IN. DE X.

PAGE

ALBINISM in hag-fishes 295

Alternation in plants . . 153

Amblystoma, Acceleration isä re-

tardation in metamorphosis of . 385

Amphipoda, Synopsi 26

Animals, Vernacular names of . . 551

Appendages of Tremataspis i223

Aquatic adaptations in senos . 651

Arboreal adaptations in mammals 731

Archonithiformes, Classification of 33

Autodax, Habits MM... .. ... Sia

BAKER, F. C. Rib variation in

Cardium. .

ny, BW. Na or “hitherto

unknown ephemerid nymphs of

as material for the P of vari

ation < 171

Biology, toot equipment hr. Md

Birds, Classification of 33

Brimley, C. S. cad on the. re-

production of certain reptiles . 261

CAMBARUS, Habits of . I

Campbell, D. H. Antithetic v ver-

sus homologous alternation . . 153

Cardium, Rib variation in 481

Case, C. E. The structure and

— of the Am

Pelycosa «99

Pasbibuplln, Biracterl ob. ee

Clark, H. L. e water snakes

of Southern Michigan 1

Eyes of Machilis

PAGE

Cockerell, W. P. A trip to the

Truchas Peaks, New Mexico

Compound eyes of Machili . 319

Conklin, E. J. Amitosis in the

egg-follicle cells of the Son . 667

Corymorpha, Development o - 579

Cricket, peys in egg- a.

cells :

e I. A. poses of ical

ized stages in some plants of the

Botanic Gardens of Harvard

University . 7. 243

Davis, B. The origin of the

Sporoph

Dean, i Albinism. par-

tial albinism and polychromism

i fi d . 295

Dipnoans, Permian - 493

Doran, E. W. Viináculir names

of animals . . 851

"dem Lo wn Classification ab 33

Dublin, L. Adaptations to

BEER: sl fossorial and

cursorial habits in mammals. Il.

Arborial adaptations

alt

EASTMAN, C. R. A peculiar modi-

fication amongst Permian Dip-

bome 2. ee s

— cells of cricket, A mito-

sis in

Electricity, Paring bath oo

Sphwend nemis New. edd «a

Eunice viridis =

iv | INDEX.

PAGE

FisHEs, Cutaneous sense organs of 313

—— Fossil from Mount Lebanon 685

Fresh-water of Mexico. . . 771

——— Generic names o A. x 3400

Sense of hearing in. . . . 185

Flowers, Colors of . -. 365, 44

Fossil fishes from Mount Lebanon 685

Fossorial bets in mammals 819

Fulgur, Studies of . 81

GASTROPODA, Studies of. II. . . 515

Grabau, A. W. Studies of Gastro-

poda. II. Fulgur and Sycotypus 515

dig Albinism in . . 295

Hargitt, C Synopsis of North

American Invertebrates. XVI.

Pt. IV. The Soyda 2331

Harris, J. A. The habits of Cam-

are. 200-1 Cd dE

Hats from the Nootka Sound re-

gon . «65

ay, O. P. So ome seiner on E ne

fossil fishes - Mount Lebanon,

Syria.

Hearing, Sense of, i in Fishes gis dis

Herrick, C. J. the morpho-

logical and physologc classifi-

cation of the cutaneous sens

organs of M "pup UNE TY)

Holmes, J. Synopses of

North American Invertebrates.

XVIII. The Amphipoda - 267

Howard, A. D. On the structure

of the outer segments of the rods

in the retina of vertebrates . . 541

KINGFISHERS, Systematic position

E s. a a or o Oe

LITHOBIUS, Variationin. . . . 299

Localized stages in plants . . .2

Lovell, J. H. The Colors of

Northern gamopetalous flowers

365. 443

MacpouGarL, D. T. Mutation in

Past 5. 5. o. 4 ra

Machilis, Compound eyes of . 319

PAGE

Mammals, Aquatic adaptations in 651

Arboreal adaptations in . . 731

Fossorial Rer d in. .—- ..81

L. par

Mark, e bath

heated by HER e. 115

Matawan formation, New species

of plants from. 27

May, A.J. A Comfibation. to the

morphology and development of .

Corymorpha pendula Ag.. . . 579

Meek, S. E. Distribution of the

fresh- water fishes of Mexic EIT

Fresh-water ika dis-

tribution of . i00 209 771

Michigan, Nase los of

Morse, M. Synopses of Nort h

American Invertebrates. XIX.

The Trichodectide . .

Fossil fishes

Mount Lebanon,

from

NEEDHAM, J. G. An out-door

equipment for oe work in

biology . pau. 4 0M

pe and Lione:

iology 71, 205, 347, 625

i 74, 7o 2106, 353, 434.

504

dis 634, 810

Exploration . . . 3 + 799

Physiology . . 626

ZoGlogy, 72, 121, 209, 549, 497, 557,

630,

Zoógeography . UP XT

Nootka Sound, hats feet

ODONTOBOLCA, Classification of . 33

Osburn, R. C. Adaptations to

aquatic, arboreal, fossoriai and

cursorial habits in mammals, I.

Aquatic adaptations . . . . 651

PACKARD, A. S. Corrections to

his * Life of Lamarck ”

Palolo worm of Sam Ta

Paraffine bath, iude by elettrio-

suas

INDEX.

PAGE

Parken G. H. The sense of hear-

ing in fis 1

Patten, W. go the PEENE

of Tremataspis . aa

—— On the essi c of the Pter-

aspide and Cephalaspide . . 827

m dandi and rela-

ionships o 85

Perforation 5 a vein n by a an artery

in the cat . . 489

Plants, taia Magos in : (243

ew species from the Mai

wan pawns " | 097

—— Mutationin . :.237

Postglacial deposits Nixcitstios

Of.

The causes of

the metamorphosis of Ambly-

stoma n rnum . cca: . 388

Pteraspide, Structure ef. à . 827

Publications received 82, 293, Stk; 645

QUATERLY record 149, 361, 727, 893

See Bigelow,

sonable dé P.

R. P.

Regeneration of body of a starfish 818

Reptiles, Reproduction of . . . 261

Retina of vertebrates, Rods in. . 541

Review:

Bailey’s Birds of the Western

U.S. - 497

Baldwin’s F neat in Phi-

losophy . 205

Desunt d woii-

tion - 347

Dvina s Relationships of

the Flounders . 126

Selenichthyes 127

Calkin’s Protozoa 4

Conklin’s Raber. of a

chio} 21

Costantin's bubaibos it

acquired characters .

Cunningham's Sexual dun

phism . . . - 349

Reviews:

Dean Sage, Townsend, Smith

and Harris's salmon and

Trout .

Driesch's Mind i in nature .

Enderlein's Wings of Telea .

Fernald's Coccide of the

World .

F — Pectoral ijpdhd-

ages o

Gardiner's ‘Maldive Hind

211,

Goette's Zoölo ur.

Griffin's Anatomy of N pee

Gurley’s Habits of fishes .

Hicker’s Autonomy of the

germ nuclei

Hatcher’s Oligticnis pidi

Patagonia narrative .

Hay’s Fossil i ema

Hertwig's Zoó

Hunter's Studies in Yobéct

ife

Ioteraatitual Úis Bot-

and Evermann’s

Fishes of Formosa . .

——— American food and game `

ihe . 3. 4.

Jordan and Fowlers Fishes

of Japan

Keyser's Birds of the Rockies

Kraemer's Pharmacognosis .

Livingston's Osmotic pressure

and diffusion in plants .

Lyon's T of the v

ledon

‘tious of

rkness

and developmen

Masterman’s Developm

Cribrella . "a o

McIlvaine d ies

Ameri ungi

Meek’s Fishes of Mexico .

Mell’s Biological laboratory

method ... . . =

» Gay

PAGE

808

560

434

I2I

. 139

. 128

vi INDEX. ;

PAGE

Reviews:

Morgan's Regeneration. . . 7I

Morse's Observations on liv-

ing.Brachiopoda a> . o 122

Mouillefert's Forestry ova, 336

SFs Neuron theory . . . 631

Parker's Hearing in fishes. . 499

Optic chiasma in teleosts, 500

Pierce's Plant physiology . . 564

bl's Development of the

ace. 73

Recent Midi on EO EEA 352

Roth’s For A pei

op and Van

s water fowl family 498

Sargent's Origin of Mono-

cotylbdona. o co s ovs 38

Scharff’s Atlantis problem . 431

Setchell and Gardner's N. W

Ape.: pi 034

Simroth’s Zoögeo inks £4 20D

Small’s Flora of the S. E

United States . . . 810

Tower's Wings in Coleoptera 809

Arengo S Kuce s. s T4

Vernon's Variation . . . . 625

Verworn's Biogen hypothesis 629

Von Fiirth’s chemical ED

ogy . 62

Von Scraps parasites and

geographical distribution . 350

Weber's Zoógeography of the

Indo- e Acai 431

Webster's Diffusion of insects 561

bbc s Animal plored

wy: te, 030

um s Pei Nautilus . 351

—— Development of Lin-

gula tae e M8

Wises of Lingula . 124

—— Habits of Lingula . . 124

Ritter, W. E. Further notes on

the habits of Autodax lugubris 883

pee Palolo worm of à . 875

medusz, Synopsis “338

F. The compound die

of Machilis . = 319

PAGE

Seffer, P. O. Examination of

organic remains in postglacial

dooa i o a. c 795

Shufeld, R. W. On the classifi-

cation of certain groups of

bh... 5.70 oL us 33

On the osteology and system-

atic position of the king-

„fishers (Halcyones) .

ations to

W. Adapta

ADU de iboral: fossoria

cursorial habits in mammals.

III. Fossorial rea :

Sporophyte, origin

Starfish, A saeco i a dy ws

Sterki, V. Notes on the Union-

ida and their classification . . 103

Synopses of invertebrates. XVIII.

Amphipoda.

“NI

—— XIV. Pt. IV. “The cna:

medusz Eit

—— XIX. The Trichodectide: 609

Sycotypus, Studies of . . . $15

TREMATASPIS, Appendages in 223, 573

be EREA Synopsis of . 609

uchas peaks, A trip to . 887

UNIONIDA, Classification of . . 103

VARIATION in ribs of Cardium 481

— — in Lithobius 299

—— in Littorina 17I

——— in water snakes I

Vein, em of, by an artery

in the

Ve mots names sol kainis soe ae

Vertebrates, Retina of . . . 6t

WATER snakes, Variation in .

Weysse, A. W. The peskissiion

of a vein by an artery in the cat 485

Williams, S. R. Variation in Lith- 289

obius forficatus . 2

Willoughby, C. C. Hats tns the

Prelim-

inary report on the Palolo worm

of Samoa. Eunice viridis (Gray) 875

pur.

VOL. XXXVII, NO. 433

| THE

AMERICAN |

NATURALIST ~

A MONTHLY JOURNAL Seo see e

DEVOTED TO THE NATURAL SCIENCES ce

IN THEIR WIDEST SENSE

CONTENTS

I. The Water Snakes of Southern Michigan

= The American Naturalist.

a 2 ASSOCIATE EDITORS:

- Jj. A. ALLEN, PH.D., American Museum of Natural History, ses York.

<E A. ANDREWS, PH.D., Johns Hopkins University, Baltimore.

WILLIAM M. DAVIS, M.E., Zarvard Sopce Con brige

ALES HRDLICKA, M.D., Mew York Cit

"D. S. JORDAN, LL.D., Stanford Universit

» CHARLES A. KOFOID, PH.D., Ünsterrity WF California, Berkeley.

_ J. G. NEEDHAM, PH.D., Lake eet University.

ARNOLD E. ORTMANN, Pu.D., Princeton University.

PF RICHARDS Y S.B., F.R.M M'S, € d Montreal.

IE York

tical summaries of progress in some

will be briefer articles on various

itorial l Peoaos on scientific questions of the

of recent oo and a final Ment for

THE

AMERICAN NATURALIST

Vor. XXXVII. January, 1903. No. 433.

THE WATER SNAKES OF SOUTHERN MICHIGAN.

HUBERT LYMAN CLARK.

THE water snakes of the genus Natrix offer some puzzling

problems, not only to the systematist, but also to the student

of geographical distribution. There can be no doubt that much

of the difficulty is due to our lack of knowledge; for, unfortu-

nately, the study of living snakes, is not especially popular,

and preserved specimens are so apt to lose their normal colors

that the proper comparison of snakes from different localities

becomes a matter of very great difficulty. The occurrence of

a red-bellied water snake in southern Michigan has been known

for many years, but its relation to the common water snake,

Natrix fasciata sipedon (L.), has never been satisfactorily deter-

mined. It was with the hope of throwing some light upon the

solution of this problem that the work was undertaken, the

results of which are embodied in this paper. During the past

two years one hundred water snakes captured in the vicinity of

Olivet, Mich., have been examined, and careful observations

have been recorded, from the living or freshly killed specimens,

in regard to sex, size, proportions, and color, and the number of

scale rows, gastrosteges (plates on belly, in front of vent), and

; I

2 THE AMERICAN NATURALIST. [Vow. XXXVII.

urosteges (plates on ventral side of tail, behind vent). Special

attention was also given to habits, food, and localities where cap-

tured. All the specimens were taken in the months of April,

May, and June. While the number of snakes thus handled is

not so large as had been desired, the results obtained are of

sufficient interest and importance to warrant publication, in the

hope that observers elsewhere will make similar records and

thus enable us to ascertain the truth in regard to our common

water snakes and their distribution.

NATURAL HISTORY.

Much of the country around Olivet is low and swampy ;

small lakes connected by more or less sluggish streams are of

frequent occurrence, and even in dry summers there is no lack

of water. No wonder, therefore, that water snakes are com-

mon, especially if one has learned when and where to look for

them. Three easily distinguished forms occur, which are

designated by Cope! as Natrix leberis (L.), N. fasciata sipedon

(L.), and N. f. erythrogaster (Shaw). The first of these is the

smallest and the least common. The largest specimen met

with was 747 mm. in length, but all of the others were less

than 700mm. There are only nineteen rows of dorsal scales,

and the olive color, with three narrow, longitudinal black

stripes on the back, and a buff lateral stripe very prominent

anteriorly, is also very characteristic. Cope says that this

species shows “very little variation in any respect," but the

few specimens I have examined show a great deal of diversity

in the distinctness of the stripes and the amount of slate on

the belly. Judging from what I have seen of its habits, this

snake is very fond of the water, as I have never found it out of

that element, except when sunning itself on branches immedi-

ately above some stream or the edge of a pond. The two

snakes which are regarded by Cope as subspecies of JV. fasciata

are much more abundant than X. Ze£eris, and all of my detailed

observations have been made upon them. Both are very aquatic

1 Cope, E. D. The Crocodilians, Lizards, and Snakes of North America.

Washington, Government Printing Office, 1900.

No. 433.] SNAKES OF SOUTHERN MICHIGAN. 3

in their habits, but erythrogaster (known about Olivet as the

* red-bellied black snake") is more often seen away from the

water, and several specimens were taken a rod or more from

the nearest stream, while szfedon is rarely seen more than a

few feet from water. Both are very active, and when once

alarmed are very shy, but erythrogaster is decidedly the more

wary and somewhat more rapid in its movements. Both swim

with speed and grace and can remain under water for some

time. Their food consists chiefly of frogs, toads, and fish; no

other animal remains were found in the stomachs. They cap-

ture fish of considerable size, —a large szpedon having swallowed,

just previous to capture, a sucker (Catostomus) a foot in length.

Fish are usually, perhaps always, swallowed head first, but

frogs and toads are taken either way. Both szpedon and

erythrogaster will strike repeatedly and savagely when cornered,

but the bite is absolutely harmless, and though the teeth may

be strong enough to break the skin and draw blood, the wound

is practically painless and heals quickly, unless the teeth, which

are very small and easily torn from the jaw, remain in the

wound. Like all of our snakes, these water snakes have a very

strong odor, especially during the breeding season. This odor

is due to a thick fluid secreted in glands situated at the base

of the tail and opening to the exterior at the sides of the vent.

These glands are 20 mm. long, more or less, and are present in

both sexes, but in the male they lie above the hemipenes and

are therefore less easily discovered. The secretion of szpedon

is yellowish or brownish in color and has an odor difficult to

describe but very characteristic ; to me it smells slightly “ burnt "

and is very disagreeable. The secretion of erythrogaster is white

or pale yellow and has a slightly acid, musky odor quite differ-

ent from that of sipedon, and, to me, not so disagreeable. Such

statements are obviously insufficient to enable another observer

to make much use of them, but it is practically impossible to

so describe an odor of this kind as to make it recognizable. The

difference between the two kinds of snake is, however, very

marked to any one smelling first one and then the other.

. Both of these water snakes are accustomed to come out of

the water, especially on sunny days, and lie, more or less coiled,

4 THE AMERICAN NATURALIST: (Vor. XXXVII.

upon the bank, on logs in or beside the water, upon the branches

of bushes overhanging the water, or upon piles of brush through

the open spaces of which they can drop quietly down into the

water below. Such piles of brush are their favorite spots, and

one often sees three or more snakes coiled up together on the

same pile. When so resting they seem rather stupid and may

be closely approached and killed with a stick; but I think this

is due, not to stupidity, but to reliance upon their protective

coloration, for when once convinced that they are observed

they will glide into the water without delay. That the colora-

tion of both forms is protective is perfectly obvious to any one

who has observed them in their natural haunts. The resem-

blance to an old stick, when they are lying motionless in the

water or on brush piles, is so great that it is often very difficult

for an unpracticed eye to detect them. They apparently fre-

quent the same spot day after day, for weeks at least, if not

too seriously disturbed. They seem to become more wary with

experience ; a fine large specimen of erythrogaster tempted me

no less than six times to a certain spot, but each time he was

more wary, and I failed in all my attempts to capture him. In

no case were sipedon and erythrogaster found on the same pile of

brush, and specimens of the two forms were never seen together

or even near each other, although males of erythrogaster

were on several occasions found mating with females of the

same form, and male szpedons with females of their own race.

The local distribution of erythrogaster is peculiar. While

sipedon is found about every pond or stream near Olivet,

erythrogaster seems to be confined to a tract of low land, lying

to the east of the village and bordering a creek which connects

Pine Lake with the Olivet mill pond. This area is about a

mile long and less than a quarter of a mile wide, but I know

of no specimens of erythrogaster having been taken elsewhere.

An advertisement was inserted in the local paper, offering a

good price for red-bellied black snakes, and thirteen were

brought to me in response, but every one was taken within the

area designated. The common sifedon also occurs in that

area but is not very frequent. It is difficult to account for

this peculiar localization of erythrogaster, as I cannot see that

No. 433] SNAKES OF SOUTHERN MICHIGAN. 5

the environment is essentially different from that offered by

other creeks near by.

DIFFERENCES DUE TO AGE AND SEx.

Soon after the collection of statistics was begun, it became

apparent that the two sexes could be easily distinguished by

external characters, and later on it was seen that the propor-

tions of the body were different in very young snakes from what

they were in the adults. (Perhaps it ought to be stated that

age is assumed to be correlated with size, and snakes less than

500 mm. in length are regarded as young.) Before we pass on,

therefore, to a comparison of sipedon and erythrogaster, it is

important to make clear the differences which are dependent

on age and sex.

1. Relative Length of Head and of Tail, and Diameter of Eye,

in Old and Young.— If comparison is made between the five

largest females and the five largest males, on the one hand, and

the five smallest females and the five smallest males, on the

other, of sipedon, it becomes clear that in young snakes the

head and tail are longer in proportion to the body, and the eye

is larger in proportion to the head, than in adults. While this

is what might be expected, it is interesting to see how consid-

erable the difference is. The form szfedon is used for compari-

son because the far greater number of specimens examined

makes the contrast more marked. This table shows that if

a snake 937 mm. long kept the same proportions when adult

that were shown when it was 505 mm. long, it would have a

TABLE I.

Length of Tail in Per- mem pi oar: a yiee Diameter of Eye in

f the Total "a "eph Percentage of Length

iine om’ | tal Plates, in Percent-| ipea

nam age of Length of Body. T

10 large sipedons, 22.3% 3.2% 18.3%

averaging 937 mm.

10 small sipedons, 23.1% 38% 20.2%

averaging 505 mm.

6 THE AMERICAN NATURALIST. | [Vor. XXXVII.

tail 6 mm. longer, a head 4.6 mm. longer, and an eye nearly

half a millimeter larger than it does have. While the difference

in length of tail is thus rather small, the difference in head and

eye is very considerable.

2. Relative Length of Tail in Males and Females. — If com-

parison be made between the males and females of either szpe-

don or erythrogaster, the difference in the proportion of tail and

body in the two sexes is very marked.

TABLE II.

Minimum Average Length

Sel S poms Average | Length of Tail | Length of Tail | of Tail in Per-

PT Te cpm Length. |in Percentage of | in Percentage of centage of

E Total Length. | Total Length. | Total Length.

Erythrogaster | ĝ 16 977 2 5.376 22.4% 23-7%

Erythrogaster | 9 8 1060 21.9% 20% 20.9%

Sipedon $ 33 657 26.7% 23.3% 24.8%

Sipedon 9 25 808 22.5% 16.3% 21.1%

The above table includes all of the water snakes examined in

which the tail was uninjured. It will be seen that while the

males average very much smaller in size, the tail is very much

longer than in the females. Indeed, in both sipedon and

erythrogaster the maximum tail measurement for a female falls

short of the minimum for a male, while the average for a

female is approximately 3 per cent less than for the male. It

thus appears that a snake 800 mm. long will be found to be

a male if the tail is over 180 mm., and a female if the tail is

less than 180 mm. Ordinarily a male snake 800 mm. long will

have a tail not less than 24 mm. longer than a female of the

same size.

3. Number of Urosteges in Males and Females. — If a compar-

ison be made between the number of urosteges in males and

females, it becomes evident that here again there is a marked

sexual difference.

These numbers refer to the urosteges of one side only, the

actual number being double the above, since these plates are

arranged in alternating pairs. Occasionally there is one more

No. 433.] SNAKES OF SOUTHERN MICHIGAN. 7

TaBLE III.

Maximum Minimum Average Num-

Species. Sex. oder Number of Number of ber of

LA tte Urosteges. Urosteges. Urosteges.

Erythrogaster ó 16 82 68 77

Erythrogaster 9 8 71 62 67

Sipedon $ 33 79 68 74

Sipedon 9 25 69 58 63

urostege on one side than on the other, but in such cases the

larger number was recorded. A water snake with more than

70 urosteges is (in southern Michigan) almost certainly a male,

while one with less than that number is almost as surely a

female, only three males, out of 49 examined, having less than

70. It is interesting to see that the number is not dependent

at all upon the size of the snake. The five largest females of

sipedon average 62.6 urosteges apiece, and the five smallest

average precisely the same, while the five largest males average

72.4, and the five smallest, 72.6.

4. Correlation between Length of Tail and Number of Uro-

steges. — It is a noteworthy fact, though quite in accord with

what might be expected, that there is a certain amount of cor-

relation between the length of the tail and the number of uro-

steges. Thus, we find that the males of szpedon which have tails

25 per cent of the total length, or longer, average 75 urosteges

apiece, while those in which the tail is less than 24 per cent

average only 71. The females which have tails 22 per cent of

the length or over average 65 urosteges apiece, while those

which have tails less than 21 per cent average considerably less

than 62. Thus, for each one per cent in the length of the tail of

sipedon there are approximately three urosteges, without regard

to sex or age. This correlation is not perfect, however, for

long-tailed snakes sometimes have a small number of urosteges

and short-tailed snakes a large number; thus, one female with

a tail just 21 per cent of the body has 65 urosteges, while

another with the tail 22.5 per cent has only 63. Moreover,

snakes with tails of the same length sometimes differ greatly

in the number of urosteges; thus in the case of two males

8 THE AMERICAN NATURALIST. | [Vor. XXXVII.

having tails 25.5 per cent of the body length, one has 78 uro-

steges and the other only 7o. The accompanying diagram (I) is

designed to show the variability in length of tail and in number of

urosteges, and also the correlation between those two characters.

5. Greater Variability of Females. — One of the most inter-

esting facts brought out during this investigation is that female

water snakes are far more likely to vary from the normal than

are males. This is not a matter of size, for many of the aber-

rations are among the small snakes, and they do not seem to be

more frequent among large specimens. Thus, of 11 females

over 9oo mm. in length, 45 per cent were normal as regards

the labial plates and number of scale rows (the points in which

the variability is most marked), while of 12 snakes less than

800 mm., less than 42 per cent were normal. In both erythro-

gaster and stpedon there are, normally, 8 labial plates on each

side of the upper jaw and ro on each side of the lower; the

normal formula therefore is AL Now, of the r9 male

erythrogasters examined, 16 possessed the normal number and

arrangement of the labials, and of 33 male szpedons, 25 were

normal; of 52 males, therefore, 41, or 79 per cent, were normal

as regards the labials. Of the 8 female erythrogasters, only 4,

and of 30 female szpedons, only 14, were normal ; of 38 females,

therefore, only 18, or 47 per cent, were normal as regards the

labials. The 20 abnormal females show 26 variations from the

normal, and of these 21, or 80 per cent, are added plates, while

the r1 abnormal males show 16 variations, of which only 9, or

56 per cent, are added plates. Granting that these cases are

too few to determine any general law of variability, they are

nevertheless suggestive. Turning now to the number of scale

rows on the back, we find additional evidence of the greater

variability of females. The number of such rows is, normally,

22, counting where they are most numerous, which is usually

about one-third of the total length, back of the head. Of the

52 males examined, 45 had 23 rows, 3 had 24, and 4 had 25;

thus 86.5 per cent were normal. Of the 38 females, 29 had

23 rows, 6 had 24, and 3 had 25; thus, 76 per cent were

normal. It ought perhaps to be emphasized that this increase

No. 433.] SNAKES OF SOUTHERN MICHIGAN. 9

in number of scale rows is not correlated with size, for although

the nine aberrant females average larger than most female

sipedons, three of them average only 650 mm. and the seven

aberrant males average 17 mm. less than the average male

sipedon. From these figures it can be easily shown that about

68 males in 100 will have the normal number of scale rows and

labials, but of 100 females only 36 can be expected to be nor-

mal in both respects. On looking over my list of 52 males

Less than 20 % |20 24.424.895 26.| More than 26%

14.57. 58 59 60| 61|62] 63| 64| 65| 66 | 67 | 68|69 | 70| 71| 72] 73| 7 5|76| 77 78 79 80 -

? g PE i aur o E

Diagram I. — To ill te the diff: in the length of tail and in the engins — in

the t f NS. sipedon, and th lati betw th

eme ength of tail in percentage of total length.

Horizontal lines represent the number of ini Vertical lines show the ae of

tail (upper row of figures) and also the number of urosteges (lower row of figures

piled from the statistics of 33 males ora 25 "adig

and 38 females, I find that this is about the proportion which

prevails; there is, however, one less normal male and two

more normal females than would be expected.

6. Greater Variability of Lower Jaw.— In comparing the

variability of the sexes, the interesting fact was brought to

light that the number of labials in the lower jaw is much more

variable than the number in the upper jaw; that is to say, it is

much more common to find 8—8 upper labials than 10—10

lower. Thus, of the 52 males, 50, or 96 per cent, have the

10 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

upper labials normal, while only 41, or 79 per cent, have nor-

mal lower labials; of the 38 females, 31, or 81.5 per cent, have

normal upper labials, while only 22, or 58 per cent, have the

lower normal. Thus, of 90 snakes, 81, just 90 per cent, have

the normal number of upper labials, while only 63, or 70 per

cent, have the lower labials normal. There is no well-marked

difference between the right and left sides of the head, though

it may be noted in passing that of 10 aberrations in superior

labials, 7 were on the right-hand side, while of 32 aberrations

in inferior labials, only 14 were on the right-hand side. It

may further be noted that of the 10 aberrations in superior

labials, all were due to added plates, while of the 32 aberra-

tions in inferior labials, only 20 were due to added plates.

COMPARISON OF SIPEDON AND ERYTHROGASTER.

Having thus made clear some of the peculiarities which dis-

tinguish males from females and adults from young, in the

water snakes under consideration, we may now pass on to a

careful comparative study of the two so-called “subspecies.”

We do not need to stop and consider points of internal anat-

omy or those external characters which are common to both

forms and serve to indicate their position in the genus Natrix.

We will therefore take up those points wherein erythrogaster

differs from szfedon, and see how marked and how constant

those differences are.

I. Szge.— There can be no question that erythrogaster is a

larger snake than szpedon. As yet I have not seen a really

small specimen of erythrogaster, while the smallest specimen

of sipedon captured must have been born only a few days

previously. The tableat the top of the opposite page will make

the difference in size clear.

The average erythrogaster is therefore considerably more than

25 per cent larger than the average szfedon. The contrast

between the sexes is much more marked in sipedon, though

even in erythrogaster the females are very decidedly larger.

The absence of small specimens of erythrogaster is one of the

most puzzling facts met with, and one for which it is difficult

No. 433.] SNAKES OF SOUTHERN MICHIGAN. II

TABLE IV.

a Number of Maximum Minimum Average Length

— A Specimens. | Length in mm. | Length in mm. in mm.

Erythrogaster ó 16 1218 760 977

Erythrogaster 9 8 1270 854 1060

Sipedon $ 33 850 257 657

Sipedon 9 25 1189 392 808

to account. Possibly the young are born later than those of

sipedon, and careful searching in the late summer or early fall

may yet reveal some of them.

2. Proportions. — When we come to compare the proportions

of sipedon and erythrogaster, we have to bear in mind the fact

that small snakes differ appreciably from large ones in the

relative lengths of head and tail, and diameter of the eye. Thus,

if we averaged all of the available szpedous and placed the

averages beside those of the available exythrogasters, we should

be led to some erroneous conclusions. For example, it would

then seem that sipedon has a distinctly longer head than ery-

throgaster. That this is not so may be made easily apparent

by taking the eight largest females and sixteen largest males

of sipedon and comparing with the eight females and sixteen

whole males of erythrogaster. This is as fair a choice as pos-

sible, since erythrogaster averages so much the larger, and

the proportions of males and females are so different. The

following table shows clearly the result of such comparison.

TABLE V.

Average |Length of Tailin | Length of Head*| Diameter of Eye

Number of i : : :

les. : : Lengthin| Percentage of |in Percentage of | in Percentage of

spun T Geet | Toal Lagh | Boby Longi | Haad Leigh

Erythrogaster | à 16 977 23:7% 3-1% 21.1%

Erythrogaster | Q 8 1060 20.9% 3-1% 19.9%

Sipedon, largest| ĝ 16 726 24-5% 3:3% 19.1%

Sipedon 9 8 989 20.8% 3.176 17-5%

I2 THE AMERICAN NATURALIST. | [Vor. XXXVII.

It will be seen that the females of the two forms agree

remarkably in length of head and tail, while the male szpedons

show only a very slight and unimportant increase over the

males of the other form, and this slight increase is doubtless

due to their very considerably smaller size. The one important

point brought out by this table is that eryzArogaster has a much

larger eye than sipedon, the average difference being over 2

per cent. This is very noticeable in living and freshly killed

16% 17% 18% 19% 20% ng 2% 23%

/ \

10 / dez

Li as N

i 3 J

/ UE

/ ji n

1 ! y

/ V ^

À V

t `

M. M

, P i q

H \ *

gi i

a T M

A FILE N.

LT ll LE) S

T E "ES

E. ee a

0 E [4

Diacram II. — To illustrate the difference in the size of the eye, between JV. f. sifedon

and N. erythrogaster

eryth Horizontal lines represent the numbe

individuals. Vertical lines represent the diameter 7 the eye in ORNK of head MI

Jj g 7 us.

snakes, but it is obvious in preserved specimens. Of the 27

specimens of erythrogaster, none had the eye less than 4.5 mm.

in diameter, while 20 had it 5 mm. or more, and in four of

these it was 6 mm. Of 63 specimens of sipedon, on the other

hand, only six had the eye more than 4 mm. in diameter, and

in only two of these did it measure 5 mm. The accompanying

diagram (II) shows at a glance the relative size of the eye in the

27 erythrogasters and the corresponding 27 sipedons (8 largest

females and 19 largest males) In this diagram, percentages

No. 433] SNAKES OF SOUTHERN MICHIGAN. 13

between 15.6 and 16.5, inclusive, are reckoned as 16 per cent,

those between 16.6 and 17.5 as 17 per cent, and so on.

It is worth noting that the three szpedons with eyes over

I9 per cent of the head length and the three evythrogasters with

eyes over 21 per cent are all large males, while the eryzArogasters

with eyes less than 20 per cent are large females. A glance

at Table V will show that there is other evidence to indicate

that males have slightly larger eyes than females. The differ-

ence, however, is hardly sufficient to be easily recognized.

3. Number of Urosteges. — Reference to Table III will show

that the male erythrogaster averages three, and the female four,

more urosteges than the corresponding sex of szfedon. This

is rather noteworthy in view of the fact that there is no appre-

ciable difference in the length of the tail in the two forms.

The same point may be illustrated by selecting a few examples

of sipedon, giving the total length and the number of urosteges,

and placing above each the corresponding specimen of erythro-

gaster of the same sex, which is, of all on the list, nearest in

size. Thus:

Erythrogaster, 9, 1182 mm. long has 68 urosteges.

« 61 m

Sipedon, 9,1189 * «

Erythrogaster, 9, 1030 * «& u yo «

Sipedon, Q, 1030. * “ í 65 &

Erythrogaster, 8, 806 * “ u 75 T

Sipedon, B * € € g u

Erythrogaster, 8, 760 * «cw yg «

Szpedon, 4, 774 " & ^ € 95 "

Although these cases were selected at random, they are purely

illustrative, and not at all decisive. Examples might be given

showing opposite conditions. Nevertheless, it must be admitted

that erythrogaster seems to have, er a few more urosteges

than sipedon.

4. Number of Gastrosteges. — In no et except color, is

there shown such a marked difference between erythrogaster

and sipedon as in the number of gastrosteges, a character

which is of great importance in distinguishing different species

of snakes. It is important to note here that there is no evident

connection between the number of gastrosteges and sex or

14 THE AMERICAN NATURALIST. [Vor. XXXVII.

size. That it is not a matter of sex is shown by the fact that

the females of erythrogaster average 151.4 gastrosteges apiece

and the males 150.8, while in szpedon the females average 141.7

and the males 142.9. These differences seem too small to

have any significance. That the number of gastrosteges is

not dependent on size is shown by the fact that the five largest

sipedons, averaging 1066 mm. in length, have only 143.8 gas-

trosteges each, while the five smallest, averaging only 440 mm.,

Pen 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157

10 | \

! ]

| \

[

} AN, IN

5 \ Ha "A

i EM, 1 ^

i i ^ 7 Y

/ N D \

1 Y V

h i

^ EN F

0 Pa X.

Diacram III. — To illustrate the di

ff in th ber of g ges | N. f. sibedon

and N. erythrogaster. The anal plate is not included

TM sipedon. ‘ythrogaster. “Horizontal lines represent the number of

individuals. Vertical lines represent the number of gastrosteges. Compiled from the

statistics of 27 erythrogasters and 63 sz/edons.

have 144.2 gastrosteges each. Comparing erythrogaster and

stpedon, without reference to age or sex, we find that the

former has on the average 151 gastrosteges, while the latter

has only 142. But the difference between the two forms is

made most apparent by the accompanying diagram (IIT).

It will be seen at once that 148 is the maximum number for

sipedon, and, at the same time, is the minimum number for ery-

throgaster. It ought to be stated that occasionally an imperfect

No. 433-| SNAKES OF SOUTHERN MICHIGAN. 15

or half plate occurs at the side between two gastrosteges.

This was the case in one female erythrogaster and in two males

and two females of szpedon. In all such cases the extra plate

has been counted as an additional gastrostege.

5. Color. — In dealing with the matter of color, we meet

with great difficulties, owing to the impossibility of stating

differences with mathematical exactness. Moreover, in no

other particular is there so much room for difference of per-

sonal opinion and so many chances for errors of judgment.

Nevertheless, since it is in the matter of color that sipedon and

erythrogaster exhibit their most constant and striking differ-

ence, it is absolutely essential to any proper understanding of

the relation of the two forms that this difference be clearly

shown. First of all, therefore, an exact description of the

typical coloration of each form in life will be given, using

the color names of Ridgway's Nomenclature of Colors.

Erythrogaster. Dorsal surface black, passing through slate

black and blackish slate to nearly slate color on sides; ventral

surface bright rufous, orange rufous, or even Chinese orange,

shading anteriorly through saturn red to deep chrome on the

throat and finally to creamy white on the chin; whole head

with a reddish tinge; upper labials nearly rufous except on

upper and anterior edges ; outer, anterior edges of gastrosteges

more or less slate color, the same shade being more or less

evident on urosteges.

Sipedon. Dorsal surface dark bister with irregular, narrow,

transverse bands of wood brown; beginning on the fifth or

sixth row of scales, and running down vertically on sides, are

broad, pale, almost whitish bands, anteriorly and posteriorly

continuous with, but for the most part alternating with, the

transverse bands on back ; between these vertical lateral bands

the scales are chocolate brown, more or less mottled with black ;

chin creamy white; gastrosteges creamy white, anteriorly with

two semicircular spots of hazel or ferruginous, the arc of the

semicircle coincident with the anterior edge of gastrostege ;

farther back additional ferruginous spots appear, and these

gradually merge together, at the same time becoming more

and more clouded with black, so that near the vent the

16 THE AMERICAN NATURALIST. [VoL. XXXVII.

gastrosteges are black with a little white on the posterior edges ;

urosteges mostly black, with inner edges white; head mottled,

light and dark brown ; lower edge of upper labials pale gray.

The colors of szpedon do not undergo a very marked change

during a few months in alcohol or formalin, simply becoming

more dull, though after the lapse of years they fade, especially

if exposed to the light. The colors of erythrogaster, however,

are completely changed in either formalin or alcohol, the black

tending to become bister or clove brown and the whole under

surface becoming pale cream color, with faint indications of

slate on the anterior edges of the gastrosteges. The twenty-

seven specimens of erythrogaster collected about Olivet showed

practically no variation in color, except that a few had the mid-

ventral line a somewhat deeper shade of rufous. In no case

was there the slightest evidence of markings on the back, or

of spots on the belly. The sixty-three specimens of sipedon,

on the other hand, show a very wide range of variation, not

only in the distinctness of the markings and in the amount of

brown and black on the ventral surface, but also in the shade

of the ground color, both dorsally and ventrally. The middle

of the ventral surface is often marked with an ill-defined longi-

tudinal area of yellowish, sometimes almost reddish yellow.

Just before the shedding of the skin, the black of erythrogaster

becomes very dull, and the ventral side a dull, almost salmon,

red, quite different from the normal shade. In captivity this

stage may last two or three weeks, but in freedom it is probably

passed through more rapidly. In sipedon the casting of the

skin causes a preliminary obscuring of the dorsal markings, so

that in cases where they are naturally faint they may be

apparently wanting.

However much specimens of sipedon varied from normal,

none of those examined showed the slightest approach to

erythrogaster, and it is very difficult to see how the coloration

of the latter could ever have gradually developed from that of

the former. While still seeking a solution of this puzzle, four

specimens of Natrix from the United States National Museum

were very kindly loaned to me by Dr. Stejneger. Of these I

shall have more to say later. Suffice it to say here that they

No. 433.] SNAKES OF SOUTHERN MICHIGAN. 17

helped me to imagine the steps by which erythrogaster might

have developed from szfedon, although it by no means follows

that such were the steps. The following ten are the stages

that I have selected, but they are of course arbitrary, and I could

easily have subdivided the sixty-three szpedons into a dozen color

varieties, would such a division have been of any service.

TABLE VI.

pont heus ew Han Ventral Markin

Markings. | Ground Color. gs.

No. Color.

I | Brown |Indistinct| Creamy or | Some brown and much black

yellowish

2 | Brown | Distinct | Creamy or | Some brown and much black

yellowish

3 | Brown | Distinct | Yellowish | Brown and black

4 | Brown | Distinct | Yellowish, | Brown and black

5 |Blackish| Distinct | Yellowish | Little brown; much blackish slate

ous

6 | Black |Indistinct | Yellowish | No brown; much slate

rufous

7 | Black |Indistinct Rufous | Muchslat terior half of gastrosteg:

8 | Blac None f Much slat terior half of g g

9 | Black None Bright Little slate on anterior edges of gastro-

rufous teges

10 | Black | None Bright Deep rufous on mid-ventral surface; very

rufous little slate

Although these are such hypothetical stages, more than

half of them occur among the ninety snakes I have examined.

The following diagram will show at a glance their relative

abundance, but it will of course be borne in mind that the first

four stages, which include all of the szpedons, might have been

divided up into a much larger number of color varieties had it

been desirable. This would not, however, have affected in any

way whatever the great gap between stages 4 and8. The only

purpose of this diagram is to show plainly that gap.

Although, of course, this diagram is not really comparable

with that showing the number of gastrosteges, since we are

dealing here with a purely artificial arrangement and not with

18 THE AMERICAN NATURALIST. | [Vor. XXXVI,

exact numerical series, yet it is interesting to see how the sepa-

ration of erythrogaster from sipedon, so evidently shown by the

diagram of gastrosteges, is emphasized by this diagram of color.

6. Variability.— It has already been shown that female water

snakes are more variable than males in the number of scale

rows and labials. It is interesting to see that in both these

0 1 2 3 4 5 6 7 8 ) 10

L N

7 i

RnS IV.—To illustrate the difference in color between X. f. —— and i ers.

gas he numbers at the left indicate the number of individua

top pe the type of coloration as given in Table VI. eoo from the records of

27 erythrogasters and 63 szfedons.

respects, as well as in color, sz?edoz is far more variable than

erythrogaster. This is clearly shown by the table on the

opposite page.

It may further be stated that there are two specimens of ery-

throgaster, one male and one female, which are apparently normal

in every respect, as they have the average number of gastro-

steges (151) and of urosteges (77 and 67 respectively), the

proper number of labials on each jaw oni and the proper

number of scale rows (23). Such a normal individual of sipe-

don is not to be found among my 63 specimens. In Table II

may be found another illustration of this same point, for it

No. 433.] SNAKES OF SOUTHERN MICHIGAN. I9

there appears that the range of variability in the length of tail

is over IO per cent in sz?edoz and only a little over 5 per cent

in erythrogaster. We have already seen that sipedon is much

more common and much less restricted in its range than ery-

throgaster, and since it is clearly much more variable, these facts

serve as an excellent illustration of the generally accepted belief

that common and wide-ranging species are the most variable.

TABLE VII.

E TE F £f d

A E z m. f» “5 Z

D 9 in M4 8 |u Tee 3 Ty EU $134

Sel 4 |a |g i Z $9 jg 2 |S [5g 4 js

Species. Sages! &$ |£ fee à |& (Fal b |& 4d] $18

92/5 = | Su isu) S | $e |vs| 3 | Ss les) = | Se

sB5|[$4| 8 | 88 lon) § | 88 len! 8 188125! 8 | S32

43852 E [$2 53 5 |ER BS E | 8 |5E| S | Es

diez & de lad) & do [Zm| d | alas & | ad

Erythrogaster | à | 19 | 18 | 95% m 16 | 84% "t 18 | 95% 15 | 79% "

Erythrogaster |Q| 8| 7 | 87.5% sins 4 | 50% 3 100% 415

Sifedon $ | 33 | 32 | 97% 25 | 76% 27 | 81% el^ 57:5% A

Sipedon Q| 30 | 24 | 80% 9% 18 | 60% 7b 21 | 70% 12 | 4

THE SYSTEMATIC POSITION OF ERYTHROGASTER.

With these facts before us, we may well consider what light

they throw on the real relationship of the red-bellied black

snake to the common water snake. . The table on the follow-

ing page will help to set before us, so that they may be readily

grasped, the points of resemblance and difference between

the two.

In the relative length of head and tail, in the number of scale

rows, and in the number and arrangement of labials, there is

evident agreement between the two forms; but, on the other

hand, erythrogaster is a larger snake than sipedon, the male

especially averaging 50 per cent more (see Table II), the eye

is very much larger, the gastrosteges are more numerous, the

urosteges slightly more numerous, the color is totally different,

the odor is distinguishably different, and the percentage of

variability is very much less. In addition to these characters,

certain peculiarities of habits and habitat help to distinguish

20 THE AMERICAN NATURALIST. | [Vor. XXXVII.

TaBLE VIII.

-= s |35 2

else sb lsg a |S [S P

= [5] ad g p v E l5 IH Color and | of Variability

S |$3 gs |o m zs 8 |2 i

to | eae s/o ngng] Se |8.lB Odor of Se- in

i S |SPESPEG SE 2 | 98 |3$\5 Color cretion of

Species. | 9 ASP BAS 2 Is a9 ZZ sg . E. TRIN EN

p o0 9 9g9|o9|5 ta og 9 Bb Postanal E

Pp Rise ED

; hic LS Ed «ien! g£2| ex i ASS g

E E AEAEE A Er R

BTA AA EA xudmm| AB «5x5 5 laal A

Erythro- \1019| 22.3%) 3.1% | 20.500, 23 PE aa 151 | 72 |Blackabove, White to |26%]| 4% |30%

gaster rer rufous be- |light yellow;

neath ; no musky,

markings (slightly acid

; 8-8

Sipedon | 733\*22.7%\*3.2% |*18.370| 23 re 142 | 68.5 Brown Yellow or |38%|24%|51%

e, brownish;

cream color| somew

beneath; “ burnt”

cross bands| and disa-

above, greeable

: blotches

beneat|

* Average of 8 largest females + 16 largest males, for reasons already given.

erythrogaster from its more common relative. Were all our

knowledge of these two snakes confined to what has been

learned about them from the study of these Olivet specimens,

the proper course would be simple and no one would hesitate

to write Natrix erythrogaster as a good species. But unfortu-

nately for the followers of such an easy course, Natrix fasciata

is a very widely distributed and variable species, and water

snakes referred to the subspecies erythrogaster have been

taken in many parts of the United States south of Michigan,

and even in Mexico. In fact, erythrogaster is regarded as

characteristic of the Austroriparian district, and its occurrence

in Michigan is looked upon as an extreme northward extension

of its range.

Through the kindness of Dr. Stejneger, to which reference

has already been made, there were sent me from the National

Museum three specimens of Natrix, which were referred to

erythrogaster by Cope, and a fourth specimen, from the Dismal

Swamp, Virginia. All are females.

these specimens carefully:

1. The specimen from the Dismal Swamp (National Museum,

No. 26,256) resembles the Olivet ezythrogasters very closely,

Let us now examine

No. 433:] SNAKES OF SOUTHERN MICHIGAN. 2I

but is smaller than any of my specimens (712 mm.) and has a

much longer head and tail proportionately. These differences

may be due, however, to the measurements having been made

from the preserved specimen, in which the body would natu-

rally have shrunk more than the head or tail. So far as can be

judged from preserved material, the color was originally the same

as in Olivet specimens. This snake clearly throws no light on the

question of relationship to szpedon, but it leaves little doubt in

my mind that the Virginia and Michigan snakes are identical.

2. The second specimen (National Museum, No. 1350) is a

small snake less than 600 mm. long, collected many years ago

by Professor Agassiz at * Lake Huron." It has only 146 gas-

trosteges, the diameter of the eye is less than 19 per cent of

the head length, and the markings on the upper surface are

those of sipedon. The tail is broken so that the number of

urosteges could not be determined exactly, and the whole speci-

men is so badly faded that it is not possible to say what the

colors or markings of the ventral surface were in life, but there

are no distinct dark markings on the gastrosteges. In spite

of this, however, the snake seems to me clearly a szfedon and

it probably never even approached erythrogaster.

3. The third specimen (National Museum, No. 1351) is from

St. Louis, Mo., and is also an old and faded specimen, but

the presence of light transverse bands, bordered with black,

across the back is very evident. Underneath the specimen is

practically unmarked, and it may have been rufous, like evythro-

gaster. The diameter of the eye is 23 per cent of the head

length, and there are 152 gastrosteges, but there are 24 rows

of scales and only 61 urosteges. The specimen is probably an

erythrogaster, with evident indications of relationship to some

stpedon-like form.

4. The fourth specimen (National Museum, No. 1341) is

from Lansing, Mich., less than thirty miles from Olivet, and

is also old and badly faded. It has the diameter of the eye

20 per cent of the head length, r51 gastrosteges, 64 urosteges,

and 25 rows of scales. It is like erythrogaster in color, except

that at intervals of 20 mm. along the middle of the back are

indications of dark transverse markings.

22 THE AMERICAN NATURALIST. (Vor. XXXVII.

In the Olivet College museum there is a specimen of erythro-

gaster, a female, taken at Olivet in the spring of 1889, which is

the largest water snake I have yet seen. The tail is broken,

but careful calculation for the lost portion shows that the speci-

men was certainly over 1300 mm. inlength. The eye is 6.5 mm.

in diameter, 22 per cent of the head length; there are 154

gastrosteges and 25 rows of scales. There are no dorsal mark-

ings of any kind, but the ventral surface is mottled with a

great deal of slate color along the sides, especially near the

vent, on the posterior gastrosteges and the urosteges. The

coloration is therefore No. 8 of Table VI.

We have here, then, three (or possibly four) snakes which

seem to be what might be considered connecting links between

erythrogaster and sipedon, or some other form of fasciata. Are

they really such? Two facts must be noted: first, all are old

specimens, the most recent having been taken thirteen years

ago, and that one is most nearly a typical erythrogaster; second,

all are females, the variable sex, and are aberrant in number of

scale rows, urosteges, or gastrosteges. They are not, therefore,

actually intermediate forms, but individuals which have varied

from the normal in color as wellas in some other particular.

The smallest of my specimens of erythrogaster, a male 760 mm.

long, was kept in captivity for six weeks, at the end of which

time he shed his skin. Although when captured his coloration

was perfectly normal, without a trace of markings, his new

suit showed along the sides faint indications of lighter, vertical

bands, visible only in just the right light. Might this not indi-

cate the ancestry, as the spots on the breast of a young robin

indicate its ancestry, without making the individual in any sense

a connecting link ?

