. (4) Those gland cells corresponding to

what I have termed “gland cells 3” in S. ei/hardi are not

egg-shaped, as in that species, but have an elongated narrow

form; in structure and in their color reaction to stains they

show a close similarity to the cells of the cephalic gland:

thus, they stain intensely with hematoxylin, and with the

triple stain of Flemming (safranin, gentian violet, orange G)

combine with the safranin and not with the violet. (5) The

978 THE AMERICAN NATURALIST. [VoL. XXXV.

stilet basis is slightly constricted medially. (6) There are ten

instead of nine longitudinal nerves in the proboscis.

These differences do not concern the general appearance,

so that the figure of S. ez/hardi given by Montgomery (95,

Taf. VIII, Fig. 17) might serve equally well as a general dia-

gram of the structure of S. asensoriatum.

OCCURRENCE AND DISTRIBUTION.

During the early autumn of 1899 Stichostemma was found

in great abundance in one of the lagoons in Jackson Park,

Chicago, which has free communication with Lake Michigan.

Since that time it has been present in the lagoon at all sea-

sons of the year. Large numbers are found in masses of

Spirogyra or other filamentous alga, and often, though less

commonly, the animals are found among Elodea. They are

most numerous along the shore where the water is only a

few inches deep. Their presence here is probably due to

the higher temperature of the shallow water near the shore

and the consequent more luxuriant growth of the vegetation

which they frequent. Their food, which consists of species of

Nais, is very abundant in the lagoon under the same condi-

tions, and probably also, directly or indirectly, constitutes a

factor in the local distribution.

The distribution about the lagoon is rather irregular and

uneven. In some spots the worms are so abundant that hun-

dreds may be taken from a single liter of the aquatic vegeta-

tion. In other spots, apparently equally well suited to these

forms, only a few individuals or none at all are found. While

these areas are rather definite during a single season, the worms

do not always reappear in the same spot in succeeding years.

It is usually necessary to hunt for the favorable localities anew

each year.

The seasonal distribution is very definite. During the later

summer and autumn the animals are usually very abundant,

young and old together. They do not disappear from along

the banks until actual freezing occurs. When the ice forms,

many of the worms are undoubtedly destroyed, though a large

No. 420.] HISTORY OF STICHOSTEMMA., 979

number of them must reach deeper water, where they continue

to live beneath the ice during the winter. The lagoon is twelve

to fifteen feet deep in many places, and no freezing occurs in

the deeper parts. There is no evidence that the worms are

capable of enduring actual freezing. After the first appear-

ance of ice in the lagoon I have chopped out and thawed

masses of the vegetation in which the worms had been abun-

dant a few days before, but which were frozen solidly into the

ice when removed. In aquaria and jars prepared from these

no worms ever appeared. During the winter of 1899-1900

a part of the lagoon remained open until about the middle of

January, although the weather was cold and the remainder

was covered with several inches of ice. Up to this time the

worms were found here in abundance. Later this area was

covered with ice a foot or more in thickness, and the bottom

was also frozen in the shallower parts, and at this time no

worms were obtained from the region.

It is possible that the worms burrow into the mud, but sam-

ples of the mud taken at various times during the winter have

never yielded a specimen. I have been unable to examine fully

the deepest parts of the lagoon, but believe that the animals

remain more or less active there during even the colder months.

The worms reappear near the shore in large numbers very

soon after the ice begins to melt. In 1900 the lagoon was

covered with from twelve to sixteen inches of ice on March 20.

By March 26 the ice had melted along the shores, leaving some

two or three feet of open water from six inches to a foot in

depth. At this time large numbers of worms were found in

this open area, nearly all of them large specimens with well-

developed eggs, whereas the worms taken in January are of

various sizes and mostly with very small eggs or none at all.

The small worms taken in January are the young of the

preceding autumn which have not attained full size. If the

worms had remained inactive until the ice melted, it is diffi-

cult to understand how their increase in size and the growth

of the eggs could have occurred in so short a time as three

or four days. I believe, therefore, that they must have spent

the month of February and the early part of March in the

980 THE AMERICAN NATURALIST. [VOL. XXXV. .

deeper parts of the lagoon, reappearing at once near the sur-

face as the temperature of the water began to rise. The small

naids which form their food can be taken at all times through

the winter from beneath the ice, so that so far as food is con-

cerned Stichostemma is well supplied during the winter months.

Apparently, then, the young of the preceding autumn reach

their full size during the winter, reappearing near the surface

with nearly ripe eggs as soon as the ice disappears.

During the winter of 1900-1901 the lagoon was wholly cov-

ered with six to eight inches of ice before the end of Decem-

ber, and the temperature during the greater part of January

and practically the whole of February was far below 32°. The

ice became exceptionally thick and did not disappear till near

the end of March. The worms were not found along the shore

until April 12, and then only a few appeared, all of them large

and filled with eggs which were about half grown. During

the early part of December, 1900, before the lagoon was com- |

pletely frozen over, the worms were more abundant than I had

ever found them before. Their late appearance and small num-

bers in April, 1901, must probably be attributed to the pro-

tracted cold weather and the thickness of the ice in the lagoon.

During the months of May and June comparatively few

full-grown specimens are found, but the young are increas-

ingly abundant as the season advances. These become sexual

about July 1 or a little earlier. By this time nearly all the old

worms which survived the winter have disappeared. During

the remainder of the summer the worms continue to increase

in numbers until freezing kills them or drives them to deeper

water.

There is no evidence that the worms possess the power of

entering upon a resting stage of any kind which is capable of

enduring freezing without injury, though there are some facts

that indicate the possibility that they may become more or less

quiescent for a short time. When the animal is brought under

conditions which are disagreeable to it, or is irritated, it secretes

a large amount of slime and in many cases becomes enclosed

in a covering of slime, which gradually contracts and becomes

more dense until it resembles a cyst. Within this the worm

No. 420.] HISTORY OF STICHOSTEMMA. 981

may remain for some weeks, either moving slowly or not at

all, or it may leave in a few days. Left in jars of clear water

without algae, the animals are likely to make these slime cov-

erings repeatedly. As will appear below, I believe that the

. fact that the animal is disinclined to leave the slime is a

thigmotactic reaction. Violent stimulation will often induce

removal, or the slime may be picked apart with needles, thus

freeing the animal, which is active and normal in appearance

in every respect. In many cases mutilated specimens or ani-

mals in impure water form these coverings and remain in them

until death.

It is possible that the worms may form these “ cysts " about

themselves during the winter. These do not, however, enable

them to resist freezing and do not serve as a protection against

desiccation, as will appear below. It is difficult to understand

how the animal is better protected within this “ cyst,” except,

perhaps, as regards predatory enemies, than it is under the

usual conditions. While this process is perhaps to be regarded

as showing the manner in which the process of encystment

may arise, I think it is certain that in its present development

this slime covering is of little value as a cyst. The causes

and method of its formation are discussed below.

MOVEMENTS.

Locomotion forward is accomplished principally by means

of the cilia with which the body is covered. During the for-

ward movement the head is turned slightly from side to side

in regular succession, and as the body follows the same course

the movement as a whole is distinctly sinuous, the body show-

ing altogether four or five curves to left and right at any given

time. It is probable that under normal conditions the sinuous

character of the movement is a distinct aid in locomotion. In

crawling through masses of algz the lateral movements of the

head will, of course, enable the animal to find the interstices

more readily, while the sinuous bending of the body affords a

firmer support and a greater power for forcing a way among the

tangled threads. The value of these movements in enlarging

982 THE AMERICAN NATURALIST. . [Vor. XXXV.

the field of direct stimulation and contact is evident. The

animal crawls along the bottom of a jar with the ventral sur-

face uppermost almost as often as in the other position. It

is unable to support itself free in the water, although it

often drops from the surface to the bottom, making the usual .

crawling movements during the descent. During all forward

movement the head is continually making rapid searching

movements in all directions.

Backward movement is accomplished in a manner entirely

different from that just described for forward locomotion, vtz.,

by muscular contractions.

The muscular activities concerned in backward locomotion

consist of a series of peristaltic waves. A contraction of the

circular muscles near the posterior end occurs, and the end is

thrust backward ; then a wave, consisting of strong contractions

of the longitudinal and relaxation of the transverse muscles,

producing a shortening and thickening of the region affected,

begins at the posterior end of the body and travels forward. `

The whole process is quickly repeated, and in continued back-

ward movements three or four of these peristaltic waves may

be visible on the body at one time. On a smooth surface like

glass these movements are rather ineffective, but it is easy to

see how, under natural conditions, the posterior end of the body

becomes more or less completely fixed as soon as increase in

size begins and the other portions are drawn back to it.

The body is extremely contractile, and different parts are

capable of independent contraction. In the reassumption of

the extended position after contraction, the extension usually

begins at the anterior end and proceeds backward along the

body as the head advances.

Movements of the proboscis within the rhyncoccel are very

common. Contraction of the circular muscles in the posterior

portions of the body causes the proboscis to be thrown for-

ward, and, conversely, extension of the head region forces it

backward. In general the proboscis, together with the fluid

of the rhyncoccel, tends to occupy the region of least pressure.

The relation of the cilia to the different movements 1s

interesting. During forward movement the cilia are in active

No. 420.] HISTORY OF STICHOSTEMMA. 983

motion and constitute the chief organs of locomotion. If,

. however, the forward movement ceases suddenly, or if the

head be slightly drawn back, the cilia in the part affected stop

their movement and remain nearly perpendicular to the body

surface. If the animal remains quiet for a short time the

cilia may begin to move slightly. It is difficult to determine

whether this movement affords any definite impulse in either

direction, but observation indicates that there is a very slight

backward current along the body. When the head and ante-

rior portions of the body are suddenly and violently drawn

backward, as often occurs, or if the animal begins to move

backward, the cilia of the contracting region are laid flat

against the body, all pointing forward, and resume their usual

position when the backward motion ceases. The body is at

all times covered with a thin layer of slime, and it is possible

that the turning forward of the cilia may be simply the result

of the backward movement of the body through the slime.

Observation of the head region, which is constantly undergoing

contraction and extension, shows a beautiful play of the cilia,

and often in the rapid but very slight contractions without

actual backward movement, the cilia over a considerable area

are all instantaneously turned forward against the body, per-

haps to reappear almost immediately. The rapidity and uni-

formity of their change of position render it probable that it

is the result of a definite stimulation similar to that causing

reversal of the cilia in many other forms.

I have not been able to find any indications of actual

reversed beating of the cilia causing backward movement of

the body. The backward movement is apparently brought

about wholly by muscular contractions, the cilia being laid

against the body and pointing forward, so that they offer no

resistance, but remaining motionless or making very slight

indefinite vibrations. These conclusions agree well with those

reached as the result of macroscopic observations on the mov-

ing animal. Stichostemma does not appear to glide backward,

but accomplishes this movement by violent muscular contrac-

tions as described above. Except when the stimulation is

very intense and sudden, it is more likely to turn the head

984 THE AMERICAN NATURALIST. [Vor. XXXV.

back upon the body and thus to change its direction of move-

ment, than it is to move bodily backward.

As in other nemerteans and in turbellaria, the body is

always covered with a thin layer of slime or mucus secreted by

the dermal glands. Any stimulation or irritation causes an

increase of this secretion to such an extent that the animal

becomes enveloped in a thick layer of the substance. The

animal is able, like many other related forms, to move beneath

the surface film, and presumably this secretion is an important

factor in locomotion here and elsewhere.

Small dishes in which a number of specimens are kept

become filled with long strands and masses of the slime, in

which the animals move about. The accumulation of the secre-

tion does not appear to injure the animals in any way. Wilson

(00) has noted the large amount of slime secreted by Cerebrat-

ulus, and his observations are paralleled by my own. Certainly

for so small a form as Stichostemma the amount of this slime

secreted is extremely large.

RESPIRATION.

Respiration undoubtedly occurs through the surface of the

body. The swallowing of water which Wilson (00) has

observed in Cerebratulus has never been seen in Stichostemma.

The animals are apparently sensitive to changes in the oxygen

content of the water, but their small size and the absence of a

cuticle probably render special respiratory organs unnecessary.