All of the evidence so far collected seems to me to show

that we have in Natrix erythrogaster a well-defined species of

water snake, probably derived from some form of fasciata,

though probably not sipedon. Possibly the separation has been

completed during the nineteenth century and the specimens in

the National Museum, referred to above, are some of the last

connecting links, though I am inclined to regard them merely

as unusually aberrant females. At any rate, what we need

No. 433.] SWAKES OF SOUTHERN MICHIGAN. 23

now is fresh evidence and much of it. Are connecting links

between erythrogaster and any forms of fasciata now to be

found anywhere? Do erythrogaster and forms of fasciata

breed together? Do the females of erythrogaster ever produce

any young that are not clearly young evythrogasters? Do the

females of any form of fasciata ever produce erythrogasters ?

Until some or all of these questions are answered in the affirma-

tive, erythrogaster is entitled to rank as a distinct species of

Natrix. But there is still much to learn as to its range and its

breeding habits.

OLIVET COLLEGE, MICHIGAN,

August, 1902.

NEW OR HITHERTO UNKNOWN EPHEMERID

NYMPHS OF THE EASTERN UNITED

OIAIIRS

EDWARD W. BERRY.

THE nymphs of these, our most primitive Neuroptera, are

especially interesting because of their varied and great speciali-

zation, each species having solved the problem of existence in

a somewhat different manner. The described nymphs are few.

That of Betisca obesa Say has been known and figured for

some years. Needham,! in 1901, describes the nymphs of

seven species, representing as many genera, from the Adiron-

dack region ; and a number of undetermined American nymphs

are described and figured in Eaton's monograph. It is singular

that these most interesting aquatic larvae are so little known.

They are very easy to rear and form a delightful addition to

any aquarium. Almost any pond or stream, of whatever size,

will furnish its quota of forms, and there is a constant succes-

sion of species throughout the year. Thus I found Blasturus

cupidus extremely common during the latter part of March and

the first part of April, while diligent search on May 17 failed

to disclose any specimens. Habrophlebia americana was

abundant during the last week in May, while on June 14 I

could not find any trace of it.

In these brief notes special attention is directed to the

structure of the gills, because they afford a convenient and

admirable criterion of the relations of the various species,

both to one another and to their environment, and will prove

exceedingly useful as a basis for phylogenetical hypotheses

when more extended observations have been accumulated.

I am greatly indebted to Nathan Banks of the Department

of Agriculture for authoritative determinations of the imagos.

1 N. Y. State Museum, Bull. No. 47.

26 THE AMERICAN NATURALIST. |. (Vor. XXXVII.

All the specimens are from Boynton's Pond, a shallow sheet of

water about one hundred feet in diameter on the outskirts of

Passaic, N.J.

Habrophlebia americana Banks mss.

This fine little species was common among the floating

masses of Spirogyra which skirt the edges of the pond; rather

sedentary in habit, but very active swimmers when disturbed.

They swim by rapid vertical movements of the abdomen, mean-

while holding it considerably elevated. The chief organs of

locomotion are the flattened abdomen with the expanded lateral

margins of its segments. The caudal sete are weak and but

thinly clothed with hairs, and can be of but little assistance in

SNP 2

e aee ——

Fic. 1. — Habrophlebia americana Banks mss.

swimming. The abdomen is not held in a raised position when

the nymphs are resting, as it is in Callibztis, for instance.

Antennz slender, about 2 mm. long, sparsely and minutely

hairy at the joints toward the base. Head wider than long,

with the eyes on the posterior lateral angles. Color dark

brown; margins of the abdominal segments and their lateral

extensions, together with the terminal third of the caudal setze,

yellowish. Abdomen flattened; segments 3 to 6 about the

same width and becoming slightly longer; segments 7, 8, and

9 rapidly narrowing, the ninth being about half the width of

the third; posterior margins of segments 6 to 10 minutely

toothed; lateral margins of all the segments produced more or

less beneath the gills, thus protecting them when swimming.

This lateral expansion increases posteriorly, and the posterior

lateral angles of segments 8 and 9 are produced into a sharp

spur.

No. 433.] UNKNOWN EPHEMERID NYMPHS. 27

Gills single, lanceolate, bilobed, similar in outline to the first

stunted pair in Blasturus cupidus; directed laterally, borne on

segments I to 7, and all alike except for the third, fourth, and

fifth pairs being slightly larger; margins of all somewhat

clothed with scattered hairs.

Caudal setz all hairy on both sides, — sparsely so, however,

and then only at the joints; angle of separation considerable;

terminal third naked and lighter colored than the basal two-

thirds. Middle seta longest, length 5-6 mm.

Legs rather small and weak, the third pair the longest; coxa

with a row of eight tiny spines ; femur abundantly spined; tibia

and tarsus spined at angles and hairy, the hairs longest on the

tibia. This species seems to walk backward or forward with

equal facility.

Total length 6—7 mm. (figure enlarged x 6); greatest width

I.I to 1.5 mm.

These were taken the afternoon of May 24 and were fully

grown, but owing to the cool weather none emerged until

May 29, when one emerged ; the next day (May 30) two others

emerged, and the first specimen molted his subimago accoutre-

ments and became a full-fledged ephemerid.

This is the only known species from the United States, and

the only recorded locality as far as I know, although Eaton in

his revision states that he has seen a specimen belonging to

this genus from New Hampshire.

Blasturus cupidus Say.

This is one of the commonest spring May flies of the eastern

states. Imagos have been recorded from the following New

Jersey localities: Fort Lee, Staten Island, Caldwell, Westville,

and Jamesburg. The full-grown nymphs were common .

beneath the floating debris around the pond margins during

the last week in March and the first week in April. On

May 17 none were to be found.

Nymphs stout, widest across mesothorax (about 3 mm.),

total length 12 to 13 mm., color dark; antennz about half as

long as body (or 6 mm.) ; legs comparatively strong, minutely

spined throughout. Comparatively slow swimmers, but active

28 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

walkers and climbers, walking with facility either forward or

backward. Lateral margins of posterior abdominal segments

produced into a point.

Outer caudal setze fringed on both sides, habitually held at

an angle of 60° to 70° (Fig. 2, e); length about 12 mm. ; joints

Fic. 2. — Blast idus Say

over seventy, short at the base, becoming elongated toward

the tip; margins of joints finely spined, hairy only at the

nodes (Fig. 2, d).

Gills exposed, leaf-like, held laterally, on segments 1 to 7,

their movement comparatively slow (about 130 per minute); on

the first segment they are simple lanceolate rudiments, forked

for over two-thirds their length into two slender sparsely hairy

branches (Fig. 2, a) ; they are double on segments 2 to 7, ovate

No. 433.] UNKNOWN EPHEMERID NYMPHS. 29

in outline, their mid-veins extended into slender filaments,

which are nearly as long as the gill leaf proper (Fig. 2, 4); the

gills gradually become smaller and more slender posteriorly,

until on segment 7 they are lanceolate (Fig. 2, c). Veins red-

dish, thick, passing abruptly into very fine branches.

Callibetis ferruginea Walsh.

Imagos of this species have been taken from Canada to the

southern states; none are recorded from New Jersey localities,

however. The nymphs are very common beneath the floating

Fic. 3. — Callibztis ferruginea Walsh (spring form).

debris and Spirogyra masses, and appears to be the most com-

mon species in the vicinity of Passaic except for an undeter-

mined species of Heptagenia from the brooks of this vicinity.

I have taken them at intervals from the first week in April

through July; they were more abundant, however, during the

springtime.

General color light brown, sometimes greenish ; eyes lateral;

body widest across the mesothorax; legs about equidistant at

base, weak, light colored, darker at the tips of the joints.

Gills exposed, on segments I to 7, simple, somewhat two-

lobed, broadly oval in outline, held vertically when at rest;

vibration intermittent but rapid (about 200 per minute). They

30 THE AMERICAN NATURALIST. . [Vor. XXXVII.

decrease regularly in size posteriorly. Fig. 3, æ represents one

of the first pair and Fig. 3, 4 one of the last pair.

Total length about 9 mm. ; antennz 5—6 mm. ; caudal setze

7mm.

Caudal setae abundantly fringed, the lateral ones on the

inner side only; fringe three or four times as long as setz is

wide, regularly abundant, not confined to nodes except towards

tip, where it is short and sparse and almost imperceptible;

fringe widest on the middle third, which is strikingly dark

colored (Fig. 3, d); lateral setze slightly longer than terminal,

Fic. 4. — Callibztis ferruginea Walsh (summer form).

bringing their tips in a line; angle between outer setz habitu-

ally about 30°; active darting swimmers, as might be imagined

from the oarlike caudal setze, and correspondingly poor walkers

and climbers. When at rest the abdomen is held much elevated,

as in Fig. 3, c.

The specimens taken on June 14 (Fig. 4) differ somewhat

from the preceding, and I imagine the former might have been

females. One of the latter which I bred was a male, but I

cannot be sure that it is the individual from which I wrote the

description.

Total length 10 mm. ; antennae 5.25 mm.; setze 7 mm.;

greatest width 2.25 mm. The habitual angle at which the

outer caudal setze was held was slightly greater than in the

No. 433.] UNKNOWN EPHEMERID NYMPHS. 31

former case; the margins of the joints more conspicuously

spined, particularly those of the abdomen, which increase in

size laterally, the last one forming a decided spur at the

posterior-lateral angle of the segment; gill veins somewhat

stouter, and gills with a much more decided basal lobe. Col-

lected on the afternoon of June 14. During the night ‘four

subimagos emerged, and the next afternoon the only living

one molted to the imago stage.

PASSAIC, NEW JERSEY.

ON THE CLASSIFICATION OF CERTAIN GROUPS

OF BIRDS.

(SUPERSUBORDERS : ARCHORNITHIFORMES ; DROMJEOGNATHE ;

ODONTOHOLC#.)

R. W. SHUFELDT.

INTRODUCTION.

For a number of years I have paid considerable attention to

the anatomy of birds, and especially to the osteology of the

class. This has been done chiefly with the view of enabling

me sometime to draw up a provisional scheme of classification

of this group of the Vertebrata. This is now well on its way

toward completion, but before finishing it I prefer to await

the appearance of other taxonomical schemes by other authors,

either now in press or coming out in parts, as well as the pub-

lication of certain memoirs of my own on the same subject.

My work on the osteology of birds came to assume such formi-

dable proportions that I failed to find a publisher either in this

country or Europe who would undertake the publication of it.

Therefore I was compelled to issue it in the form of separate

memoirs, or monographs, many of which have already appeared,

while eight others have been accepted and will be issued in

due course.

In the present contribution I offer the researches I have

made in the osteology of the Archaopteride, the Ornithure,

or ostrich forms, and the Odontoholcz. In doing this I have

gone most carefully over all the literature on the subject that

was available to me, and have examined a number of the

skeletons of the birds contained in these groups, by far the

larger share of which belonged to the department of compara-

tive anatomy of the United States National Museum, to which

institution I am especially indebted for the facilities it extended

34 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

to me. My thanks are particularly due to Mr. Lucas for his

many courtesies in bringing the material before me, and to

the free use of the collections under his charge. I am also

indebted to my friend the late Prof. E. D. Cope, the late

Prof. O. C. Marsh, and others for suggestions.

With these brief prefatory remarks given by way of explana-

tion we may next proceed to the consideration of Order I of

the class Aves, — the Saururz, and the other groups enumer-

ated above.

I. ORDER SAURURJE.

SUPERSUBORDER, SUBORDER. SUPERFAMILY. FAMILY.

Archornithiformes. Archornithes. Archæopteridæ.

So widely known is the fossil material representing the cele-

brated species of Archæopteryx that any very extended descrip-

tion of it would be quite unnecessary in this place. A great

deal has been written upon the two species of this extinct genus

since 1861, when Hermann von Meyer described the first speci-

men, which was probably nothing more than the impression of

a primary feather discovered in the lithographic slate of Solen-

hofen, in Bavaria, a deposit belonging to the Upper Jurassic.

Two years afterward Owen described the first skeletal

remains found in the same locality, it being largely the pos-

terior part of the bird now known to science as Archæeopteryx

lithographica. A far more perfect example was found in 1877,

from which the skull and the greater part of the skeleton could

be made out. The first of these specimens is now in the British

Museum, and the last one in the Museum of Berlin. It has

never been the fortune of the present writer to have personally

examined any of this material. In addition to the literature of

the subject, however, I have before me a fine photograph of

the British Museum specimen, which was secured by the Cen-

tury Company of New York City to illustrate an article of

mine in the Century Magazine (January, 1886). The majority

of those illustrations were reproductions of my own drawings

and among them a restoration I had made of Archzopteryx,

but had I this restoration to make again, it would present

a very different appearance, especially in the covering of the

No. 433.] CERTAIN GROUPS OF BIRDS. 35

body and the characters of the long tail. It favors its reptiloid

organization too much, for it is probable that the typical species

of the genus Archzopteryx were about seventy-five per cent

„bird and but twenty-five per cent reptile. If the fossil remains

of the earlier ancestral stock of this group of forms are ever

discovered we will meet with types presenting just as much

of the reptile in their organization as bird, but they will not

have developed the feathers that Archaeopteryx possessed, nor

will the hind limbs be as ornithic in structure. Some of these

long-tailed reptiloid birds were about the size of a fish crow,

while others were much larger. As is well known, they had a

long, lizard-like tail composed of twenty-one vertebra, and into

the skin that covered these were inserted twenty-one pairs of

conspicuously developed tail feathers, a pair to each vertebra.

Morphologically, these long and slender joints were distinctly

reptile in character, and doubtless had quite as much motion,

individually and collectively, as do the vertebrae in any of our

larger whip-tailed lizards of the present time. The compara-

tively small, pyramidal skull of these ancient forms was much

flattened above, with its occipital aspect truncated obliquely.

Either orbital cavity was large, and true teeth in grooves, or

sockets, armed either mandible.

Reptilian characters largely nd in the remainder

of the vertebral column of Archzopteryx, for the articular sur-

faces of such of the vertebra as have thus far been examined

and studied are flat, and the sacral ones were few in number.

According to Marsh the sternum was represented by a single

broad plate of bone, and it is likely that it developed a keel.

The shoulder girdle was very birdlike, especially the os furcula.

Pycraft, who has examined all the fossil specimens of these

Jurassic birds, says: * The dorsal ribs have been described as

wanting uncinate processes; an unsafe conclusion, since these

are often absent in the skeletons of existing birds, having been

lost in maceration. The cervical ribs appear to have been

much more slender than in modern birds, and to have remained

movably articulated throughout life. * Abdominal ribs,' resem-

bling rather those of the Crocodilia than of the Chameleonida,

appear to have been present."

36 THE AMERICAN NATURALIST. [Vor. XXXVII.

The three bones composing one-half of the pelvis were appar-

ently distinct, quite as much so as among the young of modern

birds. Of these pelvic bones the ilium is best seen, and is said

to be characteristically avian. Apart from a few transitional

characters denoting the origin of the form from reptilian stock,

the skeleton of the pectoral and pelvic limbs are almost entirely

ornithic. Whether the avian-like humerus was pneumatic has

not as yet been definitely decided. This is due to the fact that

the pneumatic fossa is still concealed in the matrix. A low

pectoral crest was developed, a feature we would naturally

look for, as doubtless these birds could fly well. Three well-

developed digits composed the skeleton of either hand ; the first,

or pollex, having two phalanges, index three, and medius four.

All of the distal or ungual phalanges supported claws. The

carpus, according to Pycraft, *probably agreed with that of

modern birds; except that the distal mass of fused bones

remained distinct throughout life, and that digit III was pro-

vided with a separate carpal bone." Every anatomist who has

examined the pelvic limbs of these fossil forms declares that the

skeleton of the pelvic limb is almost entirely avian in character,

Professor Gadow having noticed that the metatarsal to the

hallux digit was short and free, being directed backwards.

Others have observed, as Professor Dames and Dr. Hurst, the

reduction of the distal extremity of the fibula, but whether the

bone stands in front of the tibia, as in Iguanodon, and stated by

Dr. Stejneger, seems to be doubtful. The metatarsals were

elongated as in existing birds, and apparently firmly ankylosed

together, though their original separateness is easily made out

by the presence of the sutural lines between their shafts.

Including the hallux there were four toes, as in the higher bird

groups of this day, and their ungual phalanges were all armed

with a horny claw. Archzeopteryx possessed a femur departing

in general structure and appearance but very little from what

characterizes that bone in any medium-sized corvine type of the

present time, and so requires no special description. There

seemed to be, however, considerable curvature to its shaft. In

addition to the authorities I have already named as having con-

tributed to the knowledge of these Jurassic birds, may be added

No. 433.] CERTAIN GROUPS OF BIRDS. 37

the names of Huxley, Dollo, Fiirbringer, Romanes, Seeley,

Woodward, and Zittel. Each and all of these writers have

taught that, by extremely slow and gradual development in

time, our .existing birds were derived from ancestral reptilian

forms, and that the discovery of such a genus as Archzopteryx

need create no surprise, for it represents just such a type as we

would look for far back in geologic time during the earlier

transitional stages in the evolution of the class Aves.

The American Jurassic has also furnished fossil remains of

another land bird, but whether arboreal or not, as was the case

with A. “ithographica, cannot be determined from the limited

material. It was a toothed bird of some considerable size, and

was described as Laopteryx priscus by Marsh, who obtained it

from the Jurassic of Wyoming. It is principally represented

by the posterior portion of a skull, and this, it is said, presents

a somewhat struthious character. The single tooth found

near this skull was more or less like the teeth possessed by

Ichthyornis.

Beyond the fact, however, that Laopteryx probably belonged

to the same geological age as did Archaeopteryx, there is noth-

ing to indicate in the remains we have what manner of appear-

ing bird it was, much less as to whether it possessed a tail like

Archzopteryx. It is provisionally placed here in the order

Saururz for convenience only.

II. ORDER ORNITHURJE.

SUPERSUBORDER. SUBORDER. SUPERFAMILY. FAMILY.

Struthiornithes. Struthionidæ.

Rheornithes. Rheidæ.

Dromaiidæ.

Dromæognathæ. 4 Casuariornithes. Casuariidæ.

Dromornithidæ.

Dinornithes. Dinornithidæ.

Æpyornithidæ.

Æpyornithes.

Fürbringer employed the term Ornithuræ to designate his

Subclass II of birds, created to contain all existing and extinct

species of this group of vertebrates not included in Subclass I, the

Saururæ, which is represented alone by the fossil Archeopteryx

38 THE AMERICAN NATURALIST. [Vor. XXXVII.

lithographica and its allies. Gill, Stejneger, and others have

used the term Eurhipidurz for all birds in contradistinction

to the Saurure, while the last-named author throws the

toothed birds of the American Middle Cretaceous outside of it.

Now as Eurhipidurze means “ fan-tailed birds," it is a term not

strictly applicable, for neither the struthious birds nor the grebes

possess fan tails, while it is more than likely that the represen-

tatives of the ichthyonine birds did, and this very probably

was the case too with Hesperornis. As with so much that

now goes in avian taxonomy, even the lizard-tailed (Saurure)

and the bird-tailed (Ornithurz) orders can only be considered

provisional divisions. However, they can do duty until the

day comes when a fossil bird is discovered somewhat more rep-

tile-like than Ichthyornis, but presenting in the skeleton of its

tail a decided advance birdwards from Archzopteryx, even to

the point of the first stages of the formation of a pygostyle.

SUPERSUBORDER DROM/EOGNATHJE.

In this group it is intended to include all the existing and

extinct struthionine birds which are morphologically closely

allied to the ostrich (Struthio camelus). This will include the

Dinornithidz, but obviously exclude the Apterygidze, the Cryp-

turidze, and other families that are not ostriches in any sense of

the word, any more than was the American cretaceous toothed

loon, the Hesperornis. It comes about as near the expression

of true avian affinities to associate the tinamous with the

ostriches, for the reason that the posterior extremities of their

ilia and ischia have remained free, as it would be to relegate

the cassowaries to the crane group, simply because in them

those bones fuse together in the adult.

The supersuborder Dromzognathz includes the following sub-

orders, namely : (1) Struthtornithes ; (2) Rheornithes ; (3) Casua-

riornithes ; (4) Dinornithes ; (5) ZEpyornithes.

Of these the Struthiornithes are represented by the existing

African ostriches of the family Struthionide. The Rheornithes

include the South American ostriches of the family Rheidz, of

which there appear to be at least three well-defined species.

No. 433.] CERTAIN GROUPS OF BIRDS. 39

The Casuariornithes include three well-marked families, — the

Dromaiide, the Casuariidze, and the Dromornithida. The Drom-

aiidæ contain the emeus, the ostrich birds of Australia, of which

there are two species, Dromeus nove-hollandie, and D. irroratus.

The second family of the Casuariornithes or the Casuariidx

includes the cassowaries, other ostrich birds of the Australian

region, of which there are at least nine existing species of the

genus Casuarius. The cassowary of the island of Ceram is one

of the best known. It is the helmeted cassowary of science

(C. galeatus). The third family, Dromornithidze, is represented

by the extinct Australian genus Dromornis (Cat. Foss. Birds

Br. Mus. p. 355). The fourth suborder of the present super-

suborder, or the Dinornithes, has been created to contain

the family Dinornithidz, a group susceptible of being divided

into at least three subfamilies, and a number of genera (see

Trans. Zoól. Soc., London, Vol. XIII, Part XI, October, 1895,

P. 417). All these ostrich forms now appear to be extinct,

although this extinction has taken place only within compara-

tively recent time. They were the moas of the islands of New

Zealand, and were exterminated through the agency of the

inhabitants of the islands. The fifth suborder of this group is

represented by some five species of extinct ostrich forms of

the island of Madagascar, all referred to the genus /Epyornis.

This suborder has therefore been termed the ZEpyornithes, and

it has but the one family, ZEpyornithidze.

In Alfred Newtons A Dictionary of Birds under the

article * Roc" will be found an excellent article giving the history

of the discovery of these ancient ostriches, and excellent refer-

ences to the literature of the subject. In the same work are

found many other useful descriptions, the key to each of them

occurring under the title * Ratitae." Professor Newton there

says : “ According to the views adopted in this volume the sub-

class Ratitze comprehends of existing forms the orders Apteryges

(kiwi), Megistanes (cassowary emeu), Rhee (rhea), and

Struthiones (ostrich), together with the extinct /Epyornithes

(roc) and Immanes (moa). As regards the relation of other

older forms to the Ratita [as Odontornithes and Stereornithes]

it seems best at present to use reserve.”

40 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

It is safe to say that all of the ¢yfzca/ ostrich forms are

descended from some common stock. As will be seen further

on, the Apterygide are not considered to have any special

relationship with the ostriches ; they are altogether a different

kind of bird.

A complete account of the comparative osteology of the

Dromzaognathz would of itself make a large volume, so only

such information as is necessary for the purposes of classifica-

tion and to exhibit the general features of the skeleton among

these birds is presented here.

Several years since I wrote out a brief account of Struthio

camelus, with the intention of setting forth the various views

entertained by avian taxonomers and osteologists as to its sys-

tematic position since 1865, and to give the main features of

its skeleton. A good deal that I recorded was selected from

the observations of Huxley, the Parkers, and others, for many

anatomists have described more or less completely the oste-

ology of Struthio, and have held many opinions as to its affini-

ties. These opinions are becoming, however, more and more

unanimous. The researches of the ornithopalzontologists have

also greatly assisted the solution of the problem.

In my account referred to above I pointed out further that,

of all the class Aves, Struthio camelus Linn. is the largest

species of bird in existence, and it has been known, described,

and written about for ages. Of recent years some naturalists

have been disposed to recognize more than one form of African

ostrich, but the claim has not as yet been fully established.!

Suborder I. Struthiornithes.

Family STRUTHIONIDE : Struthio camelus.

Newton holds the opinion that * The genus Struthio forms

the type of one group of the subclass Ratitze, which differs so

widely from the rest in points that have been concisely set

forth by Professor Huxley (Proc. Zodl. Soc., 1867, p. 419) as

to justify us in regarding it as an order, to which the name

! Newton, A. Art. “Ostrich,” Dictionary of Birds, Pt. iii, pp. 662-666. The

alleged differences seem to be purely of a superficial character.

No. 433.] CERTAIN GROUPS OF BIRDS. 41

Struthiones may be applied ; but that term, as well as Struthi-

onidz, has been often used in a more general sense by system-

atists even to signify the whole Ratitz.! The most obvious

distinctive character presented by the ostrich is the presence of

two toes only, the third and fourth, on each foot, —a character

absolutely peculiar to the genus Struthio.?”

Huxley (Proc. Zoöl. Soc., 1867) placed the genus Struthio

alone in his first group, in the order (II) Ratitæ, while Garrod

included all the ostrich-birds, tinamous, screamers, fowls, bus-

tards, flamingoes, Musophagide, and Cuculidz, in his order

(I) Galliformes. Sclater arranges them thus:

Subclass II, Ratitæ.

Order XXIV, Apteryges.

XXV, Casuarii.

XXIV, Struthiones.

Reichenow’s scheme places them in a

Series I,

Order I, Brevipennes,

Family I, Struthionide,

including the entire assemblage of the once-called struthious

birds.

In a Subclass IV (Eurhipidurz), Stejneger classifies them

thus:

Superorder I, Dromzognathe.

Order I, Struthiones.

Superfamily I, Struthioidex.

II, Rheoidez.

III, Casuaroidez.

Family I, Dromaiide.

II, Casuariide.

Superfamily IV, Dinornithoidez,

followed by the remainder of his classification.

1 At one time it was not uncommon to include the bustards among the Stru-

thionidz !

? Remains of a true ostrich have been recognized from the Sivalik formation

in India, and the petrified egg of an apparently allied form, Struthiolithus, has

been found in the south of Russia. Among the more important treatises on this

bird may be mentioned: E. D’Alton, Die Shelete der Straussartigen Vögel abge-

bildet und beschrieben, folio, Bonn, 1827 ; Professor Mivart, “On the Axial Skele-

ton of the Ostrich ” (Proc. Zoöl. Soc., vol. viii, p. 385); M. Alix, Essai sur Pappareil

42 THE AMERICAN NATURALIST. | [Vor. XXXVII.

Fiirbringer, in an order Struthiornithes, creates a suborder

Struthioniformes, which is further subdivided into a * gens,"

Struthiones, and the family Struthionide. The Struthioni-

formes is an “order” in Dr. Sharpe's systematic arrangement

of the class, an arrangement he clearly sets forth in his Hand-

List of Birds (Vol. 1, pp. 1-8, London, 1899). And thus we

might proceed, giving one classificatory scheme after another,

each and all practically presenting the same idea as to the

position of the ostrich in the system. Reichenow’s scheme,

however, is a little antiquated now, while I hardly think that

all naturalists will agree with Garrod in placing the cuckoos

and flamingoes in the same * cohort," and these in the same

order with the ostriches.

Huxley has said (Proc. Zoöl. Soc., 1867, p. 419) the Afro-

Arabian genus Struthio is the type of one group of this order

(Ratitae) characterized by:

1. The prolongation of the maxillary processes of the pala-

tine bones forwards, beneath the maxillo-palatines, as in most

birds.

2. The thickening of the inner edges of the maxillo-palatines,

and their articulation with the facets upon the sides of the vomer.

3. The shortness of the vomer, which does not articulate

with either palatines or pterygoids posteriorly.

4. The slight, or wanting, ossification of the prefrontal pro-

cesses of the primoidal cranium.

5. Theunion of the bodies of the sacral vertebrze with the

anterior ends of the pubes and ischia.

6. The presence of two shallow notches on each side in the

posterior margin of the sternum.

7. The proportions of the fore limb. The humerus is about

equal in length to the distance between the pectoral arch and

the ilium, and is therefore much longer than the scapula. The

antebrachium is not half as long as the humerus. The manus

possesses the ordinary three digits ; and two of these, the radial

and the middle, are provided with claws.

8. The union of the pubes in a symphysis.

locomoteur des oiseaux (Paris, 1874) ; and Professor Macalister, “On the Anatomy

of the Ostrich " (Proc. Roy. Zrish Acad., vol. ix, pp- 1-24).

Fic. 1. — Skeleton of ostrich (Struthio camelus Linn.). Subadult, much reduced.

: ws :

> Lir

No. 13,806, Coll. Nat. Mus

44 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

9. The abortion not only of the hallux, but also of the distal

end of the metatarsal bone and of the phalanges of the second

digit of the foot, whence the foot is two-toed.

IO. The presence of thirty-five precaudal vertebrae.

In the same place Huxley gives the following osteological

characters for the Ratitæ, or the *struthious birds," which

* differ from all others in the combination " of these ied

ities (p. 418).

1. The sternum is devoid of a crest, and ossifies only from

lateral and paired centers.

2. The long axes of the adjacent parts of the scapula and

coracoid are parallel or identical. The scapula has no acromial

process, nor has the coracoid any clavicular process ; at most

there are inconspicuous tubercles representing these processes.

3. The posterior ends of the palatines and the anterior ends

of the pterygoids are very imperfectly, or not at all, articulated

with the basisphenoidal rostrum, being usually separated from

it and supported by the broad, cleft, hinder end of the vomer.

4. Strong * basipterygoid " processes, arising from the body

of the basisphenoid and not from the rostrum, articulate with

facets which are situated nearer the posterior than the anterior

ends of the inner edges of the pterygoid bones.

5. The upper, or proximal, articular head of the quadrate

bone is not divided into two distinct facets.

In his Anatomy of Vertebrated Animals Huxley has like-

wise pointed out that the cervical vertebrz in the Ratitze have

short transverse processes and ribs, disposed very much as in

the Crocodilia, and I find that the ultimate vertebra of this

series in the ostrich bears a pair of free cervical ribs (see

Fig. 1). There appear to be nineteen of these cervical vertebrz,

and six free dorsals, the latter developing lofty neural spines

that gradually increase in height from before backwards. Four-

teen or fifteen free caudals are also found in the chain, and

these terminate with a stumpy pygostyle.

The dorsal ribs and their haemapophyses are strong and

substantial; the later exhibit a peculiar curving, and the epi-

pleural appendages of the former are aborted in subadult indi-

viduals. Two pairs of short pelvic ribs are seen.

No. 433.] CERTAIN GROUPS OF BIRDS. 45

Sir Richard Owen, who published a great deal about the

osteology of ostriches and their kin, both living and extinct,

says of the sternum of Struthio camelus that “it is broader in

proportion to its length, and subquadrate in the ostrich,” and

that in “all these keel-less sternums ossification begins, as in

the ostrich, by a pair of centers expanding until they meet and

coalesce in the middle line, and thence, according to the stimulus

of the growth and pressure of the pectoral muscles, extending,

as a keel, into the interspace.” 1

In the pelvis of Struthio the ilia are long and narrow, their

postacetabular portion being thoroughly and widely separated

from the ischium upon either side (Fig. 1.), while in front the

preacetabular region is shorter and much concaved externally.

Huxley says: “In Struthio, alone, among birds, do the pubes

unite in a median ventral symphysis. Another, not less

remarkable circumstance, in the ostrich, is, that the 31st to the

35th vertebrz inclusively (counting from the atlas) develop

five lateral tuberosities. The three middle tuberosities are

large, and abut against the pubis and the ischium. In these

vertebra, as in the dorsal vertebrae of Chelonia, the neural

arch of each vertebra shifts forward, so that half its base articu-

lates with the centrum of the next vertebra in front; and the

tuberosities in question are outgrowths, partly of the neural

arch, partly of the juxtaposed vertebral centra, between which

it is wedged. Hence, in young ostriches, the face of each

tuberosity exhibits a triradiate suture." ?

A conspicuous propubis is developed in the case of Struthio,

and this has been figured by Owen (Anat. Vert., Vol. II, p. 36, m),

but in that figure the peculiar structure to which attention

was invited by Garrod is not shown ; this consists of a small

osseous plate attached to the pubis, that is partly surrounded

by cartilage Forbes speaks of “this paper, written in con-

junction with Mr. Frank Darwin," and points this out as the

principal point of interest, and refers toit as “a peculiar nodule

of bone lying on the center of the pubis and, in some respects,

1 Anat. Verts. vol. ii, pp. 24, 25-

2 Ibid., pp. 251, 252.

3 Garrod, A. H. Coll. Sci. Papers, p. 99.

46 THE AMERICAN NATURALIST. | [Vor. XXXVII.

similar to the * marsupial' bone of the implacental Mammalia

and its corresponding fibrous representative in certain Carniv-

ora."! As it has not yet received any special name it might

be called the suprapubic ossicle. In Garrod's figure, where it

is given, the pubis and ischium are firmly united posteriorly,

as is also the case in Owen's figure, cited above, while in the

pelvis of the ostrich in the collection of the United States

National Museum these bones are distinctly independent of

each other posteriorly (Fig. 1). Perhaps these do not unite

until the bird is greatly advanced in age, and that this specimen

is in a subadult stage of growth, which is the more likely as

the epiphyses of the proximal extremities of the tarsometatar-

sals have not as yet coóssified with the shaft.

Owen says in *the ostrich the two clavicles are distinct

from each other, but are severally anchylosed with the coracoid

and scapula, so as to form with them one bone on either side."

In the pectoral extremity the humerus of the arm is reduced

to a mere curved and slender rod of bone, with slightly enlarged

ends; while the radius and ulna of the antebrachium are even

more decidedly aborted.

In The Ibis and in The Philosophical Transactions of the

Royal Society of London (1888) W. K. Parker has given

instructive figures of the manus of Struthio camelus, and they

go to show that in the adult individual the radiale and ulnare

ossicles of the carpus are separate andin bone. The phalanges

have a most reptilian look, and the terminal joints of all three

fingers are distally armed with a free, movable claw.

The pelvic extremity of Struthio is powerfully developed, all

the bones present being massive and strong. Both the proxi-

mal end of the femur and its distal condyles are greatly

enlarged. A patella is not developed in bone. The tibio-

tarsus and tarsometatarsus are straight, and of nearly the same

length. The distal end of the latter is modified for the articu-

lation of the third and fourth digits, the only two toes possessed

by this bird.

The cnemial process of the tibia is ossified by a separate

epiphysis, in common with Rhea. Owen says the pneumatic

1 Forbes, W. A. Coll. Sci. Papers, P- 203.

No. 433.] CERTAIN GROUPS OF BIRDS. 47

foramen of the femur in the ostrich is situated posteriorly

rather than in front, as it is in nearly all other birds, while the

*epicnemial process" of the tibiotarsus “extends forward,

without rising above the level of the proximal surface, and con-

tracting to its termination, there divides into small pro- and

ecto-cnemial processes; the latter the shortest and tuberous."

As for pneumaticity, the bones of an ostrich enjoy a greater

degree of it than do those in the case of any of the true

Laridze.

Struthio camelus is, with respect to existing birds, most

nearly related to the South American ostriches, the various

species of Rhea.

The late T. J. Parker, in his admirable memoir *On the

Cranial Osteology, Classification, and Phylogeny of the Dinorni-

thidae " (Trans. Zool. Soc., London, Vol. XIII, Part XI, October,

1895), gives very complete tabular schemes comparing the

cranial characters of several of the supersuborders of the Dro-

mzognathz, and from these it will be seen that a number of

excellent characters distinguish the cranium of Struthio from

that of Rhea.

Suborder II. Rheornithes.

Family : RHEIDÆ.

In the genus Rhea, the only genus of the present family, are

contained those ostrich-like birds of South America, commonly

known in Europe as nandu. According to Newton there are

at least three species of these, vzz., K. americana, R. darwini,

and R. macrorhycha. Considerable has been written upon their

osteology, but more particularly has the skeleton of Rhea amert-

cana been described, which received the attention of Huxley,

of the Parkers, and of not a few others. Nearly all recent

authoritative taxonomers place these birds in an order, coequal

with the order occupied by the ostriches proper (Struthio).

Some of the special osteological characters of Rhea have

been pointed out by Huxley, thus:

I. The maxillary processes of the palatines are short and

unite with the inner and posterior edges of the maxillo-

palatines.

48 THE AMERICAN NATURALIST. | [Vor. XXXVII.

2. The maxillo-palatines are thin, fenestrated plates, which

do not articulate with facets on the edges of the vomer.

3. The vomer is as long as it usually is in birds, and articu-

lates behind with the palatine and pterygoid bones.

4. The prefrontal processes are little ossified.

5. The bodies of the proper sacral vertebrae do not unite

with the pubes or ischia ; and the centra of the sacral vertebrae,

which ossify late, are extremely elongated and slender.

6. The short sternum narrows posteriorly and presents a

notch in the middle of its posterior edge.

7. The length of the humerus exceeds the distance between

the shoulder girdle and the ilium, and is of course greatly longer

than the scapula. The manus has the same conformation as

that of Struthio.

8. The pubes are free, but the ischia unite beneath the

urosacral vertebrae.

9. The hallux is absent, but the second, third, and fourth

digits are complete.

10. There are only thirty-two precaudal vertebra. (Proc.

Zoöl. Soc., 1867, pp. 420-422.)

As in the case of Struthio, the skeletology of Rhea has long

been known, and Prof. Kitchen Parker has, in his famous paper

“ On the Osteology of Gallinaceous Birds and Tinamous," given

us a few of the necessary characters for the use of the taxon-

omer, they being presented in connection with what is there

done with Tinamus.

The late T. J. Parker compared the skulls of Struthio and

Rhea in his memoir on the Dinornithidze (see azzea), and showed

that the differences existing between these two birds, in so far

as that part of the skeleton is concerned, was in his estimation

of ordinal rank. These distinctions, however, in the present

work are considered to be but of subordinal value.

Suborder III. Casuariornithes.

Families : DROMAIIDE (the emeus); CASUARIIDJE (the cassowaries) ;

ROMORNITHIDEE (extinct).

Huxley has already pointed out (Proc. Zoól. Soc., 1867, pp.

422, 423) that the osteology of Casuarius and Dromzeus (emeu)

No. 433-] CERTAIN GROUPS OF BIRDS.

Fic. 2, — Right lateral view of the skeleton of Casuarius galeatus, greatly reduced.

i . S. Nat. Mus.

o. 16,964, Coll. U

50 THE AMERICAN NATURALIST. | (Vor. XXXVII.

is much alike, and as a rule these birds have been by taxon-

omers properly considered to be more closely related than are

Struthio and Rhea to each other. In the memoir just referred

to, this eminent authority grouped the Malayo-Australian genera

Casuarius and Dromzus together, and for this group gave the

following osteological definitions :

1. The maxillary processes of the palatines are short as in

Rhea.

2. The maxillo-palatines are flat, imperforate plates, which

unite solidly with the premaxillae and the vomer.

3. The vomer is long, and articulates behind with the pala-

tine and pterygoid bones. l

4. The prefrontal processes are large and well ossified.

5. The bodies of the proper sacral vertebræ do not unite

with the pubes or ischia; and the bodies of the urosacral ver-

tebræ are very large, thick, and well ossified.

6. The sternum is long and escutcheon-shaped, at first

widening and then coming to a point behind.

7. The humerus is not nearly half so long as the distance

between the pectoral arch and the ilium, and is much shorter

than the scapula. The antebrachium is not more than half as

long as the humerus. Only one digit, the median, is complete

and bears a claw.

8. Neither the pubes nor the ischia unite in the middle line

of the body. 7

9. The hallux is absent, but the other digits are complete.

IO. There are thirty-five precaudal vertebrae.

Other osteological characters to be noted in the skeleton of

Casuarius galeatus may be thus tabulated (see Fig. 2):

1. The light, spongy osseous core surmounting the top of

the skull. In life this supports the horny helmet.

2. There are nineteen cervical vertebra, the last three

bearing big free ribs that are without epipleural processes.

There are szx dorsals, the first five of which have ribs connect-

ing with the sternum by means of hemapophyses. Only the

four in the middle of the series possess epipleural appendages ;

the last pair of ribs fail to connect with the sternum, as is also

the case with the smaller pair of pelvic ribs present.

No. 433.] CERTAIN GROUPS OF BIRDS. 5I

3. The external nostril is near the apex of the bill (Owen).

4. The clavicle is anchylosed with, or rather is a continuous

ossification from the scapula; but the coracoid bone is free

(Owen).

5. “In the adult Indian cassowary (Casuarius galeatus)

there is only one carpal bone free; the manus is a solid single

piece, with only one finger (the second, or index) developed, and

this has only zwo phalanges — it ought to have Zree; and the

distal phalanx is an inch long and carries a large claw " (Parker).!

6. The pelvis resembles the pelvis in the emeu in form and in

its main characters ; but in Dromzeus the ischium is, posteriorly,

well separated from the ilium, while in the cassowary it fuses

with it in that locality.?

7. The pelvic limb is powerfully developed : there is a big

femur presenting many of the common ornithic characters,

while the other bones of this extremity are also large, save

hallux and first metatarsal, which are absent.

8. In the tibiotarsus the cnemial projections are conspicu-

ously produced and rise to some extent above the proximal sur-

face of the bone. The fibula has a massive head, while below

its articulation with the shaft of the tibia it is tapering, slender,

long, and styliform, ending in a free distal extremity.

9. The hypotarsus of the tarsometatarsus is a long, low

median crest, and the ungual phalanx of the inner toe is espe-

cially elongated, straight, gradually tapering and distally pointed.

The osseous claws of the other two toes are moderately curved,

and exhibit proportions more in keeping with the remaining

joints of their respective digits.

In T. J. Parker's memoir on the Dinornithide there is a

tabular synopsis of the chief cranial characters of Dromzeus and

1* In the ripe embryo of a specimen of the Mooruk (C. dennetti) I find four

cartilaginous carpal nuclei and three metacarpal rays: the first is very small and

feeble; the second very strong and with the normal number of phalanges Ge.

three besides the metacarpal), and the distal or ungual joint is very long and carries

a long claw; the third metacarpal is about one-sixth the size of the second, and

has no phalanges on it. In the emu (Dromeus) the second digit has two

phalanges and a long curved claw.” bid., W.K. P.

2 Compare the side view of the pelvis in the plate with Marsh’s figure of the

pelvis of the emeu (Ordontornithes, p. 7, Fig. 16).

52 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

Casuarius recorded in a comparative way that is extremely

useful (Trans. Zool. Soc., London, Vol. XIII, Part XI, October,

1895, pp. 410, 411). In closing the brief account of this sub-

order, especial attention is invited to the fact that the distal

extremities of the ilium and ischium upon either side in the

pelvis of Casuarius are firmly fused together as they are in all

adult birds of the order Ornithure, and not free as in all

ostriches known to us, either existing or extinct. Although very

unostrich-like, yet no one with a knowledge of birds will ever

question the claim of the cassowary to a place among the

existing representatives of that group.

The representatives of the family Dromornithide are all

extinct forms discovered in eastern (Dromornis) and southern

(Genyornis) Australia. They here constitute the third family

of the suborder Casuariornithes, but from the fact that they are

fossil forms not far removed from the existing ostrich types,

they will not be dwelt upon in this article. T. J. Parker has

paid no little attention to them in his exhaustive memoir cited

above. (See also Lydekker, Caz. Foss. B., p. 355, 1891.)

Suborder IV. Dinornithes.

Family : DINORNITHID4 (the moas).

Considerable literature is extant of the extinct moas of the

North and South islands of New Zealand. This is amply

referred to in an admirable article, * Moa," by Lydekker,

contributed to A Dictionary of Birds, by Newton. There

one will find a number of moa's bones accurately reduced and

figured with the remarks that * Moas are distinguished from

all existing Ratitae in having a bony bridge on the anterior

surface of the lower end of the tibia above the condyles. The

tarsometatarsus has three distal trochlez, and in most cases

(according to Capt. Hutton probably all) carried a hallux. The

beak (unlike that of the kiwis) is short and stout ; the form of

the lower jaw being either U-like or V-like. The general form

of the pelvis is very like that of the kiwis; but the sternum

differs by the absence of the superior notch, the more divergent

lateral processes, and the abortion or disappearance of the

grooves for the coracoids " (p. 578).

No. 433.] CERTAIN GROUPS OF BIRDS. 53

The late T. J. Parker, who makes three subfamilies and five

genera of the family Dinornithidz (Trans. Zool. Soc., London,

October, 1895, pp. 417 et seq.), has, among other extensive

osteological comparisons of these birds, pointed out the follow-

ing facts, which he tabulates thus :

The Skull in the Dinornithide.— Occipital plane vertical or

very slightly inclined backwards or forwards ; occipital condyle

pedunculate ; occipital crest variable. Length of cranial roof

from two to two and a half times length of basis cranii.

Mammillar tuberosities usually prominent ; basitemporal plat-

form always well defined and separated from occipital condyle

by a deep precondylar fossa.

Width at paroccipital processes from less than one and a

half to more than twice length of basis cranii.

Width at squamosals from about one and three-quarters to

one and a half times length of basis cranii.

Height of cranium about one and a quarter times length of

basis cranii.

Temporal fossa extends mesiad, to a greater or less extent,

on to parietal region ; distance between temporal ridges varies

from about width of cranium at temporal fossa to half that

width. Zygomatic process short, pointed, and nearly parallel

to median plane; auditory region of skull produced into a

strong squamosal prominence.

Width of orbit about half width of cranium at paroccipital

processes, and almost invariably less than length of basis

cranii; interorbital septum absent or greatly reduced ; a broad

supraorbital ledge, produced behind into a strong, broad, post-

orbital process.

Lacrymal ankylosed with frontal, forming preorbital process ;

no orbital process; a descending process ankylosed with outer

border of antorbital, and notched or perforated for lacrymal

duct. Mesethmoid produced into paired horizontal triangular

processes. Antorbital well ossified ; ankylosed to descending

process of lacrymal; perforated dorsally by a supraorbital

fenestra of variable size. ;

Nasal either has a slender maxillary process, or there "^

distinct maxillo-nasal bone; meets its fellow of the opposite

54 THE AMERICAN NATURALIST. . [Vor. XXXVII.

side in the middle line above the ethmoid, so that the latter

does not appear on the dorsal surface; premaxillary groove on

upper surface of nasals extends backwards to or beyond naso-

frontal suture. Premaxilla strong ; body more or less elevated,

and with a distinct prenarial septum ; palatine processes broad

and produced into more or less definite vomerine processes;

width of body always more than half and sometimes one and a

half times length of basis cranii. Maxilla short and narrow;

maxillo-palatine a short, flat plate, produced dorsad either into

an irregular shell of bone containing a large antrum, or into a

thick, oblique plate containing no, or but little, trace of the

antrum.

Vomer less than one and a half times length of basis cranii;

consists of thin paired plates meeting each other ventrad in

an acute dihedral angle, and either quite free or partially

ankylosed with one another in front; firmly ankylosed behind,

in fully adult specimens, with palatines and pterygoids.

Palatine a thin twisted plate, about one and a fifth times

length of basis cranii; pedate posterior end produced into

short mesial vomerine process; articulates at anterior end

with maxilla, and posteriorly with vomer and pterygoid, with

which, in fully adult specimens, it becomes ankylosed.

Mandible very strong ; symphysis short, more or less flattened

and ridged below; distal end more or less deflected downwards.

The best part, or an extremely useful feature in connection

with Parker's work, from which the above is quoted, is the

excellent series of plates that illustrate it. These are devoted

to the skulls of the various genera of the Dinornithidae

(Emeus, Anomalopteryx, Mesopteryx, Pachyornis, Dinornis),

as well as a number of colored figures, showing the relation-

ships to each other of the cranial bones in Emeus and

Anomolopteryx.

Suborder V. Zpyornithes.

Family : ASPYORNITHID& (AZpyornis, the roc).

This group has been created to contain the now extinct

ostrich-like birds of the island of Madagascar. Fossil and

subfossil specimens of eggs and bones were first accurately

No. 433.] CERTAIN GROUPS OF BIRDS. 55

described and named by Isidore Geoffroy-St. Hilaire in 1851,

who named this new ally of the ostrich Æpyornis maximus.

This was confirmed later by M. M. Alphonse Milne-Edwards

and Grandidier (Aun. Sci. Nat., Ser. 5, Vol. XII, pp. 167—196,

Pls. VI-XVI), and now the opinion is quite universally enter-

tained among ornithotomists that these birds were ostriches

related to the genus Struthio of the African continent or the

adjacent mainland. It has been shown, however, that the

largest species of /Epyornis thus far discovered, as indicated

by its fossil remains, was by no means as big or as tall a bird

as the larger species of the Dinornithidz of New Zealand.

The fossil remains in the hands of science of these Madagascan

ostriches are by no means abundant, consisting principally of

bones of the trunk skeleton and of the lower extremity. Max

Fürbringer ! has discussed the value of these very fully as well

as the work upon them by Edwards and Grandidier. It is

not considered necessary in this brief article to redescribe

these fragmentary remains, and there can be no question

but what the birds they represent were a group of ostriches

quite as distinct as the present existing ostriches of Africa

(Struthio).

This concludes my brief survey of the osteological characters

of the fossil and existing forms of the true ostrich birds.

Before concluding the present article, however, I should like

to call attention to a well-known fact, that it is very generally

believed that Apteryx is closely allied to the Dromzognathe,

and should be grouped with them. Many claim that the

family Apterygide, to which it belongs, is in the same suborder

with the Dinornithide, but the more attention I pay to the

phylogeny of birds the less and less do I see the glaring evi-

dences of the struthionine affinities of these birds.

It would seem that other naturalists besides myself have, or

do, entertain similar doubts upon this point. Dr. Sharpe in

his admirable work A Review of Recent Attempts to classify

Birds, in giving his ideal plan of an arrangement of birds in a

museum in order to exhibit their relationships, says on page 59,

1 Untersuchungen zur Morphologie und Systematik der Vögel. 11, Allgemeiner

Theil, pp. 1463-1465.

56 THE AMERICAN NATURALIST. | [Vor. XXXVII.

after he has grouped the ostriches together: ** A little further

afield we should come to the Apteryges, and here attention

should be drawn to the ralline tendencies of these abnormal

Ratitz,, with all those other peculiar characteristics on which

it is not necessary here to dilate at length." Just why the

Apteryx should be called “abnormal” more than any other

bird living, or extinct, I fail to see. Any puzzling form may

seem abnormal when persistent attempts are made to force it

into an assemblage of other forms where it does not strictly

belong.

Again, Fürbringer in his vertical aspect of the phylogenetic

tree of birds has the branch Apterygiformes arise from the

main trunk near the rails and far removed from any of the

ostriches. In his opinion this Apterygian branch soon forked,

however, and gave rise to the two families, Apterygida and

Dinornithida. In his lineal scheme the position given these

is in an order Alectoronithes, containing the Apteryges, the

Crypturi, the Galline, and the Opisthocomida. Many large

groups both of land and water birds in this lineal scheme

separate them from the ostrich birds, and it is very evident

from all this that Fürbringer was of the opinion that the moas

and kiwis are but very remotely related to the ostriches, the

rheas, the emeus, the cassowaries, or any of the rest of that

assemblage. |

T. J. Parker commented upon this in the following words:

* The most definite opinion I have met with as to the phy-

logeny of the Ratitze is that expressed in the elaborate genea-

logical tree which illustrates Fürbringer's great work. He

ascribes a common origin to the moas and kiwis and to the

emeus and cassowaries, but derives his four main groups of

Ratitzae — the Struthioniformes, Rheiformes, Casuariformes,

and Apterygiformes — separately from a primitive stock.

* Mivart, in his memoir on the axial skeleton of the Ratitz

(Trans. Zool. Soc., Vol. X, 1871), gives no definite opinion as

to the phylogeny of the group, but his diagram illustrating the

mutual relationships of the various genera seems to indicate

his belief in their monophyletic origin. He shows a main

stem dividing into two branches; one of these divides again

No. 433.] CERTAIN GROUPS OF BIRDS. 57

for Struthio and Rhea; the other forks a second time, one

branch dividing again for Casuarius and Dromzus, the other

for Dinornis and Apteryx.

“The monophyletic origin of the Ratitz is also supported

by Newton, who, in his luminous article, * Ornithology,' says

‘that these forms — moa, kiwi, emu and cassowary, rhea,

and finally ostrich — must have had a common ancestor nearer

to them than is the ancestor of any carinate form' seems to

need no proof.

* Professor Newton's classification indicates no closer affinity

between any of the genera except the emu and cassowary,

which together constitute his order Megistanes; each of the

other genera has an order to itself.

* A study of the skull certainly confirms the view that the

nearest ally of the Dinornithide is Apteryx, and that the four

families of Australasian Ratitz are more nearly related to one

another than is either of them to the Asio-African and South-

American forms. Struthio and Rhea differ so much from the

Australasian members of the subclass as to lend strong sup-

port to Fürbringer's view that they arose separately from a

primitive stock ; but whether the cassowaries and emus on

the one hand and the moas and kiwis on the other had a dis-

tinct or a common origin is a very complex question.

“The main difficulty lies in deciding what characters should

be considered as of phylogenetic importance and what merely

adaptive, but it appears to me that in the following particulars

the emu and cassowary show an undoubted relationship to the

moas.1

* The general,characters of the maxilla, maxillo-palatine, and

antrum in both genera.

* The general relations of the vomer, palatines, and pterygoids

in both genera.

“ The presence of a vestige of the maxillary process of the

nasal in Dromzeus.

“The well.ossified antorbital ankylosed to the descending

process of the lacrymal in both genera.

1“ As my conclusions are based upon a study of the skull, I have omitted all

reference to /Epyornis, Dromornis, Megalapteryx, and Palzocasuarinus

58 THE AMERICAN NATURALIST. [VoL XXXVII.

“The elevated body of the premaxilla with its distinct pre-

narial septum in Casuarius.

* Forbes's discovery (Zvans. Nat. Zool. Inst; Vol. XXIV, 1891,

p. 185) of a dinornithine bird which he calls Palaeocasuarinus

will, if the detailed account of his very interesting researches

bears out the opinions expressed in his preliminary note, lend

strong support to this view. The tibia upon which the genus

is founded have, as the name implies, a remarkable resem-

blance to those of the cassowary.

* On the other hand, I know of no character in the skull of

Rhea by which it definitely approaches the moas, and the

presence of a maxillary process to the nasal, the form of the

cerebral fossae, and the position of the pneumatic foramen of

the quadrate seem the only particulars in which the ostrich

comes in any way near them. Struthio and Rhea are, in fact,

sharply separated both from one another and from the Austral-

asian Ratita, as well by the characters of the bony palate as

by those of the pelvis. The characters possessed by them in

common with the other Ratitze are of two kinds: ancestral

characters, such as the form of the vomer, the basipterygoid

processes, and the single-headed quadrate, which, according to

the view taken in this paper, are accounted for by the hypothesis

of common descent from a group of generalized flying birds or

Proto-Carinatz ; and adaptive characters, such as those of the

sternum, shoulder girdle, and wing, which they share to a

greater or less degree with all flightless birds.

“The marked differences between the moas and kiwis are

certainly for the most part adaptive ; the two families resemble

one another in the increased size of the olfactory organ and the

reduced size of the eye ; but both processes have gone so much

further in Apteryx that the differences between the two, in

this respect alone, give the skulls the appearance of being more

widely separated than those of any other two ratite birds. The

real affinities underlying these differences are, however, shown

by the striking similarity of the bones of the palate in the two

forms. The absence of a maxillary antrum in Apteryx seems

at first sight a difference of great importance, but the fact that

this cavity has disappeared or become vestigial in one of the

No. 433.] CERTAIN GROUPS OF BIRDS. 59

most specialized genera of the moas seems to indicate that its

complete atrophy in the kiwi is simply to be looked upon as

an instance of the extreme specialization of that genus."

I have thus fully quoted from T. J. Parker's views upon the

relationships of the ostriches and Apteryx for the reason that

they are important and useful in the present connection, and that

they are entertained by many other naturalists. It must be

remembered, however, that these views are drawn up after an

examination of the bony skulls alone, or very nearly alone. It

must be borne in mind, too, that Marsh endeavored to make

ostriches, or ratite birds, out of Hesperornis and Ichthyornis

simply because they possessed the ancient form of palate, and

that their ilia and ischia possessed free posterior extremities.

I now pass to a consideration of the osteological characters of

the Odontoholcz, the supersuborder to which the Hesperoni-

thide belong. It will be necessary to reproduce my observa-

tions in order to properly set forth and support my scheme of

classification which, as I have already said, will be published

in the future.

SuPERSUBORDER III. ODONTOHOLCA.

SUBORDER. SUPERFAMILY

AMILIES.

Pygopoformes. Hesperornithoidea. Hesperornithide.

Enaliornithide.

Fossil remains of upwards of fifty individuals representing

extinct species of the Hesperornithide have been taken from

the Middle Cretaceous of Kansas and Colorado, where occurred

also Icthyornis (Order II) and its allies. Marsh also described

the bones of these great ancestral divers in his Odontonithes,

and left us a restoration of the skeleton of Hesperornis regalis.

This was not a difficult task for the reason that nearly perfect

skeletons of that form were deposited in the museum of Yale

College, and one of these was so complete that it lacked only

a very few unimportant bones, as the distal ungual phalanges of

two or three of the podal digits.

Hesperornis regalis had a length of about six feet, and an

adult specimen when assuming the erect attitude would have

had a height of about three feet. The distinguished Scotch

60 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

anatomist, D’Arcy W. Thompson, and myself have carefully

examined into the osteology of these extinct divers, and have

shown that they undoubtedly are among the ancestors of the

existing Pygopodes (vzde postea).

The skull of Hesperornis in its general formation resembles

closely the skull of such a diver as Urinator lumme, but exhibits

characters common to many birds belonging to widely sepa-

rated groups of the age in which it lived. For example, the

capacity of the cranial casket was small; the superior articular

head of the quadrate bone had but one articular facet ; they had

teeth in grooves, all the length of either ramus in the lower

jaw, but confined to the dentary borders of the maxillaries in

the case of the upper. The ramal symphysis was cartilagi-

nous throughout the life of the individual, and consequently

each ramus was a separate bone.

According to Marsh, the proximal extremities of the palatines

and the distal ends of the pterygoids did not articulate with

the basisphenoidal rostrum. The stout basipterygoid processes

arose from the body of the basisphenoid, and each one articulated

with a facet upon the corresponding pterygoid situated near

its proximal end. It had separate vomers. The supraorbital

glandular depressions, the holorhinal nostrils, the firmly united

intercranial sutures, the form of the quadratojugal, and many

other characters, exhibiting but little change in their now living

descendants, are each and all characteristic of the Pygopodes.

As the teeth do not belong to the skeleton, they require no

special description in an article devoted to osteology ; be it

sufficient to say that they were purely of a reptilian type.

The mandible was long and slender, and “the rami were

united at the symphysis in front only by ligament, a feature

unknown in modern adult birds. There is an imperfect articu-

lation between the splenial and angular elements, which prob-

ably admitted of some motion, and all the other sutures are

open, or distinguishable. There was apparently a mandibular

foramen. There is a well-marked shallow groove on the outer

superior margin of each dentary bone for the reception of the

maxillary teeth when the jaws were closed. The angle of

the mandible extends backward but a short distance beyond

No. 433-] CERTAIN GROUPS OF BIRDS. 61

the articular face for the quadrate, and the extremity is

obliquely truncated " (Marsh). ZZesperornis regalis possessed 49

vertebra in its vertebral column, or 23 presacrals, 14 sacrals,

and r2 caudals. Both in the articular facets of the centra and

in other characters they agree with such modern genera as

Urinator and other divers. In its caudal skeleton this great

cretaceous diver was peculiar, the free anterior caudal vertebrae

being short, with lofty neural arches and not conspicuous

diapophyses, and exhibited an opisthoceelian articulation. Great

horizontal expansion characterized the long transverse processes

of the mid and posterior caudals, while the last three or four

elements of this part of the vertebral chain coóssified together

in the adult, forming a fat, horizontally compressed, pygostylous

mass, very different from anything to be found in the form of

the terminal piece in the tail of existing Aves. None of the

presacral vertebrae united, and none of the caudals possessed

zygapophyses. It is very likely that Hesperornis used its

broad, horizontally flattened tail much as the now-existing

beaver among modern mammals employs its paddle-like caudal

appendage, — a powerful aid as a propeller and rudder to the

aquatic locomotion of this ancient pygopodous fowl.

The sternum in this genus is flat and broad and thin posteri-

orly. Anteriorly, it is rounded and projecting, while but two

shallow notches are to be seen in its xiphoidal margin. The

ribs, some of them bearing uncinate processes, resembled those

of the loons, but the shoulder girdle, with its non-united clavicles,

was weak and small in character, and the pectoral limb was

reduced to a rudimentary humerus.

The pelvis of Hesperornis was like the pelvis of our modern

loons and grebes, and Marsh observed that it resembles that

of Podiceps, being very long and narrow, as in that genus, and

in other diving birds. He also remarked that the “acetabulum

differs from that in all known birds, in being closed internally

by bone, except a foramen that perforates the inner wall, as in

the crocodiles. The ilium, ischium, and pubis, moreover, have

their posterior extremities free and distinct."

The powerful bony framework of the pelvic limbs of this great

extinct diver agrees in many characters with the corresponding

62 THE AMERICAN NATURALIST. [Vou. XXXVII.

parts in the skeleton of the legs in grebes and loons. This

has also been noted by Stejneger, and Marsh said in his

Odontornithes that the “posterior limbs of Hesperornis regalis

present an admirable example of adaptive structure. The

means of locomotion were confined entirely to these extremi-

ties, and the life of Hesperornis was probably more completely

aquatic than that of any known bird. It may fairly be ques-

tioned whether it could even be said to walk on land, although

some movement on shore was of course a necessity. Considering

the posterior limb as a whole, it will be found a nearly perfect

piece of machinery for propulsion through the water. Provision

was made for a very powerful backward stroke, followed by a

quick recovery, with little loss by resistance, a movement quite

analogous to the stroke of an oar, feathered on its return.

“Among recent birds, we have, in the genus Podiceps, the

nearest approach to the legs and feet of Hesperornis, and the

osseous structure of these parts is essentially the same

throughout in the two genera. The muscular system, also, of

this member must have been very similar in both. In many

respects, however, the bones of the posterior limbs of Hes-

perornis present evidences of a more primitive structure than

is seen in any recent diving birds."

The femur was remarkably short and stout, being flattened

in the antero-posterior direction. The large, rounded head was

supported by a short, strong neck, and excavated above by a pit

for the ligamentum teres. Trochanter major was large, and the

entire proximal end of the bone possessed an articular surface

for the antitrochanter. As in Urinator, the shaft was curved,

and exhibited strong prominences for muscular insertion.

Expanded distally, the outer condyle was the larger of the two,

and only a shallow groove divided it from the inner one.

Tibiotarsus was a long, powerful bone, by all odds the big-

gest one in the skeleton of this bird. It much resembles the

same bone of the leg in any of our typical grebes. At its dis-

tal end, as in Podiceps, there is no osseous bridge in front span-

ning the longitudinal muscular groove below.

A very large free patella was developed, and it was pierced

by a foramen to transmit the tendon of the ambiens muscle, as

No. 433-] CERTAIN GROUPS OF BIRDS. 63

in Sula. Compressed transversely, it was distinctly triangular

upon lateral aspect. |

Marsh observed that the fibula of H. regalis agreed essen-

tially with that of Podiceps, and, as in that genus, the entire

skeleton of the limb is non-pneumatic. ‘In the adult Hes-

perornis, the second, third, and fourth metatarsals are thor-

oughly coóssified into a stout, transversely compressed bone of

moderate length, but in most specimens traces of the sutures

remain. The fourth metatarsal element so greatly exceeds the

other two in size, that it forms by far the greatest part of the

entire tarsometatarsal bone.

« The first metatarsal is not coóssified with the main shaft

of the tarsometatarsal bone, but is a mere remnant, united to

the lower half of the second by cartilage [ligament?].”

Different species of Hesperornis, as H. crassipes, H. gracilis,

and H. regalis, exhibited marked characteristic differences in

the various bones of their skeletons. But they were only of

specific value.

Of the skeleton of the foot in ZZ. regalis, Marsh wrote that

the feet of “ Hesperornis resembled more closely those of the

genus Podiceps than of any other birds. The number of

digits is the same, the number of phalanges in each digit iden-

tical, but the proportions of the latter are different and quite

peculiar. In Podiceps, and the other grebes, the outer toe is

the longest, but the middle one almost equals it in length and

size, while the second is but slightly smaller. In Hesperornis,

however, the fourth or outer toe is the dominant one, being

three or four times as powerful as the adjoining one, or as the

other three combined. Again, the phalanges in Podiceps are

very elongated and slender, and the terminal ones spatulate,

while, in Hesperornis, the phalanges are short and thick, with

the terminal ones more or less pointed. The phalanges in

Hesperornis are, in fact, shorter than in most swimming birds,

and in their individual proportions remind one of the toe bones

of the penguins " (Odontornithes, pp. 99 and 109). i

Fossil remains of many other birds have been discovered in

the cretaceous formation in different parts of America, and a

number of these have been described and named by Marsh

64 THE AMERICAN NATURALIST.

and others, but the few bones thus far in the hands of science

are too fragmentary to admit of saying to what manner of birds

they belonged, much less as to the affinities of the several

forms they represent. Others have also come to light in

Europe to which the same remarks apply.

As I have already pointed out in the Journal of Anatomy

(London, April, 1892, p. 202), I consider all the species of the

genus Hesperornis as having belonged to a family Hesperor-

nithidz, and this family may possibly have been an offshoot of a

superfamily, the Hesperornithoidea, which contained forms pos-

sessing the power of flight ; and from these latter our present

Pygopodes have descended, while the offshoot-genus Hespe-

rornis died out during the cretaceous time, and left no direct

descendants.

What I have said elsewhere in regard to the characters in

the skeletons of these ancient birds not possessed by their rep-

resentatives of the present age, applies also to Hesperornis.

For example, the structure of the palate, the extremities of the

pelvic bones being free, and so forth, are derived from their

reptile ancestors just as the ostriches derived theirs, and the

last named are now existing forms that have carried them down.

Family: ENALIORNITHIDA.

Of the remains of the fossil Enaliornis I know nothing

beyond what I have learned from reading. Lydekker, in the

article “Fossil Birds,” in Newton’s Dictionary of Birds

(p. 280), has said, “In 1858 Barrett discovered in the Upper

Greensand of Cambridgeshire remains described by Professor

Seeley in 1866 (Ann. and Mag. Nat. Hist., Ser. 3, Vol. XVIII,

p. 100) under the preoccupied name Pelagornis, but in 1867

renamed Enaliornis (‘Index to Aves and Reptilia, Camb.

Mus., Quart. Journ. Geol. Soc., Vol. XXXII, p. 509). These

indicate a bird apparently allied to Colymbus, and not improb-

ably to Hesperornis.”

Fiirbringer fully discusses what is known of the Enaliornith-

ide (pp. 1152, 1153) and is satisfied of the relation of Enali-

ornis to the extinct toothed loon, Hesperornis, as well as to the

various existing Pygopodes, and classifies it accordingly.

HATS FROM THE NOOTKA SOUND REGION.

CHARLES C. WILLOUGHBY.

In the early days of the New England whaling industry the

sailors brought back as mementoes many valuable ethnological

objects from the Pacific islands and the northwestern coast of

America. Much of this material found its way into the cabinets

of the older societies of Boston, Salem, and other New England

towns.

The Peabody Museum of Harvard University has acquired a

number of these old ethnological collections, either whole or in

part, including that of the American Antiquarian Society, the

Boston Athenzum, the Boston Marine Society, the Massachu-

setts Historical Society, and the Boston Museum. The few

objects in these collections from the northwest coast are of

great value, illustrating as some of them do phases of the arts

which have become extinct or much modified.

Among the objects received from these societies are eight

hats of the type illustrated upon Plate I, a style of head cover-

ing very rarely found in museums or private collections. It is

probable that this form of hat originated among the southern

Wakashan tribes, probably the Nootkas, although Lewis and

Clark found them on the lower Columbia in 1605 at Fort

Clatsop and thus described them (p. 768).!

* We gave a fish-hook also in exchange for one of their hats.

These hats are made of cedar-bark and bear grass interwoven

together in the form of a European hat with a small brim of

about two inches and a high crown widening upwards. They are

light, ornamented with various colors and figures. . . . These

hats form a small article of traffic with the whites, and their

manufacture is one of the best exertions of Indian industry."

And again on page 777, writing of the dress of the women:

“The only covering for their head is a hat made of bear grass

1 History of Lewis and Clark Expedition. Edited by Elliott Coues. Vol. ii.

66 THE AMERICAN NATURALIST. (VoL. XXXVII.

and the bark of cedar interwoven in a cone form with a knob

of the same shape at the top. It has no brim, but is held on

the head by a string passing under the chin and tied to a small

rim inside the hat. The colors are generally black and white

only, and these are made into squares, triangles, and sometimes

rude figures of canoes and seamen harpooning whales.”’

When Captain Cook visited Vancouver Island during his

famous voyage of 1776-1780, he found the same form of head

covering worn by the Indians of Nootka Sound, and on one

of the plates in the second volume of the octavo edition

(London, 1784) is a drawing of a woman with a hat of this

form. This is reproduced at å, Plate I.

Cook writes (p. 242) that the “natives wear a hat like a

truncated cone or a flower pot, made of very fine matting, orna-

mented with a round knob or a bunch of

leather tassels, having a string passing

under the chin to prevent it blowing off,”

and on page 266, * The whole process of

their whale-fishery has been represented

. on the caps they wear."

It is probable that this kind of head cover-

Fie. 1. — Detail of ing was prevalent from Nootka Sound to the

MM Columbia River, at least in the coast region.

The peculiar manner in which the cedar-bark strips and the

grass spires are manipulated (Fig. 1) to form the design is seen

in the modern basketry of the Skokomish, and is probably found

in other basketry of the Puget Sound region. It occurs in the

basketry of the Hooper valley and other northern California

Indians, as will be seen on examination of the basket caps and

old cooking bowls made from shredded pine roots and squaw

grass. This squaw grass of the Hupa and Shasta Indians

seems to be identical with the bear grass of Lewis and Clark.

The conventional representation of the canoe shown in the

whaling scenes upon Plate I is found upon the modern basketry

of the Makah Indian of Cape Flattery, Washington, the southern-

most of the Wakashan (Nootka) family.

The hats in the Peabody Museum are all of twined weaving,

and are made principally of cedar bark and grass spires. The

No. 433] HATS FROM NOOTKA SOUND REGION. 67

construction is double, as shown in the cross-section (Fig. 2).

Each headpiece consists really of two hats, an inner and an

outer one, joined at the rim, the last few pairs of twisted woof

elements of the outer hat enclosing also the ends of the warp

of the inner. The inner hat, or lining, is coarsely but neatly

woven of cedar bark, and only in one specimen (a, Plate I) is

there a knob at the top of the lining corresponding to that of

the outer hat. Upon the under side at about three inches from

the rim each warp element is doubled upon itself, forming a

loop about three-fourths of an inch long. Through these loops

is run a strong double cord of Indian hemp. The loops are

bound together by twined weaving, and form an inner rim

edged with the cord of hemp, which fits the head snugly. To

this is fastened the thong which passes

beneath the chin of the wearer.

The exterior or outer hat is woven

principally of grass spires and cedar

bark. In most of the specimens a nar-

row strip below the knob is made of

fine cedar roots. The warp appears to

be formed of split roots, and is fine and

strong. The grass of the woof was some irseeertan ure qo

originally an ivory white, the selected

cedar bark used in conjunction with it being usually stained a

dark brown or black.

Each strand of the twisted pair of woof elements forming

the design is composed of a grass spire and a strip of cedar

bark of the same width laid side by side, the strand thus

formed being white upon one side and black upon the other.

These double strands are used not only where figures appear,

but throughout the groundwork of the design as well. The

figures are principally black upon a white ground. In forming

them the strands are simply reversed, the black sides which

were concealed beneath the grass spires in the white back-

ground being carried outward, as shown in Fig. 1. In some of

the specimens the knob at the top is woven separately and

afterwards joined to the hat. In the hat illustrated at f, Plate I,

a small wooden hoop is placed within the knob to preserve its

68 THE AMERICAN NATURALIST.

shape. The materials are selected and prepared with great

care. The designs are spirited and well executed, and the

technique is of the highest order.

The principal design upon all but one of the hats represents

the chase of the orca or killer whale. It has been harpooned,

and the harpoon line with attached floats is trailing behind.

A man standing in the bow of a canoe is about to dispatch the

animal with a lance. Other canoes, apparently empty, fill out

the design. The Indians of Nootka Sound and vicinity, as well

as the tribes to the southward, hunted the orca, but the animal

was held sacred by the northern coast tribes and was never

hunted by them.

A very different design is shown upon the hat illustrated at

f, Plate I. The mythical bird is represented four times, twice

by itself near the brim and upon opposite sides, once hovering

just above an orca, and again with the whale in its talons. A

peculiar and characteristic feature is the life line, extending

from the beak to the heart, which is represented by a light spot.

There are two small, winglike projections back of this spot on

each of the birds, which remind one forcibly of the tufts of

feathers above the neck membrane of the pinnated grouse. A

similar mythical design is etched upon an Eskimo harpoon rest

carved in ivory, illustrated on Plate LXXII of the National

Museum Report for 1895. In this drawing the bird is twice

represented hovering over the whale, and twice with the whale

in its talons.

There are doubtless many valuable and rare ethnological

objects still in the families of the old whalers in the eastern

and middle states. These should be deposited for safe keeping

in museums of standing, where they would be preserved for all

time, and be accessible to students. Otherwise their destruc-

tion or loss is inevitable.

PEABODY MUSEUM OF

AMERICAN ARCH/EOLOGY AND ETHNOLOGY,

AMBRIDGE, Mass.

«'S98vÁo A S(007) ,, uro1j ‘punog txjooN Jo ueuroA *2 |” sv oures ‘9 ! UOIZar punog vxooN IP Wo SY Sa ‘p *p—*] 31v1q

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Morgan's ** Regeneration.’’ ! — Students of experimental morphol-

ogy have been done a real service by the author of this book. He has

brought together and classified in an orderly way practically all that

is known about the subject of regeneration in animals. A not incon-

siderable part of this knowledge has been contributed by Professor

Morgan himself or by his pupils. The author is hostile to *the

hypothesis of preformed nuclear germs" and to *the theory of

natural selection as applied to regeneration," but shows no spirit

of unfairness. His work is both careful and comprehensive. It

includes chapters on the early and justly famous experiments made

by Trembly, Bonnet, and Spallanzani; on the external factors of

regeneration ; on the internal factors of regeneration; on regenera-

tion in plants ; on regeneration and liability to injury ; on self-division,

budding, etc. ; on animal grafting; on relation to regeneration of the

* germ-layer theory” and “ the law of biogenesis”; on regeneration

in egg and embryo; theories of development ; theories of regenera-

tion; etc. It is a work both of merit and of permanent value, well

worthy of a place in the excellent series of biological books in which

it is published. WEC

Inheritance of Acquired Characters. — One of the most interesting

additions to the series Scientia is Costantin's ? “ L'hérédité acquise."

In a very brief space the author gives a readable account of Weis-

mann's germ plasm theory and of the changes that it has undergone.

Then follow chapters on heredity in asexual reproduction, on artificial

section, and on certain objections to the influence of the environment.

The very interesting subject of hereditary diseases is well treated, and

the little volume ends with a chapter on germinal selection. While

in no sense an original contribution to the subject, the book forms an

excellent introduction for the beginner or even the layman.

1 Morgan, T.H. Regeneration. Columbia University Biological Series, vol. vii.

New York, Macmillan, 1902. 8vo, xii + 316 pp» 66 figs. :

2 Costantin, J. L’hérédité acquise, Scientia, Biologie, No. 12. Paris, Carré

et Naud, 1902. 86 pp.

72 THE AMERICAN NATURALIST. | [Vor. XXXVII.

ZOOLOGY.

The Habits of Fishes.! — The author contends that: Physiologi-

cally, fresh water (and probably all) fishes fall into two groups, —

those which spawn in warming water and those which spawn in cool-

ing water, and the cause of spawning is the temperature trend in oze .

direction ; structurally similar forms tend strongly to sustain similar

relations to the temperature curve, Ze, to spawn either all on its

ascending, or all on its descending limb; in at least some cases

apparent exceptions can be harmonized with the law ; for a given

species the temperature relations which determine its migrations, and

probably also its geographical distribution, are the same as those

which determine its spawning. These facts demonstrate the presence

of a temperature-responsive nerve-mechanism, which is a character of

prime importance, entitled to at least superfamily rank ; its existence

explains wy with in-cooling spawning,? is (and must be) associated

to-cooler migration? and boreal distribution (and with in-warming

spawning, to-warmer migration and austral distribution); by a work-

ing backwards from the time of most successful hatching, the time

of spawning has been determined via natural selection; that time

so fixed, by a further working backward natural selection has deter-

mined the time of precedent migration ; there are, de Jacto, beach

spawners ; in type of egg the beach spawners agree with the fresh

water, and differ from the pelagic, forms, and this difference explains

why species of pelagic genera are so rare in fresh water, and beach

spawners are now uncommon, they having mostly become ana-

dromes; for having attained to a seek-the-beach impulse, the con-

ditions on the beach were such that, natural selection not opposing,

the beach spawners must, through the mere continued action of the

temperature-responsive mechanism, have been led, step by step, into

the forming streams of a rising continent; and in the streams the

necessary accessory instincts have been evolved, all in accordance

with accepted biological principles. "The most important generaliza-

tions are : Dynamically, fishes fall into two great groups according as

they are stimulated to migrate geographically, to migrate for spawn-

ing, and to spawn, by warming water, or by cooling water ; and this

dynamic factor necessarily involves a northern limit to the range of

1 Gurley, R. R. Amer. Journ. Psychol., vol. xiii (July, 1902), pp. 408-426.

2 In-cooling spawning, spawning in cooling water; /o-cooler migration, migra-

tion from warmer water to cooler.

No. 433.] NOTES AND LITERATURE. 73

the species of the first group, and a southern limit to that of the

species of the second group, the limit in each case being the point

where the spawning temperature disappears; and the factor which

has determined whether a given species was to remain a marine, or

to become a fresh water form, has been the egg type. W. C. K.

Development of the Face. — As a contribution to the study of the

external form of developing vertebrates, Rabl!has published eight

quarto lithographic plates illustrating the development of the face in

mammals. The species chosen are the rabbit, the pig, and the

human being; and of the first as many as seventeen stages are

shown. Each stage is illustrated as a rule by three views of

the head : full face, profile, and three-quarters. The material from

which the drawings were made was selected with as much care as

the circumstances would permit, and fixed in picro-sublimate or in

platinic chloride and sublimate. It was found advantageous for

surface views to stain the embryos in Grenacher's alcoholic borax

carmine. The surface configuration of such specimens has been

rendered in the figures with exquisite fineness of touch, reflecting

great credit alike on the draughtsman and the lithographer. The

figures are not marred by descriptive lettering, but a sheet of semi-

transparent paper is attached to each plate and bears the outlines

of the figures and the lettering. The text is a running description

of the material. Problems of external morphology are not dis-

cussed in it, though the author hopes that the work may afford a

basis for the study of the laws governing the development of external

form. Three more parts are to be issued covering presumably the

other groups of vertebrates, and the whole will constitute an indis-

pensable guide not only to the specialist in the development of the

face, but to embryologists in general Great credit is due to the

author and to the publisher for the production of so beautiful a piece

of work, and also to the Imperial Academy of Sciences in Vienna

for its generous support. P.

Pectoral Appendages of Birds. — The latest published part of

Fürbringer's? exhaustive studies on the comparative anatomy of the

lRabl.C. Die Entwicklung des Gesichtes. Heft 1, Das Gesicht der Sauge-

thiere, I. Leipzig, W. Englemann, 1902. vi+ 21 pp. 8 pls.

? Fürbringer, M. Zur vergleichenden Anatomie des Brustschulterapparates

(1902), pp. 289-736, Taf. X VIII-XXII.

T4 THE AMERICAN NATURALIST. | [Vor. XXXVII.

pectoral region of vertebrates contains an account of the bones,

nerves, and muscles of the breast, shoulder, and wings of birds. In

addition to the admirably clear descriptions of the anatomy of these

organs, this part is of special interest in that it contains a full dis-

cussion of the systematic relations of the orders and families of

birds as well as an account of the relations of birds to other verte-

brates. The diphyletic origin of the birds as represented by the

Ratit: and Carinate is regarded as an untenable assumption by

Fürbringer, who argues for the racial unity of the whole group.

From this standpoint the Ratite represent either a primitive stock

from which the Carinate have sprung or, as Fürbringer believes, a

degenerate assemblage derived from the Carinatzm. The author

further believes that birds have had their origin from none of the

known orders of reptiles, but from an ancient stock among the very

oldest reptiles and of which at present no remains are known. P,

Spiders of the United States.! — As the title indicates, this book

was written to meet a popular need. The first part, which is given

up to a short introduction on structure, collecting, and habits, might

very well have been enlarged. The remainder of the book is taken

up by a systematic account of spiders.

About two hundred of the common species of the eastern United

States are described. The descriptions are short, often very short.

The illustrations are excellent. Each species has one and often four

figures to show the characteristic parts and markings. The common

round web species that are found in similar places are described

together. The photographs of the webs are especially good, and

from them the whole process of web making can be followed.

The value of the book would have been increased for amateurs by

reference to a few of the popular books on spiders, and for more.

advanced readers by the authorities of the species.

BOTANY.

The Yuccez.? — At a time when systematic activity is preponder-

atingly segregative and the tendency to publish species separately

! Emerton, James H. Zhe Common Spiders of the United States. Boston,

Ginn & Co, 1902. 8vo, xviii + 225 pp., sor figs.

?Trelease, William. Thirteenth Annual Report of the Missouri Botanical

Garden, july, 1902. i

No. 433.] NOTES AND LITERATURE. 75

has become excessive, every piece of monographic work based upon

prolonged study of a particular group and passing mature and con-

nected judgment upon all its species is a welcome addition to

scientific literature. This is especially true of Professor Trelease’s

work on the Yuccez from the fact that it treats a group of natural

difficulty, in which, as in the case of the palms, cycads, Cactacez,

and other large and succulent plants, ordinary herbarium methods

are least effective and must to a great extent be supplemented by

the slower and far more difficult process of visiting the growing plants

in their native habitats or cultivating fresh material. The paper

under consideration is an octavo of 133 pages, copiously and excel-

lently illustrated by roo plates, the latter being chiefly halftones

from photographs. The work presents “the principal conclusions

reached in an intermittent herbarium, garden, and field study extend-

ing over the last sixteen years, in the course of which nearly all of

the spontaneous species have been examined and photographed in

their native homes."

The author divides the Yuccez into five genera. The genus Yucca

is confined to those species which possess globose or broadly cam-

panulate flowers with a thin polyphyllous perianth and a short thick

or obsolete style. From Yucca, which includes twenty-seven species

and may be regarded as the central and typical genus of the group,

Hesperaloe, with two species, is distinguished by its narrow perianth,

Hesperoyucca (monotypic) by its filiform style, Clistoyucca (mono- .

typic) by its thickened perianth, and Samuela, with two species, by

its gamophyllous perianth. Yucca is divided upon the nature of the

fruit and seed into three sections: Chanoyucca (the filamentosa

group), Heteroyucca (the gloriosa group), and Sarcoyucca (the bac-

cata group).

Under each species and variety exhaustive bibliography and syn-

onymy are given. These cover not merely the botanical treatment

of the plants concerned, but also the far more involved and vague

horticultural references, and the frequency with which the mark of

interrogation accompanies the citation of synonymy is certainly

significant. Here an energetic specialist, exceptionally situated for

the thorough investigation of his group and engaged in the revision

of not over thirty or forty species, finds himself, even after some six-

teen years’ effort at the elucidation of his group, obliged to _ ne

less than ten question marks in stating the synonymy of a single

Species. A few of these doubts refer, as might be supposed, to old

and vague characterizations published by the earlier authors, who

76 THE AMERICAN NATURALIST. [VoL. XXXVII.

did not realize the complexity of the genera with which they were

dealing; but the majority relate to the botanico-horticultural species

and varieties published by authors who are still living. The great

indefiniteness which surrounds plants of this sort may well raise the

question, whether a botanist who undertakes either voluntarily or at

the solicitation of some nurseryman to give a scientific name and

botanical description to some cultivated plant of which he knows

neither the country, habitat, natural origin, nor degree of permanence,

is doing more to advance or to block the progress of botanical classi-

fication. Happily this sort of work is relatively rare in America.

In Europe, however, some botanists of rather high standing seem

ever ready to undertake this, the lowest type of taxonomic activity,

especially in the already much abused groups of succulents. Cer-

tainly Yucca has had its share of such botanico-horticultural treat-

ment, and it is therefore a pleasure to see the genus subjected to a

critical and scientific revision based so largely upon study of the

plants in their natural state.

It isto be regretted that space was not found in Professor Tre-

lease's monograph for a more complete citation of herbarium speci-

mens, especially those of standard sets, for no feature of monographic

work gives more definiteness and permanent influence to a revision,

since by fixing uniform standards in reference collections throughout

the world it is of the greatest service in bringing about harmony in

. classification.

The paper closes with a résumé of the economic uses, phylogeny,

and ecology of the Yucceæ. A series of outline maps is also intro-

duced to show the distribution of the forms known to occur in

nature. B tfr

The International Catalogue.! — The first part of the long-

expected Royal Society's catalogue of current botanical literature,

though dated in May and with the MSS. completed in January

last, did not reach the libraries of this country until the middle of

August. Aside from general prefatory matter, this first part contains

the well-known classification of botany adopted by the Council, and

an alphabetical index to the same, in English, French, German, and

Italian, followed by a topographical classification in the same lan-

guages. The catalogue proper consists of an authors’ catalogue,

1 The International Catalogue of Scientific Literature, first annual issue.

M. Botany. Published for the International Council by the Royal Society of

London. Vol.i, pt.i. 1902

No. 433-] NOTES AND LITERATURE. 77

each entry marked with a classification symbol and ‘secondary refer-

ence marks. This authors’ list, occupying eighty-four double column

pages, is followed by a subject catalogue with alphabetical arrange-

ment of authors under each principal heading. In the systematic

part, each main group is provided with an index to the contained

new genera and species. The number closes with a list of the jour-

nals abstracted, with abbreviated titles. "

Notes. — The principal articles of the Botanical Gazette for August

are: Heald, * The Electrical Conductivity of Plant Juices”; McCal-

lum, “Nature of the Stimulus causing the Change of Form and

Structure in Proserpinaca palustris”; Schneider, “ Rhizobia mutabile

in Artificial Culture Media”; Nelson, * Notes on Certain Species

of Antennaria " ; Reed, “A Survey of the Huron River Valley”;

Ramaley, “The Trichome Structures of Erodium cicutarium” ; and

Copeland, ** Two Fern Monstrosities."

The Bulletin of the Torrey Botanical Club for July contains the follow-

ing articles: Torrey, * Cytological Changes accompanying the Secretion

of Diastase "; Banker, * Historical Review of the Proposed Genera of

the Hydnaceæ”; Shear, ‘Mycological Notes and New Species”;

Durand, *Studies in North American Discomycetes: II, Some New or

Noteworthy Species from Central and Western New York” ; Lamson-

Scribner and Merrill, * New or Noteworthy North American Grasses ”;

Eastwood, * New Species of Nemophila from the Pacific Coast"; Brit-

ton, E. G., * Trichomanes radicans " ; Berry, “ Liriodendron celakovskit.”

The Plant World tor July contains the following principal articles :

Parish, « Through Desert and Mountain in Southern California” ;

Idelette Carpenter, “The Protection of Native Plants ": Pollard,

“Plants used for Cuban Confectionery”; Hay, “ A New Station for

the Gray Polypody”; and eight pages of Mr. Pollard’s “ Families

of Flowering Plants, — from Hydrophyllacez to Solanacez."

The first part of Vol. LIV of the Proceedings of the Academy of

Natural Sciences of Philadelphia contains the following botanical

articles : Meehan, “Contributions to the Life-History of Plants,

No. XVI,” and Harshberger, “The Germination of the Seeds of

Carapa guianensis?” An article by Montgomery, “ On Phylogenetic

Classification,” though written principally from the zoülogical point

of view, deals with questions of the greatest interest to botanists.

Several small papers of taxonomic interest were issued under date

of August 6, as signatures of the Proceedings of the Biological Society

of Washington.

78 THE AMERICAN NATURALIST. [VoL. XXXVII.

Rhodora for July, in addition to a large number of notes of local

interest, contains an article by Dr. Robinson on the protection of

our native flora, descriptions of several new grasses, by Merrill, and

a protest by Fernald against the substitution of Washingtonia for

Osmorhiza, apropos of the discovery of Osmorhiza obtusa in Labrador.

The August number of Rodora contains the following articles:

Fernald, “ Taraxacum palustre in America" ; Davenport, ** Notes on

New England Ferns”; Sargent, * Additions to the Flora of Mas-

sachusetts" ; Williams, * Noteworthy Carices at Sudbury, Massa-

chusetts"; Shaw, “ Carex aurea in Connecticut" ; Norton, “ New

Stations for Peltranda and Conopholis " ; Cissbendn, * Aulacom-

nium heterostichum in Maine"; Lee, “ Aguilegia canadensis var.

flaviffera in Maine”; Rich, “Juncus torreyi and E//isia nyctelea in

Massachusetts.”

Torreya for August contains the following articles: Lloyd, * Vivip-

ary in Podocarpus"; Earle, * A Key to the North American Spe-

cies of Russula ” ; Howe, * A Note on the Vitality of the Spores of

Marsilea"; Abrams, *A New Hemizonia from California"; Brit-

ton, ** An Undescribed Species of pease sts M * Notes

on Verbena ” ; Shinn, * LZunu/aria cruciata in Fru

Recent stiches not likely to meet the eye of most botanists are:

Lemmon, “ Conifers of the Pacific Slope," in Sierra Club Bulletin,

Vol. IV, No. 2, and Lemmon, “ Oaks of Pacific Slope," in Z*azsac-

tions of Pacific States Floral Congress.

Vol. III, Part IV, of J. Medley Wood’s Natal Plants, concluding

that volume, has recently been issued from the press of Bennett &

Davis, of Durban. Each volume contains one hundred habit and

detail plates with appropriate text.

Fascicle 6 of Vol. I of Mr. Howell’s Flora of Northwest America,

comprising pages 563 to 666, completes Plantaginacea; and reaches

into Pontederiacez.

The results of studies on relations of plants to electricity made

at the Harvard botanic garden, are given in a paper by Plowman

in the American Journal of Science for August.

The effects of a tornado in the Victoria (Kamerun) botanical

garden are noted by Preuss in Der Tropenpflanzer for Jul

In Country Life in America for September, H. A. Doty has an

article on the Pollination of Asclepias, illustrated by a number of

low-power photo-micrographs.

No. 433.] NOTES AND LITERATURE. 79

Messrs. Gardiner and Hill discuss the histology of the endosperm

during germination in Tamus and Galium, in the Proceedings of the

Cambridge Philosophical Society of August 13.

A leaf section of Tecoma, showing the sunken nectar gland, is

published in the Gardeners’ Chronicle for July 19.

A useful directory of agricultural experiment stations in foreign

countries is published as Bulletin No. 772 of the Office of Experi-

ment Stations, U. S. Department of Agriculture.

The tropical growth of epipthytes, which proves detrimental to

the growth of cacao and lime trees, is discussed by Howard in the

West Indian Bulletin, Vol. III, No. 2.

An article by Fawcett on the banana industry in Jamaica is con-

tained in No. 2 of the current volume of the West Indian Bulletin,

which also has an article by Hart on the preparation of essential

oils in the West Indies, and an article by Freeman on the aloe

industry of Barbados.

Country Life in America is publishing a series of articles on the

making of a country home, which, like everything else published

in that journal, are beautifully and instructively illustrated.

The growth of the famous St. Michaels pineapple is described by

Bernegau in Der Zropenpffanzer for August.

Consul Ragsdale reports on Ginseng in China, in Advance Sheets

of Consular Reports of August 23.

An article on “Coffee ; Its History and Commerce,” by Marshall,

is contained in the American Journal of Pharmacy for August.

The World’s Work for September contains an interesting article

by Howard on breeding new kinds of corn.

A comparative anatomical study of water lilies, by Chifflot, is pub-

lished as fascicle ro of the Annales de P Université de Lyon.

As a reprint from the Annals of the Carnegie Museum, Mr. Ashe

issues lena of a number of new Pennsylvanian species of

Crate

The opening double number of Vol. XVI of the Annales des Sci-

ences Naturelles, Botanique, is entirely occupied with ovule studies of

Rosacez.,

A paper on the root tubercles of Medicago and other leguminous

Plants, by Professor Peirce, constitutes No. 10 of the current volume

of botanical Proceedings of the California Academy of Sciences.

So THE AMERICAN NATURALIST.

A Revision of the Japanese Umbelliferz, by Yabe, is contained

in Vol. XVI of the Journal of the College of Science of the Tokyo

University.

The first part of Abbé Léveillé’s Monograph of the Genus Ono-

thera, illustrated with numerous photograms from herbarium sheets,

and anatomical and other details, has been issued by the author at

Le Mans. ‘Two other fascicles are promised for the early part of

1903 and 1904, respectively.

The alders of Japan are revised by Matsumura in Vol. XVI of

the Journal of the College of Science of the Tokyo University.

Habit and bark photograms of Pinus inops are published by Pro-

fessor Rothreck in Forest Leaves for August.

The Fern Bulletin for July contains the following principal articles :

Anthony, “ Fern Hunting in Nassau”; Watkins, “Some Ferns of

the Sierra Nevada Range”; Eaton, “The Genus Equisetum in

North America”; Gilbert, * Georgia Ferns”; Clute, ** Botrychium

ternatum and obiguum" ; and a continuation of Clute’s list of

fernworts collected in Jamaica.

The genus Selaginella receives a considerable addition in new

species in a paper by Hieronymous, in Hedwigia for August.

. An extensive and largely illustrated paper on the mosses of

Alaska, by Cardot and Thériot, forms a brochure of Vol. IV of the

Proceedings of the Washington Academy of Sciences, issued July 31.

The paper is based on collections made by members of the Harri-

man Alaska Expedition in 1899, and is No. 29 of the “ Papers from

the Harriman Alaska Expedition.”

The coralline algz of Japan form the subject of a largely illus-

trated paper by Yendo, in Vol. XVI of the Journal of the College of

Science of the Tokyo University.

No. 3 of the University of Maine Studies consists of a preliminary

list of Maine fungi, by Ricker.

Bulletin No. 96 of the Kentucky Experiment Station is in part

devoted to poisonous and edible mushrooms, illustrated by a num-

ber of good photograms.

Torrendia, a gasteromycetous Amanitopsis, is described and fig-

ured by Bresadola in fascicle 2 of the current volume of A/t della

I. R. Accademia degli Aggiati in Rovereto.

CORRESPONDENCE,

To the Editor of the American Naturalist.

SIR : — By an unfortunate mistake in my “Lamarck, the Founder

of Evolution," it is stated on p. 56 that he died December 28, 1829,

whereas the exact date is December 18, 1829. Therefore, on p. 56,

line 3 from the bottom, for December 28, read December 18; and on

P. 57, line 5, for December 30, read December 20.

The following is a copy of the Acte de décès :

VILLE DE Paris— Année 1829. L’an mil huit cent vingt-

neuf, le dix-huit décembre, est décédé à Paris, au Jardin du Roi,

JEAN BAPTISTE PIERRE ANTOINE De MONET DE LAMARCK, membre

de l'Institut, âgé de quatre-vingt-cinq ans, veuf.

Le membre de la Commission,

Signé: DURANTON.

Also on p. 9, line 15, for “father” read * grandfather."

A. S. PACKARD.

PUBLICATIONS RECEIVED.

BALDWIN, J. M. Development and Evolution, including Psychological Evolu-

tion, Evolution by Orthoplasy, and the Theory of Genetic Modes. New York,

The aises Company, 1902. 8vo, xvi + 395 pp. $2.60.— JORDAN, D. S.

The Blood of the Nation: a Study of the Decay of Races through the Survival

of the Unfit. aati American Unitarian Association, 1902. 82 pp. — KEYSER,

LEANDER. Birds of the Rockies, with a Complete Check List of Colorado Birds.

Chicago, A. C. McClurg & Co., 1902. 8vo, ix + 355 pp.,8 pls. and text-figs. —

MELL, P. H. Biological Laboratory Methods. New York, The Macmillan Com-

pany, 1902. Svo, xiii + 321 pp., 123 figs. $1.60.— VERWORN, Max. Die Bio-

enhypothese. Eine kritisch-experimentelle Studie über die Memes in der

lebendigen Substanz. Jena, Fischer, 1902. 8vo, iv + 114 2.50 marks.

ALEXANDER, A. B. Notes on the Boats, Apparatus, an Fishing Methods

employed by the Natives of the South Sea Islands, and Results of Fishing Trials

by the vane Ae uA S. Fish Com. for rgor. Pp. 741-829, Pls. XXX-

XXXVII. Text-figs. — N, J. 4. Mammal Names proposed by Oken in his

Lahibodk 5 Zoologie. dm Amer. Mus. Nat. Hist. Vol. xvi, pp. 373-379. —

ARECHAVALETA, J. Contribución al conocimiento de la Flora Uruguaya: varias

especies nuevas y otras poco conocidas. Ann. Mus. Nac. Montevideo, 1902.

24 pp. 8 figs. — BANKS, N. A List of Spiders collected in Aisne by Messrs.

Schwarz and Barber during the Summer of 1091. Proc. U. S. Nat. Mus. Vol. xxv,

pp. 211-221, Pl. VII. — BARBER, C. M. Notes on Little-Known Mexican Mam-

mals and irc apparently not recorded from the Territory. Proc. Biol. Soc.,

Wash. Vol. xv, pp. 191-193. — BouLE, M. Machairodus Européens. Bull. Soc.

Geol., France, Sér. 4. Tomei, pp. 551-573. 17 figs. — BOULE, M. Les créatures

géantes d necu Rev. Gen. Sci., October, 1902. 46 pp., 30 figs. — BouLE, M

Les Tan de la France Coins Livre Guide VIII Cong. COMM Geol.

36 pp., 3 pls., 28 figs. — BouLE, M. Etude paléontologique et archéologique sur

la station p Meere du Lac Karár (Algerie). Z’ Anthropologie. Tome xi,

pp. 1-21, Pls. I-II. dm text- dem — Boure, M. Equidés fossiles. Ju. Soc

Geol., Kies Sér. 3. pp- 531-542. 22 figs. d: Hovis E, M. Labri

sous Roche du Rond > Saime- Arcon- Dalier o Loire). Z’Anthropologie.

hei x, pp. 385-396. 23 figs. — Bours, M. mpte-Rendu de l'excursion dans

le Massif Central. Compt. Rend. pois Cong. iu Geol. 5 pp.— Bouts, M.

La géologie et la paléontologie de Madagascar dans l'état actuel de nos connais-

sances. Compt. Rend. VIII Cong. Internat. Geol. 16 pp., 1 map. — CALKINS,

G. N. Marine Protozoa from Woods Hole. Bull. U. S. Fish Com. for rgor.

Pp. 413-468. 69 figs. — CLARK, H. L. Paper vin the Hopkins pepe Gala-

pagos Expedition, 1898-1899. XII. Echinoderm Proc. Wash. Acad. Sei.

Vol. iv, pp. 521-531. — DouGLass, EARL. Fossil M of the White River

Beds of Montana. 7yrans. Amer. Phil. Soc, N.S. l p. 237-279,

Pl. IX. — DuERDEN, J. E. Boring Algz as Agents in the Neg tu al of Cor-

als. Bull. Amer. dne Nat. Hist. Vol. xvi, pp. 323-332, Pl. XXXII. — DUERDEN,

. E. The Morphology of the Madreporaria. II. Increase of the Mesenteries

PUBLICATIONS RECEIVED. 83

in Madreporia beyond the Protocnemic Stage. Ann. Mag. Nat. Hist., Ser. 7.

Vol. x, pp. 96-115. 13 figs. — EIGENMANN, C. H. The Eyes of Rhinema Flori-

d Proc. Wash. Acad. Sci. Vol. iv, pp. 533-548, Pls. XXXII-XXXIV. —

GoETTE, A. Lehrbuch der Zoologie. suy Wein. 1902. Svo, xii +

504 pp. 512 figs. 12 marks.— HALL, W. L. CHRENK, H. von. The

Hardy Catalpa. Bull. U. S. Dept. Agr., pete. of pO No. 37. 58 pp.