Foop.

The only food which the animal is known to take consists

of specimens of Nais which are captured and gradually drawn

into the alimentary canal. The actual capture of these animals

has never been observed, although in several cases they have

been seen in the partially digested condition in the alimentary

canal, and in one case a specimen was found with a large Nais

partly swallowed. The Nais was as long as its captor, and

although half its body was in the alimentary canal of the

No. 420.] HISTORY OF STICHOSTEMMA. 985

nemertean the free portion made active movements. The nemer-

tean was comparatively quiet, but at short intervals the mouth

was widely opened and the body extended, and in this manner the

Nais was drawn further into the alimentary canal. It could be

clearly seen through the body wall of the nemertean that diges-

tion of the annelid was proceeding very rapidly. Later a part of

the Nais was disgorged, and some portions were seen to be com-

pletely disintegrated, while others which had been in the ali-

mentary canal only a few minutes, ten to fifteen, showed distinct

evidences of digestion. This rapid digestion of the food was

also observed by Wilson ('00) in the case of Cerebratulus.

It appears extremely probable that the food is captured in

the manner which Wilson (00) has described for Cerebratulus,

though I have never observed it. Indeed, there is a striking

similarity between this form and Cerebratulus, both in the

nature of the food and in the method of taking it. In the

cases of both an actively moving annelid forms the food, being

attacked and overpowered by the nemertean, and in both

cases the prey is often as large as its captor and always more

active. I have not been able to determine whether Sticho-

stemma always begins the process of swallowing at the posterior

end of the body of its prey. In the one case of feeding actu-

ally observed, the head of the naid had been broken off com-

pletely and lay in the slime secreted by the nemertean, but

the free end of the annelid was apparently the anterior end

from which the head had been broken. In the bodies of the

naids which had been completely swallowed digestion had

always proceeded so far that it was impossible to determine

with certainty which was the anterior end.

The feeding apparently takes place only in darkness, or

in light of little intensity, and therefore occurs normally

during the night. During the day the nemerteans may

remain in direct contact with the naids in the interstices of

the masses of algae, but apparently do not attack them. The

next day, however, or after a few hours in darkness, the naids

are fewer in numbers and the intestines of the nemerteans

indicate very distinctly a recent meal. The failure to observe

the actual capture and, in all but the one case mentioned above,

936 THE AMERICAN NATURALIST. [Vor. XXXV.

the swallowing of the food, is probably due to the fact that the

nemerteans are easily disturbed by a jar or movement of the

dish containing them, and especially by the light. In such

cases they probably release the prey or disgorge it, if partly

swallowed, as in the case described above.

There is no direct evidence that the adult animal takes other

food under normal conditions. The fæces, so far as recogniz-

able as such, always contained the setze of naids. Moreover,

specimens have been kept for weeks in jars with many other

animal forms, Protozoa, Turbellaria, rotifers, copepods, ostra-

cods, etc., but no traces of any of these forms have ever been

found in the alimentary canal.

The nature of the food of the young individuals is a matter

of interest. When first hatched they are less than a millimeter

in length. Probably they do not take food immediately, but

they certainly feed when from one to two millimeters in length.

The food at this time probably does not consist of naids, for

it is difficult to understand how these small animals could

capture any except the very smallest. I am inclined to believe

that they feed upon other small forms during these early stages,

though I have no direct evidence upon this point.

I have often suspected the animals of eating each other.

In jars containing a large number of specimens there is always

a remarkable decrease in numbers during the first weeks, and

the walls and bottom of the jar do not show the whitish disin-

tegrating masses that are always found, even after several days,

in dishes where isolated specimens die. Moreover, it is the

smaller animals that disappear, those remaining being of large

size and appearing well fed. I think the circumstantial evi-

dence in favor of the view that the worms readily devour

members of their own species is very strong, but here again I

have not been able to make actual observations.

The animals are capable of living for a long time without

food. Single specimens have been kept for more than three

months in jars of clear water with no other life except prob-

ably a very few Protozoa. In one case a jar containing à

number of individuals was kept for six months. Three of

the animals survived during this time.

. No. 420.] HISTORY OF STICHOSTEMMA. 987

During starvation a great reduction in size occurs. During

the earlier stages the animal grows somewhat more slender,

but later its length is also greatly reduced. In the case men-

tioned in the preceding paragraph the three specimens found

at the end of the six months were less than five millimeters in

length, though of about the normal proportions. When placed

in the jar the specimens were of average size, Z.e., from twelve

to fifteen millimeters in length. The reduction in size was

wholly due to lack of food.

The starving forms usually acquired after a few weeks a

blackish pigmentation, granules of black pigment being depos-

ited in the body wall and increasing in abundance until in

some cases the body had lost its light yellowish color and

become a dull, dirty brown. In such cases microscopical

examination showed the whole surface of the body to be

dotted with small granules of black pigment. This pigment

is not altogether absent in newly captured and apparently well

fed specimens, but is usually not sufficiently abundant to have

any appreciable effect upon the general coloration of the

body. It undoubtedly corresponds to the pigment mentioned

by Montgomery (95, pp. 99-100) as occurring as an interstitial

tissue.

The increase in the amount of pigment is undoubtedly con-

nected with some alteration in the metabolism. It is possible

that it is not directly connected with starvation but with some

other effect of captivity; but, as far as my own experience is

concerned, it appears in specimens which have been without

food for some time. Decapitated specimens or pieces of the

body, which may live for months, sometimes become distinctly

brown or blackish.

REACTION TO STIMULI.

As regards the reactions of the animal to stimuli, it is not

intended to give here the results of extended experimentation.

Some work has been done along these lines, but the results

are reserved for a later time.

988 THE AMERICAN NATURALIST. [Vor. XXXV.

PHOTOTAXIS.

The most conspicuous reaction is the reaction to light. The

animal is negatively phototactic for even moderate light inten-

sities. The response is very distinct and definite. If a num-

ber of animals in a dish or jar be brought suddenly into diffused

daylight, each begins almost immediately to move in the direc-

tion away from the source of light. Within a few seconds

every animal in the dish is moving in this manner. Arrived

at the opposite side of the dish, the animals do not immedi-

ately come to rest, but move about restlessly, following the

sides of the dish, or in some cases even turning back toward

the light for a short distance. The final result of these move-

ments in a cylindrical jar is the aggregation of the worms in

the region of least light intensity, not at the point farthest

from the light but on the two sides at a point midway between

the points farthest from and nearest to the light, where a large

part of the light is shut off by the walls of the jar. In this

respect these animals resemble Planaria (Loeb, '93), with the

difference, however, that they are more restless and do not

form such distinct and well-marked aggregations under these

conditions as does Planaria.

In jars kept in a dark closet or in diffuse daylight on dark,

cloudy days or toward night, the worms crawl about freely ;

i.e., they are not distinctly negative for very low intensities.

Examination of a jar during the night shows the worms to be

active. They are thus evidently nocturnal in habit like Planaria.

Animals from which the whole head, including eyes and

brain, has been removed, react to light in the same manner as

do normal animals, but the reactions are slower and less exact

(Parker and Burnett, '00).

The question as to whether the reactions of Stichostemma

are strictly speaking phototactic or photopathic, whether two

different kinds of reaction occur, is perhaps not easily decided.

Certainly, so far as experiments already made indicate, there

is nothing in the reactions of the animals to light that cannot

be explained as due to the effect of different intensities with-

out relation to the direction of the rays (Holt and Lee, '01).

No. 420.] HISTORY OF STICHOSTEMMA. 989

'THIGMOTAXIS.

The thigmotactic reaction is positive and distinct. The

animal shows a strong tendency to crawl into interstices and

is usually found in masses of algae. In clear water it will com-

: monly select the angle between bottom and sides of the jar

and gradually come to rest there.

Several specimens in a jar of clear water will often aggre-

gate in a single mass, crawling over and between each other

and finally becoming nearly quiet.

A single individual, when it changes its course of movement,

does not commonly make.a curve but turns the head directly

back against its own body and retraces its course. Undoubt-

edly the animal reacts thigmotactically to its own body. Ina

similar manner on coming into contact with the body of another

individual, or indeed any object, the worm crawls along its

sides or edges, seeking the spots where the greatest portion of

its body is in contact with surfaces.

FORMATION OF **CvsTS."

Since the formation of the cyst-like slime covering, mentioned

above, appears to be more or less closely connected with the

thigmotactic reactions of the animal, it is discussed a little

more fully here.

As noted above, there is a considerable increase in the

amount of slime secreted when the animal is irritated in any

way. Any strong stimulation, e.g., strong light, handling in

a pipette, etc., appears to cause this increase in some degree.

Animals placed in a jar of perfectly clear water without algæ

or other places of concealment show a tendency to settle in

accumulations of their own slime, and it appears, in some

cases at least, that the secretion becomes more dense under

these conditions. It often happens that within a day or two

an animal becomes surrounded by a dense coat of somewhat

elastic slime, which even after removal of the animal retains

a more or less definite form. Within this slow movements

may go on continuously. Sometimes the body is so closely

990 THE AMERICAN NATURALIST. [VOL XXXV.

coiled that the covering is almost spherical, or the body may

simply be bent back upon itself at one or two points, thus

giving the covering.an elongated form. The worm often

remains in this covering for days or even for weeks, but may

leave it at any time. That the animal is not incapable of

making its way out is shown by the fact that violent stimu-

lation will usually cause movements of sufficient force to

rupture the wall of the covering and free the enclosed worm.

The most interesting fact, and one which, as I believe, throws

some light on the nature of this phenomenon, is that even

after repeated stimulation or after rupture of the wall with

needles, the animal in many cases exhibits an apparent reluc-

tance to leave the covering. The head is protruded, makes

the usual searching movements, and then is withdrawn again,

and the worm continues its slow movements within the cavity -

of its case.

I believe that the enclosure of the animal within this secre-

tion, and its apparent reluctance to leave its covering, are the

result of the strong positive thigmotaxis which other actions

show to be present. In a jar of clear water the slight accumu-

lations of slime, which are the result either of irritation by

strong stimuli or have been formed by the continuous secretion

of slime at some particular spot where the animal has rested

for a time, serve in some degree as a contact stimulus, and the

worm tends to remain in or about them. Continued secretion

leads to the formation of a more or less definite coat of slime,

within which a cavity is formed by the continuous crawling of

the worm. Various parts of the coiled and twisted body stimu-

late the sensitive head, giving the sensation of contact, and the

head may often be seen to force its way in between two moving

coils of the body, the whole body following. Thus the animal

is constantly forming new figures and arranging its body in

complex coils within the slime. From time to time, if the

animal is well fed, the faeces and intestinal mucus are added

to the wall, which thus becomes whitish and more opaque.

Within the covering, then, the impulse which leads the animal

to seek contact with surfaces is satisfied, and a strong stimulus

is necessary in order to induce emergence. This latter fact

No. 420.] HISTORY OF STICHOSTEMMA. 991

is clearly evident from the apparent reluctance noted above

to leaving the covering, even after violent stimulation. The

head being protruded finds no contact surfaces except the flat

glass, and is again withdrawn. In some cases worms which

had been removed from the mucous coverings happened to

come into contact with the latter again in their wanderings,

and after crawling about over them reéntered them, thus show-

ing in the clearest manner possible the thigmotactic nature of

their preference for these cyst-like coverings.

Another interesting habit which these worms exhibit is

closely related as regards its causes and nature to that just

described, and is also clearly thigmotactic. In many cases, in

their restless wanderings after introduction into jars of clear

water, the worms reach the surface. They may crawl any-

where upon the surface film, but prefer the margins, where the

surface of the water curves slightly upward toward the glass.

Undoubtedly the surface tension of the water affords a certain

resistance to their movements here and calls forth a contact

sensation. Into this angle between the surface film and the

side of the jar the worms push their way and then crawl com-

pletely out of the water and up the side of the jar. The slime

secreted by the surface prevents immediate desiccation, but

becomes thicker and harder as evaporation goes on, and thus

in even greater degree than the slime coverings in the water

affords the sensation of contact. The worms never attempt to

return to the water, but remain within the cases thus formed

until death ensues, which occurs within a few hours. The

whole series of reactions leading to the emergence from the

water and the failure to seek it again is thigmotactic.