30 pls. — Hav, W. P. Observations on the Crustacean Fauna of Nickajack Cave,

Tennessee and vicinity. Proc. U. S. Nat. Mus. Vol. xxv, pp. 417-439.

— Hovey, E. O. Martinique and St. Vincent: a Preliminary Report upon the Erup-

tions of 1902. Bull. Amer. Mus. Nat. Hist. Vol. xvi, pp. 333-372, Pls. XXXIII-

LI. — JouNsow, R. H. Axial Bifurcation in Snakes. 7Zrans. Wis. Acad. Sci.

Vol. xiii, pp. 523-538, Pls. XXXI-X XXVIII. —JoRDAN, D. S., and FowLER, H. W.

A Re view of the Ophidioid Fishes of Japan. Proc. U. S. Nat. Mus. Vol. xxv,

pp. —766. 5 figs. — JoRDAN, D. S., and Starks, E. C. A Review of the

Henibranchist Fishes of Japan. Proc. U. S. Nat. Mus. Vol. xxvi, pp. 57-73-

3 figs. — KELLOGG, V. L., and Kuwana, S. S. Papers from the Hopkins Stan-

ford Sie renia 1898-1899. X. Entomological Results: Mallophaga

from Birds. Proc. Wash. Acad. Sci. Vol. iv, pp. 457-499, Pls. XXVIII-XXXI.

— KisuiNoUvE, K. Some New Scyphomedusz of Japan. Journ. Coll. Sci. Imp. -

Univ. Tokyo. Vol. xvii. 17 pp. 2 pls. — Kusano, S. Studies on the Parasitism

of Buckleya Quadoriala, B. & H., a Santalaceous pum ite, and on a the Structure

of its Haustarium. Journ. Coll. Sci. Imp. Univ., Tokyo: Vol. xv

— McKenney, R. E. B. Observations on the Conditions of e OS

in Luminous Bacteria. Proc. Biol. Soc., Wash. Vol. xv, pp. 213-234. — MONACO,

PRINCE A. DE. Sur la troisième campagne de la Princesse Alice II. Compt.

Rend. Acad. Sci., Paris. Tome cxxxiv, pp. 961—964. — MosER, J. F. The Sal-

mon and Salmon Fisheries of Alaska. Report of the Alaskan Salmon Investiga-

tions of the United States Fish Commission Steamer A/éatross in 1900 and 1901.

Bull. U. S. Fish Com. for rgor. Pp. 173-401. 45 pls., maps. — OSB ORN, H. dr

LAMBE, L. M. Contributions to Canadian Paleontology. Vol. iii, pt. ii. On

Vertebrata of the Mid-Cretaceous of the Northwest Territory. por pcm

Canada. 4to, 81 pp, 20 pls, 24 text-figs. — RATHBUN, Mary J. Japanese

Stalk-Eyed Crustaceans. Proc. U. S. Nat. Mus. Vol. xxvi pp. 23-55. 24 figs.

— RATHBUN, Mary J. Descriptions of New Species of Hawai iian Crabs. Proc.

U. S. Nat. Mus. Vol. xxvi, pp.75-77- 4 figs. — RICHMOND, C. W. Birds col-

lected by Dr. W. L. Abbott and Mr. C. B. Kloss in the Andaman and Nicobar

Islands. Proc. U. S. Nat. Mus. Vol. xxv, pp. 287-314. — RICHMOND, C. V

Description of a New Subspecies of Stenopsis Cayennensis from Curaçao. Proc.

Biol. Soc., Wash. Vol. xv, pp. 159-160. — SNODGRASS, R. E. Papers from the

Hopkins Stanford Galapagos Expedition. VIII. Entomological equum ud

Schistocerca, Sphingonotus, and pte Proc. Wash. Acad. Set.

PP. 411-454, Pls. XXVI-XXVII. — STARKS, E. C. The Shoulder cide dj

Characteristic Osteology of the EA Fishes. Zro oc. v: S. Nat. Mus.

Vol. xxv, pp. 61 19-634. 6 figs. — VAUGHAN, An Addition to ite Mie

ij. Soc, Was

Fauna of the Aquia Eocene Formation of aa Proc. Bio

Vol. xv, pp. 205-256. — VAUGHAN, T. W. Redescription of £n Coral Platy-

trochus Speciosus. Proc. Biol. Soc., mE. Vol. xv, pp. 207-209. — WILDER,

H. H. Animal Classification. — WiLpER, H. H. A Synopsis of Animal Classi-

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VOL. XXXVII, NO. 434 FEBRUARY, 1903

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR. WIDEST SENSE

CONTENTS :

Page :

I. The Structure and Relationships of the American Pelycosauria . E.C.CASE 85 2

II. Notes on the Unionide and their Classification . - > - - V.STERKI 103 :

II. A Paraffine Bath heated by Electricity . - - . Professor E. L. MARK 115

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THE

AMERICAN NATURALIST

VoL. XXXVII. February, 1903. No. 434.

THE STRUCTURE AND RELATIONSHIPS OF THE

AMERICAN PELYCOSAURIA.

BE. C. CASE.

PERHAPS the greatest interest excited in the morphology of

the Permian reptiles of the order Pelycosauria centers about the

development of the temporal and quadrate regions as they seem

at present to afford the most direct phylogenetic evidence of the

origin of the mammals. The fragmentary character of the speci-

mens and the character of the matrix in which they are preserved

has rendered advance in the knowledge of the group very slow.

In previous papers, both alone and in collaboration with the late

Dr. Baur, the author has made known some of the structures

of the order, has demonstrated its Rhyncocephalian nature and

has given a new definition (Baur and Case '97 and '99). These

conclusions were based upon new material, but affected seriously

the work of Cope ; they were accepted by Cope for but one of

his forms only (Diopeus leptocephalus) with the proof that he had

already recognized the character in that form (Cope '97). The

unfortunate death of Prof. Cope rendered the type specimens

temporarily unavailable so that continued study was almost

impossible, but the collections have since come into the posses-

86 THE AMERICAN NATURALIST. Vor. XXXVII.

sion of the American Museum of Natural History in New York,

and through the kindness of the Director of the Department of

Paleontology the author was permitted to examine the types

during the past summer. As the material was made available

to me even before cataloging by the Museum, I desire to express

my thanks publicly to Dr. H. F. Osborn for his kindness in

the matter.

Unfortunately the most of the type specimens have never

been freed from the matrix, Prof. Cope contenting himself with

the description of new forms and such characters as could be

made out from the uncleaned specimens. A reéxamination of

the types in the light of better preserved and prepared material

has led me to conclusions very different from those of Cope and

of far-reaching effect upon the classification of the Pelycosauria

and the related Permian reptiles of other countries.

These conclusions may be stated as follows : —

1. All known reptiles from the American Permian, other than

the Cotylosauria, possessed two temporal arches; there is no

approach to a single zygomatic arch, as described by Cope in

some of them.

2. The Pelycosauria followed a line of development that led to

extinction while the persistent line of development was followed

in other regions, perhaps in Africa. The progress of develop-

ment, from a stage represented by Diopeus, is expressed in the

skull as follows : —

(a) The depression of the posterior angles of the skull (sus-

pensorial region) by the shortening and degeneration of the

quadrate ; the bones of the temporal arches remaining dis-

tinct but becoming very slender. This process leading to

extinction by the weakening of the articular region.

(b) The gradual elevation of the facial region of the skull

and the lessening of the transverse diameter through the

vertical growth of the maxillary and the harrowing of the

frontals and nasals.

tusks, the deepening of the diastema, the increasing con-

vexity of the alveolar edge and the development of serra-

tions on the edges of the teeth.

No. 434.] THE AMERICAN PELYCOSAURIA. 87

(d) The increase in size of the ectopterygoid process of the

pterygoid bone.

(e) Correlated with these changes the development of the

enormously elevated neural spines.

Following is a short description of the skulls of the types

representing the advancing stages of the development. The

genera Dimetrodon, Embolophorus, and Naosaurus are not

described, as the first two have been very fully described (Baur

and Case '99 and Case!) and Naosaurus is very similar to them

in the regions under discussion.

Theropleura uniformis. — Only the portion of the skull ante-

riorto the posterior edge of the orbits is preserved; but this

much is fairly free from the matrix, and the parts are easily

made out. A careful examination of the skull compels me to

differ from the description by Cope in one or two points. He

mentions (Cope '80) the presence of “at least one large incisor

tooth," and says the incisor teeth are separated from the maxil-

lary teeth by a diastema. The anterior end of the snout is

crushed, but the incisor teeth are seemingly all preserved, and

I could not. detect any one that was notably larger than the

others; they are small and sharply recurved. Neither could I

detect the presence of a diastema marked by any concavity of the

alveolar edge of the maxillary bone such as is so persistent in

the other American Pelycosauria ; it is possible that there was a

lack of teeth at this point, but as the bone is somewhat crushed

at the point it is impossible to say certainly. As mentioned by

Cope, there is no distinct maxillary canine, but the teeth increase

in size to near the middle of the series and then diminish so

that the 19 from the posterior end is the largest; the whole

tooth line and the alveolar edge of the maxilary is straight

without a trace of the convexity downward that marks the more

specialized form. As described by Cope, the skull is low and

acuminate ; the orbits are large and the nares are near the ante-

rior end of the snout on the sides. The whole skull is strikingly

Similar in general outline and appearance to the skull of Pro-

terosaurus, and in just the same characters of a low and broad

1A description of Embolophorus dollovianus, unfortunately delayed will appear

in the Journal of Geology for January and February, 1903-

88 THE AMERICAN NATURALIST. [VoL. XXXVII.

skull it differs from the specialized Pelycosauria. As the pos-

terior portion of the skull is destroyed it is impossible to say

with absolute certainty what was the condition of the temporal

arches, but from the other relations of the skull there seems no

reasonable doubt that they were as in Proterosaurus, z.e., with

two distinct arches.

Cope describes the neural spines as elevated in all the species of

Theropleura, but this is not to be taken in same sense as the ele-

vation of the spines in the specialized members of the Pelyco-

sauria. The length of the elevated spines does not exceed two

or three times the length of the centrum where in Dimetrodon

it is as much as 28 times the length of its centrum.

Diopeus leptocephalus. — The skull of this form is represented

by disarticulated fragments which are fortunately quite free from

crushing, and from the matrix. The fragment showing the tem-

poral region is made up of the quadrate and quadrato-jugal, the

united squamosal and prosquamosal; cemented to these are the

pterygoid and the epipterygoid almost in the natural position.

These fragments show that the temporal region, as figured by

Cope (Cope' 92), was very similar to the same region in Sphen-

odon which it especially resembles

in the united squamosal and pro-

squamosal. The anterior end of

the prosquamosal is missing, but

the squamosal is complete and

shows the articular surface for the

parietal. The posterior edges of

the temporal vacuities are pre-

served and show that they were

wide open and somewhat oval as

represented in figure 1.

a QE p pss, The resemblance to the quadrate

sl, diim dee region of Sphenodon is somewhat

poral; j, jugal; z, (quadrato-jugal) Superficial as the quadrate does not

seu Ae HP appear on the lateral surface but is

covered by the vertical extension of the quadrato-jugal. The

quadrato-jugal is much stronger than in Sphenodon and has

the form of a right angle with one arm vertical and the other

No. 434.] THE AMERICAN PELYCOSAURIA. 89

extending horizontally forward. The vertical arm overlaps the

outer surface of the quadrate as described and meets the squa-

mosal, the horizontal arm joins the jugal and forms with it a

strong inferior bar. Seen from the rear it is evident that the

squamosal comes down as far as the lower limit

of the quadrate and appears on the outer surface

of that bone. The pterygoid comes well up on

the inner surface so that the quadrate is prac-

tically surrounded by bones and has relatively

small vertical extent. Figure 2.

The facial portion of the skull is formed al-

most entirely by the maxillary which presents

some peculiarities. It is relatively long and the

alveolar edge is almost straight with hardly a

trace of the sharp downward convexity of the

specialized Pelycosauria. On the inner side of

the maxillary the lower edge is locally thickened

by the development of a sharp buttress which LTT ae.

accommodates the roots of the teeth; near the ph

anterior end of the maxillary there is a single greatly enlarged

canine and the buttress is correspondingly enlarged but no where

near so much as in the most specialized forms of Dimetrodon,

instead there is a strong rib developed on the lower side of the

maxillary from the base of the root of the tooth to the top of the

maxillary bone. The root does not extend into the bone as far

as in Dimetrodon. This maxillary canine is rather more pos-

terior in position than in Dimetrodon, posterior to it are 19

counted teeth and alveoli, but a portion of the bone is covered

by a fragment of the lower jaw so that there were probably from

23 to 25 teeth ; anterior to the canine are four teeth and alveoli

in the maxillary. Where the maxillary meets the premaxillary

there is a shallow depression of considerable antero-posterior

extent, but it is lined with teeth throughout so that it is but the

beginning of a diastema.

The form of the prem

trodon, there is a vertical a

anterior edge of the nares.

which is the first or inner one and poster

much smaller.

axillary is quite similar to that of Dime-

nterior process which forms the

There is but a single large incisor

ior to it there are four

go THE AMERICAN NATURALIST. | [Vor. XXXVII.

Another fragment shows the upper portion of the maxillary

and the frontal united by their inner surfaces. The two portions

of the maxillary show that it was a thin, elevated plate, but rela-

tively not nearly so high as in Dimetrodon. The frontal differs

markedly from the frontal of Dimetrodon and all the specialized

Pelycosauria, it is nearly as broad as long, showing that the top

of the head was flat and broad. The orbital region is not pre-

served, but the anterior portion of the preorbital is, and this with

the posterior end of the maxillary serves to locate its position

with fair accuracy. The front edge of the preorbital is marked

by the edge of a pit or foramen which is similar to the pits

described by Seeley in Cynognathus and regarded by him as har-

boring a gland. In figure 3 is shown a partial restoration of the

skull of Diopeus illustrating the beginnings of the features found

in the more specialized Pelycosauria.

1G. 3. Restoration of skull of Diopeus leptocephalus. x, nasal; mx, premaxillary ;

maxillary ; /; frontal; 4/, prefrontal; Z, lachrymal; 7, jugal; 7, Sec ad; pt, aal:

yn posiorbital; F parietal; sg, squamosal; 7sg, prosquamosal; gj, quadrato-jugal; g,

uadra

The pterygoid bone is also interesting as showing the simple

beginnings of the peculiar pterygoid characteristic of the Pely-

cosauria. There is the same tripartite division of the bone, but

in the Dimetrodonts the posterior limb extends only as far back

as the quadrate and is united with it by cartilage while in Diopeus

it articulates strongly with the quadrate. The middle line is

destroyed but the external or ectopterygoid process is much

more slender and thin than the same process in the Dimetro-

donts. The outer surface which abuts against the lower jaw is

slightly rugose and the lower edge carries 15 teeth and alveoli.

No. 434-]

THE AMERICAN PELYCOSAURIA. 9I

In Sphenodon there is the same ectopterygoid process but is

smaller and perhaps does not touch the lower jaw; it is made up

of the pterygoid and ectopterygoid but in the Pelycosauria there

is no distinct

ectopterygoid

that I have

ever seen, it

is either close-

ly co-ossified

with the pter-

ygoid or is be-

tween the an-

terior plate of

the pterygoid

and the max-

illary, a re

gion not pre-

served in any known specimen.

Fic. s. Edaphosaurus pogo-

nias. Side view of tem

sal; sg, prosquamosal ; 27,

quadrate; g, quadrato-jugal.

single mass and connected with a so-called zy.

4. Edaphosaurus pogonias. Side and top vi f kull. pmr,

premaxillary ; 772%} maxillary; /, frontal; 7, jugal; 766, postorbital ; £, pari-

etal; z, (quadrato-jugal) zygoma; fg, pterygoid; sz, (squamosal) supra-

temporal ; g, quadrate. After Cope.

| Seeley describes a similar proc-

ess in the African Theriodonts but says it

-is made.up of the ectopterygoid and pala-

tine.

Edaphosaurus pogontas. — This genus

differs so markedly from the other Pelyco-

sauria, especially in the presence of a clus-

ter of crushing teeth on the inner side of

the mandible, that it may have to be re-

moved from the order. There is one point

in the structure, however, that must be

mentioned. Cope describes and figures

this form (Cope '92) as having but a single

arch in the temporal region. In common

with most of the other types in the collec-

tion this specimen has not been cleaned,

and the very slightest removal of the matrix

showed that there were two temporal arches

instead of one. Instead of the lower por-

tion of the suspensorium being formed of a

gomatic bone, as

92 THE AMERICAN NATURALIST. [Vor. XXXVII.

shown by Cope, figure 4, there are several bones, the quadrato-

jugal is broken and crowded under the lower end of the suspen-

sorium, and that there was a distinct process is indicated by a

broken stump, which is undoubtedly the posterior end of the pro-

squamosal arch, figure 5. The position of this arch shows that

the upper vacuity was large and rounded and the lower narrow

and smaller, perhaps nearly closed, but there were two distinct

arches.

Clepsydrops natalis. — The posterior portion of the skull in

the type specimen is badly crushed, and the whole specimen is

covered with a thin layer of matrix, so that there is much uncer-

Fic. 6. Clepsydrops natalis. Side view of skull after Cope. ømx, premaxillary ; mx, maxil-

lary ; /, frontal ; of, postorbital; 0b, postorbital ; 2, parietal; sz, (squamosal) supratempo-

ral; 7, jugal; ? 4g, pterygoid.

tainty regarding the exact limits of each bone. The anterior

portion of the skull is strikingly similar to the same region in

Dimetrodon and Embolophorus in all the characters of extreme

specialization such as the high and narrow facial region, the

deep diastema and the enlarged canine and incisor teeth so that

there is strong presumptive evidence that the posterior arches

had an equal similarity. This is further borne out by the very

fact of the fractured condition of the posterior angles of the

skull, which are almost universally destroyed in the Pelycosauria

because of the weak arches. Certain elements of the temporal

region can be made out beneath the thin coating of matrix, but

these do not seem to me to be what Cope thought them. He

No. 434.] THE AMERICAN PELYCOSAURIA. 93

described and figured but a single arch (Cope '78, '92). The

portion marked jugal by him, figure 6, is crossed by a break that

seems to me to be very likely along the suture between the

quadrato-jugal and the jugal. Posterior to the orbit the bones

are badly broken and mixed, but certain parts seem capable of

a very different interpretation from that of Prof. Cope. The

small fragment labeled post-orbital in Cope's figure seems to me

to be a bit of bone from the base of the skull extending out of

the orbit. Posterior to the orbit Cope has figured but one piece

which he calls the supratemporal; this appears different to me

inasmuch as the anterior edge distinctly shows the rounded

excavation forming the posterior edge of the orbit and indicating

Fic. 7. Clepsydrops natalis. Side 1 fi

restoration, as it appears to the aut hor o pme,D maxillary; $ "E, maxillary; Pto, dera

id.

sq, prosquamosal ; 7, jugal; gf, pS aso 7 pt, pterygo

that it is the postorbital bone. Further back the mass bifurcates,

this would make the anterior leg of the bifurcation the lower

part of the postorbital, a consideration which is borne out by its

relation to the upper part of the jugal; the suture between the

two is hidden by the matrix and a fracture. The posterior leg

is exactly in the position of the anterior end of the postorbital-

prosquamosal arch if the arches had the same depressed form as

in Dimetrodon. These ideas are expressed in figure 7.

Further evidence that Clepsydrops had the same arch struc-

ture as in Dimetrodon is found in the greatly elevated neural

spines which are found only in the most specialized types. In

Clepsydrops they are from eight to nine times the length of the

centrum.

94 THE AMERICAN NATURALIST. (Vor. XXXVII.

For purposes of comparison the figure of the lateral aspect of

the skull of Dimetrodon is here introduced to show the most

specialized skull of the Pelycosauria. Figure 8.

The relation of the African forms to the American. — Just as the

American Pelycosauria passed through a series of specializations

which led to extinction so the related African forms (Therosuchia

or Therodontia) seem to have passed through a series of stages

which promised persistence and which if not leading directly to

the ancestry of the mammals at least illustrate by a beautiful

example of convergence the method by which the articular

region of the mammals might have been developed.

Fic. $. Dimetrodon incisivus. Side view of skull. mx, premaxillary; mx, maxillary ; 7,

nasal ; Af, prefrontal; f, frontal; Z, lachrymal ; £f, postfrontal ; fo, postorbital ; , parietal ;

J; jugal ; sg, squamosal ; 752, prosquamosal ; //, pterygoid; gj, quadrato-jugal; g, quadrate.

The most important steps in the development of the skull

structure from a form like Diopeus to such a form as Gomphog-

nathus or Tritylodon seem about as follows :

1. The degeneration to the quadrate and the development of

a definite posterior cranial wall by the extension of the parietals.

2. A tendency to the union of the bones of the postorbital

and temporal regions to form solid temporal arches.

3. The approximation and union of the two temporal arches

to form a zygoma.

4. The union of the posterior end of the zygoma with the

prosquamosal portion of the prosquamosal + squamosal and the

squamosal portion spreading out as a thin plate closely applied to

the skull wall almost at right angles to the zygoma.

5. The development of a mammalian palatal region and a

characteristic basipterygoid region from a type similar to Dime-

No. 434.] THE AMERICAN PELYCOSAURIA. 95

trodon by the loss of cartilage and the approximation of the

posterior portions of the pterygoids and the basioccipital and

basisphenoid bones.

6. The development of the bipartite occipital condyle.

Among the American forms there seems to be none which

approach the condition of a single zygomatic arch; among the

African forms none has been discovered which shows the two

temporal arches at all perfectly.! The nearest approach to the

Rhyncocephalian condition is in Procolophon and Cynognathus,

as the structure of these two clearly indicate the previous pos-

session of two arches we may assume the existence of a Diopeus-

like ancestor as indicated above.

I select as examples to illustrate the above conclusions Cynog-

nathus, Galesaurus and Gomphognathus because they are the

best preserved and described forms and because they are typical

of the three fairly distinct groups, the Cynodontia, Lycosauria

and Gomphodontia. All of these have the well developed occip-

ital crest and cranial wall, but show the advancing stages of

development in the arches and certain changes of importance in

the teeth.

In Cynognathus the postorbital and postfrontal have united

and developed posteriorly parallel to the jugal. The jugal bar

reaches far back and comes in contact with the inferior portion

of the united prosquamosal and squamosal (psq + sq) figure 9.

The quadrato-jugal is probably crowded beneath the lower end of

the psq + sq as indicated below. The union and strengthening

of the bones of the two arches is accompanied by an approxima-

tion of the two arches leaving only a small inferior temporal

vacuity between the two and by the suppression of the quadrate.

The psq + sq is described by Seeley (Seeley '95) as a thin plate

bent upon itself nearly at right angles, the point of the angle

extending outward and forming the posterior angle of the skull.

The inner half of this plate, squamosal portion, lies closely applied

to the cranial wall, and the outer, prosquamosal portion, forms

the posterior part of the upper half of the composite temporal

possessed the two temporal arches,

! In all probability Saurosternon, Huxley, ar

It may occupy a position

but it is primitive to the forms discussed in this paper.

similar to Theropleura among the American forms.

96 THE AMERICAN NATURALIST. [Vor. XXXVII.

bar. Seeley describes a vertical channel marking the outer side

of the angle of the psq + sq persisting downward to the point

. 9. Cynognathus crateronotus. Side view of skull. 7»&r, premaxillary; m2, maxillary ;

m rontal; gf, prefrontal; Z, lachrymal; 77e, postorbital; 4, parietal; sg, squamosal; 75g,

prosquamosal; 7, jugal. After Seeley.

where the bone expands to cover the quadrate, this groove prob-

ably marks the point of union of the squamosal and prosqua-

mosal.

The quadrate is figured by Seeley (figure 10) as a flattened

bone with two ascending bars be-

neath the psq + sq, the two bars

show through an evident breaking

' away of the thin plate of the cover-

ing bone. At the same time it is

noticeable that the outer part of

the lower edge of the psq + sq

comes in contact with the articular.

Seeley describes the first of the

vertical bars as “distinct from the

quadrate below and the squamosal

Fe xw Pus we d QUE above with which it is in close con-

region PEE s7 ps9, — tact"; the same fact is expressed

e pire € ‘ticular. Afta by a line on the quadrate separating

Seek. the lower part from the portion

bearing the vertical bars. It seems very ‘possible that this upper

portion is the lost quadrato-jugal which does not appear on the

side of the skull.

No. 434.] THE AMERICAN PELYCOSAURIA. 97

In Cynognathus the palate is typical reptilian, the teeth have

only small lateral tubercles and the whole dentition resembles

that of Dimetrodon in the large incisors and canine, the sharp

diastema and the convex tooth line.

In Galesaurus and in Cynognathus platyceps there is a

single temporal arch with no trace of an inferior temporal

vacuity. The bar is formed of two parallel bars, one above the

other, with no indication of distinct bones forming each bar.

The lower of these bars is called by Lydekker (Lydekker '90)

the quadrato-maxillary and the upper the squamoso-maxillary ; if

the condition of this form is derived from a condition represented

by Cynognathus crateronotus the lower bar is really the jugal,

the quadrato-jugal having disappeared beneath the psq + sq and

the upper bar is the prosquamosal + postorbital. As the prosqua-

mosal is already united with the squamosal, there is now a single

mass, the postorbital + prosquamosal + squamosal. Seeley points

out that in Galesaurus the lower portion of the psq + sq descends

below the quadrate posteriorly.

In Galesaurus the palate approaches the mammalian condition

found in the Gomphodontia and the teeth have strong lateral

tubercles. The diastema has almost entirely disappeared though

there is still a prominent canine.

In Gomphognathus the temporal region is much the same as

in Cynognathus, the region is described as follows by Seeley

(Seeley '95) *the squamosal rest upon the parietal as a thin film,

which descends laterally to the level of the occipital condyles,

when it makes a sudden angular bend upward and outward.

The lateral contour of the ascending bar of the squamosal

bone is convex, and nearly at right angles to that of the lateral

border of the occiput; both are equally thin, and the two plates

define a V-shaped squamosal notch which is open superiorly.

But the squamosal bone is much more massive than is at first

obvious, and below this thin external posterior plate, there is an

excavation in the bone, which shows it to be expanded laterally

for 1 ł inch beyond the compressed ridge at the base of the V-

shaped fold just described."

The squamosal described by Seeley is of course the psq + sq of

this article; the prosquamosal portion forms the posterior part of

98 THE AMERICAN NATURALIST. [Vor. XXXVII.

the zygoma and its anterior end is wedged into the posterior end

of the jugal. In front, the edge of the temporal vacuity is

formed by the jugal externally and the postfrontal internally.

The prosquamosal bar no longer lies parallel to the jugal bar;

the jugal bar no longer extends far back to articulate with the

prosquamosal near the suppressed quadrato-jugal, but the two

bars lie in the same plane and unite in the middle to form a true

mammalian zygoma.

In Gomphognathus the mammalian palate is fully developed

and the teeth are tuberculated and indicate probably an herbivo-

rous habit.

The significance of the development in the American and

African forms.—ln a previous paper the author has shown that

it is possible to recognize two lines of development in the

temporal region of the reptiles. -The forms showing the two

lines were described as the mastocephalous and saurocephalous

groups (Case '98) as follows: ‘From this point (the Progano-

saurian condition, Diopeus') onward the Reptilia are divided

into two groups, one with an elongate quadrate, which includes

all the modern and most of the extinct Reptilia, and one with a

depressed quadrate reaching its highest development in the

Permian, and in all probability losing its identity in the direct

ancestors of the mammalia.” It now seems possible to trace a

step farther the origin and history of the phyla.

The lower jaw is the movable element in mastication, and

in biting it acts as a lever of the third class with the power

supplied by the masseter and temporal muscles applied nearer to

the fulcrum, represented by the quadrate, than to the weight,

represented by the substance being chewed. It is evident that

where the jaws are used for such heavy work as chewing the

fulcrum region must be very strong to resist the strain imposed

upon it; in the forms which swallow their food without previ-

ous mastication, as the Pythonomorpha and Ophidia, and the

Amphibia in large measure, there would be little need of resist-

! [n the article quoted I stated that the generalized ancestor of the Pe/ycosau-

7ia had not yet been discovered, it now seems probable that that stage of evolu-

tion is represented by Diopeus, which will be seen from the description given

above to fill the necessary conditions.

No. 434.] THE AMERICAN PELYCOSAURIA. 99

ance in the quadrate region, and in all these forms it is notably

weak. The development of a chewing habit, either grinding or

sectorial, would bring a greatly increased strain upon this region.

We know little that is definite concerning the origin of the

temporal and quadrate region in the reptiles next above the

Cotylosauria, but as Baur has shown, this may well have

developed by a sort of natural trephining of the solid cranial

roof, perhaps in response to the demand for a lighter skull con-

nected with changing feeding habits, and resulting in the Pro-

ganosauria.!

In the Proganosaurian skull the quadrate was well developed

and had much the same position and proportions as in the

modern Sphenodon, but was of slightly less vertical extent. As

indicated by the character of the teeth these forms used the

jaws very little if at all for chewing. The beginning of a chew-

ing habit, either sectorial or the simpler kinds of crushing or

grinding, would demand a simple vertical motion of the jaws,

with a consequent demand on the quadrate to resist a stronger

thrust directed vertically upward. The shape and character of

the articular surfaces of the quadrate and articular bones in some

of the American forms, as in Dimetrodon, make any other motion

than a vertical one impossible.

The strengthening of the temporal and quadrate regions to

resist the increased vertical strain was accomplished in two

ways: First by an enlargement of the quadrate, especially in

the vertical direction, and its closer union with the skull wall ;

this was accompanied by a decrease in the number of bones in

the temporal arches and an enlargement of those remaining

(compare the condition in the Squamata and Testudinata).

Second by a reduction in the size of the quadrate till it finally

disappears (in the mammals or Promammalia) and at the same

time a union of the bones of the temporal arches till the articu-

! [n this connection it is of interest to note that the author discovered perfora-

tions in the roof of the skull of the Diadectidze in the Cope collection in the posi-

tion of the superior temporal vacuities, the forms are too specialized to make the

interpretation of this occurrence certain, but it may well be the first step toward

the Proganosauria type. In the line of the argument of this paper it p =

interesting to note that the Cotylosauria all had strong, crushing, or grinding

teeth and strong quadrate bones firmly enclosed in the roofing bones.

IOO THE AMERICAN NATURALIST. [VoL. XXXVII.

lation of the lower jaw finally comes at the base of a strong

buttress formed by the coóssified bones of the suspensorium.

The second of the two processes, described above, was

attempted in two different ways. The flattening of the quadrate

was common to both, but the supporting temporal arches were

disposed of very differently. In the American Pelycosaurians

the bones of the temporal arches all retained their identity and `

remained distinct one from the other, but the posterior ends of

the bones followed the quadrate down in its degeneration until

the angle of the skull was depressed and the arches were long

and slender; compare figures 3 and 8. This made a very weak

suspensorium especially as the opisthotic, the single bone which

connected the quadrate directly with the skull wall was separated

from it by the intervention of considerable cartilage. When the

jaw was compressed on food in the act of biting, the strain on

the quadrate region would be directly upward, but instead of

this strain being met by bones arranged to meet it parallel to

their length and firmest attachment, as in the modern lizards,

or against bones solidly united and joined to the brain wall, as in

the mammals, it was directed almost at right angles against the

free ends of slender bones at a point fartherest removed from

their attachment to the skull. Such an arrangement was clearly

unadapted to resist the strain imposed upon it by the force of the

fierce bites these animals were capable of inflicting. As shown

above, the advance in the slenderness of the quadrate region kept

pace with the advance in the development of the carnivorous

habits as evidenced by the increase in size of the incisor and

canine tusks and the serrations of the cutting edges of the

teeth; the latter speaking of a growing habit of sectorial masti-

cation. The two processes working in opposition.

In the second line of changes that centered around the

depressed quadrate there was a decided tendency to a coales-

cence of the bones of the temporal region ; thus the postfrontal

and the postorbital were frequently united, the squamosal and

prosquamosal the same and the quadrato-jugal either became

united with the posterior end of the jugal or, more probably

as shown below, disappeared beneath the lower end of the psq +

sq. After the union of the individual bones the two arches

No. 434.] THE AMERICAN PELYCOSAURIA. IOI

united, first partially (Cynognathus crateronotus and Procolophon)

and finally completely (C. platyceps and Galesaurus). Notable is

the position assumed by the squamosal portion of the psq + sq;

as shown above in the description of the forms it is spread out as

a thin film upon the surface of the cranial wall formed by the

developing of the parietals, so that it was firmly held by its

application to the skull wall as in the mammalian temporal and

is free only at its lower extremity. From this free extremity of

the squamosal portion the prosquamosal portion extends forward

almost at right angles, the quadrate and quadrato-jugal are

probably concealed beneath the point of union of the two and the

quadrate is so far overwhelmed that the lower end of the psq + sq

comes into articular compact with the articular.

This structure perhaps presents the best mechanism to resist

the strain imposed upon the quadrate in biting ; it is straight up

against the lower end of a bone fastened to the skull by nearly

its full length and its parallel to the strongest attachment. That

the forms in which it occurs had strongly developed biting habits

is evidenced by the tuberculated teeth and the herbivorous grind-

ing teeth of the Gomphodontia.

If the reasoning presented here is correct it is possible to

recognize two phyla among the Permian Pelycosauria; one char-

acterized by the persistence of the two Rhyncocephalian arches

and the development of a weak articular region, culminating in

the high-spined Pelycosauria and the other characterized by the

union of the arches and the development of a mammalian tem-

poral region culminating in Gomphognathus and Tritylodon,

perhaps in the Promammalia. The last branch practically

includes all of the Theriodontia — Theriosuchia.

102 THE AMERICAN NATURALIST. (VoL. XXXVII.

LIST OF REFERENCES.

BAUR, G., AND CASE, E. C.

°97. On the Morphology of the Skull of the Pelycosauria and the origin

of the mammals. Anat. Anzeig., XIII, pp. 109-120.

|799. The History of the Pelycosauria with a description of the genus

imetrodon, Cope. Trans. Am. Phil. Soc., (2), XX, pp. 1-62.

CASE, E. C.

'98. The significance of certain changes in the temporal — of primi-

tive Reptilia. Am. Nat., XXXII, p. 72.

CoPE, E. D.

'80. Second Contribution to the history of the Vertebrata of the Permian

formation of Texas. Proc. Am. Phil. Soc., XIX, p. 40.

'92. On the Homologies of the Posterior Cranial Arches in the Reptilia.

Trans. Am. Phil. Soc., XVII, pp. 12-26.

'97. Recent papers relating to vertebrate paleontology. Am. Nat.,

XXXI, pp. 315-323.

LYDEKKER, R.

'90. Catalogue of the fossil Reptilia and Amphibia in the British

Museum. Pt. IV. p. 69.

SEELEY, H. G

'95. Researches on the structure, organization and classification of the

fossil Reptilia. Pt. IX, Sec. 1. On the Therosuchia. Phil.

Trans. Roy. Soc. Vol. 185, pp. 987—1018, pl. 88.

'95. Researches *** Pt. IX, Sec. 4. On the Gomphodontia. Phil.

Trans. Roy. Soc. Vol. 186, p. 1-57, pls. 1-2.

'95. Researches * * * Pt. IX, Sec. 5. On the Skeleton in New Cyno-

dontia from the Karroo rocks. Phil. Trans. Roy. Soc. Vol. 186,

pp. 59-148.

STATE NORMAL SCHOOL, MILWAUKEE, Wts.

NOTES ON THE UNIONIDA AND THEIR

CLASSIFICATION.

V. STERKI.

Tur Unionidz are not only the most conspicuous inverte-

brates of our fresh waters, and wonderfully rich in species, but

they are also very interesting morphologically and physiologically.

The publication of Simpson’s synopsis of the Naiades, made a

new era in our knowledge of these animals. On the one hand,

Simpson has undertaken the important and difficult task of

revising the enormous number of described species by the care-

ful examination and comparison of an immense mass of material.

A large number of nominal species have been referred to those

which were believed to be valid, when reduced to synonyms and

varieties. On the other hand, he has shown that not only the

shells, but principally the soft parts, must be considered in order

to build up a proper classification and he has investigated and

sifted the attempts of earlier writers in that direction.

Having studied for years such of our Unionidz, or, more

properly, Unioninz, as were obtainable, their soft parts, propa-

gation, and embryos, I have formed some ideas of my own regard-

ing their classification, which proves to be somewhat at variance

with that given by Simpson. At first it seemed preferable to

wait, and study and compare further,’ until now, almost two

years later, I feel more satisfied than ever that these views have

a real foundation and some scientific value.

The Glochidia.—It has been found long ago, especially by

Isaac Lea, that there are considerable differences of shape in the

shells of the embryonic young of Unionida, but, so far as

known, no attempt has been made to apply this fact to classifica-

tion. The phases and changes of early and embryonal life are

more and more regarded as important in estimating the system-

' Many living specimens of different groups and species were kept under obser-

vation for shorter or longer periods.

103

104 THE AMERICAN NATURALIST. | [Vor. XXXVII.

atic position and phylogeny of all groups of animals, and must

be taken into account in studying the Unionidae. Since differ-

ences between embryos seem to be correlated with circumscribed

groups of the adult animals, they must be regarded as a valuable

factor in classification. The writer has examined embryos of

many species at different stages of development, not only in

regard to the shells but also to the soft parts showing decided

and interesting differences between the several groups and

species.! There are three main types of glochidia, so far as

known:

I. Those of Anodonta, Alasmidonta, etc. — Their valves are

large, rounded-triangular, with a long dorsal commissure, a strong

thickening along the margins, and are pointed in the middle of

the ventral side, each bearing a large, rough spine, or “hook "

(spur) bent inward so as to prevent the closing of the shell,

which remains wide open during the embryonal stage of life.

This formation is of decided importance, morphologically and

physiologically, for doubtless these strong spines facilitate the

attachment of the young mussels, after being discharged from

the branchiz of the parent, to the fins, etc., of fish, and possibly

to parts of other aquatic animals. It is to be regretted that so

little is known in regard to the early postembryonal life of the

young Naiades.

2. Those of Proptera, e. g., alata Say.— The dorsal commis-

sure is short, the dorso-ventral diameter being the largest; the

ventral margins are rather short, truncated or slightly curved ;

there is a smooth spine at the anterior and posterior ends (of the

ventral margins) of each valve, bent slightly outward and by

passing those of the opposite valves, allows the shell to close

along the ventral side. The long anterior and posterior sides are

truncate, nearly straight, and the shell is widely gaping at both

ends, a feature which is permanent, to a degree, in the adult

shell.

3. The glochidia of those species which were generally ranged

under the old genus Unio, with the exception of Proptera, have

a moderately long dorsal commissure, and are generally rounded

' It is intended to publish the results of these investigations when they are

more advanced.

No. 434.] NOTES ON THE UNIONIDA. 105

along the other margins, but otherwise of various shapes, the

dorso-ventral diameter (properly altitude) showing considerable

differences. There are no spines, the margins of the valves are

simple, or nearly so, and the shell can be entirely closed. There

are minor differences in the configuration of the shells and the

soft parts.

Lhe Hinge. — The species of the old genus Unio, with few

exceptions, have perfect ! hinges, and this seems to be a feature

characteristic for the whole group, and is of systematic impor-

tance.

In the species ranged under Anodonta, Alasmidonta, etc., on

the other hand, the hinges are generally more or less deficient,

or wholly wanting. Symphynota pressa Lea (Unio pressus) is a

real, or apparent, exception, it having rather perfect but feeble

hinge teeth. Other features of the shell, the soft parts and the

embryos, show relationships with Symphynota, such as S. costata

Raf. Natura non facit saltus.

In this connection should be mentioned the genus Margari-

tana, some species and varieties of which have perfect and others

imperfect hinges, in adult specimens.

The muscles, especially the large adductors, are generally of a

different color and texture in the Alasmidonta-Anodonta group

than in the “ Uniones.”

The Marsupia. — It is now generally known that there are

two different types of embryo-bearing branchiz, or parts of such.

In Anodonta, Alasmidonta, etc., and in a part of the Uniones (in

the older, wider sense) the ova and embryones are lodged only

in the outer, or all four branchiz, which, when not gravid, are

of the ordinary formation and appearance. In another group of

the Uniones, the marsupia are not only invariably in the outer

branchize, but also permanently differentiated, so that they

may be recognized even when not gravid. They are located in

certain parts of the branchiz, in most species, with app dug

mately fixed numbers of ovisacs for each species, or extending

over the outer branchie through their whole length, as in Ptycho-

' Only as regards the hinges of the Naiades and not in a general same the

hinge-type of the Cycladidz, for example, is more perfect, and more constantly so.

106 THE AMERICAN NATURALIST. (VoL: XXXVII,

branchus; but in the latter genus also they are permanently

differentiated in the female.

This fact has possibly not been given due weight, and Simpson

has been severely criticised for giving systematic values to physi-

ological features. This character, however, is distinctly an ana-

tomical one, and must be regarded as a factor in classification.

It is well known that in some of the Quadrula all four bran-

chiæ bear ova, or embryos, but it has not been proven that this

is so in all, especially in younger specimens, such as Q. pustulosa

Lea, etc. Neither is it known whether in some species of Unio

and Pleurobema the same condition may not be found occasion-

ally. Moreover, Simpson himself says that in Anodontoides

Jerussaciana all branchiz have been found bearing ova, and

Gysser makes the same statement in regard to Margaritana

margaritifera. This clearly shows that a distinction between

two groups cannot be founded on this feature, the more so,

since such closely allied forms as Unio, Pleurobema, and Quad-

rula are separated.

Differences of Male and Female Shells. — For some groups,

these differences have long been known, most of the genera

Lampsilis and Truncilla being familiar examples. The distension

of the female shell near the posterior-inferior end — as a rule —

is obviously the result of the demand for space for the voluminous

gravid marsupia near the posterior ends of the outer branchiz.

These differences are of various kinds and of very various degrees,

as especially among the Truncille, not only between different

groups but also between species of the same groups, and even

between different local forms of one and the same species, e. g.,

Lampsilis luteolus Lam. There are even species where we can

speak not only of a distension in the female, but of a different

formation of the entire shell, as in Plagiola securis Lea. In

other forms the differences may be very slight, as in Odov. ellipsis

Lea, although its marsupia are of the same type with those of

Lampsilis ventricosus Barnes. The members of the genus

Ptychobranchus though having voluminous, and highly differenti-

ated marsupia, show no outward differences of the shell, and some

females may be even less inflated than males of the same size,

as has often been observed by the writer. However, there is a

No. 434] NOTES ON THE UNIONIDE. 107

difference of another kind: a deep, oblique sulcus on the inside

of each valve, in the female, the space occupied by the marsu-

pium, so that the sexes can readily be distinguished on the empty

shells at least in older specimens.

It may be repeated here that in younger examples of Lamp-

silis, etc., the female shells cannot be distinguished from the

males during the first two or three years of life, and the disten-

sion is formed only from that age on, with the development of

the marsupia.

The female shell of Zyitogonia tuberculata Barnes, is less

inflated posteriorly than the male, and has a flat, thin extension

at the posterior end. Here the difference is of another kind

than in Lampsilis, a small portion of the branchia, if any, find-

ing room in the extended part of the shell, the genus evidently

ranging under another group : with Unio (s. str.), Quadrula, etc.,

without differentiated marsupia.!

In some species of Unio, Quadrula, etc. there are slight,

more or less marked differences between the male and female

shells, the latter being, generally, more inflated, and sometimes

differing in outline so that the sexes can be recognized with

some reliability. Yet these differences are not so typical and

so constant as in Lampsilis, Tritogonia, etc.

Species of another group, however, show well-marked differ-

ences, eg., Alasmidonta truncata H. B. Wright? and A. calceola

Lea. The fact was well known to the older conchologists, and

even the late Hy. Moores, three fourths blind, readily discerned

the sexes by a touch. The females are more inflated and more

voluminous posteriorly, curved down, and having generally a

stronger ridge. It is much the same with Strophitus edentulus

! So far as known, this species has not yet been seen in the gravid condition.

Its branchia, however, show no differentiated parts, and are Darren during fall

and winter, while the ovaries are charged with ova and the testes with sperm, as

in Quadrula, etc., which Tritogonia closely resembles in its soft parts and shell.

In dissecting a large female, I founda number of thin, grayish, beaded strands, of

various lengths, scattered in the outer branchiæ, in all probability rows of abor-

tive and degenerated ova. The strands were between the branchial fibrilla and

parallel with them. i

* The name is the one given in Simpson's Synopsis. Since then it has been

asserted by Fox that the Western form is the true 47. marginata of Say. (See

Nautilus, vol. xv, pp. 16, 47.)

108 THE AMERICAN NATURALIST. [VoL. XXXVII.

Say, and the sexes can generally be recognized by the shape of

the shell; and the same may be said of Symph. costata Rat.

(rugosa Barnes).

We have, then, among the Unioninz with differentiated mar-

supia, species in which the female shells are decidedly different

from the males, others in which they are slightly so, and still

others where differences are not recognizable. Those without

differentiated marsupia mostly show slight or no such differences,

while in Tritogonia, though of another type, it is well marked ;

and again there are marked differences between male and female

shells among the Alasmidonta, etc. It is evident from these

facts that much weight cannot be placed on this feature for

purpose of classification. If so, natural groups would have to be

divided, and discrepant forms united.

The Inner Branchia.— The upper edges of the inner branchize

are adherent to the abdominal sac either immediately, or by an

interposed soft membrane of varying width. In some genera,

or species, even in the same individual a part of the branchiz

may be of one, the balance of the other type. Simpson has paid

considerable attention to this feature, and it seems that there are

no constant differences available for classification.

Seasons of Propagation. — Since the publication of my obser-

vations several years ago, I have been able fully to confirm the

facts then stated. The forms with differentiated marsupia, as

Lampsilis, Ptychobranchus, etc., bear embryos during the fall and

winter, and discharge their young in spring and early summer,

rarely a part of them in autumn. The same was found to be

true with the Alasmidonta, Symphynota, etc. On the other

hand, the Unio (s. str., Quadrula, Tritogonia, etc. were invari-

ably found with the branchiz barren during fall and winter, their

ovaries filled with ova and the testes with sperm. This shows

that Lampsilis, etc., bear their young during a long period,

about eight or nine months. With Unio, etc., the charging of

the branchiz with ova, the development and discharging of the

young is all effected within a few summer months. We may

properly designate the latter as summer breeders, the former

as winter breeders.

This is a purely physiological feature, and, moreover, is proved

No. 434.] NOTES ON THE UNIONIDA. 109

only for the animals living in this latitude. But being coinci-

dent to a considerable degree with anatomical characters, it

certainly is significant, and seems to have some bearing on the

phylogenetic origin of these groups, as stated elsewhere. It

appears to point to different climatic conditions under which the

several groups developed and differentiated from their ancestors.

Reviewing the points discussed above, we find the principal

natural divisions as follows :

I. The forms with perfect hinges, typically and generally

Their embryonic shells are not pointed in the middle of the ven-

tral margin, and able to close.

2. The forms more or less deficient in, or lacking, hinges,

typically and generally. Their embryonic valves are subtrian-

gular, pointed on the ventral side, and each bearing a large spine.

Each of these main groups seems to contain the forms nearest

related. Any other arrangement would separate mr allied

forms and unite incongruous ones.

There may be one exception, however, as pointed out above,

and that is the genus Proptera. In the members of the genus

there are a number of features at variance with those of group

I, 2. e, the thin shells; the dorsal wings, anterior and posterior

to the beaks; the gaping in front and behind, probably the

umbonal sculpture; the slight, and in some species, even

deficient hinges. Adding to these the very different form of

the embryos, we have a collection of characteristics which place

these molluscs not only in a generic rank of equal standing with

that of Lampsilis plus its nearly related genera, but as a group

by itself. In my opinion they rather represent a third main

division between those of Lampsilis-Unio and Alasmidonta-

Anodonta. On the other hand they have one characteristic in

common with Lampsilis, etc., 7. e, the differentiated marsu-

pium, consisting of distinct ovisacs. Whether this feature

should be regarded as paramount in contrast with all the

others mentioned, may be questioned. For the present, how-

ever, I prefer to range them alongside of those which have

been regarded as their relatives.

Judging from certain features, there is some similarity between

Proptera and Pseudospatha Simpson, in regard to the shells, and

IIO THE AMERICAN NATURALIST. [VoL. XXXVII.

it will be interesting to compare their soft parts and embryos,

when obtainable.

If designations are wanted for the main divisions, they might

be called: (1.) Holodontes, (2.) Haplodontes.

The former again contains two main subdivisions, — one in

which the marsupia are differentiated, the other in which they

are not so. Among the first, Ptychobranchus represents a group

by itself, equivalent to the balance, owing to the exceptional

formation of its branchiz and ovisacs, and features of the shell.

A group represented by C. irrorata Lea, reflexa Raf. and

dromus Lea seems a natural one, although there are differences

in the number and size of the ovisacs, which might be regarded

as sufficient for generic distinction. And so it is with the bal-

ance, of which Lampsilis is the typical genus. The differences

between male and female shells in the several groups of Trun-

cilla are more considerable than between some of the Truncilla

and Lampsilis s. str., and so it is in regard to some other features.

Simpson ranges under the Lampsilis group also Cristaria

Schum. and Pilsbryoconcha Simpson, the hinge teeth of which

are defective or almost wanting, and the soft parts unknown, as

are also the embryos. With a knowledge of the latter, their

position will be more fully ascertained. In regard to Pseudo-

spatha we refer to what has been said above.

In the second subdivision of the Uniones, Tritogonia is

distinguished by its shell. Unio, Pleurobema and Quadrula,

constituting a very large and conspicuous part of our fresh-water

bivalves, and comprising various plastic and variable forms, are

so closely related and almost inseparably connected, that even the

highest authority is in doubt under which of these genera certain

species should be ranged. In regard to the branchiz, we refer

to what has been said on p. 108.

It may be in place here to mention that these mussels quite

generally are not able to open their shells as wide as the Lamp-

silis, etc., do. And this seems to be in close connection with the

fact that their posterior mantle openings, especially the branchial,

are much less fringed than those of Lampsilis, the fringes having

the function of rakers, preventing the entrance of coarse ma-

terials with the inward current of water, as has been seen by

No. 434.] NOTES ON THE UNIONIDA. III

actual observation, e. g., on L. ventricosus. Whenever any larger

object touches the margin of the mantle, that is the long and

partly branched fringes, they at once move inward and by inter-

lacing form a fine network. Some other species also, 'e. g, C.

irrorata, with the valves not opening wide, have the papilla only

moderately developed.