If the animals do not leave the water within the first twenty-

four hours after being placed in the clear water they very rarely

do it afterward unless they are again set in active motion by

some strong stimulus. I have kept worms in a jar of this kind

for three months ; the only specimens which left the water did

so within the first day or two, although the conditions within

the jar remained equally unfavorable during the whole period.

The fact that the animals tend to leave clear water only dur-

ing a short time after being placed in it is undoubtedly due

992 THE AMERICAN NATURALIST. [VOL. XXXV.

to temporary excitement in consequence of handling, strong

light, etc., when first introduced into the jar. For some time

they are very active and many of them reach the angle

between the surface of the water and the sides of the jar.

Some of them force themselves into the angle with powerful

movements and thus leave the water. Later the movements

become less rapid, and even when they happen to reach the

angle between the water and the side of the jar they content

themselves with following it or remaining quietly in it. More-

over, others have meanwhile come into contact with other indi-

viduals or with slight accumulations of slime and tend to remain

quiet. It is probable that there is also some degree of accli-

matization to the new conditions, so that they in themselves

fail after a time to stimulate the animal as strongly as they did

at first.

It is clear that the formation of these coverings cannot be

regarded as constituting a true encystment. The animal does

not enter a quiescent stage within the covering, nor is it enabled

to survive conditions which would otherwise cause death, except

that its life may be lengthened for a few moments by the

formation of the covering after it leaves the water, or that it

may in some cases escape the attack of predatory enemies by

formation of the slime covering.

The formation of the coverings is the result of the follow-

ing conditions: the surface of the body is constantly secret-

ing slime, and stimulation or irritation causes an increase in

the amount of the secretion, perhaps also some change in

the quality; and secondly, accumulations of this slime are of

sufficient density to call forth in some degree the positive

thigmotactic reaction.

On the other hand, it is quite possible, and even probable,

that the process of encystment might arise in some such man-

ner as this in a positively thigmotactic animal. Indeed, in the

present case a change in the quality of the secretion, rendering

it less permeable to water, appears to be the only, or at least

the principal, modification necessary for the formation of a

true cyst. It is possible that in Stichostemma the secretion

may become so modified in time that the animal will succeed

NO. 420.] HISTORY OF STICHOSTEMMA. 993

in tiding over the dry seasons by becoming encysted, and will

then become capable of living in shallow, temporary pools,

which dry up during a part of the year.

At present, however, the formation of the slime coverings

cannot be regarded as an adaptation to particular conditions.

It is merely an incidental result of certain reactions of the

animal which have no relation to lack of water. In those cases

where the animal leaves the water the result is always or nearly

always death. If the nature of the secretion should for any

reason undergo change in some individuals or races, selec-

tion of those secreting the most impermeable covering might

occur.

The question as to whether the emergence from the water

ever occurs under natural conditions requires a brief consid-

eration. It might be supposed that in nature the animals can

always satisfy the thigmotactic impulse among the vegetation

and débris beneath the surface ; but since the reaction is so

distinct, and includes so large a proportion of the animals

in clear water in the laboratory, I can scarcely believe that it

does not occur sometimes in nature, and probably very often.

Moreover, as is noted below, foulness of the water often induces

the emergence, and in the lagoon where these worms are found

the water in certain localized regions may become stagnant to

a considerable degree in consequence of the decay of the luxu-

riant vegetation. I believe, therefore, that the animals must

leave the water frequently in nature, and that these individuals

are usually eliminated. It is possible that an animal which has

left the water may occasionally find its way back to it, but

I have never been able to observe a case of this kind in the

laboratory, though I have often looked for it.

It appears probable, however, that the change from an

aquatic to a terrestrial habit may be due, at least in such forms

as the turbellarians and nemerteans, to a positive thigmotactic

reaction which leads the animal to force itself into the angle

between the surface film of the water and the substratum

and thus to leave the water in the manner so well illustrated

by Stichostemma. Perhaps Stichostemma itself will become

able in time to resist evaporation and to obtain its food under

994 THE AMERICAN NATURALIST. [Vor. XXXV.

the altered conditions and so acquire a more or less completely

terrestrial habit. At any rate this thigmotactic reaction to

the surface tension of water must be regarded as one of the

causes which may induce creeping aquatic forms to leave the

water.

CHEMOTAXIS

Among the chemotactic reactions the most conspicuous

under conditions approaching the normal are the reactions to

the composition of the water. In general the animals show

some tendency to aggregate where the oxygen content is

greatest. Placed in jars with masses of alge and other ani-

mals, and kept where formation of oxygen by the plants does

not occur in any marked degree, the animals aggregate at the

surface as the water becomes foul Under these conditions

they are found either among the algze just beneath the surface

or crawling or resting beneath the surface film or about the

margins. Furthermore, as the water becomes foul, animals

which did not leave it earlier now crawl up the sides of the

glass and die there. This reaction is in this case doubtless a

mixed negative chemotactic and positive thigmotactic reaction ;

that is, the impulse to seek the surface is given by the increas-

ing lack of oxygen or the increasing quantity of decomposition

products in the water. Having reached the surface and being

perhaps still restless in consequence of the chemical stimulus

from the composition of the water, the worm leaves the water

in response to the combined stimuli, and thus dies more quickly

than it would if it remained in the water, since it possesses à

high degree of resistance to the effects of foul water.

ENEMIES.

In general, Stichostemma appears not to suffer greatly from

the attacks of predatory animals. Mutilated specimens are

rarely found, though experiments show that even relatively

small pieces of the body may live for weeks. Montgomery

(95) stated that cyprids destroy Stichostemma eilhardi. Under

ordinary conditions I have never been able to observe that

No. 420.] HISTORY OF STICHOSTEMMA. 995

cyprids actually attacked the active, healthy specimens. Occa-

sionally they crawl over the body, but very soon leave it again,

apparently without inflicting injury. In a series of experi-

ments in which two or three specimens of Stichostemma were

placed in small jars of clear water with hundreds of cyprids, it

was found that although the worms remained alive and appar-

ently uninjured for a day or two, they were finally completely

devoured. When specimens of Stichostemma were cut in

pieces and thrown into a jar containing numerous cyprids,

they were rapidly devoured. In other experiments two or

three worms were placed in a small dish with several dragon-

fly nymphs, which were given no other food. The nymphs

occasionally attacked the worms, sometimes biting them in

two. Within two days the worms were usually bitten into

several pieces or killed, but not devoured. At the conclusion

of the experiments the same nymphs took Gammarus eagerly,

thus showing certainly that they were not satiated with food.

Their attacks upon Stichostemma were often observed. The

worms were picked up and held by the large labium for a

moment, sometimes bitten completely through and allowed to

drop almost immediately, or occasionally bitten at various points

and badly crushed, but not eaten so far as could be observed.

The worms frequently crawled over the surface of the nymphs

without exciting any response from the latter. It is of course

possible that the nymphs devour small portions, but they are

certainly not sufficient to satisfy the hunger of these voracious

feeders. .

The worms are unable to escape from predatory forms by

rapid movements, for their movements are all relatively slow.

It was very evident during the experiments that the nymphs

often saw the worms when at some little distance from them

and approached them, perhaps reaching out and grasping them

for a moment only to let them drop again. In such cases the

nymphs had not the slightest difficulty in approaching the

worms.

It would appear that Stichostemma, especially when filled

with eggs, would afford an inviting morsel for many predatory

aquatic forms, and that without some means of protection its

996 THE AMERICAN NATURALIST. (VoL. XXXV.

existence in the presence of such forms in the large numbers

that occur would be impossible. Its conspicuous color and

slow movements render it easy of discovery and approach by

those forms which use the sense of sight in finding their prey.

Nevertheless the adult, the young, and the eggs appear to

remain unharmed in most cases.

I am inclined to believe that the abundant secretion of slime

from the surface of the body and its increase under irritation

renders Stichostemma more or less undesirable as an article of

food. The slime is rather tenacious, and it is possible also that

small animals are unable to pierce it and thus reach the body

of the worm.

The bodies of the worms when cut into pieces possess a

slight but distinct fishy odor. As was determined by direct

experiment, no decided taste is present. It may be, however,

that the slime or the body possesses some quality which ren-

ders it disagreeable to at least some animals; but even if this

is not the case, the presence of the slime must afford a cer-

tain amount of protection against attacks, at least against those

of small animals. |

In connection with the possibility that the slime may afford

a means of protection, it is interesting to note that in the

experiments with the cyprids the nemerteans became very

quickly surrounded by the slime covering and remained enclosed

in it until they were artificially removed the next day. In all

probability the rapid formation of the slime “cysts” was the

result of the incessant irritation of the surface of the worm by

the cyprids crawling over it. As it formed and thickened, this

slime furnished a covering which the cyprids could not pene-

trate, or could penetrate only with difficulty after some time.

SEXUAL RELATIONS.

According to Montgomery, Stichostemma eilhardi is a pro-

tandric hermaphrodite. Stichostemma asensoriatum is hermaph-

roditic, but certainly not protandric. I have never found a

purely male specimen of any size. The gonads are visible in

the living animals, and the small odcytes can be seen very early

No. 420.] HISTORY OF STICHOSTEMMA. 997

in their history. Moreover, eggs and spermatozoa leave the

gonads together when the eggs are laid. By subjecting the

animal to pressure the spermatozoa may often be seen in

the same gonads with fully grown oócytes, and when the egg

leaves the body there are nearly always a considerable number

of spermatozoa sticking to its membrane. The contact with

the water apparently serves as a stimulus, and fertilization as

well as maturation occurs very soon after the egg leaves the

body. :

From these relations it is evident that self-fertilization is

possible. As a matter of fact it is undoubtedly the rule. In

a large number of cases specimens which had been isolated for

a week or two laid eggs, and these were always fertilized and

developed. I have often removed the eggs artificially by cut-

ting the animal into pieces, and in every case where fully

grown eggs were present, fertilization occurred. The fully

grown eggs can be distinguished by the fact that no peduncle

of attachment is present. In younger eggs the membrane is

incomplete, and the egg is in continuity with the protoplasm of

the gonad through a small stalk or peduncle. I have never

been able to obtain unfertilized eggs. Finally, sections of the

animal with full-grown egg cells always show the spermatozoa

clustered over the surface of the egg or massed in some part of

the gonad, while specimens with younger egg cells show stages

of spermatogenesis, and still younger stages show merely the

young oócytes and the sperm mother-nuclei. The spermatozoa,

or some of them, always pass out with the egg, and I believe

that each egg is fertilized by the spermatozoa of the same

bsolutely no evidence that spermatozoa

gonad. There is a

None have ever been

reach the oócytes in any other manner. |

found in the slime which encloses the egg after it is laid in

None have been found in the water in

which many specimens had been kept. But I think the strong-

est evidence in favor of self-fertilization lies in the fact noted

above, that spermatozoa can be seen upon the membrane of

each egg cell and that the penetration of these spermatozoa

can be observed immediately after the egg cell reaches the

water. Asin S. eilhardi, the genital pores are not preformed,

the normal manner.

998 THE AMERICAN NATURALIST. [Vor. XXXV.

so that spermatozoa cannot enter from the exterior before egg-

laying unless they make their way through the body wall.

That this is not the case is shown very clearly by the time of

their appearance, their relations to the remainder of the gonad,

and by the fact that fertilization itself always occurs outside of

the body. The structure and relations of the various parts of

the gonads and the oógenesis afford a large number of inter-

esting facts, but these need not be discussed in detail here.

There is strong evidence that each gonad is always hermaphro-

ditic after its original formation, and that egg cells and sperma-

tozoa almost always develop together. : i

I believe that cross-fertilization occurs very rarely in Sti-

chostemma in nature, even if it ever occurs. The egg cells

when they pass out of the body are immediately enclosed in

the slime from the surface, and the spermatozoa with them.

While it is, of course, possible that a few spermatozoa might

make their way through the slime to the water and then again

through the slime surrounding the eggs of other individuals,

the most careful observations have never given the slightest

evidence that this actually occurs. Furthermore, if it did

occur, the second set of eggs would be fertilized before such

a spermatozoón could reach them.