The systematic position of Margaritana is still uncertain. In

shape and appearance of the shell, and nacre, they resemble more

Unio, than Alasmidonta. The hinge is perfect in some and

defective in other species, in what seem to be individuals, typic-

ally perfect, and in younger specimens. It has been mentioned

that Gysser occasionally found all four branchie charged with

ova, and in all probability they are summer breeders, like Unio,

etc. That, however, must be ascertained by actual observation.

The crucial test will be found in the examination of the em-

bryos.' — I have also seen in some specimens of Quadrula kirt-

landiana Lea, very numerous, crowded, small muscle scars

scattered over the inner surfaces of the valves inside the pallial

line, described as characteristic for Margaritana.

The second main division consists of Alasmidonta, Anodonta

and their allies, and these two genera seem to represent the two

main groups. Among the former, S/rophitus edentulus Say,

shows a somewhat exceptional feature, which may entitle it to

generic rank. As regards the shell, soft parts and formation of

embryos, it stands very near to Alasmidonta, but the ova and

embryos are imbedded, in various numbers, about from ten to

twenty, in cylindrical albuminous masses placed transversally in

the outer gills. In my opinion, these masses, which have also

been compared with and called placentz, are not homologous

and equivalent with the ovisacs of Lampsilis, etc. The question

deserves more study and comparison.

Some other species and genera, e. g, Lastena lata Raf. need

more examination, in order to ascertain some of their charac-

teristics and their exact systematic positions.

The following table, based upon what is written above, will

' I should be glad to receive not only whole specimens, but gravid gills, or parts

ra Linn. and M. mondonta

of such, in alcohol, or even dried, of Marg. margaritife

ay. f.

II2 THE AMERICAN NATURALIST. Vor. XXXVII.

show more plainly a systematic arrangement which appears

natural and logical. It is not carried out to all the genera as

established and recognized by Simpson, but only to groups

showing characteristic, distinctive features. Again it may be

pointed out that Proptera, possibly with an additional genus,

might be regarded as representing a third (7. e., the second)

main division.

I. Hinge typically perfect; embryonic shell with the ventral margin

rounded or truncate, able to close.

A. Marsupia permanently differentiated in the outer branchiz (winter

breeders).

I. Shell without dorsal wings, not gaping at the anterior and

posterior ends ; embryonic shell without spines, with rounded

ventral margin, closing all around.

a. Marsupia occupying part of the branchiz.

aa. Marsupia near the posterior end of the branchiz,

bean- or kidney-shaped ; female shell usually dis-

tended near the posteriorinferior end; typical

Lampsilis

bb. Marsupia oeropriet do the whale margin of the

branchiz, or a part near the middle; shell short,

heavy ; female shell not -ucsiengs different from

e male; typical genus Cyprogenia

6. Marsupia occupying the whole: benc onii folds

when gravid ; shell elongated, not externally different in

males and females; . 1 . . Ptychobranchus

2. Shell with anterior and postedo dorsal wings (at least in

younger specimens), gaping in front and behind ; hinge teeth

feeble or defective; marsupium in posterior part of bran-

chiæ ; embryonic shells with spines at anterior and posterior

ends of truncate ventral margin, widely itid at the trun-

cate anterior and posterior ends: . . . Proptera

B. Marsupia not differentiated, the outer, or both pairs of branchiz

serving as brood pouches (summer breeders).

z. Female shell with a flat extension at the posterior end; shell

with crowded warts all o Tritogonia

2. Male and female shells Ph or not idiliesind: tipica genus:

Unio

JI. Hinge typically more or less defective or wanting; no differentiated

marsupia; embryonic valves rounded-triangular, pointed on the

ventral side, each with a large spine

A. Hinge more or less defective (rarely perfect, but feeble); beak

No. 434.]

JVOTES ON THE UNIONIDA. II3

sculpture usually consisting of simple, concentric ridges ; male and

female shells more or less different.

I. Ova and embryos imbedded in cylindrical albuminous masses

lying transversally in the outer branchie: . . Strophitus

2. Ova and embryos free in the branchiz ; typical genus:

Alasmidonta

Hinge teeth wanting; beak sculpture of undulating or two-looped

ridges, (animal, at least in some species, T typical

genos. <2 ERU. : . . Anodonta

CONTRIBUTIONS FROM THE ZOOLOGICAL LAB-

ORATORY OF THE MUSEUM OF COMPARA-

TIVE ZOÓLOGY AT HARVARD COLLEGE,

E. L. MARK, DIRECTOR. — No. 139.

A PARAFFINE BATH HEATED BY ELECTRICITY,

E. L. MARK.

Unper the stimulus of disastrous explosions of gas in other

museums and some unpleasant experiences in our own, it was

decided somewhat over a year ago to replace, as far as practica-

ble, heating by gas in the Museum laboratories with heating by

electricity. The greatest danger from the use of gas is incurred

where two or more lights are kept constantly burning in the

same room. All of our constant burners for heating water-

baths, warm ovens, etc., have been for many years supplied with

the Koch automatic cut off, so that in case of accidental extin-

guishing of the flame, the cooling down of the burner would

automatically shut off the gas supply. But the lever of the

Koch burner will not always work, even though loaded with a

weight greater than that which it carries when it comes from

the maker; moreover, the metal, on the expansion and contrac-

tion of which the tripping apparatus depends, after a time loses

to a certain extent its expansive properties, so that it fails to

release the lever and cut off the gas.

The Departments of Botany and of Zoólogy were already in

possession of a number of copper water-baths for heating paraf-

fine and warm ovens of various constructions, which it was

desirable to retain if they could be provided with a suitable

electric heating apparatus. With the aid of suggestions from

Professor Sabine of the Department of Physics and the coópera-

tion of constructors of electrical apparatus, a plan was finally

worked out which answers fairly well the requirements of the

ordinary paraffine water-baths.

115

116 THE AMERICAN NATURALIST. [Vor. XXXVII.

There were two chief problems to be solved: first, to ascer-

tain the minimum heating capacity of the heating coil necessary

for each bath; secondly, to devise an automatic regulator to

control the electric heating current. To reduce the amount of

heat lost, each bath was provided with a felt jacket,— covering

all parts except the top,— enclosed in a canvas cover. The

felt was about half an inch thick, and the canvas cover after

1G. 1.— Paraffine water-bath and automatic electric heating apparat The rso-ohm tele-

graphic relay sat is mounted on the top of box containing a pee battery. Plug

ceptacle

in place in the . At the right the = of another hearg apparatas removed

from oes Bo hung on a hook. Att eft a detached h

wire cable hanging on the wall. œR., Cites T., thermometer ; W., wires of heating

circuit; W., wires of relay circuit.

being slipped over the felt was laced up on one side, as is shown

above the faucet in Figure 1. The manufacturers of the heat-

ing apparatus employed (the Simplex Electric Co. of Boston and

Cambridge) then determined empirically the proper resistance

and length of coil required to maintain a given temperature in

each of the different forms and sizes of baths. To insure uni-

form distribution of heat, the coil should make at least one whole

No. 434.] PARAFFINE BATH. r17

turn. As constructed for our paraffine baths the coil is a some-

what flexible cylinder, about half an inch in diameter, with cop-

per covering. With its attached insulated wires such a coil is

shown hanging on the wall at the left in Figure 1. The resist-

ance metal used in the coil was a copper-nickel alloy. In the

case of the paraffine baths this coil was simply shoved into the

bath through an inch hole made in the top of the bath. The

knob with connecting wires protrudes outside, and the coil rests

on the floor of the bath, immersed, of course, in water. In the

case of the thermostats used for incubating purposes, etc., a hole

was cut in the side or bottom of the water reservoir, and the

coil after being introduced was soldered in place so as to close

the hole.

To devise a regulator was more difficult. The use of an ordi-

nary rheostat proved to be impracticable, because it was not possi-

ble to make sufficiently fine gradations of resistance for different

temperatures, nor by it to provide against fluctuations in the

initial current.

The method finally adopted utilizes a one-cell relay battery to

magnetize an electro-magnet. The heating current is made to

pass through the armature, which is pulled into contact with a

vertical post by means of a delicate spring. When the armature

is drawn away from the post by the magnet the heating current

is broken. Into the circuit of the relay battery is put the regula-

tor; when the relay circuit is closed at the regulator, the mag-

net operates on the armature and breaks the heating current ;

when the relay circuit is broken at the regulator, the armature

is drawn back by the spring to its first position thus closing the

heating circuit.

Our Reichert gas regulators (2), already in use for the pur-

pose of regulating the gas supply to the burners, were adapted

by very slight changes to the new requirements. The tubular

glass stopper carrying the gas inlet was removed; in its place

was put a cork, bored to receive one of the copper wires ( W’)

from the single-cell battery of the Leclanche type (Samson cell

No. 2). The copper wire terminates with a No. 20 platinum

wire (Fig. 3, P.) which is about an inch long; the height of the

lower end of this wire above the mercury in the column can be

118 THE AMERICAN NATURALIST. [VoL. XXXVII.

roughly adjusted by moving the copper wire through the cork

and wedging it in place by a small wooden wedge occupying

with the wire the hole in the cork. The contact of the mercury

with the platinum wire serves to close the circuit of the battery,

the other wire from the battery being connected to the mercury

by means of the screw and piston in the side tube used to adjust

the height of the mercury column. Into this single-cell circuit

is put a standard 150-ohm telegraphic relay machine of the pat-

tern used by the Western Union Telegraph Co.

——

EEA

—————

—

z SA

E EEA

E———

| —————

Fig.3

Fic. 2. — Diagram to show the connections of heating coil and circuit with relay battery, electro-

magnet and mercurial regulator. A., armature; B., battery; C., heating coil; M., electro-

magnet; P., post in the heating circuit; R., regulator; S., spring; W., wire of the heat-

ing circuit; W 7., wire of the relay circuit.

TT vb wd A | 1 Reich

Fic. 3

vt? 1

5 E

-— Upper p g modified — drawn to larger

scale than in Figure 2, P*., platinum wire.

The closure of this single-cell circuit magnetizes the core of

the electro-magnet (77.), which, pulling against the delicate spring

(S.), overcomes it and moves the armature (4.) away from the

post (P.), thereby breaking the heating current.

When, owing to the interruption of the heating current, the

bath cools, the mercury in the regulator recedes from the plati-

num point, thus breaking the relay circuit, the electro-magnet

becomes demagnetized, and the delicate spring pulls the armature

into contact with the post and thus closes the eating current.

The finer adjustment of the distance between platinum point and

No. 434] PARAFFINE BATH. 119

mercury in the regulator is effected, as in gas regulation, by the

screw and piston working on the mercury in the horizontal arm

of the regulator.

The points on armature and post at which the heating current

(110 volts alternating) is made or broken require to be made of

heavy platinum wire (No. 15), for otherwise there is danger that

the metal will fuse.

The paraffine baths of the form shown in Figure 1 are about

19 inches in diameter and 5 inches deep. The current required

to heat such a bath is approximately equal to that of four 16-

candle-power lamps. This could be much reduced by enclosing

the bath in a glazed frame, one side of which would have to be

movable to permit access to the cups.

The whole apparatus, except the heating coil, was furnished

and installed by Clark & Mills, 23 Church St., Cambridge, and

543 Boylston St., Boston. It cost, including the heating coils,

between $25.00 and $30.00 for each bath.

In determining the resistance to be used in the heating coil,

one should make it as low as possible consistent with the maxi-

mum heating capacity required ; for with greater heating capacity

the heating current will be in operation a shorter time than with

less heating capacity, and consequently the 7z/ay current —

which is in operation whenever the heating current is not — will

be required to work longer, and therefore the battery will become

exhausted sooner. When the battery is so far exhausted that it

will no longer magnetize the core sufficiently to overcome the

spring and break the current, the regulation fails and too high a

temperature results. However, a single cell will, with proper

care, last for several months without renewal. To guard against

the danger of too weak a relay current, one should test the cur-

rent from time to time with a voltmeter.

There is one difficulty with this mercurial regulator, due to

the oxidation of the mercury at the time of making and breaking

the relay current. This in time causes a deposit on the platinum

wire of oxide, which acts as an insulator and thus prevents sharp

contact. The deposit may be removed, however, by immersing

the platinum wire from time to time in weak nitric acid, and if a

layer of high-test kerosene oil covers the mercury in the regu-

lator the oxidation may be prevented.

NOTES AND LITERATURE.

ZOOLOGY.

Development of Cribrella. — The early development of the star-

fish Cribrella has been made the subject of exhaustive study by Dr.

A. T. Masterman. Segmentation, which is very variable, always

culminates in a morula of equal cells. This becomes a blastula,

after which gastrulation takes place. The blastopore closes and the

archenteron divides into a mensenteron, and an anterior ccelom, and

a posterior caelom. The embryo then escapes as a free ciliated larva.

The posterior caelom becomes the hypogastric ccelom of the post-larval

stages. The anterior ccelom becomes differentiated into a central

cœlom, and a right, and a left lateral coelom. The central ccelom

becomes the “dorsal sac”; the right lateral becomes the epigastric

ceelom ; and the left lateral the hydroccele which eventually gives off

five radii. The bilateral larva loses its exact symmetry through an

enlargement of its left side. Fixation takes place and the unsym-

metrical larva is converted into a young starfish. The larval sagittal

plane corresponds to the discal plane of the starfish, the left side of

the larva giving rise to the oral, the right to the aboral side of the

adult. 'The bilateral stage suggests a bilateral ancestor similar to

Balanoglossus.

Studies of Recent Brachiopoda. — Seldom has a single year

brought so many valuable additions to our knowledge of recent

Brachiopoda as are represented by the five papers that form the sub-

ject of this review. In fact, the only period that at all compares with

it is the year 1873, which saw the pioneer embryological work of

Morse and Kovalevsky.

The first of these papers, by Dr. Conklin,’ has to do with the same

species * Terebratulina septentrionalis! that afforded Prof. Morse

' Masterman. A. T. The Early Development of Cribrella oculata (Forbes)

with Remarks on Echinoderm PASER Transactions Royal Society of Edin-

burgh, vol. xl, pp. nyan Pls. 1902.

* Conklin, E. J. The Sea of a Brachiopod, Terebratulina septen-

trionalis, Coiithouy: db the Zoological Department of the University of

Pennsylvania. Proc. Amer. Philos. Soc., vol. xli, No. 168, pp. 41-76, pls. I-X,

1902.

122 THE AMERICAN NATURALIST. [Vor. XXXVII.

material for the first investigation ever undertaken of the entire devel-

opment of a brachiopod, and is a very careful reinvestigation of the

early stages of this species by the aid of modern methods of prepara-

tion and microscopical technique. The early cleavage forms are

found to present considerable variation, commonly in the produc-

tion of very unequal divisions. These forms do not especially

resemble those of the molluscs and annelids, but are more like those

of the Bryozoa and Phoronis. Gastrulation is by invagination. The

suggestion is offered that certain deep-staining granules seen at

the free ends of the gastrula cells are associated with the cilia of the

embryo. The archenteron at first possesses one constriction instead

of two as in Cistella. The proper orientation of the embryo shows

that it belongs to the hypogastric type, an important difference from

the Chatognatha with which the Brachiopoda are sometimes sup-

posed to present close relationship. The larva shows three regions,—

cephalic, mantle, and peduncular, which, however, are not true seg-

ments. The mantle at first covers the peduncular region and is sub-

sequently folded up over the cephalic portion. Dr. Conklin compares

the embryo of Terebratulina with the trochophore and with the

embryos of Phoronis and the Bryozoa, and concludes that the two

latter groups and the Brachiopoda should go together in a single

phylum, and that their relations with the Chetopoda and Cheetog-

natha are not especially close.

* Observations on living Brachiopoda" by the veteran student of this

class, Prof. E. S. Morse, is a timely contribution to a much neglected

subject. Occasional references to the habits of brachiopods may be

found in various works, as those of Davidson, CEhlert, Woodward,

etc., but Prof. Morse's paper contains by far the most extensive and

valuable series of observations on the subject yet published. The

title, however, does not convey an adequate idea of the extent of the

work, for the author has given a large amount of anatomical detail,

especially in regard to the mesenteries and perivisceral bands, sense

organs, “hearts of Hancock," accessory hearts, etc., which is not

confined to observations on the living object. The species chiefly con-

sidered are: Glottidia pyramidata, Lingula lepidula, and L. anatina,

though in addition observations have also been made upon Discinisca

lamellosa, and D. stella, Terebratulina septentrionalis, Terebratalia

coreanica, T. minor, Laqueus rubellus, Hemithyris psittacea, H. albida,

and Dallina grayi. The tenacity of life in brachiopods, especially

1 Morse, E. S. Observations on living Brachiopoda. Mem. Boston. Soc. Nat.

Hist., vol. 5, No. 8, pp. 313-386, pls. 39-61, July, 1902.

No. 434.] NOTES AND LITERATURE. 123

in the Inarticulata, has often been a subject of remark. Prof. Morse

succeeded in transporting live specimens of Lingula from Japan to

this country. “One cannot help,” he says, “associating this remark-

able vitality of these genera with their persistence through geological

horizons from the Cambrian to the present day almost unchanged in

character.” The freedom and extent of movement of the valves of

Lingula, in locomotion and in burrowing, are quite at variance with

Hancock’s notion of the operation of the muscles, and much more in

accord with what one would be led to expect from the general con-

formation of the valves. The action of the seta of the anterior

mantle margins of Lingula, in forming tubes for the passage of incur-

rent and excurrent water, is pointed out for the first time, though

perhaps slightly anticipated by an observation by Yatsu. Another

fact of much interest is the relation between the length of the sete

and the mobility of the valves upon the pedicle. When there is great

mobility the setze are long, and vice versa. Attention is called to the

manner in which the ridges (stria) of the shell of Terebratulina

coincide with the sete, though it is doubtful if there was any such

connection as Morse suggests between the setae and spines of such fos-

sil brachiopods as Productus. The movements of the brachia are

described as most graceful and varied. The two parts of the brachia

are always moved simultaneously and symmetrically. The “heart

of Hancock” is shown to exhibit none of the properties of a propel-

lant organ, and is probably connected in some way with reproduction.

This paper is profusely illustrated by outline drawings and one

plate of colored drawings, which will be a revelation to those stu-

dents who have never had the opportunity of viewing living brachio-

pods.

Mr. Naohidé Yatsu deal with Japanese

The three papers by pane

Lingulas. The development of Lingula anatina! is a contribution

of the highest importance to our knowledge of brachiopod embry-

ology. The earliest stages of development of this type are here

detailed for the first time. Even the discharge of the sexual elements

is noted. Females with ripe eggs will not discharge them if isolated

from the males, so that probably the discharge of the sperm acts as a

stimulus to the female to discharge her eggs. The breeding season

lasts about one and one half months in Japan — from the middle of

July to the end of August. The gastrula is formed by invagination.

tina. Journal College

! Yatsu, Naohidé. On the Development of Lingula ana

art. 4, pp. 1-112, pls.

of Science, Imperial University of Tokyo, Japan, vol. xvii,

I-VIII, 1902.

124 THE AMERICAN NATURALIST. [Vor. XXXVII

The whole development of Lingula is preéminently direct, the mantle

lobes growing over the cephalic region a initio. There is conse-

quently no inversion of the mantle lobes as in Cistella, Thecidium, etc.

'The embryo is also two lobed instead of three, as in the latter

genera, the caudal lobe being absent in Lingula. “The shell is

formed at first as a circular lamella folded double along one of its

diameters, and is secondarily divided into two valves along the poste-

rioredge." This is in marked contrast with the method of shell origin

that obtains among the Articulata. In conclusion, attention is called

to the resemblance between the mode of cleavage in Lingula up to

the 32-celled stage and that of some species of Phylactolaemata.

This is in harmony with the observations of Dr. Conklin above

referred to, as is also the fact that in Lingula there is no true seg-

mentation of the embryo.

Mr. Yatsu's paper on the Histology of Lingula! gives a detailed

account of the several kinds of bodies found in the ccelomic fluid,

namely, the blood corpuscles, leucocytes, and spindle bodies. An

extended account is given of the latter — their form, occurrence,

development, and significance. They are shown to be metamor-

phosed blood corpuscles, “a cel whose nucleus has degenerated and

whose cytoplasm has turned into a pores structure.” Their function

is excreto

A ibi | paper by the same author? deals with the habits of

Lingula, and is extremely interesting. These forms live in mud flats

which are exposed at low water, and though ordinarily no trace of

them can be seen, their presence is sometimes detected by three

small holes in the mud. These holes must be evidently produced by

thé setal tubes mentioned by Morse. The cirri of the brachia,

though not the brachia themselves, can be protruded from the front

of the shell. The life of Lingula may be as long as five years.

Yatsu mentions the extreme tenacity of life in this genus. In one

instance an influx of sediment that proved fatal to all the lamelli-

branchs of the locality had no effect upon the Lingulas. The plates

which accompany these three papers by Mr. Yatsu certainly testify

to his own and the lithographer's skill. They are models of clear-

ness and beauty.

! su, Naohidé. Notes on the Histology of Lingula anatina Bruguiére.

ps College of irse Imperial University of Tokyo, Japan, vol. xvii, art.

5+ PP- 1-29, € :

? Yatsu, Nao ohidé. On the Habits of a Lingula. Annotationes Zoo-

logice Japanensis, vol. iv, pl. 2, pp. 61-67, 1

No. 434.] NOTES AND LITERATURE 125

Salmon and Trout.'— This is one of the most desirable of recent

publications on angling. The three sections of which it is composed

are unequal in quantity and to some extent in quality yet they effec-

tually cover their field. The first section, of 149 pages, by Dean

Sage, on the Atlantic Salmon, is a most excellent piece of work, how-

ever regarded. ‘The dearest interests of the angler, habits, localities,

tackle, capture, and the like, are admirably and thoroughly treated.

Discussing but a single species, future changes of names and position

can have little effect on this essay. Literary merit, accuracy, and

inclusiveness combine to establish it as one of the most permanent

contributions on the subject.

The second section, of about 40 pages, on “ The Pacific Salmons,”

by Messrs. Townsend and Smith, is filled with information about

species not as well known as Salmo salar and which apparently do

not lend themselves as readily to the purposes of the sportsman.

The third section occupies more than 200 pages and, treating of

“The Trouts of America,” it deals with a considerable number of

species. The author, Wm. C. Harris, is one of the first of living

authorities on his topics and, so far as the matters of most impor-

tance to anglers are concerned, there is probably no one more com-

petent. A veteran and an enthusiast he has the experiences of

many years from which to draw material that is always full of life

and entertainment. On whatever most directly pertains to angling

and tackle or its manufacture his work has its greatest value. The

technical science, in which he appears to take some pride, introduces

elements that make for less of permanence. For the classification

and nomenclature, and in great part knowledge of distribution, are

only approximations, unsatisfactory and more or less discredited by

the authorities, liable to be modified or superseded in the near

future.

Opportunities for improvement on revision are not lacking. T he

following from page*194 will serve as an instance: “In 1486, six

years before the discovery of America, Wynken De Worde, among the

first of English printers, published that famous work, ‘ The Booke of

St. Albans’ on ‘ the dyssporte of fysshyng’ by Dame Juliana Berners

or Barnes, the Prioress of Sopwell in England ; it was the first book

on fish and fishing printed in the English language.” This would

make it appear that the Book of St. Albans and the Treatise of

Fishing were identical; but in fact the “ Treatyse of Fysshynge wyth

! Dean Sage, Townsend, C. H., Smith, H. M. and Harris, William C. New

York : Macmillan, 1902. Svo. pp. 417, illustrated.

126 THE AMERICAN NATURALIST. (VoL. XXXVII.

an Angle,” the first book on the subject of fishing printed in England,

was published by Wynkyn De Worde in 1496, and the first edition of

the Book of St. Albans, that of 1486, did not contain it. The cornea

of the eye of fishes is said to be flat on page 307, the tecth on the

head of the vomer of chars are badly treated on pages 274 and 288,

the explanation of color on page 304 is not well done, and various

items have been overlooked in proofreading.

The book is beautifully illustrated and printed; it contains a large

proportion of all that is known of American Salmon and Trout and

concerning their capture.

Boulenger on the Relationship of the Flounders. — In the

Annals and Magazine of Natural History (Vol. X, pp. 295-304)

Dr. Boulenger has a very suggestive discussion of the origin of

the group of flounders or flatfishes.

He rejects entirely the idea that these fishes are related to the

codfishes, with which group they agree in the absence of fin spines

and in little else.

They are obviously more nearly related to the ordinary spinous-

rayed fishes, showing a general similarity in the structure of skeleton,

especially the shoulder girdle, the pelvis, and the tail. ‘The

increased number of ventral rays and other characters show real

affinity with Zeus. Dr. Boulenger regards the John Dory, Zeus

Jaber, as the nearest living ally of the flounders. In the Eocene

rocks is found a genus, Amphistium, the type of a family Amphi-

stiidæ, regarded by Boulenger as clearly intermediate between Zeus

and the rhomboid flounders, which are the earliest known represen-

tatives of that group. The three families, Zeidæ, Amphistiida, and

Pleuronectidz, are joined together by Boulenger to form a new

division of spiny-rayed fishes, which he calls Zeorhombi. ‘The

Amphistiidz differ from the flounders almost solely in the sym-

metrical head and eyes, and have essentially the structure cf the

flounder larva, so far as the skeleton is concerned.

Incidentally Dr. Boulenger discusses the suggestion of the present

writer that the notable fact of the smaller number of vertebra in

tropical fishes is due to the specialization of natural selection, a

process less rapid in the cold regions, the fresh waters and the

open seas. Dr. Boulenger claims that the form with twenty-four

vertebra, characteristic of the tropics, are at the same time the more

primitive, and that the prevalence of this number of vertebrz in ‘so

many different groups simply indicates their common descent from

No. 434] NOTES AND LITERATURE. 127

some Cretaceous or Eocene group of spinous fishes with like numbers

of vertebrae.

This view is probably correct. Certainly paleontology and tax-

onomy agree in regarding the tropical flounders, percoids, scor-

penoids, and blennies, with few vertebra, as on the whole more nearly

primitive than the cold-water or fresh-water forms which have many

vertebra. At the same time, these tropical forms are the most

highly organized, the individual parts of the skeleton being most

` highly developed.

We may perhaps regard the tropical forms as having better main-

tained their primitive character of a highly developed skeleton,

while the arctic and fluviatile forms have become degraded, their

parts less developed and increased in number through repetition,

this being due to less severity of selection and perhaps the demand

for flexibility rather than strength of body.

In any case, the progressive increase in numbers of vertebrz in

various groups, as we leave the coral-reef region, is an unquestion-

able fact, and must have some cause potent among all fishes. The

only cause yet suggested is that of the demands of natural selection

in the tropics, with its cessation or reversal elsewhere. But in many

groups it is certain that the forms with many vertebra were not as

nearly primitive as the others.

Boulenger on Selenichthyes. — In the Annals and Magazine of

Natural History (Vol. X, pp. 147-153) Dr. G. A. Boulenger takes up

the relationship of the large pelagic fish known as the opah or moon-

fish (Lampris luna).

This species has been usually placed, without evident reason, with

the mackerel-like forms. It has, however, the very archaic number

(15 to 17) of rays in the ventral fins, and these fins are subabdomi-

nal in position, although placed well forward. It has the clavicle

very large, and behind it, attached to the hypercoracoid, is a very

large, flat plate, called the infraclavicle, apparently corresponding

in Boulenger’s opinion to the interclavicle of sticklebacks. The

small hypercoracoid above this plate is on the level of the hypo-

coracoid, and out of its normal position.

Dr. Boulenger makes this fish the type of a new division called

Selenichthyes, moon-fishes. This he regards as nearest allied to the

Hemibranchii ; and for the two groups, with the Lophobranchii and

Hypostomides, he proposes a new suborder, Catosteomi, character-

ized by the development of interclavicles.

128 THE AMERICAN NATURALIST. [Vor. XXXVII.

As to this, it may be urged that it is not clear that any close

affinity exists between Lampris and the others (sticklebacks, pipe-

fishes, sea horses and sea dragons), referred to Catosteomi. The

retention of various archaic characters constitutes the sole bond of

union excepting the presence of interclavicles. Furthermore, accord-

ing to Mr. E. C. Starks, interclavicles are wanting in Centriscidz and

Macrorhamphosidz, both families of undoubted hemibranchs. He

regards the interclavicle as a mere backward or downward extension

of the hypercoracoid, not as a separate bone. "There is no evidence

that the infraclavicle of Lampris is homologous with this structure: It

is very different in form and place from the interclavicle of the stickle-

back, and it may be the ordinary hypercoracoid simply enlarged.

There is no doubt of the validity of the group Selenichthyes, what-

ever its relation to the other Catosteomi. The present writer had

defined it as a distinct suborder, under another name, in a paper now

in press.

In the same paper Dr. Boulenger defines the families of Catosteomi,

adding a new one, Protoryngnathidz, based on fossil sticklebacks,

with tubiform snout, free ribs, and the first vertebra enlarged.

Meek on Fishes of Mexico. — One of the very best of recent

faunal papers is Dr. Seth E. Meek's * Contribution to the Ichthy-

ology of Mexico."! It is based on the largest collection of Mexican

fishes yet made. ‘This was obtained in the spring of 1901 by Dr.

Meek and Mr. Frank E. Lutz in the lakes and streams of the central

table-land of Mexico. Ninety-seven species were obtained. Many

of these had been secured in the previous expeditions of A. J. Wool-

man and of Jordan and Snyder to the same region, but an unex-

pectedly large number of new ones were also secured. New genera

are Zoogoneticus, based on P/atypæcilus quitzeoensis; Chapalichthys,

on Characodon encaustus; Skiffia, allied to Characodon, based on

Skiffia lerme ; and Melaniris, based on Melaniris balsanus, allied to

Chirostoma; Xenendum proves to be inseparable from Goodea.

The new species are Rhamdia oaxace, from Oaxaca; Catostomus

sonorensis, from Chihuahua; Algansea rubescens, from Ocatlan, Lake

Chapala; Gia minace, from Chihuahua; Aztecula mexicana from

Queretaro ; Notropis robustus, from Chihuahua ; Notropis santarosaliea

from Chihuahua; Ævarra tlahuacensis from Lake Chalco, Mexico;

Fundulus oaxace, from Oaxaca; Zoogoneticus diazi, from Lake Pátz-

! Publications Field Columb. Mus., Zoól. Ser., vol. iii, No. 6.

No. 434] NOTES AND LITERATURE. 129

cuaro ; Zoogoneticus miniatus, from Lake Chalco; Skifia lerme, from

Lake Pacino and Rio Lerma; Skéfia variegata, from Lake Zira-

huen; Heterandria lutzi, from Oaxaca; Xiphophorus jalape, from

Jalapa; Chirostoma attenuatum, from Lake Patzcuaro; Chirostoma

labarce, from Rio Lerma; Chirostoma patscuaro, from Lake Patz-

cuaro; Chirostoma zirahuen, from Lake Zirahuen; Jeaniris balsa-

nus, from Rio Balsas; Lepomis occidentalis, from Chihuahua; Cichla-

soma eigenmanni, from Pueblo; Gobius parvus, from Vera Cruz;

and Gobius claytoni, from Vera Cruz. These two species are appar-

ently referable to Ctenogobius. Chirostoma lucius is identified with

C. crystallinum, not with C. lerme.

The most remarkable feature of this fauna is the extraordinary

number of closely related species of Atherinidze, alike in ~ color,

and appearance, and living in the same waters.

Jordan and Snyder found, in 1895, six such species, each about a

foot long, in Lake Chapala. To this list Dr. Meek makes further

additions. All these fishes are excellent as food and all locally

known alike as Pescado Blanco de Chapala. Dr. Meek unites the

small genera Eslopsarum (with large scales) and Lethostole (trans-

lucent, with crenate scales) to Chirostoma. The genus as thus con-

stituted is known only from the table-lands of Mexico.

The species are distributed as follows ;

Lake Chalco (City of Mexico): Chirostoma jordani, C. humbold-

tianum.

Aguas Calientes, Æ. arge.

Lake Chapala, with L. Zirahuen and Rio Lerma: C. bartoni,

C. labarce, C. zirahuen, C. chapale, C. grandocule, C. promelas, C.

lucius, C. sphyrena, C. lerme, C. ocotlane, and C. estor (= C. album).

Lake Patzcuaro : C. attenuatum, C. patzcuaro, C. humboldtianum,

C. grandocule, and C. estor.

Dr. Meek has several interesting suggestions concerning geo-

graphical distribution. These isolated rivers and lakes have fish

faunas to be compared with those of rivers on different islands, sep-

arated by the sea. But the barriers of ocean are often more easily

passed than those of the Sierra Madre. The new species are all

well figured. s 18]

Fishes of Formosa. — In the proceedings of the United States

National Museum, Vol. XXV, pp. 315-368, Jordan and Evermann

give an account of the Formosan fishes contained in museums of

Japan. Two collections were studied, — the one made by Mr. T.

130 THE AMERICAN NATURALIST. [Vor. XXXVII.

Tada of Osaka for the Imperial University, the other by Japanese

officers for the imperial school of fisheries; 186 species in all were

examined. Seventeen new species are described and figured. With

them are two new genera, — Zacco (Cyprinidz), based on Opsariichthys

platypus and Evenchelys (Murenide), based on Gymnothorax

macrurus. The summary shows that the fauna of Formosa is essen-

tially similar to that of the region about Hongkong, and that it bears

much closer relation to that of India than to that of southern Japan,

while the fish fauna to the north of Tokyo contains very little in com-

mon with that of Formosa.

One of the new species deserves additional comment. ‘The sys-

tematic position of the family of sand launces, or Ammodytide, has

been long in question. Early writers placed it among the jugular

fishes as an ally of the cusks and pearlfishes. It has no ventral fins

at all, but as there are no spines in any of the fins, it was presumed

that the ventrals, if present, would be few rayed and jugular in

position.

More recently the resemblance in general structure of the sand

launces to the silversides and other groups called Percesoces, transi-

tional forms between soft-rayed and the more recent spiny-rayed

fishes, have led to a reconsideration of this opinion, The Percesoces

have abdominal ventrals and the spines little developed. ‘Two argu-

ments in favor of this view have seemed to have value. In 1811

Pallas described an Ammodytes septipinnis from the Aleutian Islands.

'This species, not since recognized, is said to be an Ammodytes, or

sand launce, with the ventral fins eight-rayed and abdominal. For

this species Dr. Gill has suggested the generic name of Rhynchias.

In the Oligocene rocks of Puy-de-Dóme Dr. Gervais has discovered

a fossil fish, now called Codtopsis acuta, which resembles a sand

launce in most respects, and is referred to the Ammodytide by

Boulenger. In this species the ventrals are six rayed, and abdomi-

nal, The dorsal fin, unlike that of Ammodytes, is rather shorter than

the anal and opposite to it. There are no fin spines. The long dorsal,

without fin spines, the numerous vertebra and abdominal ventrals with

six or eight rays, appear also in the extinct family of Crossognathidze

of the Cretaceous. This family is certainly allied to the Percesoces.

This evidence seemed conclusive, and the sequence of families,

Crossognathidz, Cobitopsidz, Ammodytide, and Atherinide, seemed

a natural one.

The tropical Ammodytidz, having normal scales and fewer verte-

bra, have been referred to the genus Bleekeria. In the collection

No. 434.] NOTES AND LITERATURE. I3I

from Formosa is a new species, Bleckeria mitsukurii, which differs

from the other species of Bleekeria in the presence of ventral fins.

These are very small, jugular in position, and composed of a slender

spine and three rays. The scales in this species are very small,

about 115 in a longitudinal series, this count being, by a slip of the

copyist, omitted in the published description. This species shows

conclusively that the Ammodytidz are not related to the Percesoces,

are not derived from Cobitopsis or Crossognathus, and that their

real place is with the ophidioid fishes and Fierasfer. The Formosan

species is the type of a distinct genus, characterized by the presence

of ventralfins. For this, the name Embolichthys Jordan and Ever-

mann (éuBodos, a hint) has been elsewhere proposed. DS 4

Fishes of Japan.— The series of monographic reviews of the

fishes of Japanese waters is continued by Jordan and Fowler, and

Jordan and Snyder, in the Proceedings of the United States National

Museum. (Vol XXV). There is included: 1. A “Review of the

Salmonoid Fishes,” the Salmonidz, (ro species); the Argentinide,

(4 species), and the Salangide, (2 species). Four salmon

(Oncorhynchus masou, O. keta, O. kisutch, and O. nerka) are found

in Japan, one of these, O. masou, not yet known from any other

region. One salmon trout, Sa/mo perryi, is found in all streams of

middle and northern Japan. A large pikelike trout, Hucho blackistoni,

common in northern Japan, finds its only analogue in the huchen

(Hucho hucho) on the Danube. There are three charrs in Japan, —

one common, Salvelinus pluvius; the other two, S. kundsha and S.

malma, confined to the northern islands. The ayu, Plecoglossus

altivelis, is found in all rivers. It is one of the finest food fishes in

the world, — a sort of dwarf salmon with peculiar dentition.

Of the smelt, Osmerus dentex, Mesopus olidus, and Mesopus

japonicus are described, besides a new species Argentina kagoshime.

Besides the diminutive and fragile Japanese ice-fish, Salanx microdon,

a second species, Sa/anx ariakensis is described from manuscripts

of Dr. Kishinouye.

The part of this paper of popular interest in Japan is condensed

in an article, “ The Salmon and Trout of Japan,” in the “ "Annotationes

zoologice Japonenses,’ published by the Imperial University of

Tokyo.

2. A “Review of the Labroid Fishes and Related Forms” includes

45 species: of Pomacentridz, 11; Labride, 31; and Scaridz (3).

The new species are Stethojulis psacas, S. ferina, S. trossula, and

132 THE AMERICAN NATURALIST. [Vor. XXXVII.

Halicheres tremebundus. All these and several other species are

figured.

3. The Chetodontide and related families are next discussed.

Twenty-seven species being represented in Japanese waters. ‘The

new species are Cy//opsis itea, Chuetodon dedalma, Holacanthus ronin,

and Coradion desmotes. The first-named species, ;/ea, should not

have been placed in Cyttopsis, as it has the ventral rays I, 9, and

the breast broad and flat, with feeble plates. It is made elsewhere

the type of a new genus, Zen Jordan, its name becoming Zen itea.

4. A discussion of the Blennies. "This interesting group of fishes is

well represented in all the rock pools of the Japanese islands, —

the elongate species, with many vertebrz in the north; the short-

bodied, tropical forms to the southward. Forty-four species are

described, representing twenty-four genera. Of these genera, the

following — Zacalles, Azuma, Zoarchias, and Abryois — are here

characterized for the first time. Twenty species are described as

new, most of these represented by great numbers of specimens, the

outlying rocks of Hakodate and Misaki proving especially rich in

fishes of this type. The plates are the work of Mrs. E. C. Starks’

and of Capt. Charles B. Hudson, and deserve especial commendation

for their accuracy and neatness.

5. The Balistide and Ostraciide. Twenty-four species are

described, two of them being new. These are Brachaluteres ulvarum

and Rudarius ercodes. Rudarius is a new genus allied to Monacan-

thus. BI

Notes on Recent Fish Literature.— In the Bulletin of the Museum

of Comparative Zoology (Vol. XXXIX, No. 3) Dr. C. R. Eastman

gives a valuable series of notes on extinct cestraciont and acantho-

dian sharks.

The extraordinary species of Edestus and other extinct forms are

thought by Eastman to be consolidated whorls of teeth of cestra-

ciont forms. These extraordinary structures have formed a standing

puzzle, it being uncertain whether their nature was that of teeth, of

fin spines, or, as conjectured by Karpinsky, of a coiled horn at the

tip of the snout. The critical study of these structures by Dr. East-

man leaves little doubt that these structures in Edestus, Campyloprion

and Helicoprion are really teeth.

Dr. Eastman describes a number of fin spines, apparently cestra-

ciont, referable to the genus Ctenacanthus. Two new species of

Acanthodes, 4. marshi and A. beecheri are described from the rich

No. 434] NOTES AND LITERATURE. 133

beds of Mazon Creek, Illinois. A series of excellent plates illustrate

the species under discussion.

In the Bulletin of the United States Fish Commission for 1901, pp.

131-159, Dr. B. W. Evermann and E. L. Goldsborough describe the

fresh water fishes collected by E. W. Nelson and E. A. Goldman in

Yucatan and neighboring regions, besides a few small collections

from other points of Mexico. The new species are as follows:

Conorhynchos nelsoni from Rio Usumacinta, Notropis santamaria

from Chihuahua (Lake Santa Maria), Notropis lerme from Lake

Lerma, Cichlasoma teape from Teapa in Tobasco, and Batrachoides

goldmani from Rio Usumacinta. The fresh water drum Aplodinotus

&runnicns was found in Rio Usumacinta. It was never before noticed

south of the Rio Grande.

In the Proceedings of the United States National Museum (Vol. XXV,

PP. 79-81), Jordan and Snyder describe two small sharks allied to

the Dog-fish from deep waters of the coast of Japan. These species

are black in color, and one of them Z/mopterus lucifer, has a glandular

substance in the skin of the side of the belly, which is said to be

luminous in life. The other sharklet, Deania eg/antina, differs from

Etmopterus in its bristly surface. It is made the type of a distinct

genus named for Dr. Bashford Dean. Jordan and Fowler give also

a review of the Stone-wall-Perches, Ofegnathide of Japan. Two

species are described.

In the Report of the United States Commissioner of Fish and Fish-

eries, for rg01, Evermann and Goldsborough catalogue the fishes

and mollusks of Lake Chautauqua, 31 fishes are enumerated, the

Chautauqua Muskallunge, Esox oA/oezszs, being much the most impor-

tance. — Evermann and Kendall publish notes on the fishes of Lake

Ontario, 73 species, on the fishes of Lake Champlain, 54 species, and

on the fishes of St. Lawrence River, 71 species, giving the known

localities and the common names of each species. — In the same

report Mr. William C. Kendall reviews the silver-sides or brit of the

East Coast of the United States, belonging to the genus Menidia.

These little fishes are excellent as food, and invaluable as food of

the larger species. Mr. Kendall gives figures of most of the FE

nized species, peninsule, audens, baryllina, and menidia, and describes

two new varieties, Menidia peninsule atrimentis from Titusville,

Florida, and Menidia beryllina cerca from Waquoit Bay and other

localities.

134 THE AMERICAN NATURALIST. [Vor. XXXVII.

In the Journal of the Imperial Fisheries Bureau of Japan Dr.

Kamakichi Kishinouye presents a monographic review of the

Japanese Tai or species of the genus Pagrus. The Zaz is per-

haps the most valuable fish of Japan, always common, always excel-

lent. The fish god, Ebisu, is always represented in Japanese draw-

ings as bearing a red Tai, “ Akadai,” Pagrus major under his arm.

Dr. Kishinouye rejects the supposed species Pagrus ruber and

describes three valid species, Pagrus major, Pagrus cardinalis and

Pagrus tumifrons. Later Dr. Kishinouye (in lit) has announced the

discovery that the last named species is a Deutex. Deutex tumifrons

has been described by Blecher as Deutex hypselosoma. The descrip-

tions are accompanied by excellent colored plates, the work of Mr. J.

Urata.

In the Denkschriften of the Academy of Vienna Dr. Franz Stein-

dachner gives an account of the fishes and reptiles collected by the

naturalist, Princess Therese of Bavaria, on her trip from Martinique

to Guayaquil around the coast of South America. Eight new species

are described and most of them figured. These are Prionodes or

Serranus huascarii from Payta, Pomadasis schyri from Guayaquil,

Pontinus dubins from Payta, Mugil charlotte from Guayaquil, Pimelo-

della yunceusis from Pacosmayo, Pygidium quechuorum from Arequipa,

Loricaria aurea from Bodega, and Leporinus muyacorum from Santan-

der in Colombia. Steindachner adopts the name Doydixodon levi-

Jrons, referring other nominal species to its synonymy.

Dr. George A. Boulenger, in the Proceedings of the Geological Society

of London discusses the young of the ten known species of Polypterus,

with figures of the seven species found in the Congo, showing the

peculiar external gill which looks not unlike an *archipterygium."

In the young of Polypterus /apradii this gill is half as long as the

body, extending backward parallel with the pectoral.

In the Actes Soc. Scient. du Chili, Dr. Federico T. Delfin, writes of

the voracity of the Chilian hag-fish (Epzatretus dombey). He finds

that one example having free opportunity to destroy fishes devoured

in seven hours 18 times its own weight of their flesh. "This amount

was not assimilated but passed through the straight alimentary canal

of the parasite, most of it little changed. The species fed to the

hag-fish is in another paper described as a new genus of Sciænidæ

under the name of CZ/as montti.

No. 434] NOTES AND LITERATURE. 135

In a “Report on the Collections made by the Southern Cross,”

Dr. Boulenger records species of fishes taken in Antarctic regions.

In this is given a useful synopsis of genera and species of Noto-

theniidz, a family of fishes especially characteristic of that region.

Pleuragramma antarcticum a leptoscopoid fish, was taken at Lat.

78°35’ south, the southernmost fish yet known.

In the Popular Science Monthly Mr. Cloudsley Rutter gives the

results of elaborate studies in the Natural History of the salmon of

the Sacramento River.

Hay on Fossil Vertebrates. — Under the head of Bibliography

and Catalogue of the Fossil Vertebrate of North America, in the

Bulletin of the United States Geological Survey (No. 179), Dr. Oliver

Perry Hay has published a work of immense practical value to the

student of Zoólogy. It is a conscientious and laborious compilation

of the kind that wins gratitude rather than fame, although amply

deserving both.

The synonymy of each name of group, genus and species is given,

with a reference to the original type of each genus and the type.

locality of each species. The rules of nomenclature of the American

Ornithologists’ Union are adopted and consistently applied, and the

general sequence and classification is that approved by American

authors. Of fossil fishes, about 1000 species are enumerated, nearly

one third as many as now inhabit the region (North America) under

discussion.

The series begins with the Ichthyotomus sharks, Ichthyotomi being

regarded as a “ Superorder,” including the Pleuropterygia and the

Acanthodii as well as the Pleuracanthine sharks. As against the Ich-

thyotomi the other sharks are set off as a second superorder called

Euselachii. The generic name, Acanthoéssus on account of priority,

is substituted for the familiar name Acanthodes. It is claimed by

Bashford Dean that the species on which Acanthoéssus is based is

not certainly identified. Unless this plea is maintained, Acanthoéssus

must stand.

A new family, Tamiobatidz, is established for Eastman’s genus,

Tamiobatis, from palzozoic rocks in Kentucky. The name Pisces is

defined so as to include all fishes except the sharks; Gill's name,

Aspidoganoidei is used instead of the preoccupied name of Ostraco-

dermi and Cope's later substitute of Ostracophori. The name “ Aspi-

doganoid " seems unfortunate, as these fantastic creatures have little

in common with ganoids. For a group containing the Arthrodira

136 THE AMERICAN NATURALIST. [Vor. XXXVII.

and Dipnoans, Dr. Hay proposes a new subclass, Azygostei, the

Crossopterygians, Ganoids and ordinary fishes forming a third sub-

class, Teleostomi. Apparently the relative position of Dipnoans,

Crossopterygians, Arthrodira and Aspidoganoids is not yet clear

enough to render this arrangement inevitable. The Arthrodira may

be allied to the Aspidoganoids; the Aspidoganoids may be modified

sharks, or even modified lampreys. The Crossopterygians may be

ancestors of Dipnoans on the one hand and of Ganoids and bony

fishes on the other, and there are numerous other elements of uncer-

tainty. Under the head of the superorder Placodermi, Dr. Hay

removes the Antiarcha (Asterolepis etc.) from the Aspidoganoids,

and places them alongside the Arthrodira, which are regarded asa

second order of Placoderms. The other superorder of Azygostei is

that of Dipnoi. Pterichthyodes is necessarily substituted for the

familiar but preoccupied name, Pterichthys ; Phlyctznaspis is need-

lessly substituted for Phlyctzenius, on account of the earlier name

Phlyctznium. Naturalists must sooner or later come to the rule

that a name is constituted by its spelling, not by its etymology.

Words spelled differently are different words. Puer is a definably

different creature from ?ue//a, whatever the likeness.

Under the subclass Teleostomi, we have two superorders, Crossop-

terygia and Actinopteri. The name Ganoid disappears, the different

types forming three orders, Chondrostei, Pycnodonti and Holostei,

at the base of the series of Actinopteri. The name Megalichthys

replaces the later Rhizodus, and Parabatrachus is applied to the

genus formerly wrongly called Megalichthys. Palaoniscum is restored

as the original spelling of Palzoniscus, and Lepisosteus as the origi-

nal, though unclassic orthography of Lepidosteus. Redfieldius is

used for the genus, wrongly called Catopterus, by J. H. Redfield.

Ginglymodi is restored as the name of the suborder of Gars, and

Halecomorphi for the relatives of Amia. The name Eugnathus, pre-

occupied, is replaced by Isopholis; Erisichthe and Pelecopterus are

united with Protosphyrzena and placed with the Pachycormidze among

the Halecomorphi.

The Nematognathi are separated from the Plectospondyli and

placed before the Isospondyli, where apparently they do not belong.

The Suckers are reduced to a subfamily of Cyprinidae. A new order,

Phthinobranchii, is proposed to include the Hemibranchii and the

Lophobranchii, two groups not fundamentally distinct. The Cato-

steomi of Boulenger corresponds to this group, except for the inclu-

sion of Lampris, a genus of peculiar and primitive structure, but surely

No. 434] NOTES AND LITERATURE. * 137

not related to the stickle-backs. Another new order, Mesichthyes,

is proposed to include the Haplomi, Synentognathi and Percesoces.