The only possible conclusion is that self-fertilization is the

rule in S. asensoriatum.

As a rule only one egg. is formed in each gonad, but excep-

tions are not uncommon, though more than two egg cells in a

single gonad have never been observed.

EGG- LAYING.

The normal process of egg-laying has not been observed in

Stichostemma, although numerous egg strings have been found

in the jars in which the animals were kept. The extrusion of

eggs from the body in consequence of irritation or injury has

often been observed. If an individual containing fully grown

eggs be cut into several pieces, the eggs, or many of them, will

often be extruded after a short time, apparently as the result

of contractions following the injury. The first step in the

No. 420.] HISTORY OF STICHOSTEMMA. 999

process is the breaking of the body wall at a definite point,

T.e., the appearance of the pore. Immediately after this the

egg begins to flow out. The diameter of the pore is very

small, so that the egg is greatly distorted during its passage.

Indeed, it appears to flow out like rather thick fluid. At first

a small, rounded, transparent protuberance appears at the outer

opening of the pore; this is the egg membrane. Then the

yolk granules and cytoplasm may be seen flowing out through

the pore, and the part outside increases continually in size as

more and more of the egg substance reaches it. The part of

the egg outside the pore and that still within the body are

connected by the slender band of egg substance which is pass-

ing through the body wall. The nucleus usually remains in

the body until at least half of the volume of the egg has flowed

out. The small size of the genital pore presents an obstacle

to the passage of the nucleus, and often the passage of the egg

ceases until some contraction of the body forces the nucleus

through the narrow duct and it suddenly pops out through the

pore. After the passage of the nucleus the remaining portion

of the egg flows out rapidly, and the whole egg assumes almost

immediately its spherical or nearly spherical form. The sper-

matozoa on the egg membrane can often be seen clearly during

the passage of the egg, and it is evident that their movements

begin as soon as they reach the water.

It is difficult to understand how definite areas of differen-

tiated protoplasm could persist or maintain their proper posi-

tion during this outflowing of the egg. To all appearances, at

least, the egg flows out like a thick liquid under some pressure,

and it appears scarcely possible that its parts maintain the

same relative position before and after its emergence. The

whole egg is enclosed, however, in a very thin layer of trans-

parent protoplasm, an ectoplasm, and if the pressure is too great

this layer is ruptured, and the cytoplasm and yolk flow out asa

viscid, granular fluid, which tends to reassume the spherical

form. It is evident that the cytoplasm is elastic and that the

reassumption of the spherical form by the egg is due to this

elasticity, Eggs in which the ectoplasmic layer has been rup-

tured do not develop, or at least development is confined to

IOOO THE AMERICAN NATURALIST. [Vor. XXXV.

the uninjured portion. All such cases of partial development

die in early stages. When the yolk is exposed it appears to `

absorb water very rapidly, the egg or the injured part of it

increases in size and becomes more transparent, and sections

show vacuoles in place of the yolk gran-

ules. The death of the egg always

accompanies or follows this absorption

of water. Apparently the presence of

the ectoplasmic layer prevents this absorption of

water by the yolk. The egg is very sensitive to inju-

ries involving the rupture of the ectoplasmic layer.

All attempts to kill a part. of the egg ended in the

death of the whole a little later.

The eggs are laid normally in strings of about

the length of that portion of the body containing

gonads. The eggs are deposited in two rows, each

row representing the eggs from one side of the body.

All the eggs are enclosed as they emerge in slime

secreted by the dermal glands, and then the worm

crawls out from between the two rows of eggs which

remain attached to the substratum. As shown by

the figure, the eggs at one end of the string are

placed irregularly, as if the secretion enclosing the

eggs had been stretched. This end is undoubtedly

the anterior end of the string, ż.e., the end at which

the worm leaves after laying, and its form is due

to the fact that some of the slime adheres to the

body of the worm, and together with the enclosed

eggs is dragged after it as it moves away.

Ess d The cleavage of the eggs is typically spiral, all the

“mm cells being nearly equal in size during the earlier

stages. The development is of the direct type. The surface

of the embryo becomes ciliated, and it moves about within the

egg membrane for a day or more before leaving it. The spher-

ical embryo begins to elongate after three or four days, leaving

the membrane when about one-half millimeter in length. The

young animal possesses at first four eyes, the posterior pair

of the adult being formed at a later stage than the others, thus

No. 420.] HISTORY OF STICHOSTEMMA. IOOI

probably indicating their later origin in the species. I have

not been able to ascertàin whether the food of the young

is the same as that of the adult.

THE BREEDING SEASON AND ITS RELATION TO THE

ENVIRONMENT,

The breeding season of Stichostemma extends from May to

November or December, according to the temperature of the

water. Practically every specimen taken soon after the ice

melts in the spring contains the growing oócytes in the gonads,

and these reach their full size, and egg-laying begins as soon as

the shallow water along the shores becomes sufficiently warm.

During the whole of the season from May to November the

very young worms are abundant in sizes, varying from a milli-

meter up, and the eggs are often found in the alge with the

worms.

Egg-laying occurs freely in the laboratory. Often when the

algze containing worms are brought in and allowed to stand for

twenty-four hours, a large number of egg strings will be found

near the surface and on the sides of the jar. Apparently the

eggs are always laid during the night or in darkness when the

animals are moving about freely. Even in jars of clear water

and without food the eggs are often deposited abundantly dur-

ing the first few days of captivity, but individuals which do not

lay eggs during that time are not likely to do so later, though,

as experiment shows, they contain fully grown oocytes.

Eggs can be obtained in the laboratory at any time during

the year when the worms can be found in moderate numbers.

During December many individuals with oócytes at least half

grown are found. In nature these eggs are probably not laid

until the following spring, but if the worms are kept at room

temperature in the laboratory a number will usually deposit

eggs within a week or two, and this will occur even if the

animals are kept without food. By these observations and

those made upon animals under natural conditions during

nearly two years, I am convinced that the limits of the breed-

ing season are determined chiefly by the temperature of the

IOO2 THE AMERICAN NATURALIST. [Vor. XXXV.

water. Probably food is actually an important factor, but its

abundance is also more or less dependent on the temperature

of the water, so that the latter must be regarded as the most

important factor in determining the time of egg-laying.

That food is not of prime importance is clearly shown by a

large number of observations in the laboratory. In animals

which contained only small oócytes when taken, and which are

kept in clear water without food, the growth of the oócytes

will continue, and within a week or two eggs may be laid.

This has been found to be the case even during the winter,

and it shows very clearly that the important factor is the

degree of temperature which permits the metabolism necessary

for the formation of the yolk and the growth of the odcyte

in general. In such cases the oócytes complete their growth

either at the expense of the other tissues or some store of

nutritive substance within the body. The body of the animal

may even decrease somewhat in size during the growth of the

oócytes. In some way the energy of the body is applied

to the completion of these important products, even at the

expense of the remainder.

If an individual containing young oócytes of small size je

cut transversely near the middle of the body, and the pieces

placed in clear water without food, both will usually live for

months. The posterior piece does not regenerate, but the

oócytes grow much more rapidly in it than in the anterior

half, and may attain their full growth and be capable of

fertilization and development before those contained in the

anterior half have reached more than half or two-thirds of

their full size. I can explain this fact only on the basis of

the difference in activity of the two halves. The anterior

half, possessing a head and brain, behaves like a normal

animal, moving about actively, while the posterior half, being

without a brain and never regenerating a head or brain, is

much less active and less sensitive to stimuli. In other

words, much of the potential energy of the body which is

employed in the anterior piece in producing the various move-

ments, and probably in a more intense general metabolism,

is directed in the posterior piece to the completion of the

No. 420.] HISTORY OF STICHOSTEMMA. 1003

growth of the oócytes. These cases afford a most instructive

example of the relation to each other of some of the various

so-called vital activities. In the presence of the brain and

anterior portion of the nerve cords the bodily movements

appear to take precedence over other functions, even though

the sexual products are not completed. But when the anterior

portions of the nervous system are absent the animal, being

without the power of regenerating the lost parts, is less affected

by stimuli, and a larger portion of the potential energy

is devoted to the elaboration of the sexual products. This

process must, as a matter of fact, be regarded as the function

next in importance to that of carrying out the normal move-

ments and responses to stimuli.

Mention has been made of the fact that animals kept in cap-

tivity without food are not likely to deposit their eggs after the

first week or two of captivity, even though artificial removal of

the egg cells may show them to be capable of fertilization and

development. In such cases the fate of the egg cells is quite

remarkable. They may remain unchanged in appearance for

two or three weeks, but after a time evidences of alteration in

their structure are visible even in the living animal. The egg

cells become less opaque in appearance and a clear space appears

about each, apparently the result of thickening and swelling of

the membrane. Later many of the eggs break up into round

which sections show to be merely disin-

masses and granules,

But the more remarkable

tegration products and not cells.

fact is that, as the changes go on, many OF all of the eggs pass

out of the gonads and come to lie within the alimentary canal,

where they serve as food and are actually digested. Continued

observation from day to day of jsolated specimens leaves no

doubt that this actually occurs. In many cases the egg cells

are still intact when they enter the alimentary canal and can

clearly be seen there, moving to and fro with the movements

and contractions of the body. Disintegration and digestion of

the egg goes on rapidly in the alimentary canal, however, and

in a day or two the whole intestine may be filled with a gran-

ular mass which represents the cytoplasm and yolk of the egg.

In case disintegration of the egg has begun before it enters

1004 THE AMERICAN NATURALIST. [Vor. XXXV.

the alimentary canal, it very rapidly becomes a granular mass

recognizable as an egg only through its known origin.

That these disintegrating eggs actually serve the animal as

food is evident from the fact that in a number of cases the

passage of the eggs to the alimentary canal and their diges-

tion was followed in a few days by the appearance and partial

growth of new oócytes in the gonads, each individual worm

being isolated meanwhile in clear water and without other

food during the whole period. In cases where the eggs were

normally laid by similarly isolated specimens, new oocytes

appeared only when the first set had been laid almost imme-

diately after the beginning of the experiment, so that the

animal was comparatively fresh and probably still contained a

certain surplus of nutritive material. Moreover, these oócytes

never reached a large size and usually underwent disintegra-

tion at a later period.

The manner in which the disintegrating egg cells reach the

alimentary canal is somewhat obscure. Rupture of the walls

of the gonad and the intestine is necessary. I believe that

this rupture is brought about chiefly in consequence of the

increasing disproportion between the size of the body and the

size of the egg as the starving animal decreases in size, and

that finally the egg becomes too large for the lateral regions

of the body and is forced into the alimentary canal by the

movements and contractions of the body. A study of sec-

tions reveals a decreasing thickness in the walls of the intes-

tine and gonads, as well as other histological changes, but

these need not be discussed here.

The passage of the egg cells to the intestine occurs in pieces

which have been deprived of head and brain, as well as in

normal individuals.

ABSENCE OF FISSION.

Benham (97) has described a process of spontaneous fission

in Carinella which probably serves as a means of distribution

of the sexual products, and Wilson (00) states that Cere-

bratulus fragments spontaneously at the close of the breeding

No. 420.] HISTORY OF STICHOSTEMM A. IOO5

season, apparently as a means of getting rid of a portion of

the body which has become of little use.

I have never observed spontaneous fission or fragmentation

in Stichostemma, and have never found pieces or fragments

which might be the result of such fission. Occasionally speci-

mens are found which show slight constriction at some points,

but isolation and continued observation of these individuals

have always been without result. These constrictions appear

not only in individuals with mature sexual products, but in

other stages, and usually disappear after a few days. More-

over, they are frequently irregular, being more pronounced on

one side of the body than on the other. I am inclined to

believe that they are merely the results of slight injuries,

for slight cuts in the sides of the body produce results which

are indistinguishable from those found in freshly collected

specimens.

HULL ZoóLoGiCAL LABORATORY,

UNIVERSITY OF CHICAGO, July, 1901.

LITERATURE.

'97 BENHAM, W. B. Fission in Nemertines. Quart. Journ. Mir. Sci.