But while these groups are closely related, they differ almost as much

among themselves as the Haplomi differ from some Isospondyli or

the Percesoces from some Percomorphi. It is not clear that the

Phthinobranchii really differ much from the Percesoces, especially

since Mr. Starks has shown that the interclavicle, or infraclavicle,

supposed to distinguish the former, is merely an expansion of the

Aypocoracoid, and that it is wanting in Macrorhamphosus, Centriscus

and Aeoliscus among the Hemibranchs. The arrangement of these

transitional fishes in distinct orders, or suborders, offers very great

difficulties, because the groups, adopted though natural, are not set

off by strong characters.

The Chetodonts, Labroids and Pomacentrids are placed first

among Percomorphous fishes— for no evident reason, as the Berycidx

are more primitive in structure and earlier in time. Surely Chaeto-

dipterus does not belong to Chætodontidæ, nor Priscacara to the

Pomacentridz, nor Platax to the Carangide.

Erismatopterus, Amphiplaga, Asineops and ‘Trichophanes are

placed in the Aphredoderida. Near the Aphredoderidz, would be

safer. The suborder Pareioplite replaces the preoccupied name

Loricati for the mailed-cheek fishes.

Dr. Hay has earned the lasting thanks of his brother ichthyologists

for the pains he has taken in this work, and the portions treating of

the groups higher than fishes will doubtless be found as carefully

done and as helpful.

Davip STARR JORDAN.

Notes. — Ikeda (Journal of the College of Science, Tokyo, Vol.

XVII) has made an extended series of experiments to determine the

mode of closure of the blastopore and the position of the embryonic

body in amphibian eggs. He shows that the results obtained by

puncturing eggs and by other similar methods can never be depended

upon to reveal the normal course of development, for widely differing

results are obtained depending upon the position of the puncture.

He believes that the embryonic body in Amphibia may be formed

at any position on the egg surface, and that many authors have

overlooked or underestimated the varying rate at which different

parts of the blastoporic lip enclose the yolk-mass, a rate which

determines the final closing point of the blastopore and consequently

the position of the embryo.

138 THE AMERICAN NATURALIST. [Vor. XXXVII.

Kishinouye has recently published (/our. Col. Sci. Tokyo, Vol.

XVII) descriptions of five new species of Japanese Scyphomeduse.

All represent new genera and one a new family of the Stauromedusze,

Stenoscyphide.

The octopod genus Amphitretus has thus far been known only

through a single specimen collected by the “Challenger” and

described by Hoyle. A second specimen taken by the collector of

the Missaki Marine Laboratory in the deep water of Sagarni Sea,

Japan, in 1897, is now described by Ijima and Ikeda (Annotationes

Zoologice Japonenses, Vol. IV). The animal, which was nearly half

a foot long, was bell-shaped, semigelatinous and more or less trans-

parent. The more important internal parts could be discerned through

the outer gelatinous layer in which chromatophores were embedded.

Unlike all other Cephalopods the mantle is fused with the siphon in

the median plane so that there are two branchial openings into the

branchial cavity. A colored figure of the appearance of this remark-

able animal during life accompanies Ijima and Ikeda's description.

The osteology of the shoulder girdles of the hemibranchiate fishes

is the subject of a paper by E. C. Starks in Vol. XXV of the Proc.

U. S. Nat. Mus.

Brief accounts of the development and degeneration of the eyes

in the blind fish, Amblyopsis, and of the structure of the degenerate

eyes in the amphisbzenian lizard, Rhineura, have been given by C. H.

Eigenmann in the Proc. Zndiana Acad. Sci. for 1901.

A summary of the questions concerning the propagation of the

common eel forms the subject matter of an address by the President

of the American Microscopical Society, C. H. Eigenmann. ‘The

address is published in the Transactions of the Society.

The increase of mesenteries in the madrepore corals has been

studied by J. E. Duerden (dun. Mag. Nat. Hist, Ser. 7, Vol. X,

August, 1902), who finds that in most polyps of the genus Madrepora

only the six bilateral pairs of primary mesenteries are developed. In

any colony a few large polyps may possess a greater number of

mesenteries, in which case the new mesenteries are added as bilateral

pairs at only the two axial extremities of the polyp, the enterocoels of

the dorsal and ventral directives. ‘The mesenterial increase is early

associated with fission of the stomodaum and in the end probably

with complete fission of the polyp in which half the mesenteries of

each fission polyp are derived from the primary twelve of the original

polyp and the other half are new formations.

No. 434] NOTES AND LITERATURE. 139

The same author, (Bu. Amer. Mus. Nat. Hist, Vol. XVI, pp.

323-332) calls attention to the great importance of boring alge as a

factor in the disintegration of corals. These grow most vigorously

in quiet lagoons and thus contribute to the rapid decay of corals

known to take place in such situations. They are, therefore, to be

reckoned among the numerous elements concerned with the formation

of coral islands.

The crustacean fauna of Nickajack Cave, Tennessee, has been

investigated by W. P. Hay (Proc. U. S. Nat. Mus., Vol. XXV, pp.

417—439). A new species of Czcidotea, closely related to C.

nickajackensis Packard, a new species of Gammarus, and two new

subspecies of crayfishes are described.

Notes on the structure and development of a species of barnacle,

Dichelaspis miilleri, found parasitic on the gills of crabs, have been

published by R. E. Coker (Bull. U. S. Fish Com. for 190r, pp. 399—

412).

BOTANY.

McIlvaine and Macadam's ** American Fungi." ! — A second

edition of this book, which appeared first in 1900, has been brought

out, and differs from the original edition in the addition of a supple-

ment of twenty-five pages, including one full plate and several cuts.

There is no equally full and fully illustrated book on the edible and

poisonous fungi of this country, and though bulky and somewhat

inconvenient for use, it is much lighter than the original issue and it

ought to be in the hands of all who collect fungi for the vg

W. T.

Notes.— Postelsia, the new yearbook of the Minnesota Seaside

Station, makes its appearance in a very attractive form, in the issue

for 1901, recently distributed. This volume consists of seven chap-

ters, or lectures, illustrated. :

A new periodical, Annales Mycologici, under the editorship of

H. Sydow and devoted to Fungi, is announced to appear at the

opening of 1903, through the book house of R. Friedländer and

Sohn, of Berlin.

1 McIlvaine, Charles, and Macadam, Robert K., One Thousand American Fungi.

Revised edition. Indianapolis, The Bowen-Merrill Company, 1902. Quarto,

xxxvii + 729 pp. Numerous plates, some of them in color, and line cuts. $5.00.

140 THE AMERICAN NATURALIST. [Vor. XXXVII.

Since October, 1901, an Agricultural Bulletin of the Straits and

Federated Malay States, edited by H. N. Ridley, has been issued

monthly from the Government Printing Office at Singapore.

The Proceedings of the Indiana Academy of Science, for 1901, con-

tain a number of botanical papers, several of them referring to the

flora of the state.

Rhodora for September contains the following articles: Jones,

“ The Pringle and Frost Herbaria” ; Collins, F. S., ^ An Algologist’s

Vacation in Maine”; Collins, J. F., “Distinctive Features of Z7zzs

Hookeri” ; Grout, “Notes on Vermont Mosses”; Leavitt, * Seed

Dispersal of Viola rotundifolia”; Bacon, “ Anagallis arvensis and

cerulea in Vermont”; Fernald, * Aster undulatus x Novi-Belgii” ;

Webster, “ A form of the Bitter Boletus” ; Hoffmann, “ Virulence of

the Wild Parsnip,” and reports on the Josselyn Society and the

Vermont Botanical Club.

The Botanical: Gazette for September contains the following

articles: Copeland, “ The Rise of the Transpiration Stream,” (part) ;

Chandler, * Revision of the Genus Nemophila” ; Worsdell, “ Evolu-

tion of the Vascular Tissue of Plants "; MacMillan, “Suggestions on

the Classification of Seeds”; and Fairchild, “ The Sensitive Plant as

a Weed in the Tropics.”

The Bulletin of the Torrey Botanical Club for September contains

the following: Piper, “ Notes on the Biennial and Perennial West

American Species of Lappula ” ; White, “Some Mt. Desert Fungi ” ;

Hill, “ Notes on Migratory Plants”; Burt, “Some Hymenomycetous

Fungi from South America " ; and Cotton, * Three new plants from

Washington."

Torreya for October contains the following articles: Hollick, * Fos-

sil Ferns from the Laramie Group of Colorado”; Howe, “ Ca/oglos‘a

leprieurti in Mountain Streams”; Earle, “ Key to the North Ameri-

cal Species of Lactarius, II"; Cockerell, “Some New Mexico

Plants”; Harper, R. M., * A visit to Okefinokee Swamp in Southern

Georgia"; Rydberg, “Is the White Fruited Strawberry of Pennsyl-

vania a Native Species? "; and Earle, * A Much Named Fungus.”

The Popular Science Monthly for October contains two articles of

considerable botanical interest; Toumey, * A Study in Plant Adap-

tation ” ; and Cook, “ The American Origin of Agriculture."

No. 434.] NOTES AND LITERATURE. 141

Of the leaflets of Proceedings of the Biological Society of Washing-

ton issued under date of October ro, several refer to botany.

A Fora Arctica,in English, under the editorial care of C. H.

Ostenfeld, is being issued from Det Nordiske Forlag, of Copenhagen.

The first part, dealing with the Pteridophytes, Gymnosperms, and

Monocotyledons, by Gelert and Ostenfeld, occupies 134 well illus-

trated pages.

Vol. XXXIV of the Transactions and Proceedings of the New

Zealand Institute, issued in July last, has 166 pages devoted to

botany, chiefly referring to the island. Perhaps the most generally

interesting paper is one on the prothallus of Phylloglossum.

An important * Flora of Koh Chang," by Schmidt, with the assist-

ance of a number of specialists, is being published in current numbers

of the Botanisk Tidsskrift.

A further study of the revegetation of Krakatoa, by Penzig, is

contained in the concluding number of Vol. XVIII of the Annales du

Jardin Botanique de Buitenzorg.

A discussion of the southeastern United States as a center of geo-

graphical distribution of both animals and plants, is reprinted by

C. C. Adams from the Biological Bulletin, Vol. III, No. 3.

Some of the Crassulacez of South Africa are being revised by

Schonland and E. C. Baker in current issues of the /ourna/ of Botany.

A voluminous study of the Myricacez, by Chevalier, occupies a

large part of Vol. XXXII of the Mémoires de la Société nationale ot

Cherbourg.

Separates of a paper on the Carices of Japan, by Léveillé and

Vaniot, have been separately printed from the Bulletin de l'Académie

internationale de géographie botanique.

An extensive and well illustrated paper on the fern-worts of Kouy-

Tcheou, by Christ, forms a large part of the Bulletin de P Académie

internationale de géographie botanique for August-September.

An important paper on the structure and development of the stem

in the Pteridophyta and gymnosperms, by Dr. Jeffrey, is separately

distributed from Vol. CXCV of the Philosophical Transactions of the

Royal Society of London.

Fibro-vascular studies on the ferns, by Bertrand and Cornaille, are

contained in the Comptes Rendus de 1 Académie des Sciences of Paris,

142 THE AMERICAN NATURALIST. [Vor. XXXVII.

for 1901 and 1902, and the Buletin de la Société botanique de France

and the Procés-Verbaux de la Société d'histoire naturelle d’ Autun, for

the present year.

A paper on the life history of Vittaria lineata, by Elizabeth G.

Britton and Alexandrina Taylor, forms the concluding number of

Vol. VIII of the Memoirs of the Torrey Botanical Club.

Descriptions of two new species of fossil algz, of the genus Butho-

trephis, from Indiana, are separately issued by David White from the

Proceedings of the United States National Museum.

The Journal of Mycology for October, with portrait of J. C. Arthur

as frontispiece, contains the following articles: Morgan, “ Notes on

North American Fungi”; Atkinson, “Preliminary Note on Two

New Genera of Basidiomycetes”; Durand, * The Genus Angelina” ;

Atkinson, “ Preliminary Notes on Some New Species of Fungi” ;

Kellerman, “Ohio Fungi” (fascicle v); Ricker, “ Notes on Some

West American Fungi" ; Clinton, “North American Ustilaginez ” ;

Harshberger, “Notes on Fungi”; and Kellerman, “Notes from

Mycological Literature, II."

The first part of the A/onographia Uredinearum, by P. and H.

Sydow, has been distributed from the Borntrzeger press of Leipzig,

and is almost entirely occupied with the Puccinias of Composite, of -

which 298 are described, not a few of them new.

By a typographical error, a paper on Uredinez, by Pennington,

has been separately issued from the Anales de la Sociedad Cientifica

Argentina, Vol. LIII, under a cover reading “ Erudineas recolectadas

en las islas del delta del Parana."

Van Bambeke considers the crystalloids of Autobasidiomycetes, in

the Bulletin de la classe des sciences, Académie Royale de Belgique,

No. 4, for 1902.

An illustration of the need of care in the determination of plants on

which morphological studies are made, is given by Rehder in an open

letter in the Botanical Gazette for September.

Volumes II and III of the Archives de l Institut botanique de P Uni-

versité de Liége consist of five long papers on plant anatomy.

A study of the pith diaphragms of dicotyledons, by Mágócsy-Dietz,

is found in Vol. VII of the Mathematische und Naturwissenschaftliche

Berichte aus Ungarn.

No. 434] NOTES AND LITERATURE. 143

Van Tieghem discusses the classificatory value of the embryo in

Ochnacez in No. 3 of the current volume of the Bulletin du Muséum

@histoire naturelle, of Paris.

An article on “School Gardens in Cities,” by Helen C. Putnam,

containing the first general review of school gardening in this country,

with a synopsis of what is done abroad, is contained in the Rhode

Lsland School Report for 1902.

A short account, with illustration, of the public gardens at Shang-

hai, is contained in the Gardener's Chronicle of September 27.

An instructive and handsomely illustrated report of the Secretary

of Agriculture on the forests, rivers, and mountains of the southern

Appalachian region, with the President’s message transmitting it to

Congress, has recently been issued from the Government Press.

An attractive little pamphlet on Vigna sinensis, the cowpea of the

South, has been issued by the experimental farm of the North Caro-

lina Horticultural Society, at Southern Pines.

The Agave cultivated largely in the Mexican state of Jalisco for the

manufacture of the distilled beverage known as Tequila, is described

by Weber, under the name 4. tequilana, in the Bulletin du Muséum

@ histoire naturelle, of Paris, No. 4 of the current volume.

An account of the Peen-to peach, and a series of varieties that have

been selected from it in Florida, by Professor Hume, constitutes

Bulletin No. 62 of the Florida Agricultural Experiment Station.

Dr. Fairchild publishes an account of Spanish almonds and their

introduction into this country, as Buletin 26 of the Bureau of Plant

Industry, of the Department of Agriculture.

A well illustrated article on the cultivation of coffee has been pub-

lished by Sajo in recent numbers of Prometheus.

An account of the China aster and its diseases, by R. E. Smith,

constitutes Bulletin 79 of the Hatch Experiment Station of Massa-

chusetts.

Sunn-hemp, Crotalaria juncea and C. tenuifolia, as grown and used

in India, is discussed in Der 77 vopenpffanzer for October.

a new species from

An illustrated description of Juncus textilis,

t volume of the

California, is distributed by Buchenau from the curren

Abhandlungen of the Bremen Naturwissenschaftlicher Verein.

144 THE AMERICAN NATURALIST. [VoL. XXXVII.

A subject list of works on the textile industries and wearing apparel

in the library of the British patent office, has recently been issued

from that office.

No. 5 of the current volume of Anales del [Instituto Médico Nacional,

of the City of Mexico, contains the following articles of botanical

interest; Villaseñor, Preliminares al estudio de las resinas; Armen-

daris, Algunas consideraciones sobre las propiedades fisilógicas de

algunas plantas que contienen saponina; Noriega, Curso de histo-

ria de drogas, 1.

The Botanical Gazette for October contains the following papers : —

Land, A morphological study of Thuja; Copeland, The rise of the

transpiration stream; Snow, Some notes on the ecology of the Dela-

ware coast; Greenman, A new western Camassia.

‘The November number of the Bulletin of the Southern California

Academy of Sciences contains the following botanical articles : — David-

son, Spherostigma erythra; Parish, The southern California species of

Calochortus, II; Greata, Tribal character in the separation of the

style-branches in the Composite. The portion of Dr. Yates’ Prehis-

toric California contained in this number is also largely devoted to

fossil plants of that state.

The Fern Bulletin for October contains the following articles : — -

Davenport, Early fern study in America; Clute, A ten years' retro-

spect; Underwood, Some features of fern study; Druery, British

fern culture; Smith, New Zealand ferns and fern study; Miyake,

Notes on Japanese ferns; Gilbert, Historical sketch of the Linnean

Fern Chapter; Eaton, A new Equisetum (Æ. Azema/e intermedium),

and a biographical sketch of Mr. Clute, with portrait.

Part I of Vol. XXVII of the Journal of the Royal Horticultural

Society, dated September, 1902, contains a number of illustrated

articles of more than passing botanical interest; among them, one by

Dr. Cooke on Fungoid pests of the garden, one by Professor Carr on

Plant communities, one on Weeds of the garden, by Mrs. Boyle, and

one on Pelargonium disease, by Mr. Massee.

The New Phytologist, of October 30th, contains a discussion of Ele-

mentary university courses in botany, and a paper on Pryocystez, an

obscure group of algz, by Blackman, as well as a continuation of

Blackman and Tansley's Revision of the classification of the green

alge.

No. 434.] NOTES AND LITERATURE. 145

Hefts 10 and 11 of Engler’s Das Pfanzenreich consist, respectively,

of the T'ropzeolaceze, by Buchenau, and the Marantacez, by Schumann.

The Flant World for September contains the following principal

articles: — Safford, Extracts from the note-book of a naturalist on

the Island of Guam; Pollard, Frank Hall Knowlton ; Cook, A decid-

uous tropical tree; Pammel, Our vanishing wild flowers; Hill, The

etymology of Columbine; Rice, A carnivorous plant; Hopkins, A

rare freak of the Trillium; and Kaufman, A carnivorous bog.

Rhodora for October contains the following articles: — Fernald,

Two northeastern Veronicas ; Graves, Valerianella in New England;

Knowlton, Empetrum in Franklin County, Maine; Pease, Hieracium

Prealtum at Andover, Mass., Webster, J. R., Crepis virens in Massa-

chusetts; Shaw, New station for Polypodium vulgare cambricum;

Bailey, Plant stations in Rhode Island; Scorgie, /asione montana in

Massachusetts; Webster, H., A new mushroom for the market;

Deane, Calluna vulgaris in New Hampshire; Rich, Lists of New

England plants, IX., Polygonacee.

The following articles of botanical interest appear in Vol. 7 of the

second series of the Transactions of the Royal Society of Canada,

recently issued : — Laflamme, Jacques-Philippe Cornuti — Note pour

servir a l'histoire des sciences au Canada; Matthew, A backward

step in palzo-botany; MacKay, Botanical bibliography of Canada,

1900.

A careful analytical account of the flora of the Galapagos Islands,

by Dr. Robinson, with the collaboration of specialists, is published as

No. 4 of the current volume of Proceedings of the American Academy

of Arts and Sciences, and constitutes No. 24, n. s., of the Contribu-

tions from the Gray Herbarium of Harvard University.

No 22 of North American Fauna consists of a report on a biologi-

cal investigation of the Hudson Bay region, by E. A. Preble, of

some botanical and a great deal of zoological interest.

In current issues of the Anales del Museo Nacional de Montevideo,

Professor Arechavaleta is describing and figuring a considerable

number of new phanerogams, many of them belonging to genera

which are also represented in North America.

As No. 6 of the papers issued by the Botanical Seminar of the

University of Nebraska on the botanical survey of that state, a thesis

146 THE AMERICAN NATURALIST. [Vor. XXXVII.

by G. G. Hedgcock has recently been published on The relation of

the water content of the soil to certain plants, principally mesophytes.

A bulletin on dissemination, under the title ** Plant travelers," by

Professor Weed, is published as Nature Study Leaflet No. 3 of the

New Hampshire College Agricultural Experiment Station.

A paper by Matruchot and Molliard, on the Effects of frost on the

structure of plant cells, is contained in the Revue Générale de Bota-

nique for October r5th.

A lecture on Health and disease in plants, by F. S. Earle, is

published in the Journal of The New York Botanical Garden for

November.

Studies on cell division in cambium, by Schoute, have been issued

from the Verhandelingen der Koninklijke Akademie van Wetenschappen,

of Amsterdam, under date of October, 1902.

A paper on the Cactacez of Costa Rica, by Dr. Weber, separately.

printed from No. 6 of the current volume of Bulletin du Muséum

d'histoire naturelle, contains several new species of Cereus, Phyllo-

cactus, Rhipsalis and Pereskia.

Echinacea purpurea and its varieties are considered in the Revue

Horticole of November rst.

An exhaustive local treatment of Salix, with reference to the forms,

including hybrids, which occur about Regensburg, by Anton Mayer,

is published as Heft VII of the Berichte des naturwissenschaftlichen

Vereines zu Regensburg, for 1898—9.

An article on ius lambertiana is contained in La Feuille des

Jeunes Naturalistes, of November rst.

A paper on the Ferns of the Pacific Coasts, by Dr. Yates, is pub-

lished in Popular Science News for November.

An illustrated monograph of the Ulothricacez and Chaetophoraceze

of the United States, by T. E. Hazen, constitutes Vol. XI, No. 2, of

the Memoirs of the Torrey Botanical Club.

No. 5 of the current volume of the Bulletin du Jardin Impérial

Botanique, shows that at the beginning of the present year 35141

forms of plants were cultivated in the famous St. Petersburg Gardens,

the herbarium of which is said to contain more than a million and a

half of dried plants and the library to contain 29520 books compris-

ing 14608 titles.

No. 434.] NOTES AND LITERATURE. 147

Some interesting notes on the Banksian Herbarium are contained

in the Journal of Botany for November.

The Report of the committee on school gardens and children’s

herbariums, contained in No. 2 of the Zransactions of the Massachu-

setts Horticultural Society for 1901, recently issued, is a very instruc-

tive document for those interested in nature work in the public

schools.

Some of the results reached by an award for home and school

grounds, Colorado Springs,.Colo., are shown in Park and Cemetery

for October.

A well illustrated and practical paper on Shade trees for city

streets, by Murrill, is published as Buletin 205 of the Cornell

University Agricultural Experiment Station.

A paper on the Colors and aromas of flowers, by Cadevall y Diars,

constitutes Vol. IV, No. 27, of the Memorias de la Real Academia de

Ciencias y Artes de Barcelona.

A paper in the Bulletin of Miscellaneous Information, No. 36, of

the Botanical Department of Trinidad, states that Nepenthes is

profitably grown in the West Indies among orchids as a means of

catching cockroaches, which are said to be destructive to these plants.

An interesting Essay on the cultivation and curing of Vanilla, con-

stitutes No. 35 of the Bulletin of Miscellaneous Information, issued

by the Botanical Department of Trinidad.

The September number of the Agricultural Bulletin of the Straits

and Federated Malay States is largely devoted to India rubber.

Economic fruits and seeds are taken up in Lieferung ro of Wies-

ner's Rohstoffe des Planzenreiches, which is intended to be completed

in one more Lieferung.

The cold storage of apples, with a consideration of the influence

. of cold storage on the decay of apples, and the chemical changes

which occur during storage, by Morse, is published as Bulletin 93 of

the New Hampshire College Agricultural Experiment Station.

An illustrated paper, by Hartley, on injurious effects of premature

pollination, with general notes on artificial pollination and the setting

of fruit without pollination, has been published as Bulletin No. 22 of

the Bureau of Plant Industry, U. S. Department of Agriculture.

I 48 THE AMERICAN NATURALIST. | (Vor. XXXVII.

Recent economic papers published by the Department of Agricul-

ture are: — Husmann, the manufacture and preservation of unfer-

mented grape must, (Bulletin No. 24, Bureau of Plant Industry) and

Bond and Keeney, Irrigation of rice in the United States (Buletin

No. 113, Office of Experiment Stations).

A paper by Emerson, giving a preliminary account of variation in

bean hybrids, is separately printed from the 7574 Annual Report of

the Agricultural Experiment Station of Nebraska.

A new forestry publication, Zhe Forestry Quarterly, has been

started from the New York State College of Forestry, at Cornell

University. The first number appeared in October.

A practical analysis of farm forestry, by Chamberlain, is published,

with numerous illustrations, in Country Life in America for November.

Papers on the Wilt disease of the Cowpea and its control, by

Orton, and a Cowpea resistant to root knot, by Webber, are pub-

lished as Bulletin No. r7 of the Bureau of Plant Industry, U. S.

Department of Agriculture.

Oudemans and Koning have distributed, in the form of separates

from the Archives Neerlandaises des Sciences Exactes et Naturelles, a

prodromus of a mycologic flora obtained by cultures on gelatine

media of material derived from humus at Spanderswoud near Bus-

sum. The paper is illustrated by 3o plates, mostly devoted to new

and brightly colored species of molds.

A German variety of Anthurus borealis is described and figured

by Hennings in the September-October Bezblatt zur “ Hedwigia.”

No. 105 of the Proceedings of the Linnean Society of New South

Wales consists of several articles on Bacteria associated with the

sugar cane, as well as other matter of botanical interest.

The Gardeners’ Chronicle, of November ist, contains description

and figure of a convenient little vest-pocket micrometer for handy

measurements, devised by Sir Joseph Hooker.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS, AND DEATHS.

EDUCATIONAL GIFTS.

Chicago University, $1,500,000 from John D. Rockefeller; $526,000 from

other sources.

Columbia University, $18,800 from various donors.

Fairmount (Kansas) College, $25,000 from Dr. D. K. Pearson.

Fargo College (North Dakota), $50,000 from Dr. D. K. Pearson.

Hamline (Minnesota) College, $250,000 from various donors.

Harvard University, $100,000, for a professorship of comparative anatomy,

from James Stillman.

Illinois College, $50,000, from Dr. D. K. Pearson.

Johns Hopkins University, $25,000, from Dr. and Mrs. C. A. Hester.

Lafayette College, $2,500, by the will of Benjamin Barge.

Princeton University, $140,000, by the will of Mrs. Susan Dod Brown;

$10,000, from Morris K. Jesup.

Radcliffe College of Harvard University, $116,000 from several donors.

Teachers College of Columbia University, a conditional gift of $500,000

from John D. Rockefeller; $274,509, from Mr. and Mrs. B. Everett

acy.

Tulane University, the residuary estate, estimated at about $1,000,000, of

the late A. C. Hutchinson.

University of California, $111,000, for archeological purposes, from Mrs.

Phoebe Hearst.

University of Georgia, a conditional gift of $50,000, from George Foster

Peabody.

University of Pennsylvania, $100,000, from Dr. E. W. and C. H. Clark, for

the chair in Assyriology.

University of Rochester, $10,000, from Mrs. Esther B. Steele.

Wesleyan University, $40,000, from Mr. Van Vleck, for an observatory.

Wooster (Ohio) University, $400,000, from various friends.

Yale University, $50,000, by the will of Mrs. Lena Courrier; $80,000, by

the will of Benjamin Barge; $171,000, as residuary legatee of the

estate of Edward Wills Southworth.

APPOINTMENTS.

E. B. Bailey, geologist of the Scottish Geological Survey.— E. D. Bell,

professor of animal biology in the Utah Agricultural College.— Dr. Antonio

Berlese, professor of zoólogy in the Agricultural School at Portici, Htaly.—

149

150 THE AMERICAN NATURALIST. | (Vor. XXXVII.

Dr. Vincenz von Borbas of Budapest, professor of systematic botany in the

university at Klausenburg.— H. W. Britcher, instructor in zoólogy in the

University of Maine.— Dr. Henry Coutiere, professor of zoólogy in the

school of pharmacy at Paris.— Miss Clara Eaton Cummings, professor of

botany in Wellesley College.— Dr. Eugen Dubois, professor of paleontology

in the university at Amsterdam.— Dr. Durig, docent for physiology in the

University of Vienna.— Dr. Martin H. Fischern, associate in physiology in

the University of California.— Frances E. Foote, instructor in zoólogy in

Wellesley College.— J. S. Gardiner, demonstrator in animal morphology in

the University of Cambridge.— Dr. C. H. Gordon, instructor in geology and

eography in the University of Nebraska.— R. P. Gregory, demonstrator in

botany.in the University of Cambridge.— Dr. Ludwig Heim, professor of

bacteriology and hygiene in the university at Erlangen.— Dr. Addine

Hewson, assistant professor of anatomy in Jefferson Medical College.—

Dr. Lorenz Hiltner, director of the newly established Agricultural-Botanical

Institution at Munich.— Dr. P. P. C. Hoek, director of the Station for

International Research at Copenhagen.— J. Allen Howe, curator and libra-

rián of the Museum of Practical Geology.— Dr. J. A. Ippen, docent for

mineralogy and petrography in the university at Graz.— Dr. O. Juel, pro-

fessor of botany in the university at Upsala.— J. Graham Kerr, professor of

natural history in the University of Glasgow.— Dr. Karl Ritter von Keissler,

assistant in the Botanical section of the court museum at Vienna.— Dr.

Jacques Loeb, professor of physiology in the University of California.—

George Grant McCurdy, curator of the anthropological collections of Yale

University.— Dr. Siegfried Mollier, second conservator of the anatomical

institute at Munich.— Benjamin Moore, professor of biological chemistry in

University College, Liverpool.— Dr. Willibad A. Nagel, professor extraordi-

nary of physical physiology in the university at Berlin.— Dr. Joseph

Nusbaum, professor of comparative anatomy in the University at Lemburg.

— Dr. Oberhummer, professor of geography in the university at Vienna.—

Dr. F. Wilhelm Pfaff, district geologist at Munich.— Dr. Charles Queva, pro-

fessor of botany in the faculty of sciences at Dijon.— Dr. Karl Reichinger,

assistant in the botanical section of the court museum at Vienna.— Dr. Otto

M. Reis, district geologist at Munich.— Dr. Charles D. Rogers, assistant in

physiology in the University of California.— Dr. Wladislaw Rothert, pro- -

fessor of botany in the university at Odessa.— Dr. Johannes Rückert, first

conservator of the anatomical institute at Munich.— A. Schwagger, district

geologist at Munich.— Dr. Hans Strahl, professor of anatomy in the univer-

sity at Tübingen.— Dr. Emil Lietze, director of the royal geological survey

in Vienna.— Dr. J. B. de Toni, professor of botany in the university at

Modena.— Dr. F. Vejdovsky, transferred from the chair of embryology and

comparative anatomy in the university at Prag, to that of zoólogy.— Dr. I.

G. de Vries, director of the Zoological Station at the Helder.— Dr. Friedrich

Vierhapper, assistant in the Botanical Garden and Museum at Vienna.—

C. F. Myers Ward, lecturer in physiology in Charing Cross Hospital Medi-

No. 434] GIFTS, APPOINTMENTS, RETIREMENTS. ISI

cal School.— Alice V. Wilcox, instructor in zoölogy in Wellesley College —

Dr. Ernst Wilczek, professor of botany in the university at Lausanne.— Dr.

F. A. Wilder, state geologist and professor of geology in the University of

North Dakota.

RETIREMENTS.

Dr. Anton Fric from the chair of zoólogy in the university at Prag.—

Dr. Guido Stache from the directorship of the Austrian Geological Survey.

DEATHS.

Homer Franklin Bassett, student of Hymenoptera at Waterbury, Conn.,

June 28, aged 76.— Dr. Andreas Nikol Betekow, formerly professor of

botany in the Universities at Charkow and St. Petersburg, July 14.— Mr.

R. A. Blair, geologist, at Sedalia, Missouri.— Ladislava Celakowsktho, pro-

fessor of botany in the Bohemian University at Prag, aged 69.— Vincent

Leche Chesnevieux, geologist and traveler, aged 86.— Thomas Comber,

botanist, in Blackpool, England, January 24, 1902— Dr. J. G. Cooper,

zoölogist, in Haywards, California, July 19, aged 72.— Augustin Alexis

Damour, mineralogist, in Paris, September 21, aged 94.— M. Dehérain,

professor of plant physiology in the Paris Museum of Natural History.—

uise Brisbin Dunn, tutor in botany in Barnard College, December 18.—

Dr. Rudolf Finkener, professor of mineral analysis in the mining academy

in Berlin, September 14, aged 68.— Dr. A. Frenzel, mineralogist and orni-

thologist, in Freiburg in Sachsen, in August.— Mr. William Gunn, district

geologist of the British Geological Survey, October 23, aged 65.— Dr.

Theodor von Heldrich, professor of botany and director of the botanical

gardens in Athens, September 7.— Dr. Johann Janko, director of the ethno-

graphical section of the Hungarian Museum at Budapest, aged 34.— Ludwig

Kumlein, naturalist of the Howgate expedition, in Milton, Wisconsin, "

December.— Dr. Carl von Kupffer, professor of anatomy in the university at

Munich, December 16, aged 73.—Dr. Eduard Lehmann, student of the

flora of Livonia, May s, aged 61.— Major James C. Merrill, M. D., orni-

thologist, aged 49.— Dr. Ernst Meynert, professor extraordinary s -—

omy in the university at Halle, aged 39.— Mark Micheli, botanist, in

Geneva, July 10.— Dr. Millardet, professor of botany at Bordeaux— Dr.

Ludwig Molendo, bryologist, in Munich, July 25, aged 68.— T. A. ume

Müller, coleopterist, in Dresden, August 16, aged 74.— Dr. William ve

Ord, formerly tutor for physiology in St. Thomas Hospital, Londón, aged

68.— P. Anselm Pfeiffer, naturalist, in Kremmunster, July 7, aged 54-7 id

Samuel Leopold Schenk, formerly professor of embryology in the university

at Vienna, August 18, aged 62.— Dr. A. R. C. Selwyn, formerly director of

I52 THE AMERICAN NATURALIST. (Vor. XXXVII.

the Canadian Geological Survey, at Vancouver, October 19, aged 78.—

Franz Sikora, a natural history explorer of Madagascar and Réunion.—

Dr. Franz Tappeiner, anthropologist, in Meran, Tyrol, August 19, aged 82.—

Dr. Adolfo Targione-Tozetti, formerly professor of comparative anatomy at

Florence, September 18, aged 79.— Dr. Rudolf Virchow, since 1856 pro-

fessor of pathology in the university at Berlin, September 5, aged 80.— J.

B. Williamson, student of Lepidoptera, in Slough, England, June 21, aged

74.— Ur. Oliver Willis, botanist at White Plains, N. J., April 27, aged 88.

— Rev. Dr. Wiltshire, geologist, in London, October 25.

(No. 433 was mailed January 20.)

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THE

AMERICAN NATURALIST

VoL. XXXVII. March, 1903. No. 435.

ANTITHETIC VERSUS HOMOLOGOUS ALTER-

NATION 2

DOUGLAS HOUGHTON CAMPBELL.

THE nature of the alternation of generations in Archegoniates

is a question of fundamental importance in dealing with the

morphology of the vascular.plants, and necessarily has been the

subject of much discussion, with a corresponding divergence of

opinion. Both the antithetic and homologous nature of the

alternation of generations have been supported by such eminent

advocates that one hesitates to speak too confidently on either

side. As I cannot, however, after long and careful study, agree

that the argument is equally strong for both sides, and it seems

to me that proper weight has not always been given to some

of the evidence, it has seemed worth while to review the direct

evidence on both sides as completely and as fairly as possible.

I shall not enter into a detailed account of the controversy, as

that has already been done quite recently. I have myself given

fully my own reasons for supporting the antithetic theory?

1 Read before Section G. at the Washington meeting of the A. A. A.S.

*Coulter: The Origin of the Leafy Sporophyte. ot. Gaz. July, 1899.

? Campbell: Mosses and Ferns, pp. 510-514, 1895.

154 THE AMERICAN NATURALIST. [Vor. XXXVIL

and it seems to me that Professor Bower’ has effectually

refuted the arguments of Dr. Scott, who has been the special

champion of the homologous theory in England. More recently

Mr. W. H. Lang has reviewed the subject,? being more or less

non-committal, but rather leaning to the homologous view ; and

in this country Professor Coulter 3 has assumed a somewhat sim-

ilar attitude.

Owing to their perishable nature, the simpler green Algze and

Bryophytes have left very meagre fossil remains, so that their

geological history is very imperfect, and we are perforce driven

toa study of the living forms, as practically our only means of

tracing the ancestry of the higher plant forms. Of the vascular

plants there are abundant fossil remains which throw much light

upon the relationship of the Pteridophytes and seed-plants, and

the succession of forms in geologic times, but help but little in

determining the lower forms from which the former originated.

It has been urged that inasmuch as ferns, and even seed-

plants, can be traced back to the Devonian, and possibly even

further, it is hopeless to expect the secret of the origin of

the vascular plants can ever be solved. However, as many

extremely primitive forms have undoubtedly survived to the

present time, we can learn very much from a comparative study

of these with the higher plants, which must have come from

forms very similar to them. Of the forms which are of special

importance in this connection are the simpler green Algz, and

the generalized liverworts.

Zoólogists are in much the same position with regard to the

origin of the vertebrates, as botanists are concerning the vascular

plants. The former sub-kingdom is certainly as old and probably

older than any land-plants — and yet we do not find that the

zoologists consider the question of the origin of vertebrates

entirely hopeless.

I shall not attempt here to discuss the monophyletic or poly-

phyletic origin of Pteridophytes, but shall mainly concern myself

with the class which at present is the predominant one, the Ferns.

LVature, Nov. 17, Nov. 24, Dec. 1, 1898.

* Annals of Botany, 12: 585-592, 1898.

3 Joc. cit.

No. 435.] ANTITHETIC VERSUS HOMOLOGOUS. I55

According to the antithetic theory of alteration, the ferns have

originated from forms very similar to the simpler existing liver-

worts, the leafy sporophyte being an elaboration of the non-

sexual sporophyte. The homologous theory maintains that

Bryophytes and Pteridophytes have nothing to do with each

other, the latterarising quite independently from algal ancestors.

The latter hypothesis was first suggested by the alga-like pro-

tonema of the mosses, and the somewhat similar prothallia of

certain ferns, especially Trichomanes.

Opposed to this assumption is the fact that the filamentous

prothallia of such ferns as Trichomanes, or Schiz@a pusilla, are

obviously secondary developments, in the former case, at least,

associated with excessive moisture. The prothallia of most ferns

grown in water, or kept excessively wet, and poorly lighted, tend

to assume a filamentous form. Among both Hymenophyllacez

and Schizzeacez, the great majority of forms studied possess

the normal flattened prothallium of the ordinary ferns. The

filamentous protonema of the true mosses is also, if we are to

trust the evidence of comparative morphology, a secondary

development from the liverwort-like thallose protonema of forms

like Sphagnum. The true mosses and Hymenophyllacez are

probably very far from being primitive types.

From a comparison of the fossil and living ferns it is certain

that the so-called eusporangiate types are much older than the

leptosporangiate forms which are now predominant. The Marat-

tiaceze, of which only a small number of tropical species survive,

are especially well represented in the Palaeozoic rocks. Now

on the assumption that the gametophyte of the ferns is descended

directly from algal ancestors, and is not of bryophytic origin,

we should expect the gametophyte of these primitive ferns to

be more alga-like than that of the more recent and specialized

Leptosporangiates. The gametophyte of the existing Marat-

tiaceze which is well known, is a fleshy, relatively large and long-

lived thallus, closely resembling that of the lower liverworts,

being much more like these, both as regards the vegetative

growth of the thallus and the reproductive organs, than are the

prothallia of the common ferns.

The gametophyte of the Ophioglossacez, the second order of

156 THE AMERICAN NATURALIST. [Vov. XXXVII.

the Eusporangiatz, is also very massive, and as far as possible

from any known algal form. Of course in the Ophioglossaceze,

the saprophytic nature of the gametophyte has doubtless to some

extent modified its structure.

The archegonia of the Marattiaceze are extraordinarily like

those of the Anthocerotacez, and the antheridia also offer cer-

tain suggestions of a similar origin to those of the latter order.

Indeed, were it not for the biciliate spermatozoids of the Antho-

cerotacez, and their peculiar chromatophores, I should not hesi-

tate to assume a direct connection between the latter order and

the eusporangiate ferns.

The extraordinary uniformity in both structure and develop-

ment of the archegonium throughout both Bryophytes and

Pteridophytes, including such a marked character as the ventral

canal-cell; the great similarity in the origin and development of

the spermatozoids, and the details of fertilization, certainly are

very strong arguments for a common origin for all Archegoniates.

The theory that these resemblances are merely parallel develop-

ments can only be accepted on the production of very much

weightier evidence than has yet been brought forward.

When to the obvious resemblances existing between the game-

tophytes of the ferns and liverworts (of course the lower types

. like the anacrogynous Jungermanniales and the Anthocerotaceze

being understood) there are added the numerous resemblances

in the development of the sporophyte, the probability of a genetic

connection between the two phyla of the Archegoniates becomes

enormously greater. The fallacy of Dr. Scott's argument, that

the assumption of the antithetic theory involves the creation of

a structure de novo, without any apparent ancestry, has been

perfectly refuted by Professor Bower. There is no claim that

the sporophyte is an entirely new structure. It starts with the

zygote, which on germination produces a greater or smaller

number of spores, thus increasing the number of plants result-

ing from a single zygote, an obvious advantage. The rudimentary

sporophyte of Coleochzete, which there is no difficulty in homol-

ogizing with the zygote-product of CEdogonium, may equally well

be compared with the sporophyte of Riccia, as has often been

done, whether we assume that the sporophytes of the two are

genetically related or not.

No. 435.] ANTITHETIC VERSUS HOMOLOGOUS. 157

As yet we have no evidence beyond mere hypothesis as to the

way in which the motile zoóspores, arising from the zygote of

the algal ancestors of the Bryophytes, gradually were replaced

by the tetrads of non-motile spores which characterize all Arche-

goniates. The same difficulty is met with in tracing the origin

of the sexual organs. In both respects there is much greater

difference between the humblest liverwort and the highest alga

than there is between the former and the most highly organized

fern. It must be confessed that the gap between all existing

Algæ and Archegoniates is very great.

When we compare the two series of Archegoniates, the case

is very different. The structure of the archegonium and spores

is identical throughout, and the early stages of the sporophyte

agree very closely, this being especially true of the more primi-

tive types of Pteridophytes. In these the young sporophyte

remains very much longer dependent upon the gametophyte and

the external organs which characterize the Pteridophytes, are

relatively late in making their appearance. Both of these facts

point to a nearer approach to the bryophytic type in the lower

Pteridophytes, a fact which is not readily explicable on the

assumption that they are in no way connected with the

Bryophytes. In a number of the lower Pteridophytes, e. £g.

Marattia Lycopodium, Botrychium, the young sporophyte may

remain attached to the gametophyte for months, or even years,

long after it has passed beyond the embryonic stage.

If we compare the gametophyte and sporophyte of any typical

Archegoniate, we note a very significant difference in their rela-

tion to the water-supply. The gametophyte is always, to a

greater or less degree, an aquatic organism, never quite emanci-

pating itself from the life conditions of its algal ancestors. The

sporophyte, on the contrary, at least in its earlier stages, is never

exposed directly to the water, although a few groups of Pteri-

dophytes have developed, secondarily, aquatic sporophytes.

Almost from the first the sporophyte is provided with a special

massive absorbent organ, the foot, which is later superseded

by the true roots of the Pteridophytes, a much more efficient

means of obtaining water than is pro ided by the rhizoids of

the gametophyte; and the unlimited capacity for growth of

I 58 THE AMERICAN NATURALIST. | [Vor. XXXVII

the true roots of the vascular plants allows the development

of a root system to keep pace with the growth of the aerial part

of the sporophyte. There is thus developed for the first time

a plant-body strictly terrestrial in its character, and capable of

independent growth.

The gradual elaboration of the sporophyte is easily traced in

the liverworts from the simple capsule of Riccia to the large

and almost independent sporophyte of Anthoceros, or in another

direction to the elaborate sporophyte of the true mosses.

Whether or not we agree entirely with Professor Bower's

beautifully worked out theory of the sterilization of potential

sporogenous tissue, the fact is patent that there has been a

gradual elaboration of the originally purely sporogenous and

parasitic structure resulting from the zygote, in the direction of

an independent plant, with a corresponding subordination of the

spore-producing function to the vegetative life of the sporophyte.

This culminates among the Bryophytes in such highly specialized

types as Polytrichum and other similar mosses. In these there

is early developed a foot, which supplies the materials needed

for the growth of the long-lived sporophyte. In the more highly

organized forms there is present a special strand of water-con-

ducting tissue, which may be directly compared to the fibro-

vascular bundles of the higher plants. The outer tissues of the

long slender seta, also, assume the character of mechanical tis-

sues which give it the necessary strength to support the large

and complicated capsule, in which only a very small amount of

tissue is devoted to spore-formation.

At an early period the outer tissues of the sporophyte develop

chlorophyll, and there may be formed a distinct assimilative

organ, the apophysis, provided with green lacunar tissue com- .

municating with the external ‘atmosphere by means of stomata,

entirely similar to those upon the green organs of the vascular

plants. The highly specialized sporophyte of the true mosses

has not probably given rise to any type of vascular plants, but in

_the equally developed, but much more generalized sporophyte

of Anthoceros, we have a structure that may well represent the

characters of the sporophyte from which originated the leafy

sporophyte of the ferns.

No. 435] ANTITHETIC VERSUS HOMOLOGOUS. 159

Perhaps the strongest argument in favor of a common origin

for the sporophyte in all Archegoniates is the absolutely uniform

method of spore production. However ignorant we may be of

the transition from the motile zoospores resulting from the

germinating zygote of the green Algz to the immobile tetrad-

spores of the Archegoniates, there is no question as to the prac-

tical identity of the spores, both as to origin and structure

throughout not only the Archegoniates, but the seed-plants as

well. This extraordinary uniformity is perfectly comprehensible

if we accept the antithetic view, since the beginning of the sporo-

phyte must have been a simple mass of such spore-tetrads; and

in Riccia we still have a sporophyte which has scarcely advanced

beyond this stage. In the more highly developed sporophytes

of the higher Archegoniates, the spores have retained their primi-

tive characters, but a continually increasing amount of the sporo-

phytic tissue has been devoted to purely vegetative purposes.

That this formation of spore-tetrads should invariably have taken

place in structures which are secondarily spore-producing, as is

demanded by the theory of a non-sexual origin for the sporo-

phyte in the Pteridophytes, requires something more than mere

hypothesis to give it much credence. This is all the more the

case if a polyphyletic origin for the different phyla of Pteridoph-

ytes is assumed.

Of all existing Bryophytes there is no question that Anthoce-

ros offers the nearest approach to the Pteridophytes, although

this by no means implies that the latter are directly derived from

the former. Were it not, however, for the form of the chroma-

tophores and spermatozoids in the Anthocerotacez I should not

hesitate to assume this to be the case.

The sporophyte of Anthoceros is characterized by a long-

continued basal growth that adds to the tissues of the sporophyte,

which thus may reach a length of several centimeters. The

origin of the sporogenous tissue is like that of the Pteridophytes,

arising from sub-epidermal tissue, differing from all other Bryo-

phytes except Sphagnum in this respect. Moreover the sporo-

genous tissue is not continuous, but is more or less regularly

divided into sporogenous areas by intervening sterile tissue sug-

gesting an approach to a very simple fotm of sporangium. The

160 THE AMERICAN NATURALIST. Vor. XXXVII.

highly developed chlorophyll-tissues, and the central strand of

conducting tissue, apparently the true homologue of the primary

vascular bundles of the Pteridophyte embryo, are certainly sug-

gestions, if not forerunners, of the corresponding pteridophytic

structures. To my mind there is far less difference between the

sporophyte of such a simple Pteridophyte as Ophioglossum or

Phylloglossum and that of Anthoceros, than there is between the

latter and Riccia.

When Professor Coulter, for example, says, “In contrasting

the sporophytes of Bryophytes and Pteridophytes, they seem to

have nothing in common except that they are usually derived

from the oospore and represent an asexual generation," we feel

that the question has not been fairly stated. As we have tried

to show, the sporophytes of the higher Bryophytes and Pteri-

dophytes agree closely in the following important particulars:

The early divisions of the embryo; the development of a special

absorbent organ, the foot, thus inaugurating the terrestrial habit

of the sporophyte; the gradual subordination of the spore-func-

tion, and even an approach to the formation of sporangia in

Anthoceros; the development of special chlorophyll tissue and

in some cases of an assimilative organ, the apophysis ; the devel-

opment of special conducting tissue; and finally the absolute

identity in the character and formation of the spores, with the

characteristic reduction in the number of chromosomes. These

may be all mere parallel developments, and not genuine homolo-

gies ; but. it will require a great deal of direct testimony to make

this in the least degree probable, especially when to these obvious

resemblances in the sporophyte are added the equally remarkable

correspondences in the structure of the gametophyte of the

liverworts and ferns.

Again, when Professor Coulter says that the sporophyte of

the Bryophytes never develops lateral members, and has nothing

comparable to sporangia, his statement is open to question.

While the complicated apophysis of a Polytrichum or Splachnum

may not be strictly an appendicular organ, it may assume a

flattened, leaf-like form in S. Zuteum, comparable to a perfoliate

leaf, and is as truly a photosynthetic organ at least in its younger

condition, as is the leaf of a fern. Of course I do not mean to

No. 435] AWTITHETIC VERSUS HOMOLOGOUS. 161

intimate that the apophysis of Splachnum is genetically related

to the leaves of the Pteridophytes; but it is no more associated

with spore-production than they are, and shows the capacity of

the sporangium of the Bryophyte to produce special vegetative

organs in no way connected with spore-production. The segre-

gation of the spore-masses, already spoken of in connection with

Anthoceros, is at any rate a hint of the origin of the sporangia

of the Pteridophytes. As to the overwhelming tendency to

spore-production in Bryophytes, while this is undoubtedly true of

the simpler types, it may well be questioned whether it can prop-

erly be asserted of the most highly developed ones. It would

be an interesting problem, for instance, to compare the relative

output of spores in Polytrichum and Osmunda cinnamomea.

While it is inconceivable that such an extremely specialized

structure as the sporophyte of the higher Mosses could have

given rise to the leafy sporophyte of the Pteridophytes, it is

quite conceivable that both types may have originated from a

common ancestral form, which may very well have been not very

different from Anthoceros. That the latter is “hopelessly

specialized” is very far from being the case. On the contrary,

the sporophyte of Anthoceros is a remarkably generalized

structure. I mean by this, that it has not a single character

which is peculiar and cannot be duplicated elsewhere.

Pringsheim, the first advocate, I think, of the homologous

theory of alteration, based his conclusions upon the behavior of

various mosses in which the protonemal filaments may arise

directly from the sporophyte. Pringsheim believed that the

protonema was not essentially different from the vegetative

tissues of the sporophyte, from which they arose in such cases.

This reasoning is not entirely convincing. The protonema arises

normally’ from special sporophytic cells, the spores, and it is

difficult to see why, under abnormal conditions, it might not

arise from other sporophytic tissue. The same reasoning will

apply to apospory in the ferns where the gametophyte may

arise directly from the leaf-tissue. poe

The strongest argument in favor of homologous alternation is

the occurrence of apogamy, or the origin of the sporophyte as a

vegetative bud from the gametophyte. So far as I am aware

162 THE AMERICAN NATURALIST. [Vor. XXXVII.

(I make this statement with some reserve) all cases of apogamy

yet observed have been in cultivated ferns. At any rate, much

the larger number of observed cases have been under artificial

conditions, either intentional or otherwise. This suggests at once

that apogamy is a pathological phenomenon. In most cases it is

induced by preventing fertilization, which would otherwise take

place, except in a very small number of instances. Exposure

to strong sunlight has also been found to be a factor in inducing

apogamy. It is also noteworthy that most cases of apogamy

occur in varieties which differ from the normal in being crested,

or that show other indications of exuberant vegetative growth

which may certainly imply some connection between this redun-

dant growth and apogamy in the gametophyte.

Finally, as Professor Bower has pointed out, all cases of

apogamy recorded occur in leptosporangiate ferns, admittedly

the most recent and most specialized members of the class. No

Bryophytes nor eusporangiate ferns have yet been found which

exhibit apogamy. If we are to consider apogamy as in any

sense a reversion to a primitive condition, it is hard to see why

it should be confined to these highly specialized modern types,

and be entirely absent from the presumably much older and

more primitive ones.

Mr. Lang! has given a very clear account of what he thinks

may have been the course of development of the sporophyte,

according to the homologous theory. He assumes that the

primitive form of the gametophyte was a flattened thallus, pre-

sumably much like the existing liverworts, although he does not

make this comparison. He supposes that this thallus under

stress of circumstances, owing to an insufficient water-supply,

may have given rise to spores, the spore-stage following the sex-

ual stage, but being an integral part of the gametophyte, and

not produced from the egg. It is assumed that in connection

with this special spore-producing function, the leafy sporophyte

gradually assumed its definite form, and later, but this point is

not quite clear, became replaced by the similar structure arising

from the zygote. Why the spores produced by this asexually

produced structure should be identical with those developed by

lloc. cit.

No. 435] ANTITHETIC VERSUS HOMOLOGOUS. 163

the non-homologous, sexually developed sporophyte of the

Bryophyte, is not explained.

Mr. Lang’s hypothesis does not claim to be based upon any

experimental evidence, and it may be of interest to see whether

there is any actual evidence bearing upon the natural behavior

of the gametophyte when exposed to conditions similar to those

assumed by this hypothesis.

There are many Bryophytes which are regularly subjected to

complete drying up, and I should like to call attention to their

behavior. Most leafy liverworts and mosses which grow upon

the trunks of trees, or upon exposed rocks, simply dry up com-

pletely, and revive promptly so soon as water is furnished them,

behaving thus very much as many algae do. In California, and

probably the same is true of other similar semi-arid regions,

nearly all of the terrestrial liverworts are perennial, and pass

through the long, dry summer unharmed. This is well illus-

trated by several common species of middle California, such as

Fimbriaria californica, Targionia hypophylla, Fossombronia

longiseta, various species of Riccia, etc. In all of these species

the apex of the shoot remains alive, being usually more or less

perfectly protected by overlapping scales or hairs. As soon as

the first autumn rains fall, the plants at once resume growth,

and in a surprisingly short time develop their reproductive

organs. Indeed it is quite possible that the young organs are

sometimes already formed at the time the thallus ceases its

growth in the spring.

In mosses it is not uncommon to find bulbils developed, these

being merely arrested vegetative shoots, developed from the

protonema, or rhizoids.

Among various liverworts, there have been found tubers of a

somewhat different nature, and it is likely that when the thallose

forms of semiarid regions are further studied, these will be

found to be commoner than has been supposed to be the case.

In the case of the peculiar genus Geothallus of Southern Cali-

fornia, the growing point of the thallus, with the tissue adjacent,

becomes transformed into a tuber, with a large amount of

reserve food developed in the central tissues. This tuber,

which is buried in the earth, remains dormant through the

summer, but germinates promptly when supplied with water.

164 THE AMERICAN NATURALIST. (VoL. XXXVII.

In none of the species studied is there the slightest tendency

shown to develop anything resembling the spores borne upon

the sporophyte. It is true that some leafy liverworts develop

single-celled gemmze from the leaf-margins, and multicellular

gemmze are formed in various species; but these are not in any

way associated with lack of water, nor are they in the least like

true spores.

It is interesting to find that the gametophyte of -some ferns

has also developed the power of resisting drought. In California

it is not uncommon to find large prothallia of Gymnogramme

triangularis which have survived the summer, and the sporo-

phyte of this fern, as well as that of various xerophilous species

of Selaginella, can absorb water through the leaves, very much

as is done by the leaves of mosses and liverworts. Goebel? has

called attention to the behavior of G. leptophylla, where the

gametophyte is perennial, and may develop tubers much like

those found in the thallose liverworts. In this species the

sporophyte is annual.

‘The point which I wish to emphasize is this: that whereas

there are many cases where the gametophyte is subjected to the

conditions which according to Mr. Lang's hypothesis should

induce spore-formation, in no cases observed has this been the

result, but the devices for surviving drought have been of a very

different nature.

Dr. Coulter? thinks that the determining factor in the devel-

opment of the leafy sporophyte has been photosynthesis or

*chlorophyll-work." He sees no reason why such a structure

as the leafy sporophyte may not have arisen non-sexually in

response to the need for chlorophyll activity, and quite apart

from the production of any form of reproductive bodies.

In support of this view he instances the development of the

gametophoric branches in Marchantia, and the leafy game-

tophoric shoots of the true mosses. It is*hardly likely that Dr.

Coulter would derive the latter from the former, although such

might possibly be inferred from his statement. The accuracy of

his statement that *the erect structure laid hold of the game-

1 Outlines, p. 200.

? loc. cit.

No. 435] ANTITHETIC VERSUS HOMOLOGOUS. 165

tophore, rather than the sporophyte,” may be questioned. With

comparatively few exceptions, the sporophyte of the Bryophytes

is erect, while the shoots bearing the sexual organs, especially

among the liverworts, are very often prostrate. Moreover there

is in most of the latter class no development of distinct game-

tophoric shoots in the sense that such occur in the more special-

ized Marchantiacez and mosses. The transition from strictly

thallose forms like Aneura or Pallavicinia to genuine leafy forms

like the typical Acrogynee is extremely gradual, the leafy shoots

of the latter forms being in no sense buds from a thallose game-

tophyte, but direct transformations of the whole body of the lat-

ter. We are, therefore, perfectly justified in homologizing the

leafy moss plant including of course the protonema, with a thal-

lose liverwort or with the prothallium of a fern.

Dr. Coulter thinks that the spores would find more favorable

conditions upon a leafy shoot than upon a thallus, which is

doubtless true; but why this leafy shoot should not develop

gradually from the sexually-produced sporophyte of the Bryoph-

ytes, as there is the strongest evidence that it has done, he does

not make clear. The development upon the leaves, of spores of

the same type as those of the lower Archegoniates, is entirely

comprehensible, if we assume that the leafy sporophyte of the

ferns is descended from a leafless bryophytic sporophyte; but it

is hard to understand if we assume that the spores of the ferns

have no genetic connection with the absolutely similar ones of

the Bryophytes.

According to Dr. Coulter's hypothesis, the leafy sporophyte

originates as a vegetative shoot from the gametophyte in a man-

ner analogous to the production of the gametophoric shoots in

the mosses, or the apogamous origin of the sporophyte in some

ferns. Upon the leaves, which originally were purely organs for

photosynthesis, were developed secondarily the sporangia. The

germination of the non-sexual spores and of the zygote are

assumed to have been entirely similar, giving rise first to a thal-

lus, from which secondarily the spore-bearing leafy shoot arose.

If such has been the course of development, it is strange that

all trace of the thallose portion has been lost in the sexually

produced sporophyte. One would expect to find some trace of

166 THE AMERICAN NATURALIST. [Vor. XXXVII.

it in the embryo, at least of the lower types, but nothing which

can be so interpreted is ever found, unless we might consider

the suspensor of the Lycopods as of this nature, which probably

no morphologist would be likely to do.

The statement that. it is no more difficult to imagine the

gametophyte producing spores, than the spore giving rise to the

gametophyte, can hardly be admitted. The spores of all Arche-

goniates, if we assume the antithetic theory of alternation, are

the direct descendants of those produced by the germinating

zygote in the ancestral forms, which on germination give rise to

the gametophyte. This is perfectly demonstrable, while any-

thing like the production of spores, at least of the type produced

by the sporophyte, is absolutely unknown in any gametophytic

structure. The supposed cases of the production of sporangia

upon the gametophyte have been shown to be merely a greatly

reduced case of apogamy.

Of course, if we should admit that the sporophyte originated

apogamously in the first place, it would follow that foliage leaves

are older than sporophylls, and that Pteridophytes and Bryoph-

ytes have nothing to do with each other; but the weight of evi-

dence is very much against such a supposition.

That chlorophyll-work has been a very potent factor in the

evolution of the plant-body is of course beyond dispute; but its

bearing upon the origin of terrestrial plants is not so clear. All

green plants, whether aquatic or terrestrial, must make provision

for photosynthesis, and we find the arrangements for the most

favorable display of green tissue developed in various ways.

Leaves are by no means confined to land-plants, many Algze,

especially the larger Phzophycez, having foliar organs which,

although simple in structure, are often of great size, and effi-

cient organs for photosynthesis. There is abundant evidence,

also, that leaves have been developed.more than once among the

mosses and liverworts.

It is rather an exaggeration to say that the greater part of the

chlorophyll-work in any liverworts is relegated to the gameto-

phoric branches. Surely the amount of chlorophyllous tissue in

the thallus of all the Marchantiacez is greater than that in the

gametophores. It may also be questioned, moreover, whether

No. 435] ANTITHETIC VERSUS HOMOLOGOUS. 167

the elevation of the receptacles, which usually does not begin

until after the fertilization of the archegonium, is not more

intimately associated with spore-dispersal than with the need of

chlorophyll-work. In most Marchantiacez the seta is very little

developed, and the elongated stalk of the receptacles probably

takes its place.

If one were looking for gametophytic structures which most

nearly simulated the organs of the leafy sporophyte it would be

among the lower thallose .Jungermanniales, and not among the

true mosses, that one should look. The extraordinarily fern-like

aspect of certain thallose liverworts like certain species of Aneura

and Hymenophytum! recalls at once some of the Hymeno-

phyllaceze with fan-shaped, dichotomously veined leaves. These

fern-like liverworts develop a rhizome-like stem, having flattened,

dichotomously branched green shoots, curiously like a small fern

leaf. © Structurally, however, they are in all respects like the

prostrate thallus of other species.

Now it is conceivable that when the sporophyte reached the

stage when it first developed a leaf, the latter should tend

to assume a form suggesting the expanded, dichotomously

branched lamina, so characteristic of nearly all the lower liver-

worts, which I assume represent the ancestral T from which

the ferns have been derived.?

When Professor Coulter speaks of the subordination of chlo-

rophyll-work to spore-formation among the Bryophytes, I sup-

pose he is referring to the sporophyte. The rarity of sexual

organs, and consequently of the sporophyte with its spores, is a

familiar phenomenon in many mosses and liverworts. Thus

whole Sphagnum. swamps without a sporophyte, and Lunularia

multiplying almost exclusively. by vegetative buds, illustrate this.

It cannot, therefore, be that lack of vegetative energy has pre-

vented the Bryophytes from becoming predominant terrestrial

forms. It is perfectly clear that purely vegetative shoots may

be produced; but that these ever gave rise to the leafy sporo-

phyte is quite another question.

The suggestion s that the archegonium may have been derived

! Goebel, Organographic der Pflanzen, second part, p. 251.

? Campbell, loc. cit., p. 509.

? Coulter, loc. cit., p. 58.

168 THE AMERICAN NATURALIST. [Vor. XXXVII.

from a group of oógonia, protected by sterile tissue, is a novel

one, but it is hard to see upon what evidence it is based. Of

the algal forms, the structure of the oógonium of the Characez

resembles most nearly that of the archegonium ; but that this is

anything more than an analogy is questionable, and at present

it must be confessed that the origin of the archegonium is

extremely obscure.

As to the significance of apospory and apogamy, both of

these phenomena may, I think, fairly be compared to the vari-

ous types of adventitious budding. We know that among both

ferns and seed-plants, adventitious shoots may arise from almost

any portion of the plant-body. The whole sporophyte may

develop as an adventitious bud upon the root, leaf, stem, or

even from the sporangium, shown by the budding of the nucel-

lus in several cases of polyembryony, or the replacing of the

sporangium by a shoot in Isoetes. Surely no morphologist

would claim that because in Camptosorus or Cystopteris the

sporophyte may arise as a bud upon the leaf; or in Populus or

Ailanthus may spring as a bud from the root; that these facts

indicate that such was the original origin of the sporophyte,

and that the latter is directly homologous with the organ from

which it arises.

I think, therefore, that we must fall back upon the question

of water-supply as the real explanation of the peculiarities of

the leafy sporophyte. All mosses remain to a certain extent

aquatic, most of them absorbing water at all points much as an

alga does, and depending only to a limited degree upon the

rhizoids as a means of water absorption. Moreover the rhizoids

are entirely inadequate to supply a plant body of large size,

which could not, of course, absorb sufficient water for its needs

from the atmosphere. Nature has, apparently, made numerous

attempts to adapt the essentially aquatic gametophyte to a

terrestrial environment, with very imperfect success.

The sporophyte, at first a purely spore-producing structure,

has been from the beginning essentially aerial in habit, never

being directly in contact with water, but getting its water-supply

indirectly, at first through the cells of the gametophyte, but

soon developing a special massive absorbent organ, the foot, the

No. 435.] ANTITHETIC VERSUS HOMOLOGOUS. 169

forerunner of the root developed later, which puts the sporophyte

into direct communication with the earth, thus rendering it

independent of the gametophyte. With the establishment of a

true root-system, capable of unlimited development to keep pace

with the growth of the aerial portions of the sporophyte, there

began a new era in the history of the vegetable kingdom, which

has culminated in the myriad types of vascular plants which now

cover the earth.

ES.

A UE RIS)

MONTRE

déco atl

ON THE SHELL OF LITTORINA LITOREA AS

MATERIAL FOR THE STUDY OF VARIATION.

R. P. BIGELOW AND ELEANOR P. RATHBUN.

THE common periwinkle, Littorina litorea, seems at first

especially suitable for a statistical study of the effect of a new

environment upon type and variability. For the best evidence,

according to Morse ('80) and Ganong (’86), goes to show that

this species has been introduced from Europe into Nova Scotia

and New England within the last fifty years. It was first

reported from Halifax by John Willis in 1857. Previous to this

date no mention is found of the species in any list of the shells

of Nova Scotia, New England, or the Gulf of St. Lawrence.

Furthermore the shell is not found fossil in these regions, and is

not found in the prehistoric shell heaps. The species seems

therefore to have been introduced into America somewhere about

the middle of the nineteenth century.

After its introduction it spread gradually southward along the

coast of New England as far as Long Island Sound. In every

place where it has appeared north of Cape Cod it has increased

enormously and in many cases nearly driven out the indigenous

species of Littorina, until it is now probably the most abundant

gasteropod on the coast. But south of the Cape it is only fairly

abundant in certain stations.

Thus we have in Z. Zorea a species which has been intro-

duced into a new environment within a comparatively short time.

One might expect a priori that this species on migrating into a

new area would find new conditions and be subjected to new

factors of natural selection, which would tend to establish a new

type, or else that the species, having escaped, perhaps, from

some of the checks that limited its variability in its former

environment, might show a greater range of variation under

the new conditions. By a statistical study of Z. Z/forea one

might hope to determine whether one or both of these supposi-

17I

172 THE AMERICAN NATURALIST. [Vor. XXXVII.

tions are true, and whether evolution is taking place in this

introduced species. These results, if trustworthy, would be of

very considerable value at this time, because the number of

quantitative determinations of the present rate of evolution that

have been made heretofore are very few indeed, and students

of the subject are seeking to accumulate sufficient data to

remove the theory of evolution from the realm of inference

to that of observed fact.

It was for this reason that the authors planned a statistical

study of ZL. “torea. We proposed to compare variability and

type in two generations of shells which had been subjected to

their environments for different lengths of time. We hoped to

find material for this purpose in measurements of some charac-

ter in young shells and of the same character in the upper part

of fully grown specimens, as Weldon ('or) had done in his study

of Clausilia.

Thus by comparing the type and variation of the same charac-

ter in the present young shells and in the survivors of a former

generation of young shells we might expect to discover any

selection that may be taking place with respect to that charac-

ter and to determine whether the result is progressive evolution

or is simply periodic, provided of course, that the character

chosen is known to remain constant during the life of the indi-

vidual. The only available character to measure seemed to be

the angle at the apex of the shell.

For this investigation a large number of living specimens of

all sizes were collected at Winthrop, Mass., from spaces between

the rocks at low tide, during the month of February, 1902.

Many of the oldest shells were plainly much eroded at the

apex and unfit for our purpose. But all of the smaller shells and

many of the large ones appeared to be in good condition. It was

noticeable, however, that the larger shells were generally less

acute at the apex than the very small ones. The suspicion that

this fact aroused was further increased by examination with the

hand lens, which showed that near the apex of all the shells the

surface was roughened with fine depressions like minute pin-

holes. Evidently some process of erosion was at work, and it

would be useless to proceed with our investigation unless it

No. 435] SHELL OF LITTORINA LITOREA. 173

could be shown that the amount of erosion was so slight as to

be negligible. It occurred to us that possibly the amount of

erosion, on the contrary, might be even greater than would be

indicated by the condition of the surface. If the loss of mate-

rial from the surface should be balanced by the deposition of

new shell within the cavity, this might mask the erosion to such

an extent as to make it impossible to determine from the appear-

ance of the surface whether any given shell was much or little

eroded. With such an uncertainty any results obtained by the

comparison of the apexes of old shells with those of young ones

would be utterly worthless.

We proceeded, therefore, to determine the amount of erosion

in shells of various sizes from the smallest to the largest. The

method employed was the study of thin sections made through

the apex of the shell in the plane of the columella.

The sections were made in the following manner: — The

soft parts having been removed, the rounded part of the shell

was first ground from each side on an ordinary grind-stone used

for the sharpening of instruments. A slice was thus obtained

through the middle of the shell. One side of this was ground

with emery on a grinding machine used for making sections of

minerals! until a surface passing through the middle of the

columella was obtained. This surface was made perfectly even

by polishing with a paste of diatomaceous earth on a ground

glass plate. The polished side was then cemented to a glass

plate with Canada balsam, after which the other side was ground

down and polished in the same way until the section was suffi-

ciently thin. The section was then cleaned and mounted in

Canada balsam in the usual manner under a cover glass.

In this way about a dozen slides were made. Figs. I, 2, 3,

and 4 are camera lucida drawings of sections of four shells of

different sizes. Thus we have sections of two young shells, Figs.

1 and 2, a shell which is not quite fully grown, Fig. 3, and one

of full size, Fig. 4. The openings in Fig. 1 are numbered,

beginning at the top, and the corresponding openings in the

other sections are given the same numbers. Of course, the

1 The authors are much indebted to Dr. C. H. Warren for helpful suggestions

and assistance in using this machine.

174 THE AMERICAN NATURALIST. (VOL: XXXVII.

holes in any two sections will not correspond exactly, as the

planes of the sections are not made accurately at the same angu-

lar distance from the origin of the spiral, but it can easily be

seen which holes are of about the same size. These openings

are, of course, the places where the lumen of the shell lies in the

plane of the section and serve as a convenient index of erosion.

In each of the older shells we can see the outline of the younger

stages. The line a—? in each section represents the lower bound-

ary of the stage corresponding to Fig. 1.

The shell represented by Fig. 1 is about 3 mm. long and 2.5

mm. broad, and is the smallest one that we were able to cut.

On the body-whorl were three or

four ribs somewhat darker in color

than the rest-of the shell. This

marking extended into the next

whorl, but toward the apex of the

acute spire the shell was smooth.

In the section the lumen of the

shell is cut distinctly eight times,

leaving eight openings in the sec-

tion, and at the apex there is an

indication of another half turn of

the spiral. Each half turn of the

Camera drawing of a median Spiral, or opening in the section,

is numbered consecutively, begin-

ning at the apex.

Microscopic examination of the section, Fig. r, shows that the

main part of the shell is composed of a translucent material of

a pearly white color with fine darker lines of growth or fracture

lying at more or less acute angles to the sides of the lumen.

This material may be called the primary shell. Under the ribs

it is stained a dark yellowish brown color for nearly its whole

thickness, and in the upper part of the shell the whole columella

is colored. Around each opening of the section the boundary

of the primary shell is clearly defined by a very fine but per-

fectly distinct dark line. Within this boundary the lumen of the

shell is more or less completely lined by a layer of pearly mate-

rial that we may call secondary shell. This may be white and

Fig. 1. a

longitudinal section of a young shell.

X 35.

No. 435.] SHELL OF LITTORINA LITOREA. 175

clear or it may show a yellowish stain or fine lines of growth

parallel to the surface of the lumen. The secondary shell has

evidently been deposited after the formation of the primary shell,

and there is a considerable amount of it, even in this very young

specimen. In the upper part of the spire the secondary has

nearly filled the original cavity of the primary shell, so that open-

ings 2 and 3 are almost obliterated, and No. 1 is quite so.

Moreover the lumen at openings 2 to 4 is eccentric, the thick-

est part of the secondary shell being on the outer side. It will

be noticed, also, that the outlines of the primary lumen at open-

ings I, 2, and 3 are incomplete, being cut by the outer surface

of the shell. Evidently erosion has been at work already to such

an extent as to have removed the whole thickness of the original

outer wall of the first whorl, and, if no secondary shell had been

secreted, there would be an openmg from the y to the exte-

rior at this place.

Fig. 2 represents a similar section of a shell about half again as

‘large as Fig. 1, that is, 5 by 4.5 mm. The specimen is in good

condition with well marked dark ribs upon the body-whorl, and

with an acute apex, but the apical angle is greater than in Fig. 1.

Assuming that Fig. 2, has passed through a stage correspond-

ing to Fig. 1, a comparison of the size of the opening indicates

that the increased size has been attained by the addition of another

whorl. But the total number of whorls in the two shells remains

the same. This paradoxical condition may be explained by sup-

posing the first whorl, openings 1 and 2, to have entirely disap-

peared as the result of erosion. That this is the true explana-

tion is rendered highly probable by microscopic examination of

the section. In the first place, the primary outlines of openings

numbered 5 to 9 agree in their dimensions very closely with

the correspondingly numbered openings of Fig. r; and in the

second place, the evidences of erosion are abundant. Within

the primary shell between openings 9 and 11 there is a fine

wavy black line (4) that was evidently at the surface of the body

whorl during the stage of Fig. i. The distance between this

line and the primary outline of opening 9 is greater than the

distance between this outline and the present outer surface of

the whorl. The difference between these two distances gives a

I 76 THE AMERICAN NATURALIST. [Vor. XXXVII.

measure of the amount of erosion that has taken place since the

time when the wavy line was covered by the present body whorl.

Similar wavy lines are seen below openings 8 and 7 and there

the differences are greater, showing an increased amount of

erosion toward the apex, and the loss by erosion is still greater

at openings 6 and 5 where the secondary shell is exposed at the

surface.

The same thing is shown more clearly in Fig. 3, a normal,

nearly full grown shell about 15 mm. long. Between openings

Fig. 2. Similar section of a somewhat larger shell. X 16.

II and 13 there is a wavy line that forms a continuous curve

with the surface of the shell at r1. Here the erosion has pro-

ceeded only so far as to smooth off the ribs, while below openings

10 and 9 the wavy lines end abruptly almost at right angles to

the surface, and the exposed secondary shell at these openings

furnishes further evidence of erosion. At opening 9 the second-

ary shell itself is entirely removed on the outer side. But the

lumen of the shell is prevented from opening to the exterior by

a thick layer of zerziary shell, which has been deposited within

No. 435.] SHELL OF LITTORINA LITOREA. 177

the secondary and exhibits lines of growth parallel to its own

surface and unconformable with those of the secondary shell.

All of the openings above 13 in this specimen show a layer of

tertiary shell, which becomes progressively thicker toward the

apex. The numbers assigned to the openings in this specimen

are obtained by comparison with Fig. 2, and, if the reasoning

g. 3. Section of a nearly full-grown shell in good condition. The lumen is cut ten times but

only seven of the openings are shown in the drawing. X 16.

applied to that figure will hold for this one, the first five openings,

that is two whorls and a half, have disappeared entirely.

Fig. 4 is from one of the largest of the specimens collected,

about 20 mm. long. But somewhat larger shells are not rare.

To the naked eye the rounded apex and white color of the apical

region in this specimen indicated a considerable but not excessive

amount of erosion. The difference in color is shown by the

section to be due to the exposure of a large amount of tertiary

178 THE AMERICAN NATURALIST. [Vor. XXXVII.

shell, which is colored only in the inner layers. Comparison of

this section with Fig. 2 indicates that the two upper whorls of

this specimen contain openings 8, 9, ro, and 11, as shown in

Fig. 6. The uppermost whorl has almost disappeared and the

second one has lost the entire outer wall of both primary and

secondary shell. So that of the shell formed during the stage

of Fig. 2 there remains in this specimen only the columella of

. the two lower whorls and a portion of the lip region. While of the

stage of Fig. 1 there remains only the merest fragment of the

body-whorl (Fig. 5).

Fig. Camera drawing of the upper part of a spon Mani a shell about 17 by 17 mm. In the

whole section the lumen is cut only nine times. X 1

The actual shortening of the shell and the changes in shape

due to erosion cannot be measured directly. But we can estimate

them by comparing sections of younger and older stages, as in

Figs. 5 and 6. By this method it is estimated that between

the stages of Figs. 1 and 2 there has been a shortening of about

0.25 mm. In Fig. 3 this has increased to 0.6 mm. and in Fig.

4to r.4 mm. Or considering Fig. 1 as the initial stage of all

the larger shells, the length of that portion of the shell has been

reduced in the three subsequent stages by 8.3%, 20.0%, and

No. 435.] SHELL OF LITIORINA LITOREA. 179

46.7% respectively. This amounts to a reduction in the total

length of the shell at each stage of 5%; 4% and 7% respec-

tively.

Assuming that, if there were no erosion, the outline of the

section at the stage of Fig. 4 would be represented approximately

by the combined continuous and dotted outlines in Fig. 5, it is

Fig. 5. Outline of Fig. t placed on Fig. 4 so that the corresponding openings coincide.

evident that there has been a considerable increase in the magni-

tude of the apical angle. This would be shown still better by

combining Figs. 1 and 6. The difference between the angles

of Fig. 1 and Fig. 4 is 18°, or 19% of the apical angle of Fig. 4.

The loss of height in the shell of Fig. 4 is estimated at about

7%, while there is no evidence of any corresponding decrease in

180 THE AMERICAN NATURALIST. [Vor. XXXVII.

the breadth of the body-whorl. So the ratio of breadth to length,

or the ventricosity of the shell, is a ratio of a slightly modified to

a much modified character. The actual length of the shell in

Fig. 4 is about 20 mm., its breadth, 17 mm., giving a ratio of

85%. But without erosion the length is estimated to have been

21.4 mm. giving a ratio of 79%. Thus the shell is now more

Fig. 6. Oatline of Fig. 2 placed on Fig. 4 so that corresponding openings coincide.

ventricose by 6 degrees. In Fig. 3 the length is r5 mm.,

breadth, 13 mm., ratio, 8775; but the length without erosion

would be about 15.6 mm. and the ratio would be about 83 46,

an increase of ventricosity of 4 degrees.

Thus within the limits of our studies we find that in all stages

of the growth of the shell of Z. Zforea there has been a change

No. 435.] SHELL OF LITTORINA LITOREA. 181

in both size and shape resulting from erosion. While the absolute

amounts of these changes seem small, they are relatively large

enough to falsify any comparisons that may be made between

the variabilities and types at different localities or at different

ages, unless a correction for erosion can be applied.

For our measurements would show, not the normal variability

of growth, but the combined effect of variability of growth,

variability in power of resistances to erosive agents, and perhaps

variability of these agents themselves. To obtain data for the

correction of our results would involve an investigation of the

rate of erosion that would be too laborious to be undertaken.

Our attention was drawn to this species especially by the very

interesting work of Bumpus (98), who made a comparison of

the variability and types in shells from three English localities

with shells from ten stations on the New England coast. He

concluded that in all the characters studied the shells from all

the American localities (that is, those in the comparatively new

environment) were more variable than those from any of the

English localities. And he stated also that the American type is

* more enlongated, ! lighter in weight, more bulky, and the color

markings are less pronounced.”

His tests of variability were:— (1) Ratio of breadth and

length, (2) Comparison of the extremes of this ratio, (3) Com-

parison of curves for different ages, (4) Weight, (5) Bulk, and

(6) Color. Our results show that the first five of these tests of

variability are rendered of doubtful value by the erosion. Or at

any rate, the factor of erosion must be taken into account in the

discussion of these results, for, as has been shown, the ratio of

breadth to height is very considerably affected by the loss in

height, the difference is the ratios amounting to as much as .05

or .06 ; while the total range of variation found by Bumpus was

.17 in the British and .24 in the American series. And the dif-

ference between the means of the two series was only .o14, much

less than the probable effect of erosion in an apparently normal

shell such as is represented in Fig. 3.

! This e an eint for the average of the English mean ratios of breadth

to height is 89.6 and the average of the American mean is 91. That is the

American type is more ventricose.

182 THE AMERICAN NATURALIST. (VoL. XXXVII

It is in a comparison of ratios in shells of different ages, such

as Bumpus made, that the effect of erosion would be most seri-

ously felt. He found an increase of ventricosity in six localities

and a decrease in seven. Doubtless the former would have been

very much diminished and the latter augmented in the absence

of erosion.

Weight and bulk are also affected by the combination of

erosion without and deposition within. Color is the only

remaining test, and that is rather an unsatisfactory measure of

variability in this species.

Various explanations have been offered to account for the

erosion of molluscan shells, which according to Cook (’95) is

much more common among fresh-water forms than in those

inhabiting the sea. Many years ago Lewis (’59) pointed out

that the shells in a stream near Mohawk, N. Y., below the point

where alkaline wastes were discharged into the water were much

eroded, while above the point of pollution they were quite healthy

and free from defect. Jeffries (65) quotes the suggestion of

J. R. Grove that the patches and irregular hollows on the surface

of Z. /itorea may be due to electrolytic action arising from lack

of homogeneity in the shell substance. Schrubsole (86) found

that certain English fresh waters containing from one half to

four grains of lime per gallon had a strong erosive action on

shells, while no such action was noticed in water containing eight

and one third grains.

That filamentous plants penetrate many shells of molluscs and

brachiopods, as well as numerous corals and other forms, was

pointed out by Kölliker (59). Later Bornet and Flahault ('89)

summarizing the results of previous observers, described and

figured eight genera of algæ and two fungi having the shell-

boring habit, and they pointed out that these plants play an

important part in the destruction of shells, especially in quiet

waters. Recently, Duerden ('o2) has drawn attention to the

action of algae in the disintegration of corals in the West Indies.

We have not been able to discover however, any evidence of the

presence of algz or other plants upon the shells of Littorina

collected during the winter months, and have no explanation to

offer as to the cause of the erosion that we have observed.

No. 435] SHELL OF LITTORINA LITOREA. 183

This is, nevertheless, so great that we are reluctantly forced

to the conclusion that Lztterina Litorea, which on account of its

comparatively recent migration to our shores, and its great

abundance and variability seemed an especially favorable object

for statistical study, is after all not suitable for this purpose.

In order to use this material we must either assume that the

causes of erosion have a uniform intensity for all individuals, or

else we must devise some easy method for determining the actual

loss of substance in each individual. But even these precautions

would not enable one to use this material for the solution of the

special problem that we had in view.

THE BIOLOGICAL LABORATORIES.

MASSACHUSETTS INSTITUTE OF "TECHNOLOGY.

January, 1903.

184 THE AMERICAN NATURALIST. [Vor. XXXVII.

BIBLIOGRAPHY.

':89 BonNET E. and FLAHAULT, C. Sur quelques plantes vivant dans le

test calcaire des mollusques. Bul. Soc. Bot. de France. Vol. 36.

pp. cxlvii-clxxvi, pl. vi-xii.

'98 Bumpus, H. C. Variations and mutations of the introduced Litto-

rina. Zool. Bulletin. Vol. 1, pp. 247-259.

'95 CooKE, A. H. Molluscs. London. Macmillan. p. 276

'02. DUERDEN, J. E. Boring Alge as agents in the disintegration of

corals. Bull. Amer. Mus. Nat. Hist. Vol. 16, pp. 323-332.

'86 GaNnonGc, W. F. Is Littorina litorea introduced or indigenous?

Am. Nat. Vol. xx, pp. 931-940

'44 GOULD, A. A. Report on the Invertebrata of Mass. Cambridge.

"JO GOULD, A. A. Report on the Invertebrata of Mass. Ed. by Binney.

Boston

'65 Jeyeutya, Í. G. British conchology. Vol. 3.

'59 KOLLIKER, A. On the frequent occurrence of vegetable parasites in

the hard structures of animals. Proc. Royal Soc. Vol. 10, pp. 95-

99.

'59 Lewis, J. Erosion in shells. Proc. Bost. Soc. Nat. Hist. Vol. 6,

149.

'80 Morse, E. S. Gradual dispersion of certain molluscs in New Eng-

land. Bull. of Essex Inst. Vol. xii, pp. 171-176.

':86 SCHRUBSOLE, G. W. Erosion in certain fresh water shells. Jour. of

Conchology. Vol. 5, p. 66.

°0l1 WELDON, W. F. R. A first study of natural selection in Clausilia

laminata (Montagu). Biometrika. Vol. i, pp. 1

':80 VERRILL, A. E. Rapid diffusion of Littorina littorea on the New

England coast. Am. Jour. of Science. Vol. xx (series 3), p. 251.

THE SENSE OF HEARING IN FISHES:!

G. H. PARKER.

THE sense of hearing is unusual in several respects. Unlike

the other senses, it is restricted to comparatively few groups of

animals ; for, though many experiments have been tried, there

is no conclusive evidence, so far as I know, that a sense of hear-

ing is possessed by coral animals, jellyfishes, worms, starfishes,

crabs, oysters, snails, and their allies. It is true that the older

naturalists described for many of these animals organs that they

called ear-sacs and that were supposed to act as organs of hear-

ing, but the experimental work of the last fifteen years has

shown that these organs are without doubt means of controlling

the equilibrium of the animals, and not organs of hearing. The

only animals in which a sense of hearing may be said without

reservation to be present are the higher arthropods, particularly

the insects, and the four higher classes of vertebrates, the

amphibians, reptiles, birds, and mammals. Excepting the arthro-

pods and the vertebrates, it seems probable that the other ani-

mals cannot hear, that while they may be influenced by contact

with the world about them, by its light, its odors, etc., they are

uninfluenced by its sounds; in other words, they live as though

surrounded by perpetual silence. |

The sense of hearing is not only restricted to a very few

groups of animals, but the animals possessing it are always the

more highly organized members of their groups. Thus, among

the arthropods, a group which includes the crabs, lobsters, myri-

apods, spiders, and insects, the sense of hearing, if not absolutely

restricted to the most highly organized class, the insects, is at

least best developed in them. So, too, in the vertebrates, though

the frogs, toads, turtles and their like have a sense of hearing,

the efficiency of this sense is low compared with that which it

attains in the birds and particularly in the mammals, the highest

! A lecture delivered before the Department of Zoólogy of the Brooklyn Insti-

tute of Arts and Sciences, March 5, 1903.

185

J 86 THE AMERICAN NATURALIST. (VoL. XXXVII.

vertebrates. It is thus evident that the sense of hearing is

coupled with high organization.

When a survey of the whole animal kingdom is made, it is

found that most sense organs are not restricted as the organs of

hearing are ; for instance, eyes, often of a complicated structure,

occur in such lowly organized animals as jellyfishes, starfishes,

and worms, and these animals react with great precision to light.

Not only do many of these lower animals possess eyes, but they

also have organs of touch, taste, etc. ; in fact, the only important

sense organ lacking in them is that of hearing. Since the sense

of hearing is found only in the more highly organized animals

and the other senses occur in the lower as well as in the higher

animals, it follows that hearing is probably the most recently

acquired of the senses.

If in accordance with these facts we endeavor to form some

idea of the evolution of the sense organs in the animal series, we

must picture to ourselves an early origin for all the important

senses except hearing. This sense was undoubtedly the last

important one to be differentiated and the reason for the lateness

of its appearance I hope to make clear to you toward the close of

this lecture.

It is a remarkable fact that the ear in the higher vertebrates,

probably their latest important sensory acquisition, is an unusu-

ally perfect mechanism. Much has often been said about the

perfection of the human eye and this organ undoubtedly dis-

plays a marvelously delicate construction, but in my opinion the

efficiency of the ear as an organ of sense is as much beyond that

of the eye as a modern chronometer is beyond an old-fashioned

sundial. Evidence of the truth of this opinion can be seen in

the many defects that are present in the eye as contrasted with

the ear. For instance, what we call white light is well known

to be any one of many possible mixtures of colored lights.

Thus, when red light is combined with bluish green, the result-

ant is white; orange and sky-blue likewise produce white, as do

also yellow and violet, green and pink, etc. To all these com-

binations, which are totally distinct from a physical standpoint,

the eye answers with but one sign, that for white light.

Color in light corresponds to pitch in sound, for both depend

No. 435°] SENSE OF HEARING IN FISHES. 187

upon wave lengths. Thus, the keys of a piano when struck in

sequence give rise toa series of tones that differ one from

another much as the colors of the spectrum do, namely, in the

lengths of their waves. Although atrained eye cannot distin-

guish between white lights made by mixing different pairs of

colored lights, even the unpracticed ear can distinguish between

any pairs of tones when sounded on the keyboard, and, while

the person may not be able to name the keys, the actual dis-

crimination is easily accomplished. Thus white light is a meas-

ure of the deception due to our eyes, a deception that the ear

ordinarily never gives rise to. Could we see the colors in

ordinary light as we hear the tones in sound, the work of the

most extreme French colorists would be dull in comparison with

reality. The eye, therefore, fails to give much information

about light that is obtained from the ear about sound.

The occurrence of well-developed organs of hearing only among

the insects and higher vertebrates suggests, since these animals

are air-inhabiting forms, that possibly the sense of hearing is

capable of development only in an organism surrounded by air.

To test this supposition one naturally turns to the nearest aquatic

relatives of the groups possessing this sense. With the insects

these relatives are probably the somewhat distantly related crus-

taceans, but with the vertebrates they are the fishes, a class

closely allied to the other members of the vertebrate group. If

the sense of hearing can originate only in air-inhabiting animals,

no traces of it should be found among fishes. If it can arise in

other situations, the fishes are probably the class in which its

beginnings in the vertebrate series are to be sought for. The

problem of hearing in fishes, therefore, is a general one dealing

with the possible origin of one of the most important senses of

vertebrates, a sense probably the most recently developed of all

and yet in many respects the most efficient.

Hearing in fishes can best be approached from the standpoint

of the human ear. The ear in man has long been known to be

composed of three parts, the external, the middle, and the inter-

nal ear. The external ear consists of a complicated fold of skin

known as the concha and a somewhat twisted tube leading

inward to the eardrum. The middle ear is a cavity in the head

188 THE AMERICAN NATURALIST. Vor. XXXVII.

lying immediately internal to the ear-drum and connected with

the mouth by the Eustachian tube. Through the cavity of the

middle ear a bridge of small bones, the ear ossicles, passes from

the ear-drum to the opposite wall of the middle ear where the

innermost ossicle abuts against the cavity of the interna! ear.

The internal ear, which is situated somewhat deeper in the head

than the middle ear, is a complicated fluid-filled sac with three

semicircular canals and a spirally twisted portion, the cochlea.

The nerve concerned with the sense of hearing ends in the walls

of this sac. When the ear is normally stimulated, the sound-

waves from the surrounding air beat against the ear-drum and

set it in vibration. These vibrations cause the chain of ossicles

to vibrate and thus the motion is handed on to the fluid of the

internal ear. This fluid vibrates in turn and by some process

not clearly ascertained stimulates the nerves on the walls of the

ear-sac. As is well known the deafness caused by injuries to the

external and the middle ear may often be relieved by various

mechanical contrivances, but injuries to the internal ear are of

an absolute kind and not open to amelioration, for the internal

ear, as its nerve connections show, is the true organ of hearing,

the middle and external ears being only means of conducting

sound-waves to the true sense organ.

The form of ear just described is found only in the higher

vertebrates ; in the lower ones this organ presents a somewhat

simpler structure. Thus, in the frog, there is no external ear,

but the ear-drum is exposed directly on the surface of the head

and the whole auditory apparatus consists of only the middle and

the internal ear. In fishes a still further reduction takes place

in that the middle ear as such is not developed. Thus the fishes

possess only the essential part of the organ of hearing, the

internal ear, and even this is in an altered form, for their ear-

sacs, though complicated in outline and usually provided with

three semicircular canals, lack almost all traces of the spirally

twisted part, the cochlea. ;

That fishes possess a sense of hearing seems to have been

generally admitted by the older observers. , Thus Isaac Walton,

in his “ Complete Angler” (1653), when asked if trout can see

at night replies “Yes, and hear and smell too. He then

No. 435.] SENSE OF HEARING IN FISHES. 189

describes some experiments made by Sir Francis Bacon to show

that sound can pass through water, experiments that led Walton

to crave the pardon of one whom he had laughed at for affirming

that carp would come to a certain place in a pond to be fed, at

the ringing of a bell or the beating of a drum. He thereupon

declares that * it shall be a rule for me to make as little noise as

I can when I am fishing, until Sir Francis Bacon be confuted,

which I shall give any man leave to do," and he finally resolves

* to advise anglers to be patient and forbear swearing, lest they

be heard, and catch no fish."

The internal ears of the higher fishes were also known to the

older observers. So far as I am aware they were first described

by Casserius in 1610 and were studied in some detail in the fol-

lowing century by Geoffroy, Scarpa, Comparetti and the cele-

brated British physician, Hunter. The attitude taken by many

of thesé early workers in the question of theability of fishes to

hear is well illustrated by a quotation from a paper on the organ

of hearing in fishes published by Hunter (1782). This paper

contains the following statement (1782, p. 383): “As it is evi-

dent that fish possess the organ of hearing, it becomes unneces-

sary to make or relate any experiment, made with live fish

which only tends to prove this fact; but I will mention one

experiment, to shew that sound affects them much, and is one of

their guards, as it is in other animals. In the year 1762, when

I was in Portugal, I observed in a nobleman's garden, near

Lisbon, a small fish-pond, full of different kinds of fish. Its

bottom was level with the ground, and was made by forming a

bank all round. There was a shrubbery close to it. Whilst

I was laying on the bank, observing the fish swim about, I

desired a gentleman, who was with me, to take a loaded gun, and

go behind the shrubs and fire it. The reason for going behind

the shrubs was, that there might not be the least reflection of

light. The instant the report was made, the fish appeared to be

all of one mind, for they vanished instantaneously into the mud

at the bottom, raising as it were a cloud of mud. In about five

minutes after they began to appear, till the whole came forth

again."

This passage shows very clearly that in the opinion of Hunter

I9O THE AMERICAN NATURALIST. [VOL XXXVII.

the internal ears of fishes, like those of the higher vertebrates,

are organs of hearing. Without further experimental evidence,

this view was accepted by the celebrated physiologist Müller

(1848, p. 1238) in his well-known chapters on the physiology of

the senses, and by many other eminent authorities such as Owen

(1866, pp. 342 and 346), Günther (1880, p. 116), and Romanes

(1892, p. 250). To these investigators the presence of the

internal ear, seemed, as it did to Hunter, sufficient ground for

assuming that fishes could hear.

Within recent years, however, this opinion has been called

in question or even denied. Some of the grounds for this

change of view may be stated as follows. The English zcólogist

Bateson (1890, p. 251) in some investigations on the sense organs

and perception of fishes, observed that the report from the blast-

ing of rocks caused congers to draw back a few inches, flatfishes

like the sole, plaice, and turbot to bury themselves, and pouting

to scatter momentarily in all directions. Other fishes seemed to

take no notice of the report. When the side of a tank containing

pollack or soles was struck with a heavy stick, the fishes behaved

as they did toward the report of the blasting. Pollack did not

respond, however to the sound made by rubbing a wet finger

on the glass window of an aquarium or to the noise made by

striking a piece of glass under water with a stone, provided

the means of producing the noise was not seen by the fishes.

Bateson concluded that, while it may be regarded as clear

that fishes perceive the sound of sudden shocks and concus-

sions when these are severe, they do not seem to hear the

sounds of bodies moving in the water but not seen by them.

Without knowledge of Bateson's observations, the Viennese

physiologist Kreidl (1895) carried out a series of experiments

with the view of testing the powers of hearing in the goldfish.

This species was chosen because of the ease with which it could

be kept in the laboratory and further because it is one of those

fishes that have long been reputed to come at the sound of a

bell. After an extended series of experiments Kreidl (1895, p.

458) concluded that normal goldfish never respond to sounds

produced either in the air or in the water, though they do react

to the shock of a sudden blow given to the cover of the aqua

No. 435.] SENSE OF HEARING IN FISHES. I9I

rium. To test whether such responses were dependent upon the

auditory nerves, Kreidl removed these nerves and the attached

ear-sacs from a number of goldfishes and subjected them to stim-

ulation by sound. In all cases they were found to respond pre-

cisely as the animals with ears did. Kreidl, therefore, concluded

that goldfishes do not hear by the so-called ear, but that they

react to sound waves by means of an especially developed skin

sense, or, to put the matter in other words, the goldfish feels

sound but does not Zear it (Kreidl, 1896, p. 581). This condi-

tion is not so difficult to imagine as at first thought it seems, for

we can not only hear sounds under water but we can also feel

them, as anyone can prove by placing his hand under water near

a loudly sounding body.

After Kreidl had reached his conclusion concerning goldfishes,

he was led to take up a specific case of the response of fishes

to the sound of a bell and an opportunity of doing this was

found at the Benedictine monastery in Krems, Austria. Here

the trout of a particular basin were said to come for food at the

ringing of a bell. Kreidl (1896, p. 583), however, found that

they would assemble at sight of a person and without the ringing

of the bell. If they were not then fed, they soon dispersed and

no amount of bell-ringing would induce them to return. If,

however, a pebble or small piece of bread was thrown into the

water, they immediately swam vigorously toward the spot where

the disturbance had occurred. Moreover, if a person approached

the basin without being seen and rang the bell vigorously, the

fishes did not assemble. From these facts Kreidl (1896, p. 584)

concluded that the assembling of the fishes was brought about

through sight and the skin sense and not through hearing, and

that the conclusion reached with the goldfish might be extended

to other kinds of fishes.

While the problem of the auditory function of the ears of

fishes was thus being investigated, a wholly different view as

to the functions of these organs had been gradually opened up.

Through the researches of Loeb (1888), Kreidl (1892), Bethe

(1894), and Lee (1898) it became clear that whether the ears of

fishes were auditory organs or not, they were, beyond doubt,

organs for the control of the equilibrium of these animals, and in

192 THE AMERICAN NATURALIST. (VoL. XXXVII.

this respect they partook of the nature of the otocysts of the

lower animals. My own few observations are fully in accord

with this conclusion. When the nerves to the ears of the green

killifish (Fundulus heteroclitus) are cut and the animal attempts

rapid locomotion, it loses its bearings completely and swims in

any position in spirals or even in circles. Thus the ear is in

some way essential to the continued equilibrium of the fish.

Although this conclusion has no direct bearing on the question

of hearing in fishes, it makes it no longer necessary to assume

that the presence of the internal ear in a given fish implies the

ability of this fish to hear; hence the argument used by the older

investigators is shown to be fallacious.

The conclusion arrived at by Kreidl that the ears of fishes

like the goldfish, trout, etc. are not organs of hearing "was

supported and extended by the observations of one of our

American physiologists, Dr. Lee of Columbia University, who

studied the reactions of several species of fishes to such sounds

as the human voice, the clapping of hands, and the striking of

stones together in air and under water. In all his experiments

Lee (1898, p. 137) obtained no evidence whatever of the exist-

ence of a sense of hearing as the term is usually employed ;

though he found that the fishes were exceedingly sensitive to

gross shocks, such as the jarring of their tank or concussions

upon its walls. From the observations and experiments of

Bateson and of Kreidl and from his own work, Lee (1898, p.

138) believed that the conclusion is justified beyond doubt that

fishes do not possess the power of hearing, in the sense in

which the term is ordinarily used and that the sole function of

the ear in fishes is equilibration. According to this view then,

fishes resemble many of the lower aquatic animals, in that their

so-called ears are not organs for hearing, but for controlling the

equilibrium of their bodies; and, if they respond to sounds at

all, they do so through the skin.

This general conclusion seemed to me not wholly in accord

with certain well-known facts in the natural history of fishes.

Most important of these is the undoubted ability on the part of

some fishes to make sounds. Although it is conceivable that

fishes, like some totally deaf persons, may produce vocal sounds

No. 435.] SENSE OF HEARING IN FISHES. 193

that they themselves cannot hear, this conclusion is not probable

and anyone that has ever heard a young swellfish (Chilomycterus

schoepfi) make its characteristic sound when attacked by a

hungry scup (Stenotomus chrysops) cannot but receive the

impression that both fishes hear. Moreover it is very difficult

without assuming hearing to understand the economy of sound

production where, as in the squeteague or weakfish (Cynoscion

regalis), this function is limited to one sex, in this instance the

males. These habits raise the suspicion that notwithstanding

the experimental evidence thus far brought forward, fishes may

hear. It was chiefly because of this suspicion that last summer

at the invitation of the United States Fish Commission I under-

took to investigate this question.