Vol. xxxix.

'98 BónwrG, L. Beiträge zur Anatomie und Histologie der Nemertinen.

Zeitschr. f. wiss. Zool. Bd. lxiv, Heft 3.

01 Hott, E. B., and LEE, F. S. The Theory of Phototactic Response.

Am. Journ. Physiol. Vol. iv, No. 9.

'90 Lors, J. Der Heliotropismus der Thiere und seine Uebereinstim-

mung mit dem Heliotropismus der Pflanzen. Würzburg.

'03 Lors, J. Ueber künstliche Umwandlung positiv heliotropischer Thiere

in negativ heliotropische und umgekehrt. 4 rch. f. d. ges. Physiol.

'95 Pea T. H. Stichostemma eilhardi zov. gen., nov. sp.

Zeitschr. f. wiss. Zool. Bd. lix.

'96 MONTGOMERY, T. H. Stichostemma asensoriatum nov. sf. a

Fresh-Water Nemertean from Pennsylvania. Zool. Anz. Bd. xix,

No. 515.

1006 THE AMERICAN NATURALIST.

"97 MONTGOMERY, T. H. On the Connective Tissue and Body Cavities

of the Nemerteans, with Notes on Classification. Zool. /Jahri.,

Abth. f. Anat. und Ont. Bd. x.

'00 PARKER, G. H., and BURNETT, F. L. The Reactions of Planarians

with and without Eyes to Light. Am. Journ. Physiol. Vol. iv,

No. 8.

'85 SILLIMAN, W. A. Beobachtungen über die Süsswasserturbellarien

Nordamerikas. Zeitschr. f. wiss. Zool. Bd. xli.

'97 VERWORN, M. Allgemeine Physiologie. Jena.

'00 WiLsow, C. B. The Habits and Early Development of Cerebratulus

Lacteus (Verrill); A Contribution to Physiological Morphology.

Quart. Journ. Micr. Sci. Vol. xliii, pt. 1.

AN EXTRAORDINARY ANT-GUEST.!

WILLIAM MORTON WHEELER.

Since Sir John Lubbock's discovery of the association of

certain Phoridze (Phora formicarum Ver. and Platyphora Lub-

bocki Ver.) with ants,? several interesting additions to the

number of known myrmecophiles and termitophiles belonging

to this family of Diptera have been recorded. Among others

a singular group of wingless Phoride (“ Stethopathidz’’) has

been described by Wasmann? from specimens captured in the

nests of termites, and Brues has recorded the occurrence of

several somewhat similar forms in the nests of certain Texan

ants belonging to the genera Eciton and Solenopsis.* Very

recently Pergande® has published an entertaining note on the

habits of a phorid (Apocephalus pergandei Coq.) which causes

the workers of the common wood-ant (Camponotus pennsyl-

vanicus) both figuratively and literally to “lose their heads,"

for the interior of this important portion of the ant's body

furnishes the fly-larva with food and a retreat for pupation.

Far less injurious to its hosts is a myrmecophilous phorid

which I propose to describe in the following pages. Unfortu-

nately I have seen only its larval and pupal stages, but these

are passed under such unusual circumstances that the discov-

ery of the imaginal insect could scarcely supply details of very

great additional interest. As the insect is not very common,

many months may pass before I am able to complete my study

1 Contributions from the Zovlogical Laboratory of the University of Texas,

à Lubbock, Sir John. Ants, Bees, and Wasps. 1881.

8 Wasmann, E. Termitoxenia, ein neues flügelloses, physogastres Dipteren-

genus aus Termitennestern, Zeitschr. f. wiss. Zool. Bd. xlvii, No. 4 (1900),

XXIII

pp. 599-617, Taf. X .

4 Brues, C. T. Two New Myrmecophilous Genera of Aberrant Phoridz from

Texas, Am. Nat., vol. xxxv, No. 413 (May, 1901), pP- 337-356, 11 figs.

5 Pergande, Theo. The Ant-decapitating Fly, Proceedings Ent. Soc. Wash.,

vol. iv, No. 4 (1901), pp- 497-502, 2 figs.

1007

1008 THE AMERICAN NATURALIST. [Vor. XXXV.

of its life history. There is some excuse, therefore, for pub-

lishing these fragmentary observations.

On October 27 of last year I made a short excursion to

Mt. Barker, which is hardly more than an hour's walk from

the university at Austin, Texas. The woods about the base and

on the slopes of the elevation are favorite nesting grounds for

the large black ponerine ant, Pachycondyla harpax. In Octo-

ber this ant is rearing its second brood of larvz and pupa, hav-

ing completed the education of its first brood during June and

July. Wishing to continue some observations on the habits

of Pachycondyla, I dug up one of the largest nests I could find

and carried it home in a bag. On transferring it to a Lubbock

nest I took the census of the colony and found it to comprise

25 worker ants,.13 cocoons, 8 mature larve, 7 immature larvae,

and a packet of eggs. While counting the larvae, which are

shaped like the well-known cucurbitaceous product known as

the ** crook-necked squash," and covered with hairy tubercles,

I noticed that six of the largest and one of the smallest pre-

sented an unusual appearance. Each of these seemed to wear

about its neck a huge collar, —a sort of Elizabethan ruff, —

consisting of a curled larva. That this could not be another

ant-larva was apparent from a moment's examination. In all

cases it almost completely encircled the ant-larva in the region

of the first abdominal, or in some cases the metathoracic, seg-

ment. The posterior end was provided with a sort of disk,

which adhered so tightly that both larvae could be killed in

alcohol without separating. The collar-like larva was broad

behind the middle, but tapered anteriorly to a very slender

thoracic region and head provided with small jaws. These

were supported by a chitinous frame-work of such character-

istic structure as to show that the adult form must be a true

dipteron fly. The very smooth and tense integument, which

was armed with some short, hooked bristles, was very trans-

parent, so that the peristaltic movements of the viscera were

clearly visible. One of the adult Pachycondyla larvze with its

living collar is shown in Fig. 1.

! For an account of the habits of this ant, see my paper, A Study of Some

Texan Ponerinz, Biol. Bull., vol. ii, No. 1 (Oct., 1900), pp. 1-31, Figs. 1-10.

No. 420.] AN EXTRAORDINARY ANT-GUEST. IOO9

As soon as the ants had been transferred to the Lubbock

nest they were given a number of young larva of Camponotus

sansabeamus. These they soon proceeded to malaxate with

their mandibles, twirling the morsels about in the mean time

with their fore legs and lapping up the exuding juices with

their tongues. Finally they deposited the crumpled and pulpy

remains of the Camponoti on the trough-like ventral surfaces

of their larva, which had been previously placed on their

backs in a rough chamber dug in the earth of the nest. This

chamber was immediately under the glass

roof-pane, so that further developments

could be closely observed with a pocket

lens. Each ant-larva at once stretched

forth its head eagerly and began to devour

the viands with which it had just been

provided. At the same moment the dip-

teron larva, too, as if sniffing the odor of

the fresh food, unwound its tapering neck

from the ventral surface of its host, and,

without shifting the attachment of its

posterior end, at once plunged its man-

dibles into the food. Under the lens both

larvae could be seen greedily dining side

by side till the last particle of the Campo-

notus larva had been consumed or prema- dpi

turely removed by the worker ants. Fic. r, — Adult larva of Packy-

When the ant-larve were huddled close eic byes Fesi

together, a collar-like larva was sometimes

observed to reach over and help itself from the food supply of

a neighbor ; but even when thus compelled to crane its neck

to the utmost, it never shifted the attachment of its caudal end.

Sometimes when there was no food within reach it would tweak

with its sharp little jaws the sensitive hide of a neighboring

ant-larva, till the latter squirmed with pain. It would some-

times even tweak its own host, as if to make it wriggle and

perhaps thereby incite the worker ants to bring a fresh supply

of provisions. i '

The following day two living myriopods (Lithobius) were

IOIO THE AMERICAN NATURALIST. [VOL. XXXV.

placed in the nest. During the morning hours they were

killed by the Pachycondyla workers, shorn of their many legs,

cut up into pieces of convenient length, malaxated, and fed to

the larva as on the preceding day. And again I was able to

witness the strange banquet — the dwarf reaching down from

the shoulders of the-ogre and helping himself from the charger

formed by the trough-like belly of his host. The same obser-

vation was repeated on several successive days. Pieces of

various ant-larvae, beetle-larvee, Lithobius, Scutigera, Oniscus,

— all were served up to the ant-larve and partaken of with

great relish by the dipteron larvae as well. There could be no

doubt that the latter were true commensals, — perhaps the

most perfect commensals, in the original meaning of the term,

to be found in the whole animal kingdom !

As one of the smallest Pachycondyla larva, scarcely one-

fourth grown, bore a very small dipteron larva, it is, perhaps,

safe to say that the ant-larva acquires its commensal at a very

early age. The two then grow up together, so that there is

always a certain relation between the two kinds of larvae —

large Pachycondyla larva bearing large commensals, and vice

versa. The worker ants lick and cleanse the commensals at

the same time that they are caring for their own larva. This

is usually done just after meals. Since, during this operation

of cleansing, the ants spend no more time over the commensals

than they do on a similar area of the body surface of their own

larvae, it would seem that they are not even aware of the exist-

ence of the commensals. To these nearly blind ants, which

must rely almost exclusively on their senses of touch and smell,

the larvae bearing commensals, if distinguished at all from indi-

viduals without such attendants, would probably be perceived

merely as having unusually protruding necks. But there is

nothing to indicate that these insects are really capable of per-

ceiving such differences in their environment.

At this time an amusing interlude in these observations was

furnished by introducing into the nest a few living * pill-bugs "

(Armadillidium sp.). For two whole days the Pachycondyla

workers hunted these isopods from one corner of the nest to

another. An Armadillidium would scurry along rather quickly

No. 420.] AN EXTRAORDINARY ANT-GUEST. lOII

and, when overtaken, escape death by rolling itself up into a

hard ball, which the ants tried in vain to perforate with their

mandibles. As soon as they desisted the Armadillidium would

unroll itself, and the chase would begin again. Finally I was

compelled to remove the crustaceans, as the larvze seemed to

be in danger of being injured in the excitement of the chase.

On the sth of November I obtained satisfactory evidence

that the dipteron larva is not obliged to remain always with the

same Pachycondyla larva. During the night one of the large

larvae had moved and attached itself to the first abdominal seg-

ment of an ant larva which already bore a commensal around

its metathoracic segment. The two larva were oriented in

opposite directions, z.¢., with their heads reaching around oppo-

site sides of their host. Subsequently one of these commensals

moved to an unoccupied Pachycondyla larva. I was not present

when the change occurred, nor was I able to determine whether

it was the originally stationary or the adventitious larva that

moved. Although this observation makes it certain that the

dipteron larvae can shift their position from one host to another,

I am convinced, nevertheless, that they must do this with great

reluctance and only under urgent circumstances, such as extreme

hunger, the death of the larva to which they are attached, or,

perhaps, when fully mature and about to pupate.

As the days passed, the mature ant-larvee spun their brown

cocoons one by one, and one by one the mature commensals

disappeared. Did they also pupate and for this purpose conceal

themselves in the soil of the nest? or had the ants at last

detected the villains and converted them into food for the

larvae which had not yet pupated? or did they stick to their

hosts and pupate within the cocoons? In order to ascertain,

if possible, the true state of affairs, I transferred the whole

colony to a fresh nest and examined the soil of the old nest

with great care. There were no traces of the missing commen-

sals. The only remaining resource was to open the cocoons.

Several of the cocoons which had been taken with the nest

October 27 had hatched, but by November 10 there were still

thirteen cocoons in the nest, as several of the larvae had pupated

in the mean time. Five of these were opened, and in two,

IOI2 THE AMERICAN NATURALIST. (Vor. XXXV.

which contained pseudonymphs of Pachycondyla and were, there-

fore, of recent formation, commensals were found! Having

shared the table of their host, they had come to share its bed as

well. The dipteron, too, had pupated after the manner of its

kind — forming a puparium ; z.e., instead of spinning a cocoon

like the ant-larva, the dead larval skin, somewhat shriveled and

contracted, was used as an envelope, and within this the pupa

proper was formed.

This puparium, represented in Fig. 2, is from 2.25 to 2.5 mm.

long and clearly of the cyclorhaph type. It is elliptical, much

flattened dorso-ventrally, especially

along the edges, which are thin and

hyaline andalmost alate. The brown-

ish dorsal surface is thrown into deli-

cate and irregular transverse wrinkles.

Anteriorly, in the thoracic region,

there is a pair of short black res-

piratory tubes. The ventral surface

is very glabrous and distinctly paler

than the dorsal surface. From its

small size, cyclorhaphous character,

and resemblance to the puparium of

a typical Phora bred by one of my

: assistants, Mr. C. T. Brues, from

Fic. 2.—Puparium of phorid com- Some spider's eggs, I conclude that

mensal from ae cocoon of Pachy- . . .

debis Mer the insect under consideration must

be one of the Phoridæ.

Subsequently, several other cocoons were opened and two

more were found to contain the puparia of commensals. In all

four cases the puparium was invariably located in the caudal

pole of the cocoon, just to one side of the black blotch of ejecta

deposited by the ant-larva before becoming a pseudonymph.

At this point the puparium was immovably stuck to the wall of

the cocoon by means of its smooth ventral surface. Its anterior

end was directed towards the cephalic pole. As there is always

considerable space, especially at the posterior pole, between

the walls of the cocoon and the enclosed ant-pupa, the much-

flattened fly puparium did not in the least crowd its host.

No. 420.] AN EXTRAORDINARY ANT-GUEST. 1013

Reflection shows that the position of the puparium in the

posterior pole of the cocoon, though the reverse of the position

of the larval commensal with respect to its larval host, is the

only one which can be maintained by the commensal with per-

fect safety. Like other ants, the Pachycondyla leaves its cocoon

through a rent in the anterior pole. This rent is certainly made

by the mandibles of the hatching ant, and it is possible that the

callow insect may succeed in making its way out of the cocoon

without any assistance from the workers. I have hitherto

failed, however, to surprise one of these ants in the act of

hatching. But even if the obstetrical aid of the workers is

necessary, as it is in the more highly specialized Camponotinz,

any position for the commensal puparium, except at the poste-

rior pole of the cocoon, might be fatal, for the struggling jaws

and legs of the emerging ant and the jaws of the assisting ants

would certainly be very liable to cut into any delicate object

attached to the anterior or median walls of the cocoon. This

danger, of course, would not exist if the fly hatches before the

ant and is able to perforate the cocoon and escape. It is alto-

gether more probable, however, that the ant hatches before the

fly in the manner suggested below.

It would be interesting to know what the commensal larva is

doing while the ant-larva is weaving its cocoon. Does it move

about to avoid the swaying jaws of the spinning larva? or does

it take up its position from the first at the posterior end of

the larval ant and there remain motionless while the posterior

pole of the cocoon is being completed? It is very difficult to

answer these questions. The fact that the posterior poles of

all the cocoons containing puparia were somewhat distorted,

being broader, more obtuse, and more irregular than the normal

cocoons, would seem to indicate that the ant-larva may modify

this end of its cocoon for the better accommodation of the com-

mensal. I am inclined to believe, however, that the distortion

may be produced by the dipteron larva while attaching itself

just before pupating to the newly woven and still plastic cocoon.

At this point my observations end. The phorid puparia

were kept for several weeks in what I supposed to be the

proper conditions of warmth and moisture, but to my intense

IOIA4 THE AMERICAN NATURALIST. [Vor. XXXV.

disappointment they failed to hatch. During the autumn and

winter frequent and diligent search was made for more of the

commensals in all the Pachycondyla nests I could find, but in

vain. Finally, during the latter part of May of the current

year, I discovered in a very different locality two Pachycondyla :

nests which contained a few phorid larva. This discovery

proved that the phorid is double-brooded like its host. But

the larvae were very small and attached to such very young

ant-larvee that I despaired of being able to raise them in my

artificial nests as far as the imaginal stage. It remains, there-

fore, for the future to fill this gap in my observations.

As a small contribution towards filling this gap I venture to

advance the following conjecture concerning the circumstances

under which the phorid fly probably hatches. I assume that

the ant must hatch before the fly. Now very soon after the

former hatches the useless cocoon is always carried by a worker

and placed on the refuse heap, which in the natural nest is

often almost entirely made up of the empty cocoons of from

one to several broods of ants, and lies in a rather dry and well-

ventilated spot immediately beneath the stone covering the

nest. Along with the cocoon is carried the phorid puparium

still adhering to the wall at its unopened posterior end. Thus

after a privileged existence as free pensioner and bedfellow to a

generous host, it is unwittingly carried away in the worn-out

bedclothes and consigned to the family rag pile. Here the

small and probably very active fly hatches, leaves by the wide-

open front door of the cocoon, and, after mating, either returns

to lay a few eggs in the galleries of its former host, or flies

away to oviposit in some other Pachycondyla nest. Thus the

simple assumption that the fly hatches later than the ant

renders it unnecessary to suppose that the fly possesses some

peculiar means of perforating the tough wall of the cocoon,

and also accounts for the position of the puparium in the pos-

terior pole, where it would be completely concealed from the

workers even after the escape of the callow ant.

In conclusion the reader’s attention may be directed to cer-

tain particulars of special interest in connection with the life

history of the Pachycondyla commensal:

No. 420.] AN EXTRAORDINARY ANT-GUEST. IOI5

First, the peculiar habits of the phorid show clearly that the

ponerine method of feeding the larvae with comminuted insects

is not only the typical but the only method employed by these

ants, for such a commensal would certainly starve if the Pachy-

condyla larve were carefully fed like the larvae of Camponotus

and Formica, by regurgitation of liquid food from the mouths

of the workers. The phorid profits by a peculiarity in the

behavior of its host, and thereby demonstrates — by one of

Nature's experiments — that Pachycondyla harpax cannot feed

its young by regurgitation.

Second, in the peculiar symbiotic relationship existing

between the phorid and the ants, the adaptations are all on

the part of the former, whereas the latter pursues its ancient

and well-established mode of life uninfluenced by and apparently

in complete ignorance of the very existence of its little guests.

Even the distortion of the cocoon may be due, as I have sug-

gested, entirely to the activity of the phorid. These strikingly

unilateral adaptations are probably to be explained on the -

ground that the phorid in the Pachycondyla nest is as careful

and conservative — if such terms may be used — of the life

and welfare of its host as Pergande's Apocephalus is ruth-

less and destructive, The small amount of food consumed by

the former can hardly be a serious drain on the provisioning

instincts of the Ponerine. The larvae bearing the commensals

were certainly as large and healthy as any others in the nest,

and produced perfectly normal pupa, which in the cases

observed all lacked the imaginal disks for the wings and were

therefore of the worker type. Whether the presence of the

commensals by reducing the amount of food even to a slight

degree could inhibit the development of queen larva and thereby

convert them into workers, involves a problem as interesting

as it is difficult to solve in the present state of our knowledge.

Third, the commensal of Pachycondyla, together with the

Phoridæ mentioned in the introductory paragraph of this

paper, shows that the flies of this family are well MORE care-

ful study. While the Stratiomyide, Syrphidze, Midaide, etc.,

certainly make more attractive cabinet specimens and pret-

tier monographic plates, it is very doubtful whether the life

IOI6 THE AMERICAN NATURALIST.

histories of most of these conspicuous Diptera can compare in

interest with those of the tiny, dull-colored Phorida. That

the study of the smaller and inconspicuous Diptera, like the

Culicidz, has been unduly neglected is clearly shown uy recent

medical researches on the malaria parasite.

COLEBROOK, CONN., August 16, 1901.

FRIEDENTHAL'S EXPERIMENTAL PROOF OF

BLOOD-RELATIONSHIP:!

HERBERT W. RAND.

IN recent years important evidence in favor of Darwin's

theory of the descent of man has increased. Eugen Dubois

found in Pithecanthropus erectus the much-sought-for missing

link between man and living anthropoid apes. Selenka has

shown that, of the catarrhine apes, the living anthropoid apes

resemble man in having a placenta discoidalis capsularis, while

all other catarrhines have a placenta bidiscoidalis. Ernst

Haeckel has accumulated evidence showing that man has

descended from a line of extinct catarrhines whose immediate

ancestors belonged to the group of tailless anthropoid apes,

and whose more remote ancestors belonged to the Cynopithe-

cini. The evidence’ afforded by comparative paleontology,

comparative embryology, and comparative anatomy agrees in

supporting Darwin’s hypothesis.

To all this evidence is now added evidence depending upon

the similarity in chemical composition of the blood of closely

related animals. Comparative chemical analyses of blood are

attended with great difficulties because of variations in the

blood depending upon the varying conditions of nutrition in

the animal. Landois, in his researches on the effects of

animal blood transfusions upon human diseases, attributed his

failures, so far as the beneficial effects of the blood transfusion

upon the disease-were concerned, to the dissolving of the red

blood corpuscles of the injected blood by the blood of the

recipient. Experiments upon lower animals, in the way of

ome animal of a remote species, led to

injecting blood from s :

The animals

the same results as the experiments upon man.

by Hans Friedenthal: Ueber einen experi- -

1 An extended abstract of a paper Gees DLE PU

mentellen Nachweis von Blutverwandschaft, Archiv für Anatomie u Dy f1-

ologie, physiologische Abtheilung, Hefte 5 und 6, 1900.

1017

IOIS THE AMERICAN NATURALIST. (VoL XXXV.

exhibited pathological conditions with development of fever, or

very often died soon after the transfusion. The hemoglobin of

the foreign erythrocytes appeared in the bladder immediately

after the transfusion, and often more hemoglobin was given

off than was contained in the injected blood. The dissolving

of red blood corpuscles by the serum of foreign blood was first

observed under the microscope by Creite.

Landois found, in some cases, that the dissolving of the

injected blood elements did not occur. In transfusions between

the horse and ass, wolf and dog, rabbit and hare, no hemoglobin

appeared in the urine; the animal, even after the injection of

a large amount of blood, showed no pathological symptoms, but

behaved precisely as after an injection of blood from one of its

own kind. Landois concluded that only animals of very closely

related species can exchange blood with impunity. The question

then arises, How closely related must animals be in order that

their blood may be “ physiologically identical ” ?

The method of blood transfusions is not adapted to extensive

comparative researches. But the *globulicidal" action of a .

blood serum can be observed to advantage in a test-tube. To

10 c.cm. of the serum of some mammal let there be added three

drops of foreign blood from which the fibrin has been removed,

and let the mixture be kept at 38? for fifteen minutes. The

mixture is at first opaque, because of the added erythrocytes,

but at the end of the fifteen minutes the fluid regains its trans-

parency and acquires a bright red, owing to the dissolving of

the coloring matter from the added red corpuscles. Buchner

found that if the serum is heated to 55? its power of dissolving

foreign corpuscles is quite lost. Buchner also found that serum

would dissolve the white corpuscles of foreign blood. 3

The loss of the dissolving power of serum as a result of

raising the temperature shows that the dissolving power

depends upon chemical and not upon physical factors, since, by

heating, the osmotic tension of the serum is not changed.

The blood of cold-blooded animals does not respond to the

test so readily as the blood of mammals, the nucleated cor-

puscles evidently possessing greater resistance. Non-defibri-

nated mammalian blood requires longer for the dissolving of its

No. 420.] PROOF OF BLOOD-RELA TIONSHIP. IOIQ

EA than if the fibrin be removed. Control experiments

with superheated serum show that the dissolving, which does

at length occur, is not due to the action of bacteria developed

in the serum. On the contrary, the dissolving power of the

serum is inhibited by the increase of bacteria.

Thus far the dissolving action of serum has been demon-

strated only among vertebrates. So far as tried, the blood of

invertebrates (Cancer, Arenicola, sea- urchin) has no globulicidal

effect upon the erythrocytes of vertebrates (gull, rat). Among

cold-blooded vertebrates the action is so slow as to be best

observed under the microscope. Only the serum of Anguilla

acted so rapidly as to make macroscopic observation easy. It

was noted that sera which are particularly poisonous, such as

the serum of Anguilla, the domestic fowl, and cat, act most

rapidly upon corpuscles. This suggests that both effects are

due to chemical substances of the same class.

The serum of Anguilla dissolves the corpuscles of mammals,

birds, reptiles, and amphibians, and also that of other fishes.

The blood of Acanthias vulgaris is quickly dissolved by Anguilla

serum, and so is the blood of other teleosts, as Labrus macu-

latus. The effect of Anguilla serum upon the blood of other

Murzenidz was not tried. The serum of Acanthias dissolves

the erythrocytes of the gull, mouse, and of teleosts (Labrus,

Anguilla. It is not entirely indifferent toward the blood of

other elasmobranchs (Raja batis).

Among Amphibia, Anura are easily distinguished from

Urodela by difference in blood. Frog corpuscles are dissolved

by the serum of Anguilla, the gull, and cat. The dissolving

power of amphibian blood is less in animals that have been

kept in captivity for some time and poorly fed.

Among reptiles, the serum of some snakes was found to

exceed that of amphibians in globulicidal action. Yet more

powerful is the action of bird serum. This fact is parallel

to the fact that the serum of birds is especially poisonous to

other vertebrates. This common quality of the serum of rep-

tiles and birds is correlated with similarities in anatomical

structure which have caused reptiles and birds to be classed

together as Sauropsida. The serum of the domestic fowl-

IO20 THE AMERICAN NATURALIST. [Vor. XXXV.

dissolves the erythrocytes of animals from all other classes.

of vertebrates, and also of other birds.

The results of the experiments with sera of mammals agree

entirely with the results of Landois's experiments with trans-

fusions, but they do not agree with the results obtained by

Ehrlich and Morgenroth, Bordet, and Gürber. This lack of

agreement is doubtless due to difference in methods. The

investigators named above washed out by isotonic salt solution

all the serum from the erythrocytes of one animal and added

large quantities of the erythrocytes to the serum of another

animal. By this method in many cases no globulicidal action

can be observed, while, if a small quantity of simply. defibri-

nated blood be added to the serum, the erythrocytes are quickly

dissolved. The importance of adding only small quantities of

blood to the serum lies in the fact, observed by Buchner, that

in the mingling of two different sera the globulicidal action may

be either increased or diminished. In transfusion experiments

the more blood injected the better.

The carotid arteries of a cat and an ocelot were connected

so that an exchange of blood took place from one animal to the

other. After a short time the blood of each animal was sup-

posably well mingled with that of the other. No hemoglobin

appeared in the bladder of either animal. The blood of the

cat and of the ocelot is physiologically equivalent. If a cat and

a rabbit be connected in the same way, both animals die in a

few minutes from the poisonous effects of the foreign blood

upon the central nervous system. Death occurs before any

globulicidal action takes place. Two rabbits connected in this

way exhibit no pathological symptoms.

The experiments upon mammals lead to the conclusion that

among animals of the same family there are no marked differ-

ences in blood ; but the blood of animals of different suborders

is not physiologically equivalent, while the blood of animals of

different orders exhibits very marked mutual globulicidal action.

The blood of the mouse and blood of the rat are mutually inac-

tive. The blood of the hare and of the rabbit is equivalent ;

but rabbit serum dissolves corpuscles of the guinea-pig, and

rabbit corpuscles are dissolved by serum of the guinea-pig,

No. 420.] PROOF OF BLOOD-RELA TIONSHIP. IO2I

these animals belonging to different families. The blood of

the rabbit is equivalent only to that of the hare, of all the

animals experimented upon.

Of Perissodactyls, serum of either the horse or ass is inac-

tive toward corpuscles of the other, but horse serum dissolves

corpuscles of the rabbit, guinea-pig, calf, lamb, and man.

Of Artiodactyls, the serum of either the ox or swine dis-

solves corpuscles of the other, and both dissolve corpuscles of

the dog, cat, horse, rabbit, and man.

Of Carnivora, the blood of the dog, fox, and wolf is equiva-

lent, but serum of either the dog or cat dissolves corpuscles of

the other. The serum of either the cat, ocelot, or jaguar is

inactive toward corpuscles of the other two animals, but cat

serum dissolved the corpuscles of all the other mammals

tried.

Finally, among primates, human serum dissolves the cor-

puscles of fishes, frog, snakes, pigeon, fowl, night-heron, horse,

swine, ox, rabbit, guinea-pig, dog, cat, hedgehog, and lemur.

The effect of human serum was tried upon the blood of six

species of apes (the platyrhines, Pithesciurus sciureus, Ateles

ater; the catarrhines, Cynocephalus babuin, Macacus Sinicus,

Macacus cynomolgus, and Rhesus nemestrinus, at the Berlin

Zoólogical Garden). In all cases the human serum dissolved

the ape corpuscles. The serum of Macacus had no effect upon

the blood of some persons, while the blood of others was quickly

dissolved by it.

Among the true anthropoid apes is first found blood which

is physiologically equivalent to that of man. To 5 c.cm. of

transparent human serum was added a drop of blood from the

finger-tip of an orang-outang, and in another case from a

gibbon, both young animals from the Berlin Zoólogical Garden.

After twelve hours the red corpuscles were separated from the

serum by the centrifuge, apparently having suffered no change,

while the serum remained free from color. In three experi-

ments, where 10 to 20 c.cm. of fresh, defibrinated human

blood was injected into the veins of Macacus cynomolgus or

a small quantity of hemoglobin appeared

Macacus sinicus, only à

m ás a

in the urine, — scarcely more than is found in the seru

IO22 THE AMERICAN NATURALIST.

result of the defibrinating process. The animals suffered no

ill effects from the operation. Twenty-five cubic centimeters

of human blood was injected, into a ten- year-old chimpanzee.

For two days the urine was tested and showed no signs of

hemoglobin or albumen. The injected blood apparently pro-

duced no effects whatever. It has been shown by successful

blood transfusion experiments that the blood of such widely

separated races as the negro and white is physiologically

equivalent.

These blood comparisons, as well as the embryological

researches of Selenka, justify placing man and the anthropoid :

apes together in the same family, or at least in the same

suborder, rather than isolating man in a suborder of pri-

mates, coórdinate with the suborders of the platyrhines and

catarrhines.

The chemical similarity of the blood of morphologically simi-

lar animals is not surprising. The thing inherited through the

ovum and spermatozoón is not “innere Impulse," * Iden," or

“ Entwickelungsmóglichkeiten," but a certain definite chem-

ical composition, of the molecule. Development, form, and

the nature of the metabolic processes are as closely dependent

upon the chemical composition of the molecule as any chemical -

reaction is dependent upon the chemical composition of the

reagent causing it. Similarity in the chemical composition of

blood is but one factor in the chemical similarity of closely

related organisms. The chemical similarity of reproductive

cells must be regarded as an epitome of all the chemical simi-

larities of the adults.

It is well known that the horse and ass, dog and wolf, rabbit

and hare readily cross. It would be a valuable experiment to

attempt, by means of artificial fertilization, a cross between the

rat and mouse, or between the domestic cat and the ocelot.

The physiological similarity of the blood of either pair of ani-

mals points toward the possibility of a successful crossing.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RESIGNATIONS, AND DEATHS.

EDUCATIONAL GIFTS.

Alameda, California, $35,000, from Andrew Carnegie, for a public library.

Amherst College, $2000, by the will of Professor Herbert B. Adams.

_ Armour Institute, $250,000, from Mrs. Philip D. Armour and J. Armour.

Arnold Arboretum of Harvard University, $126,485, from various sources,

for its endowment fund.

Barnard College, $1500, for fitting up the zoólogical laboratory.

Berea 1 ae College, $50,000, by the will of Stephen Ballard of

peikar iaa College, a conditional gift of $25,000, from Dr. D. K.

Pearson.

Burlington, Vermont, $50,000, conditional from Andrew Carnegie, for a

public library.

Carson-Newman College, lai os, from John D. Rockefeller; $15,000,

from other sources.

Chatham, N.Y., $15,000, from Andrew Carnegie, for a public library.

Colorado College, $100,000, from an anonymous donor.

Columbia University, $100,000, from an anonymous donor, for a depart-

ment of Chinese; $10,000, for the purchase of books; $5000, for a

historical reading room; $5000, for general purposes, from an

anonymous donor.

Cornell University, a conditional gift of $250,000, from John D. Rockefeller H

an astronomical observatory and equipment, from General A. C. Barnes.

Dartmouth College, $100,000, from Amos Tuck, for a building for the —

School of administration and finance.

Des Moines (Iowa) College, a conditional gift of $25,000, from John D.

Rockefeller.

Detroit, Michigan, $750,000, from Andrew Carnegie, for a public library.

Drury College (Missouri), $25,000, from Dr. D. K. Pearson; $25,000,

from other donors, for a science building.

Eureka (Illinois) College, $60,000 in land, from Mrs. D. Bandy. ge

i (Kansas) College, a conditional gift of $25,000, from Dr. D. K.

. Pearson.

Fargo “Cole a conditional gift of $50,000, from Dr. D. K. Pearson. F

Georgia State Normal School, a conditional gift of $13,000, from G. F.

Peabody.

1023

1024 THE AMERICAN NATURALIST. [VoL. XXXV.

Hamline University, $50,000, from James J. Hill; $30,000, from M. G.

Norton

Harvard University, $1,000,000, from J. Pierpont Morgan, for buildings

for the medical school.

Illinois College, a conditional gift of $50,000, from Dr. D. K. Pearson.

Kenyon (Ohio) College, $50,000, from Senator Mark Hanna.

Leadville, Colorado, $100,000, from Andrew Carnegie, for a public library.

Massachusetts Institute of Technology, $100,000, from the alumni, for a

gymnasium; $10,000, from George A. Gardner.

McKendree (Illinois) College, a conditional gift of $50,000, from Dr. D.

. Pearson.

Middlebury (Vermont) College, $12, 500, from Ezra J. Warner; $5000

from Dr. M. en. Starr.

Milliken University (formed by the union of Lincoln College and the

Decatur Industrial College), $150,000, from James Milliken.

New York Botanical Gardens, $3200, from Misses O. E. P. and C. P. Stokes.

Princeton University, $50,000, for a library fund ; $10,000, for a fellowship ;

$150,000, from the alumni, for a gymnasium; $10,000 for a fellowship

in biology, from Mrs. Maule.

Radcliffe College, about $200,000, by the will of Susan Cabot Richardson ;

$25,000, by the will of John Sweetser.

Riverside, California, $20,000, from Andrew Carnegie, for a public library.

Rockefeller Institute for Medical Research, $200,000, from John D.

Rockefeller.

San Francisco, California, $750,000, from Andrew Carnegie, for library

purposes.

Smith College, $28,000, by the will of Mrs. Louise F risbie ; a conditional

gift of $100,000.

Stevens Institute of Technology, $50,000, from President Henry Morton.

Syracuse University, about $40,000, by the will of Justice George N. Kennedy.

Tulane University, $50,000, for library purposes, from Mrs. Caroline

Stannard Tilton.

University of Chicago, $3000, by the will of Marie J. Mergier ; $75,000

additional, from A. C. Bartlett.

University of Southern California, $100,000, from various donors; a con-

ditional gift of $40,000, from Mrs. Anna Hough.

University of Virginia, an annual income of $11,000, from Mrs. Mary

ustin Carroll, of Boston.

Vassar College, $28,000, by the will of Mrs. Louise Frisbie.

Washington and Jefferson (Pennsylvania) College, $100,000, from J. V.

hompson

Wellesley College, $28,000, by the will of Mrs. Louise Frisbie.

Whitman College, a conditional gift of $50,000, from Dr. D. K. Pearson.

Yale University, $10,000, from Irwin Rew, for the Sheffield Scientific

School; $25,000, from John S. Newbury, of Detroit, for an organ.

No.420.] GIFTS, APPOINTMENTS, RETIREMENTS. 1025

APPOINTMENTS.

. Dr. Lujo Adamovit, professor of botany in the University at Belgrade.

— Dr. Ludwig von Ammon of Munich, honorary professor. — M. Eugen

Autran of Geneva, botanist to botanical gardens at Buenos Ayres.—

^. M. Bain, professor of botany in the University of Tennessee, —

Dr. Charles R. Bardeen, associate professor of anatomy in Johns Hopkins

University. — Arthur W. Bean, assistant in histology and embryology in

Cornell University. — Professor H. C. Beeler, state geologist of Wyoming.

— Dr. Louis Beushausen, professor of geology in the mining school at

Berlin. — Professor Franz Beyschlag, second director of the Prussian Geo-

logical Survey. — Dr. G. Bodlander, professor of geology in the technical

school at Braunschweig. — Dr. J. Bóhm, custodian of the collections of the

mining school and geological survey in Berlin. — H. W. Britcher, assistant

in zoólogy in the University of Maine. — W. E. Britton, state entomologist

of Connecticut. — E. R. Buckley, state geologist of Missouri, — Dr. A. H.

R. Buller, lecturer in botany at Birmingham University. — Charles W.

Bunker, assistant in histology and embryology in Cornell University. —

Louis R. Cary, assistant in biology in the University of Maine. — Professor

Fridiano Cavara, professor of botany in the University at Catania. —

Dr. Frederic E. Clements, adjunct professor of botany in the University of

Nebraska. — Professor H. W. Conn, lecturer on agricultural biology at the

Connecticut Agricultural College. — Rheinhart Parker Cowles, fellow in

zoólogy in Johns Hopkins University. — Dr. Max Cremer, professor extraor-

dinary of physiology in the University at Munich. — Dr. R. A. Daly of

Harvard, assistant on the Geological Survey of Canada. — Dr. Percy M.

Dawson, associate in physiology in Johns Hopkins University. — Earl

fellow in biology in Princeton University. — Mr. R. Evans of

Oxford, curator of the Museum at Demarara. — Mr. M. N. Fenneman,

professor of geology in the University of Colorado. — Bruce Fink, professor

of geology and botany in Drake University, Iowa. Tr J. . Flett,

- assistant in petrology on the geological survey of the United Kingdom, —

Dr. Joseph Marshall Flint, professor of anatomy In the University of sane

fornia. — Dr. Shepard I. France, instructor in physiology in Dartmout

College. — Adolf Fritze; professor of zoólogy in the University of Tokyo.

instructor in natural history in Trinity Col-

in the University of

Pennsylvania. — H. H. Grau, : d

.— Dr. Emily Ray Gregory, proie à

paer P ^ eigen SE Haanel, of Syracuse, superintendent of

Strassburg. — Dr. Charles M. Hazen, pr | gy

College, Virginia. — Dr. Tracy F. Hazen, director of ia —

Museum of Natural History at St. Johnsbury, yu ue te z ip ti

docent for ethnography in the University at Vienna. — Dr. Herbert,

1026 THE AMERICAN NATURALIST. [Vor. XXXV.

for zoólogy in the University at Heidelberg. — Bert R. Hoobler, assistant

in histology and embryology in Cornell University. — F. W. Hodge,

assistant in charge of office in the Smithsonian Institution. — Dr. Charles F.

Hottes, instructor in botany in the University of Illinois. — J. Allen Howe,

geologist on the English staff of the Geological Survey of the United

Kingdom.— Dr. Alois Jentió, assistant in vegetable physiology in the

University at Vienna. — Dr. Duncan S. Johnson, associate professor. of

botany in Johns Hopkins University. — S. P. Jones, assistant state geolo-

gist of Georgia. — Dr. Ernst Kittl, docent for geology in the technical

school at Vienna.— Dr. Alfred Koch, professor. extraordinary of bacteri-

ology in the University at Góttingen. — Dr. F. Kolbeck, professor of min-

eralogy in the mining school at Freiburg i. S.— Dr. A. L. Kroeber,

instructor in Indian anthropology in the University of California. —

Dr. August Langhofer, professor extraordinary of botany in the University

at Agram, Austria.— Dr. Ralph S. Lillie, assistant in physiology in

Harvard Medical School. — Dr. Walther von Lingelsheim, director of the

bacteriological station in Benthen, Germany.— George H. Lyman, pro-

fessor of botany in Dartmouth College. — Dr. Florence M. Lyon, associate

in botany in the University of Chicago. — Dr. Albert F. Matthews, assist-

ant professor of physiological chemistry in the University of Chicago. —

Dr. Hermann Meerworth of Hamburg, assistant in the Natural History

Museum in Braunschweig. — Adam Miller, fellow in biology in Princeton

University. — Benjamin LeRoy Miller, fellow in geology in Johns Hopkins

University. — Dr. Siegfried Mollier, professor extraordinary of anatomy in

the University at Munich. — Dr. G. T. Moore of Dartmouth College,

algologist to the Department of Agriculture. — H. B. Muff, geologist on

the Scottish staff of the Geological Survey of the United Kingdom. —

r. Miller, director of the newly established zoological garden in Halle. —

- Bogumil Nemec, head of the new institute of vegetable physiology in

e Bohemian University at Prag. — S. P. Orth, professor of natural

science at Buchtel College, Ohio. — William Osburn, instructor in zoólogy

in the University of Cincinnati. — Dr. J. B. Overton, professor of biology :

in Illinois College. — Vládimir J. Palladin, professor of vegetable anatomy

and physiology in the University at St. Petersburg. — Dr. Henry F. Reid,

professor of geological physics in Johns Hopkins University. — Ralph W.

Richards, fellow in biology in Tufts College. — Professor I. C. Russell,

state geologist of New Jersey, to take the position next year. — Dr. Wilhelm

Salomon, professor extraordinary of stratigraphy and paleontology in the

University at Heidelberg. — Dr. E. Schellwein, professor extraordinary of

geology and paleontology in the University at Kónigsberg. — Dr. Edward

C. Schneider, professor of biology in Tabor College, Tabor, Iowa. —

Dr. G. Senn, docent for botany in the University at Basel. — W. T. Shaw,

assistant entomologist in the Iowa Agricultural Experiment Station. —

Daniel Naylor Shoemaker, fellow in zoólogy in Johns Hopkins University.

— Dr. F. L. Stevens, instructor in biology in the North Carolina College

eo 2

No. 420.] GIFTS, APPOINTMENTS, RETIREMENTS. 1027

of Agriculture. — H. H. Thomas, geologist on the English staff of the

Geological Survey of the United Kingdom. — John J. Thornber, professor

of botany in the University of Arizona. — William C. Thro, assistant in

histology and embryology in Cornell University. — William George Tight,

president of, and professor of geology in, the University of New Mexico

at Albuquerque. — Dr. O. Uhlworm, editor of the Botanisches Centralblatt,

librarian of the Royal Library in Berlin and chief of the German Bureau

for International Bibliography. — C. E. Van Orstrand, physical geologist

of the United States Geological Survey. — Dr. W. Voigt, custodian of the

Natural History Museum in Bonn.— Dr. Franz Wáhner of Vienna, pro-

fessor of mineralogy and geology in the German technical school at Prag.

— Professor Dr. Eugen Warming, director of the Danish Geological Sur-

vey. — Dr. Sho Watase, professor of zoólogy in the University of Tokyo. —

Dr. Thomas L. Watson, professor of geology and botany in Denison Uni-

versity, Granville, Ohio. — Dr. Karl Wenle, professor of anthropology in

the University at Leipzig. — Gersham F. White, assistant in histology and

embryology in Cornell University. — Dr. Edwin M. Wilcox, professor of

biology in the Alabama Polytechnic Institute, Auburn, Ala. — Dr. A. W.

G. Wilson, assistant on the Geological Survey of Canada. — William F.

Wismar, assistant in histology and embryology in Cornell University. —

W. B. Wright, geologist on the Irish staff of the United Kingdom. —

Dr. Rudolf Zuber, professor of geology in the University at Lemberg.

RESIGNATIONS.

Dr. E. B. Copeland, professor of botany in the University of West

Virginia, legislated out of position by the regents. — Dr. C. L. Herrick,

from the presidency of the University of New Mexico, on account of ill

health. — F. W. Hodge, ethnologist in the Bureau of American Ethnology

at Washington. — Professor Carl Wilhelm von Kupffer, from the director-

ship of the Anatomical Department of the University at Munich. —

Dr. E. Suess, from the professorship of geology at Vienna, after nearly |

fifty years’ service. — Dr. H. Topsóe, director of the Danish Geological

Survey.— Dr. Arthur Willey, from the curatorship of the Museum at _

Demarara.

DEATHS.

Dr. G. A. Asp, professor of anatomy in the University at Helsingfors,

April 25, aged 66. — Rev. Dr. Willis H. Barris, geologist, at vais

Iowa, in May, aged 79.— Alexander Becker, a Russian botanist an

entomologist, April 16, aged 82. — Professor M. G. Bleicher of vios

known for his researches on the geology of the Vosges. — Dr. ret-

schneider, author of several works on Chinese botany, in St. Petersburg. —

Dr. Gino Ciaccio, professor of comparative anatomy in the University of

Bologna. — Dr. E

E. W. gd ai sometime professor of pon in Buchtel dE

1028 J THE AMERICAN NATURALIST.

College, Akron, Ohio, and lately professor of geology in Throop Institute,

at Long Beach, Cal, August 17, aged 66.— A. Constant, student of

Lepidoptera in Golfe: Juan, France, May 13, aged 71. — Maxime Cornu,

professor of botany in the Jardin des Plantes, at Paris, April 4, aged 58.

— Adolf Fick, professor of physiology in the University at Würzburg,

August 21, aged 71.— H. W. Harkness, botanist, in San Francisco,

May ro, aged 8o. — W. Hartwig, curcinologist, in April, in Berlin. —

Rev. Moses Harvey, who added much to our knowledge of the giant squid,

at St. John's, Newfoundland, September 3, aged 82.— Professor Felix

Joseph Henri Lacaze-Duthiers, the eminent French zoólogist, aged 8o. —

Professor Gustav Lindstróm, paleontologist of the Royal Museum at

Stockholm, May 16, aged 72. — Dr. Otto Lugger, entomologist of Minne-

sota since 1887, May 21.— Dr. P. Calvin Mensch, professor of biology

and chemistry in Ursinus College at Collegeville, Pa., July 3o. — Dr. Joshua

Miller, archeologist of Arizona. — Dr. Charles Mohr, botanist, at Ashe-

ville, N.C., July 17. — P. G. von Móllendorff, a writer on the natural

history of China, April 19, aged 53. — Dr. Adolf Erik Nordenskjéld, the

well-known arctic explorer and naturalist, August 13, aged 68.— Miss

Eleanor A. Ormerod, the British entomologist. — Dr. Antonio Piccone,

algologist, in Genoa, May 21, aged 57. — Dr. Otto vom Rath, well known

for his studies of myriapods and cytology, in Cologne, April 23, aged 43. —

Miss Eva M. Reed, botanist and indexer in the Missouri Botanical

Gardens, killed by a train July 7. — Dr. A. F. W. Schimper, professor of

botany in the University at Basel, September 9, aged 45. — Dr. Domenico

Stefanini, professor of bacteriology in the University at Pavia, aged 80. —

Professor C. A. Tenne, custodian of the Berlin Mineralogical Museum at

Bad Nanheim, aged 48. — Friedrich Tiemann, conservator of the Zoólogical

Museum in Breslau. — Anthony Wilkin, an English archeologist and eth-

nologist, at Cairo, Egypt, May 17, aged 24.— Martin F. Woodward,

demonstrator in biology in the Royal School of Science at South Kensington,

drowned September 15.

CORRESPONDENCE.

Editor of the American Naturalist :

Sik :—— A slight correction should be made in my “ Notes on

Living Nautilus,” which appeared in the October number of the

American Naturalist. I notice that on p. 828 the block of cuts 6, 7,

and 8 has been set so that the animals seem to be looking somewhat

upward. To appear accurately, the three figures should be rotated

downward, clock-hand fashion, each about thirty degrees.

BASHFORD DEAN.

DEPARTMENT OF ZOOLOGY,

COLUMBIA UNIVERSITY, NEW YORK,

November 12, 1901.

(No. 419 was mailed November 18.)

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