Fig. 1.

The apparatus used (Figs. 1 and 2) consisted of a large

marine aquarium one end of which was replaced by a deal sound-

ing board. On this board a bass viol string was stretched so

that its vibrations could be transmitted through the board to the

194 THE AMERICAN NA TURALIST. [Vor. XXXVII.

water in the aquarium. By frequent adjustment the string was

kept at forty vibrations per second, corresponding to.a tone

almost as low as any used in music. Within the aquarium was

suspended from distant supports a small glass cage for the reten-

tion of the fishes to be experimented upon. The end of the

cage toward the sounding-board was open except for a fine net,

which, though it restrained the fishes from escaping from the

cage, made no serious interference with the entrance of sound.

Fig. 2.

In testing fishes for the sense of hearing it must be evident

from what has already been said that several precautions are

necessary. First of all, the sounds used should be sustained

and of relatively low intensity rather than of the kind that might

produce a concussion or a shock. Next, care must be taken to

determine whether these sounds influence the fish through its

ears or through its skin; for if the ears are not stimulated, a

sense of hearing cannot be said to be present. After some pre-

liminary trials I finally resolved to use for my experiments the

green killifish (Fundulus heteroclitus), a minnow common in the

No. 435.] SENSE OF HEARING IN FISHES. 195

waters about the United States Fish Commission Station at

Woods Hole, Massachusetts, where my work was done. This

fish proved very hardy and in every way satisfactory for the

work.

Three classes of killifishes were tested; first, those that were

entirely normal; secondly, those in which the nerves to the ears

had been cut; and, thirdly, those whose outer surfaces had been

rendered insensitive by cutting the nerves to the skin. The fishes

were easily etherized and cutting the nerves to the ears proved to

be a simple though delicate operation. To render the skin insen-

sitive, it was necessary to cut the fifth and seventh nerves, the

lateral line nerves and the spinal cord, but from this apparently

severe operation the fishes almost invariably recovered, fed well,

and continued normally active for several weeks.

When a normal fish was placed in the cage and the bass viol

string set in vibration, any one of four responses might be

observed: first, the rate of the gill movements increased for a

brief period; secondly, the pectoral fins, if quiet, were set in

motion, or, if in motion, their rate was increased; thirdly, the

tail fin often vibrated ; and, finally, the whole fish might give a

spring as if startled by the sound. Of these four reactions, the.

one most satisfactory for observation was the movement of the

pectoral fins, and I subsequently used this almost exclusively in

testing the fishes. Ten normal fishes when subjected each to

ten tests with the sounding apparatus responded with the pec-

toral fins ninety-six times in the total hundred. This number

may be taken as a basis of comparison for the fishes upon which

operations had been performed.

To ascertain the importance of the ear in these responses, I

next tested ten fishes in which the nerves to the ears had been

cut and from ten observations on each fish I found that the

number of responses to sound in a total of one hundred trials

was only eighteen, and some of these eighteen were of doubtful

character. It thus appears that cutting the nerves to the ears

brings about a large reduction in the number of reactions and

my results, therefore, are very different from those of Kreidl,

who, it will be remembered, observed no difference in this

respect between normal and earless goldfishes. Although at

196 THE AMERICAN NATURALIST. [Vou. XXXVII.

first sight it might seem that these experiments proved conclu-

sively that the ear of the killifish is, if not the only organ of

hearing, at least the chief organ for that sense, it is nevertheless

conceivable that the reduction in the number of reactions shown

by the earless fishes may be due not to the loss of the ear as a

sense organ but to the severity of the operation that the fishes

have undergone.

To determine how much weight should be given to this objec-

tion and at the same time to ascertain the part played by the

skin in these reactions, I tested ten fishes in which the skin had

been made insensitive but in which the ears were intact. Ina

total of one hundred trials, ten on each fish, there were ninety-

four responses to sound and six failures; in other words, these

fishes, though they had undergone more severe operations than

the earless ones, reacted almost exactly as normal fishes did.

These observations placed beyond a doubt the conclusion that

the ears of the killifish are stimulated by the disturbances set up

by the vibrating bass viol string, and the only question we have

to answer before a final conclusion can be safely drawn is,

whether these disturbances are really sound vibrations.

If the aquarium is observed closely when the bass viol string

is made to vibrate, not only will a sound be heard but the whole

aquarium will be seen to vibrate and ripples will be noticed pass-

ing over the surface of the water. Is it not possible that the

fishes respond to the motion of the aquarium as a whole or to

the disturbance indicated by the ripples rather than to the true

sound-waves? To determine this point, I substituted for the

bass viol string an electric tuning fork that gave a tone of 128

vibrations per second. The fork was so placed that its base

could by a slight movement be brought into contract with the

sounding-board and thus, without jar or disturbance to the aqua-

rium, the sound could be delivered to the contained water.

On testing fishes under these conditions, it was found that the

earless ones never responded to the tones from the tuning fork,

while those with ears very usually did respond. I, therefore,

believe that I am entirely justified in drawing the final conclu-

sion that the ears of the killifish are stimulated by sound-waves,

that is, that this fish hears. It will be recalled that in the

No. 435.] SENSE OF HEARING IN FISHES. 197

experiments with the bass viol string the earless fishes responded

eighteen times in the hundred trials. I believe that the tuning

fork experiments, in which no earless fishes responded, make it

probable that these eighteen responses were due not to the

stimulating effect of sound on the skin of the fishes, as might

be inferred, but to some other cause such as the trembling of

the whole aquarium. I do not wish to imply, however, that the

skin of a fish may not be stimulated by sound.

Although the conclusion that a fish hears is directly contrary

to that arrived at by Lee, it is not at all necessary to suppose

that the observations of this investigator and those of Kreidl

should be regarded as incorrect. Neither Kreidl nor Lee

worked on the killifish and the ears in this species may be dif-

ferent from those in the fishes studied by these two investigators.

In fact, in my own work I tried on the smooth dogfish (Mustelus

canis) the same experiments as those that I have just described

for the killifish, but without obtaining the least evidence of hear-

ing in this species. I am, therefore, quite prepared to believe

that there are fishes in which the sense of hearing is undeveloped,

and these may have been the very forms with which Kreidl and

Lee worked; but that there are fishes that do hear I feel per-

fectly certain.

The ear is related to other sense organs in a way unusually

well seen in fishes, and before closing I wish to call attention

briefly to this aspect of the subject. The sense organs con-

cerned are the skin and the lateral-line organs. Everyone is

familiar with the skin as an organ of touch, but the lateral-line

organs are less generally known for at least one obvious reason,

namely, that man possesses none. Lateralline organs occur

only in the true aquatic vertebrates, the fishes and the amphib-

ians, and in the latter only in those stages in which the animals

inhabit the water. Thus in the frog, lateral-line organs are pres-

ent while this amphibian is a tadpole, but, as soon as it takes

on the adult form and emerges from the water, these organs

disappear. Lateral-line organs, then, are in some way intimately

associated with the water habit.

These organs form regular lines on the skin of an amphibian

or a fish, though in the latter they are more frequently in grooves

198 THE AMERICAN NATURALIST. [Vor. XXXVII.

in the skin or in tubes that lie immediately under the skin and

whose courses are indicated by numerous openings, the lateral-

line pores. In many fishes three such lines of pores can be

recognized on the side of the head, one above the eye, a second

immediately below the eye, and a third on the lower jaw. From

the union of these three near the back of the head, a fourth line

passes posteriorly along the side of the fish, the lateral line, from

which the whole system takes its name.

Modern embryology has shown that the ear is closely related

to the lateral-line system. The deep situation of the ear-sac at

first sight seems to preclude this, but the ear-sac does not form

in this situation. In fishes, as in higher vertebrates, this sac

forms as a pocket of skin pushed into the head as it were from

the outer surface and in fishes it can be shown that the skin

which is thus infolded is a portion of the lateralline system.

After the sac has been formed, it sinks into the deeper part of

the head, generally loses its connection with the outer skin, and

gradually takes on its final complicated shape by producing semi-

circular canals, etc. Thus the internal ear may be regarded as

a modified part of the lateralline system. This system in turn

develops from the skin, and when its organs lie in tubes, as they

do in most fishes, the tubes are formed from groove-like depres-

sions of the skin. Thus the lateralline organs are specialized

sense organs from the skin.

These facts suggest at once interesting relations between the

three sets of organs mentioned ; for, as the lateralline organs

may be regarded as derived from the undifferentiated sense

organs of the skin, so the ear may be conceived to have been

derived from the lateral-line organs. Thus, we are dealing with

what may be called three generations of sense organs: the skin

representing the first generation, and giving rise to the lateral-

line organs, the second, which in turn produce the third, the ears.

This view of the relations of these three sets of organs accords

well with what I have been able to make out about their func-

tions in the killifish. It has already been shown that only such

fishes as have their ears intact respond to the sound of a tuning

fork. Consequently we may conclude that such sounds do not

stimulate either the lateralline organs or the skin, but that they

are a stimulus appropriate to the ear.

No. 435.] SENSE OF HEARING IN FISHES. 199

The lateral-line organs can also be shown to possess a stimulus

peculiar to themselves. These organs were originally supposed

to be merely slime glands for the production of the slimy cover-

ing so characteristic of the skins of fishes. In 1850 Leydig

demonstrated their sensory character, and since then they have

given rise to much speculation among naturalists. Since they

occur in fishes that have the usual five senses, Leydig (1868)

regarded them as organs of a sixth sense quite distinct from any

that we possess. Schulze (1870) thought they served to detect

mass movements of the water as when a current passes over the

surface of a fish or the fish swims through the water. Merkel

(1880) believed that they were simply organs of touch, and

Emery (1880) and P. and F. Sarasin (1887—1890) regarded

them as accessory ears. Fuchs (1895) brought forward evi-

dence to show that they were pressure organs, and Richard

(1896) believed they were in some manner connected with the

production of gas in the swim-bladders of fishes. Bonnier

(1896) was of opinion that among other things they oriented the

fish in reference to centres of disturbance in the water, and Lee

(1898) thought them organs of equilibration. Thus, much dif-

ference of opinion prevails as to the supposed function of these

organs.

In the killifish the stimulus for the lateral-line organs is easily

found. If an aquarium in which a normal fish is sporting is made

to undergo a slight inaudible oscillation by some means not visi-

ble to the fish, the latter will dart at once to the bottom. This

happens even when the oscillation is scarcely perceptible to the

observer. After the nerves to the lateral-line organs have been

cut, the fish will no longer respond to these slight movements,

but will continue sporting about even when the aquarium is made

to tremble considerably. The responses obtained from killifishes

under these conditions are so invariable that I conclude that their

lateral-line organs are stimulated, much as Schulze suspected, by

a slight but inaudible movement of the whole mass of water, a

movement that is too delicate to stimulate their skins.

The skin I found to be stimulated by the movements of the

water in surface waves and in currents. If a fish in which the

lateral-line organs have been rendered inoperative swims into a

200 THE AMERICAN NATURALIST. [Vor. XXXVII.

region where surface waves affect it, it will swim downward out

of the reach of these, though it does not go to the bottom of the

aquarium as a fish with lateralline organs does. If it is placed

in a current of water, it will swim as vigorously against the cur-

rent as a normal fish will. Thus the skin, though not stimulated

by sound from a tuning fork or the slight inaudible trembling of

the whole mass of water, is stimulated by currents and by sur-

face waves.

Hence the three sets of sense organs under consideration

may be regarded as having slightly different kinds of stimuli:

the skin being affected by surface waves and by currents; the

lateraldine organs by slight inaudible movements of the whole

mass of water; and the ears by the still more delicate vibrations

of water particles, sound. These three sets of sense organs,

therefore, are not only genetically connected in that they repre-

sent so to speak three generations of organs, but their evolution

has been toward more and more delicate means of stimulation.

From this standpoint the lateral-line organs of the fishes and the

amphibians may be said to be delicate organs of touch and even

the ear as an organ of hearing may be looked upon as an exqui-

sitely refined apparatus of much this same kind. Hearing, then,

is a most delicate form of touching, and the organ of hearing has

developed late in the animal series because its processes are not

original but are derived from those of the more primitive sense,

touch. Many fishes possess at once the complete series of sense

organs leading from touch to hearing, and in these animals the

activities dependent upon such organs form doubtless a more or

less homogeneous whole; but with us touching and hearing are

very distinct things, a condition, however, that we can easily

understand, for we have lost the intermediate sense, that of the

1 At first sight it might seem that this view ignored completely the equilibration

function of the ear, a function which may be the only one possessed by this organ

in some of the lower fishes, but there are no facts so far as I know that prevent

one Ron assuming that the stimulus for the lateral line organs may not be much

the same as that for the ear as an equilibration organ and thus this function

might indicate a sensory activity somewhat intermediate between that of the

lateral line organs and of hearing. At least unless this is shown not to be the

case, the equilibration function of the ear offers no real obstacle to the acceptance

of the opinion expressed above as to the physiological relations of the skin, lateral

line organs, and ear.

No. 435.] SENSE OF HEARING IN FISHES. 20I

lateral-line organs. Thus the consideration of the sense of hear-

ing in fishes leads us to an understanding of a natural group of

sense organs whose genetic connections would never have been

suspected had we not been able to investigate them in these primi-

tive aquatic vertebrates, the fishes.

Postscript. Since this lecture was prepared Professor Tull-

berg’s interesting paper (1903) on the functions of the ears of

fishes has reached me. In this paper it is concluded (p. 20)

that the ears of fishes are not organs of equilibration and though

they may perhaps be to a certain extent organs of hearing, their

original and principal function is to receive stimuli from the

movements of the surrounding water, especially from currents.

Vibrations probably stimulate particularly the maculæ acusticæ

of the utriculus, the sacculus, and the lagena; currents affect

especially the cristæ acusticæ of the ampullæ. Thus the ear is

held to be an organ directly concerned with the reactions of a

fish to a current of water (rheotaxis).

My own experiments have been directed to test the sense of

hearing in fishes rather than to ascertain what other functions

the ears in these animals may have, but some of my observations

bear on the questions raised by Professor Tullberg and I, there-

fore, call attention to them here. Professor Tullberg believes

that the ears of fishes are not concerned with equilibration

because the disturbances in the equilibrium of fishes that follow

many operations on the ears and that have been taken to indi-

cate this function, sooner or later disappear, and must, therefore,

be regarded as shock effects and not due to the loss of a special

sense organ. I have observed much the same condition in Fun-

dulus heteroclitus as that described by Professor Tullberg in

that two or three days after the eighth nerve has been cut

the fishes often swim quite normally. If, however, they are

made to swim very rapidly, they invariably lose all orientation

and move in irregular circles and spirals. I, therefore, believe

that the loss of the ear may have a /asting effect on the orien-

tation of fishes in certain forms of locomotion, for these irreg-

ular movements were always observable in earless Fundulus

even up to the time of death, in some instances six weeks after

the cutting of the eighth nerve. The apparent recovery after

202 THE AMERICAN NATURALIST. (Vor. XXXVII.

the operation I attribute to the increased use of the eye as an

organ for orientation and though this is an assumption on my

part, it is one well supported by evidence from the invertebrates,

and, until it is shown to be false for fishes, it seems to me to

make Professor Tullberg's conclusion that the fish ear has noth-

ing to do with equilibration at least premature.

My observations on the lateral line organs of Fundulus

(Parker, 1903, pp. 59-62) are in entire agreement with those of

Professor Tullberg (1903, pp. 8, 15) on these organs in other

fishes in that I have found no evidence that these organs are

essential to the normal swimming of a fish against a current.

But since in Fundulus I could not persuade myself that the ear

was not in some degree connected with equilibration, I was

unable to devise a satisfactory experiment that would test in the

absence of the ear, other organs of sense, such as the skin, that

might be stimulated by water-currents. The only evidence I

was able to obtain was that when fishes whose spinal cords and

lateral-line nerves had been cut several days before, were held by

the head near a gentle current of water the tails bent toward

the current even though the action of the current on a cordless

fish similarly held was to force the tail in the opposite direction.

I am, therefore, entirely certain that the skin of the trunk of

Fundulus heterochtus is stimulated by water-currents, though

I cannot say that these may not also stimulate the ears. I must

confess, however, that Professor Tullberg’s evidence (1903, pp.

II, 14,) on this point, namely, that after cutting both horizontal

semicircular canals the fish no longer orients to a current, does

not seem to me wholly conclusive, for this operation may bring

about the observed condition by a slight interference with the

equilibration function of the ear. In Fundulus swimming against

a current of water is dependent in my opinion upon at least two

factors: first, the retention of the normal position of the fish

which is chiefly accomplished, I believe, by means of the ears

as equilibration organs ,and, secondly, directive locomotion which

is dependent upon the stimulating effect of the current on the

skin of the fish. Under these conditions an interference with

the ear might well give rise to a loss of rheotaxis though the

primary stimulus for this form of response might be received by

No. 43:.] SENSE OF HEARING IN FISHES. 203

the skin. Since the evidence that I have gathered from Fun-

dulus seems to me to show conclusively that this fish uses the

ears as organs of equilibration and since the skin in this species

is stimulated by water-currents, I believe that the rheotaxis of

this species probably is primarily dependent upon the skin,

though it may be profoundly influenced by impulses originating

in the ear either through a disturbance of equilibrium, as I have

already suggested, or possibly directly, as Professor Tullberg

believes.

BIBLIOGRAPHY.

1890 BarrsoN, W. The Sense-organs and Perceptions of Fishes; with

Remarks on the Supply of Bait. Jour. Mar. Biol. Assoc. United

Kingdom, new ser., vol. i, pp. 225-256, pl.

1894 BETHE, A. Ueber die Erhaltung des aaga: Biol. Cen-

tralbl., Bd. 14, pp. 95-114, 563-582.

1896 BoNnNIER P. Sur le sens latéral. C. R. Soc. Biol., ser. 10, tom. 3,

PP- 917-919.

1880 Ewrnv, C. Le Specie del Genere Fierasfer nel Golfo di Napoli e

Regiori limitrofe. Fauna u. Flora Neapel. Monographie 2.

Leipzig, 76 pp., 9 Taf.

1895 Fucus, S. Ueber die Function der unter der Haut liegenden

Canalsysteme bei den Selachiern. Arch. ges. Physiol. Bd. 59, pp.

454—478, Taf. 6.

1880 GüNTHER, A. C. L. G. An Introduction to the Study of Fishes.

Edinburgh. 8°, xvi + 720 pp.

1782 Hunter, J. Account of the Organs of Hearing in Fish. Phil.

Trans. dn Soc. London, vol. 72, pp. 379-383.

1892 KREIDL, A. Zur cepa rane - des Ohrlabyrinthes.

Near Centralbl., Bd. 11, pp. 2

189 Ueber die Perception der Sean bei den Fischen. Arch.

ges. Physiol., Bd. 61, pp. 450-46

1896 .— — in weiterer Versuch iba das angebliche Hören eines

Ein

Glockenzeichens durch die Fische. Arch. ges. Physiol. Bd. 63,

pp. 581-586.

1898 Lrr,F.S. The Function of the Ear and the Lateral Line in Fishes.

Amer. Jour. Physiol., vol. i, pp. 128-144.

1850 Lrypic, F. Ueber die Schleimkanüle der Knochenfische. Arch.

Anat. Physiol., wiss. Med., Jahrg. 1850, pp. 170-181, Taf. 4, Fig. 1-3.

1868 .— — Ueber Organe eines sechsten Sinnes. Dresden, 4°, 108 pp.,

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1888 Lorn, J. Die Orientirung der Thiere gegen die Schwerkraft der

Erde. Sitzb. phys.-med. Gesell. Würzburg, Jahrg. 1888, pp. 5-10.

1880 MERKEL, F. Ueber die Endigungen der sensiblen Nerven in der

Haut der Wirbelthiere. Rostock, 8°, 214 pp., 15 Taf.

1848 MiLLeR, J. The Physiology of the Senses, Voice, and Muscular

Motions, with the Mental Faculties. Translated by W. Baly.

London, 8°, xvii + pp. 849-1419 + 32 + 22 pp.

1866 Owen, R. On the Anatomy of Vertebrates. Vol. I. London, 8°,

xlii + 650 pp.

1903 PankER, GG. H. Hearing and Allied Senses in Fishes. Bull. U. S.

Fish Comm., 1902, pp. 45—64, pl. 9

1903* — — Sense of Hearing in Fishes. Science, new ser., vol. 17,

P- 243.

1896 RICHARD, J. Sur les functions de la ligue latérale du Cyprin doré.

. R. Soc. Biol, ser. 10, tom. 3, pp. 131—

1892 ROMANES, G. 5 Animal Intelligence. Internat. Sci. Ser, No.

44. New York, 8°, xiv + 520 pp.

1887-1890 SanasiIN, P., und SARAsIN, F. Zur Entwicklungsgeschichte

und Anatomie der ceylonesischen Blindwühle Ichthyophis cum,

Ergeb. naturwiss. Forschung Ceylon. Bd. 2, 263 pp., 24

1870 SCHULZE, F. E. Ueber die Sinnesorgane der Seitenlinie bei

Fischen und Amphibien. Arch. mikr. Anat., Bd. 6, pp. 62- 88, Taf.

4-6.

1903 TULLBERG, T. Das Labyrinth der Fische, ein Organ zur Empfin-

dung der Wasserbewegung. Bihang till K. Svenska Vet.-Akad.

Handlingar, Stockholm, Bd. 28, No. 15, 25 pp.

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Recreation. London

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Baldwin’s * Fragments." ! — The Fragments are twenty-four phil-

osophical and psychological papers which appeared originally in Zhe

Psychological Review, The Presbyterian Review, The New York Inde-

Pendent, and other magazines. The author accounts for their

republication in book form by the fact that they relate to larger topics

on which he has published, or intends to publish, more extensively

in separate works.

For those who have not read these essays as they first appeared it

may be said that the author’s point of view is idealism, but an ideal-

ism whose chief aim is to interpret and to harmonize the empirical

sciences. ^ Philosophy is a new reading of science" (p. viii). This

idealism, moreover, finds the universe of science to be “a cosmos

which is not only true but also beautiful, and in some sense good,”

and “it is true and good decause it is beautiful” (p. ix). In fact

“the esthetic principle . . . . represents the point in our conceptions

where worth and truth coalesce and become one.” Thus truth, it

would seem, gets worth only when it is beautiful.

This proposition gives the secret of the whole book. ‘The ascetic

mind which desires only truth from science or philosophy, that is,

the person whose sole aim is to eliminate contradiction from his view

of the world, to make it strictly logically consistent, will get very little

from the Fragments. Professor Baldwin does not specially aim at

simplicity or precise definition, and often lets a metaphor bridge a

gap in the argument; as thus — *....no philosophy is true which

leaves out of its reckoning any degree on the arc which measures the

mutual relation between personality and nature " (p. 9).

From the same esthetic ease of attitude springs the most serious

defect of the philosophical essays, — the error of supposing that the

PSychological explanation of an idea solves epistemological problems

connected with the meaning of that idea. A renowned example of

this is, of course, Hume's explanation of causality, as the case in

E Baldwin, J. M. Fragments in Philosophy and Science. New York, Scribner's

Sons, 1902. 8vo. pp. xii, 389.

* The italics are Prof. Baldwin's.

205

206 THE AMERICAN NATURALIST. (VoL. XXXVII.

which one thing seems to cause another because the idea of the first

readily calls up in the mind the idea of the second.’ ‘I'he same error

is often made in these essays. Thus in discussing The Cosmic and

the Moral, Professor Baldwin says “The sense of ought, then, from

my point of view, is the anticipation of more experience, not yet

treated under the rubrics of description”; (p. 73). “....I think

the matter described by ‘is’ is the inadequate content of that which

we feel ‘ought’ to be; and the description of what ‘oughted’ to be,

i. e., that which was the object of description of a part ‘ought,’ is

what ‘is.’ In short, the ‘ought’ is a function of a mental content, of

a descriptive ‘is, — a motor function, I think.” If such a solution

really touched the moral problem, it would follow that what one

expects to do is what one ought to do; and any man’s virtue would be

his shrewdness at prognostication. That Professor Baldwin really

means this psychologically which he thus substitutes for ethics, is

clearly shown by the phrase “ motor function.”

Similarly the author writes: “....we rationalize nature" (p. 141),

and again: “ Now what we mean s by ‘reality’ is just a group of

experiences normally organized in a certain way; and we believe in

realities when we recognize this tendency of our experiences to fall

into certain characteristic forms of organization. We do the organiz-

ing, and so assert the reality as being there to be organized” (p.

341). Such a view is just now often expressed both in this country

and England, but when analyzed it smacks of Hume’s error, and is in

fact the same, above-mentioned psycho-epistemological fallacy. That

(psychological) process by which our ideas become organized is sub-

stituted as a solution for the epistemological problem of the organiza-

tion and significance of reality. It is strange that such a view can

be so widely entertained at precisely the moment when all scientists

are confining themselves in laboratories patiently experimenting, well

aware that they themselves neither know nor do anything except to

await the oracle.

Professor Baldwin’s essays in psychology which are reprinted in

the Fragments are not intended (with the exception of six short

papers which are purely experimental) for the professional psycholo-

gist. In method they are like the philosophical essays, and like them

necessarily sacrifice technical precision to the purposes of popular

exposition. ‘Thus concerning the logarithmic law of Lechner, one

reads, —“ This affords a groundwork .... which, in so far as the

! Of course Hume made here a second error as well, a circle, since the phrase

“calls up ” means itself nothing else than “causes.”

No. 435.] NOTES AND LITERATURE. 207

experiments are reliable and the sources of error known, is not to be

damaged by a hundred objections such as the a priori impossibility

of the measurement of psychic magnitudes” (p. 154), and so forth.

This will please the novice, but its flippancy will certainly mislead

him. If psychic facts cannot be measured, then they have not been

measured, and their measure is not given by the psycho-physic form-

ula. It therefore behooves the serious scientist to enquire either

what it is whose magnitudes the law does express, or else, what error

there was in the “a priori” reasoning which declared all measure-

ment of the psychic to be impossible. The flat contradiction must

not exist; above all it should not be exhibited for the delight and

mystification of the readers of Zhe Presbyterian Review.

Among the experimental papers the best is the well-known one on

Types of Reaction, reprinted from The Psychological Review. Here

Professor Baldwin and Mr. W. J. Shaw offer a different interpretation

of sensory and motor reaction times from that suggested by Lange,

the originator of those terms. Lange (and Wundt) believed that the

motor reaction time is shorter because for it the nervous paths are

subexcited beforehand and so prepared for swift discharge. Where-

as Professor Baldwin and Mr. Shaw prove that the motor reaction

time is in many persons not shorter, but longer than the sensory,

and that this latter is itself very variable, according as one or another

sense receives the stimulus. Thus the distinction is not between

motor and sensory but between the various sensory, reaction times

(visual, acoustic, olfactory, e/ ce¢.). The shortest reaction time is

given by that sense, which furnishes the images most used in the per-

son's thought, and which therefore gives habitually the cues for motor

discharges. Thus with a visualizer, visual stimuli will give the most

rapid reaction time; with an auditeur, auditory stimuli. The so-

called motor reaction is one given by a moteur, whose movements are

habitually initiated by motor images: to whatever sense the stimulus

is given, the impulse has to go to the motor-image-centres, and then

to the muscles. Thus its course is indirect, but since in a moteur it

cannot be shortened (that is, all his reactions are really motor), the

increased speed of discharge due to the subexcitation of the motor-

image-centres in the reactions which are avowedly motor will give

these the advantage in time over the reactions which are (errone-

ously) assumed to be sensory, that is, in which the motor centers are

not subexcited beforehand. Lange found the motor type of reaction

to be swifter, because presumably these authors say, he had mostly

moteurs to experiment on (the commoner type) and because he

208 THE AMERICAN NATURALIST. [VoL. XXXVII.

called ambiguous all results which gave a shorter sensory reaction

time. Professor Baldwin also very interestingly shows, in another

essay, that his theory fits in well with the facts of sensory and motor

aphasias. l

The other experimental papers are of no great importance. The

Fragments close with five short papers on miscellaneous subjects.

The volume is admirably printed and bound, but on the whole the

essays themselves are of such slight permanent value as hardly to

justify a second publication in this form.

Theoretical Biology. — Professor Reinke of Kiel has written ! an

introduction to theoretical biology which treats of the fundamental

biological problems in their metaphysical aspects. The work is thus

predominantly philosophical in its purpose and form. He discusses

vitalism and materialism, “ Naturphilosophie,” teleology, and adap-

tation, its content, origin and inheritance. The concepts of matter,

energy, force, law, and form in their relation to organization as seen

in the living world are discussed at length. The author introduces

the idea of dominants or “unbewusst intellegenten Kräfte” as sup-

plementary to the energy which operates by chemical processes the

highly differentiated mechanism of the living organism. Dominants

are not identical, in the author’s view, with the old vital force, but

like the mechanism have their basis in the structure of the organism.

The biological sections of the work all of which are correlated with the

fundamental problems discussed elsewhere deal with the attributes

and functions of protoplasm and the cell and with the development

and differentiation of organisms. Incidentally the problems of sex-

uality, fertilization and heredity are touched upon. In closing, the

author calls attention to the similarity of his views and those of

Johannes Müller. The illustrative matter is drawn largely from

botanical sources. The scope of the work and the clearness with

which it is written make the book of general interest and value.

1 Reinke, J. Linleitung in die theoretische Biologie. Pp. xv, 637. Mit. 83

Abb. in Text. Berlin, Gebrüder Pzetel. 1901. M. 18. —

No. 435] NOTES AND LITERATURE. 209

ZOOLOGY.

Goette’s Zoology.'— In its general appearance and make up this

work is of the familiar type. There are few novelties in it, and it is

largely morphological and systematic in character. It begins with

an Introduction which, in the inadequate space of twenty-five pages,

discusses the basis of classification, analogy and homology, cells and

protoplasm, the history of zoólogy and the theory of evolution. The

rest of the volume is devoted to the systematic discussion. In this

part the usual treatment is adopted and there are few features which

call for comment. It is a pleasure to note that the author has not

been ultra conservative. Thus we find a recognition of the close

relationships of Crustacea and Arachnida, these being united in a

group Cephalothoracica. The Phyllopoda are regarded as the primi-

tive Crustacea, and the group of Chordata is admitted, without how-

ever including the Enteropneusta. Even in details the work seems up

to date as for instance (p. 343) in the denial of the vertebrate gono-

tome, and in the recognition of two kinds of ribs.

The language is simple and easy, and the illustrations, largely

original or redrawn by the author, are clear and illustrative. One

feature, however, demands criticism. The author is apparently one

of those who thinks it a gain to science to rename things already well

named. ‘Thus we find Monoplastida and Polyplastida for Protozoa

and Metazoa, Eucephalica for Insecta, Pinnata for Pisces, Digitata

for Tetrapoda, Ditremata for Placentalia, etc. Surely such increase

in synonymy can serve no useful end. The Metazoa are divided into

Radiata, including sponges and ccelenterates, and Bilateralia, and the

latter group into Hypogastrida, including the Vermes, Mollusca and

Arthropoda, and the Pleurogastrida, composed of Vermiformia

(Sagitta, Balanoglossus) Echinodermata, and Chordata. The work

is devoted almost exclusively to recent forms and extinct groups

receive but slight notice.

A New System of Zodgeography has been invented by Prof.

Simroth of Leipzig,? which, however, is not altogether original with

him, P. Reibisch: having first propounded the fundamental idea.

The latter is as follows.

! Lehrbuch der Zoologie von Dr. Alexander Goette. Leipzig 1902. 8vo, pp xii4-

[e]

?Simroth, H. Ueber das natuerliche System der Erde, VerAamd/. deutsch.

Zovlog. Gesellsch. 1902. p. 19-42

3 Reibisch, P. Ein Bene der Erde, 27. Jahresber. Ver. f. Erdkunde

esden

1901.

210 THE AMERICAN NATURALIST. [Vor. XXXVII.

There exists ove largest axis of the earth, ending at two points

under the equator, and this is due to the fact that the diameter of

the equatorial belt, which is larger than any other, is increased by

high mountains situated at these points, which are in Ecuador on the

one side, and in Sumatra on the other.

These two points, according to Reibisch and Simroth, were situated

under the equator throughout all of the earth’s history, while all other

points of the present equator oscillate toward the north and south,

that is to say, they shift their position in consequence of a slow oscil-

lation of the poles in a direction marked by a meridian that goes

through Bering Sea on the one side, and through Europe and Africa

on the other.

Reibisch tries to support his views by a collection of facts show-

ing repeated subsidence and elevation of certain parts of the earth’s

surface: this demonstration, however, is entirely insufficient to found

upon it a theory of this importance, and moreover, the burden of the

proof does not rest with Geology, but with Physics and Astronomy.

The latter branches have been entirely neglected by Reibisch as well.

as Simroth, and thus their theory of this peculiar “ oscillation” of the

Poles lacks proper support, we may even say, has no support at all,

since the geological evidence introduced by Reibisch is by far too

scanty and superficial, and even in some cases incorrect.

Now, this assumption of an oscillation of the Poles in a straight

line, the North Pole moving alternately toward Europe and toward

Bering Strait, of which movement the earth-axis from Ecuador to

Sumatra forms the pivot, is sufficient for Simroth to explain many

problems of the earth’s history and of the distribution of animals.

It is hardly worth while to discuss Simroth's ideas zz extenso, since

the zoógeographical material introduced by him is far from being

adequate. Generally, we may say that the facts quoted are indiffer-

ent with respect to his theory, sometimes the facts are misunderstood,

and in other cases they are directly incorrect or even absurd. The

same is true of the facts quoted from Geology or Physical Geography,

of which branches Simroth seems to possess hardly the most super-

ficial knowledge, and his ideas as to the descent of some animals are

sometimes ridiculous in the highest degree.’

1 [ mention only his view that Africa does not belong properly to the earth, but

a former second moon that fell upon it. Another instance is that Simroth

Feli there is a continuous mountain chain from Ecuador k

? The worst case is, that he believes the porpoise of the Amazonas river to be a

descendant of some Chironectes-like marsupial.

No. 435.] NOTES AND LITERATURE. 211

To sum it up, Reibisch has propounded a theory that appears very

improbable from the points of view of Physics and Astronomy. It is

supported chiefly by imaginary evidence that exists exclusively in the

fancy of the author, the few instances that might be relied upon being

so scarce and so ambiguous that they cannot furnish any base for a

theory of so far-reaching consequences. Thus it is impossible at all

to seriously discuss this theory.

Simroth has used this absurd theory for the creation of a zoógeo-

graphical scheme. His demonstration that the distributional facts

correspond to his assumptions is a complete failure, due chiefly to a

lack of adequate knowledge of the respective facts. And further,

Simroth’s paper is written in so superficial a way for so important a

subject, that we must express our astonishment at this lack of truly

scientific spirit: we do not want scientific dreams and fancies, but

actual, solid and faithful work.

Gardiner’s Maldives.' — The third part of J. Stanley Gardiner's

Fauna and Geography of the Maldive and Laccadive Archipelagoes

is at hand. It contains six memoirs.

The first is by F. Jeffrey Bell on the “ Actinogonidiate Echino-

derms.” He describes five species of Antedon, 5 of Actinometra,

13 Asteroidea, r2 Ophiruoidea and 15 Echinoidea. The collection

is of the ordinary coral-reef type and consists very largely of young

forms. Many of the Ophiruoides have lost the upper surface of the

disc and Bell suggests that the gonads are set free by the separation

of the disc.

The Orthoptera are described by Malcolm Burr. Almost all are

cosmopolitan or widely distributed oriental species, probably all

occurring also in India or Ceylon. The 24 species include one of

the Forficularia, five of the Blattodea. one of the Mantodea, seven of

the Acridiodea, four of the Locustodea and six of the Gryllodea.

The commonest grasshopper, Fpacromia famulus Fabr., is found

abundantly throughout the Oriental region.

Borradaile gives an account of “The Xanthide and some other

crabs” and this, like his earlier papers. is an invaluable contribution

to the bionomics of species which have hitherto been known chiefly

from alcoholic material. Although he is not able to explain on the

ground of adaptation every feature of the different species Borradaile

‘Gardiner, J. Stanley. The Fauna and Geography of the Maldive and Laccadive

Archipelagoes, Vol. i, Part iii. Plates XIV-XVII and text illustrations 41-77,

223-346.

212 THE AMERICAN NATURALIST. (Vor. XXXVII.

is evidently convinced that they can probably be so explained. ‘The

Xanthids are “Essentially crabs of the reef surface and the coral

stocks.” A typical Xanthid, e. g. Xantho, “is of a transversely oval

shape, heavy and compact, with a hard cuticle and short legs which

can be folded up close under the body." The massive, compact

form of the body and the strong cuticle are, no doubt, adaptations to

a life spent in positions where they are surrounded with hard stony

objects, and exposed at times to surf and strong currents, and at

others to enemies which they are unable to escape by swimming like

the Portunidz..... Such genera as Chlorodius and Pseudozius are

flatter and lighter built, with longer legs, though the cuticle is still

strong, and they are more often to be found in situations like the

interstices of coral stocks, where they must maintain their position

by the use of their limbs, rather than by their weight or by wedg-

ing themselves fast like the heavier genera." A bionomically inter-

esting species is Melia tessellata, Latr., which lives among the living

branches of coral stocks. It holds in each claw a small sea anemone,

which would have to be dropped whenever the crab took food with

its chelipeds. ‘The actinians, which are firmly grasped around the

middle below the tentacles, may be useful, by means of their stinging

cells, either for defence or to “fish” for food with, or perhaps for

both purposes." The crabs of the subfamily Trapeziinz find shelter

on branched corals but only on the living branches. "They do not

injure the latter but derive protection from their stinging cells. The

end joints of the rather slender legs are curiously modified to secure

a firm foothold on the branches. The species show color varieties,

especially yellow, dark brown, banded and spotted. A new genus,

Cazecopilumnus, contains a species whose orbits are so reduced that

the animal appears to be absolutely blind. Finally there was found

on the reef Cryptochirus coral/iodyfes, first described by Heller from

the Red Sea and mentioned by Semper (Animal Life, pp. 221-223)

which takes up its abode in the coral by which it becomes enclosed

although not imprisoned.

Of the fishes, C. Tate Regan notices 65 species from the lagoons

of the atolls and six species from fresh water pools. Many of the

former seem to be wide ranging. ‘The latter “are mostly marine

fishes which are known to ascend estuaries into fresh water or are

allied to fishes with this habit." One species, Barbus vittatus Day, is

a true fresh water fish.

The marine Turbellaria are described by Frank F. Laidlaw. This

work is necessarily chiefly morphological. A new family of the

No. 435] NOTES AND LITERATURE. 213

Cotylea — Pericelidæ — is erected. The most abundant family is

that of the Pseudoceridz. Two species found in the Maldives occur

also in the tropical Pacific and two species, Zeptoplana tremellaris

and ZZysanozoon brochi are widespread, occurring in north tem

perate as well as tropical seas.

Finally Gardiner himself has a paper on * Special observations and

work relating to the formation and growth of coral reefs," still incom-

plete. He brings additional evidence, based on an increase in depth

and extent in the lagoons since 1836, for believing that they are due

to solution. The results of dredging at depths from 16 fathoms to

50 fathoms, show that the reef corals do not flourish luxuriantly

below 25 fathoms, but there are other genera of corals which “ flour-

ish just beyond where the surface forms cease to exist." To test the

rate of growth of reefs Gardiner studied the size of corals gathered

from a channel that had been cleaned out three years before. He

concludes that the reef grows at about the rate of 16 fathoms in

1000 years. He suggests that “if Falcon Island, erupted to a height

of about 250 feet in 188 5 and now a mere shoal, be cut down to 25

fathoms by the end of this century its place might well be marked

by surface reefs, perhaps even by a perfect atoll, considerably before

the year 3000 A. D." Finally the action of boring and sand-feeding

organisms is discussed. The boring forms include alga of the

genus Achyla, Cliona and a Myxospongid, the mytilid Lithodomus,

Sipunculoidea, a cirripede Lithrotrya and, especially, the Polychaeta,

above all the Eunicide. Of the sand feeding organisms the Holo-

thuroidea, the Echinoids, the enteropneust Ptychodera, Sipunculus

and 'lhalassema are important. All these organisms help in the

dissolution of the coral rock by which the lagoons are made and

kept open. C. B

A Book on the * Birds of the Rockies" ' might be either a care-

ful scientific hand-book of that region, or an introduction to popular

bird study, or a pieasant account of ornithological rambles. The

first Mr. Keyser’s book does not of course claim to be; if it is to be

judged, as the second or third, we must look for enthusiasm, for a

certain amount of scientific training, and for literary skill. There

is a wide field open to writers who have been trained in the school

of Burroughs and Torrey, who can write introductions or invitations

to popular bird study in the less well-known regions of this country.

! Keyser, L. S. Birds of the Rockies. 8vo, pp. 355- Chicago, A. C. McClurg

Co.

214 THE AMERICAN NATURALIST. [Vor. XXXVII.

The visitor to Franconia or to Florida who is interested in birds

takes Mr. Torrey's sketches as part of his outfit.

To serve as ornithological herald to Colorado was not perhaps

Mr. Keyser's ambition; an enthusiast such as he needs no other

incentive to write than the pleasure he feels in communicating his

experiences. Mr. Keyser certainly has adequate enthusiasm.

Such expressions as “rapture,” “transports,” occur so frequently

that we question whether a more subtle expression of his delight

would not be more likely to impress the reader. Our second desid-

eratum, scientific training enough to wrest from a new region some

fresh interesting matter is hardly shown in this book. ‘The titles of

some of the chapters, “ Bald Peaks and Green Vales," * A Rocky

Mountain Lake,” “ Ho ! for Gray's Peak!" etc., show that it is rather

a series of rambling sketches of the Colorado bird-landscape, so to

speak, than a collection of serious studies.

When we come to the literary quality of the book the less said the

better. It seems as if a book like this could justify its existence only

by attaining a fairly high standard of literary excellence. This should

not be hard in these days when Burroughs, Torrey, Muir and Roberts,

to name but a few of the leaders, have furnished abundant models.

Imagine any of these authors describing an indigo bird as an “ ani-

mated chunk of blue," (page 154), saying that a woodpecker “has

the habit of soaring out into the air and nabbing insects on the

wing" (page 162), or writing of pipits that “their semi-musical

calls drop and dribble from the turquoise depths of the sky " (page

239).

Of the plates by Mr. Fuertes, those in black and white exhibit that

artist's well-known charm and vigor of drawing, but those in color

are with one exception very disappointing. Why are the birds in one

of Mr. Horsfall’s charming little scenes (p. 139) called Brewer's

Blackbirds, when three at least are Yellow-headed Blackbirds ?

RH.

Protozoa. — It is a full score of years since Kent's “ Manual of the

Infusoria” appeared and Butschli's monumental monograph of the

Protozoa was begun. No comprehensive résumé of progress in

protistology has appeared in the years that have since elapsed.

This fact renders Calkin's! treatise peculiarly opportune and assures

it a welcome from many readers.

Calkins, G. N. Columbia Biological Series VJ. The Protozoa. Pp. xvi, 347.

153 Figs. New York, Macmillan & Co., 19or.

No. 435] VOTES AND LITERATURE. 215

The relegation of the systematic side of the subject to a few brief

appendices and the extensive treatment of structural, developmental,

and physiological phenomena reflect the spirit of the past decade in

zoólogical science, while the especial attention to cytological prob-

lems are in keeping with the author's contributions to the subject.

The introductory chapter contains a historical review, a discussion

of the modern classification of the group, of the distinctions between

plants and animals, and of spontaneous generation. We note the

omission of the Bathybius incident in the historical discussion,

though the Eozoón ghost is laid. The general morphology and physi-

ology of the Protozoa are treated in the second chapter, which con-

tains an admirable discussion of the structures and functions of the

protozoan body. ‘The subject of colonial organization is but partly

developed and that of economic aspects lacks perspective. The

agency of the Protozoa in causing odors in potable water are for

example fully treated but no reference is made in this connection to

the pebrine of the silk worm or to Texas fever.

The four chapters which follow deal with the structure, develop-

ment, and relationships of the Sarcodina, Mastigophora, Sporozoa,

and Infusoria. The revival of interest in the Sporozoa in recent _

years makes the review of the literature of these parasites very

opportune.

In the chapter upon the sexual phenomena in Protozoa the author

concludes that there is no instance of nuclear reduction in the Flagel-

lidia and that the maturation of forms in other divisions of the Proto-

zoa shows no genetic relation to analogous processes in Metazoa,

but are “independent” expressions of the same unknown vital forces

which cause the formation of polar bodies, or the double division of

tetrads. ‘The so-called maturation is, moreover, a phenomenon of

degeneration. No conclusive evidence, it seems, can as yet be found

in the Protozoa which enables us to trace the phylogeny of nuclear

structures of the Metazoa, though Calkin's division-center is a step in

that direction. The closing chapter on some problems in the physi-

ology of the Protozoa gives a brief insight into this most promising

and rapidly developing line of research.

The work is adorned by a number of excellent new figures and

concludes with a bibliography of about 600 titles, mainly of morpho-

logical and physiological papers. The omission of Schewiakoff's

Monograph (Russian) of the Aspiroticha and of the recent excel-

lent papers of Roux is to be deplored, for while in the main system-

atic or faunistic they are of great value to every student of the

216 THE AMERICAN NATURALIST. [Vor. XXXVII.

Protozoa and also contain data of general interest, the latter for

example would have provided some connecting links between the

Mastigophora and Ciliat for the discussion of the inter-relationships

of these groups. This book should help materially in stimulating

and fostering the study of this most interesting group by American

students by whom it has been much neglected during the past decade.

BOTANY.

Notes.— The Proceedings of the Pittsburgh meeting of the Society

for the Promotion of Agricultural Science include the following botani-

cal papers: — Bessey, Notes on the poisonous plants of Nebraska;

Voorhees and Lipmans, Individualitv of plants an important factor

in plant nutrition studies; Arthur, The Æcidium as a device to

restore vigor to the fungus; Lyon, The importance of considering

previous environment in conducting variety tests; Bolley, Preliminary

efforts to develop a continuous process of seed disinfection by means

of formaldehyde vapor; Bolley, Use of the centrifuge in diagnosing

plant diseases; Beal, Observations by the roadside; Rane, Plant

depredations and plant culture; Weems and Hess, Study of the food

value of some of the edible fungi of Ames [Ia.]; Chester, Bacterio-

logical analysis of soils; and Lazenby, Some notes on the water con-

tent of certain plants.

The Botanical Gazette for December contains the following articles :

— Frye, A morphological study of certain Asclepiadacee ; Leavitt,

The root-hairs, cap, and sheath of Azolla; Stevens, Studies in the

fertilization of Pliycomycetes; Berry, Notes on Sassafras; Fairchild,

A date-leaf boat of Arabia; and Cockerell, A variable larkspur.

The Bulletin of the Torrey Botanical Club, for December, contains

the following articles : — Cannon, A cytological basis for the Mende-

lian laws; Vail, Notes on the genus Rouliniella; Underwood, Review

of the genus Danza; Rydberg, Studies on the Rocky Mountain Flora,

IX; Rusby, Enumeration of the plants collected by Dr. H. H. Rusby

in South America, 1885-6, XXXII; and Knowlton, Report on a

small collection of fossil plants from the vicinity of Porcupine Butte,

Montana.

No. 435.] NOTES AND LITERATURE 217

The Ohio Naturalist for December contains the following botani-

cal articles : — Schaffner, Observations on self-pruning and the for-

mation of cleavage plains; Schaffner, The flora of Little Chicken

Island; Burr, Compass plants of Ohio; and Simkins, Record of

observations on the dandelion.

The Plant World for November contains the following articles : —

Longenecker, Mushrooms; Niles, Origin of plant names; King, a

summer outing in Iowa; Harvey, The dwarf Mistletoe at Mount

Ktaadn; Pammel, An old sphagnum bog in La Crosse county, Wis-

consin; and, in the supplement, pages 247-253 of Mr. Pollard's The

families of flowering plants.

The Plant World for January contains the following articles:—

Waters, “ Obtusilobata " forms of some ferns; Messenger, The pres-

ervation of our native plants; Collins, Dimorphism in the shoots of

the Ginkgo.

Rhodora for December contains the following articles : — Fernald,

Variations of American Cranberries; Andrews, Bryophytes of the

Mt. Greylock region, II; Deane, Remarkable persistence of the

button-bush; Floyd, Cristate form of JVzpArodium marginale;

Cheney, Rare plants in Centreville, Mass.; Jewell, Some ferns of

Franklin County, Me.; Gutterson, Cuphea procumbens at Andover,

Mass.; Clark, Erodium meschatum in Connecticut; Mann, An Amer-

ican occurrence of Centaurea diffusa; Pease, Two new stations for

Arceuthobium ; and Bissell, Some species of Crepis and Leontodon.

Torreya for December contains the following articles: — Lloyd,

Vacation observations, II; Earle, Key to the North American spe-

cies of Cortinarius, II; and White, Petiolate connation in 77ifolium

pratense.

The gth volume of the Proceedings of the Towa Academy of Sciences

contains the following articles of botanical interest: — Shimek,

Forestry in Iowa; Pammel and King, Vascular cryptogams of Iowa

and adjoining parts of southeastern Minnesota and western Wiscon-

sin; Pammel, Preliminary notes on the flora of western Iowa,

especially from the physiographical ecological standpoint; Hitch-

cock, A list of plants collected in Lee County, Florida ; and, Hume,

Ustalaginez of Iowa.

Like earlier parts, no. 106 of the Proceedings of the Linnean Society

of New South Wales contains a number of botanical articles.

218 THE AMERICAN NATURALIST. (VoL. XXXVII.

Papers on Chantransia efflorescens — by Lehmann, derithrifying

bacteria — by Baur, and the structure of Beggiatoa mirabilis — by

Hinze, are contained in the Kiel Abtheilung of n. F. VI, of the

* Wissenschaftliche Meeresuntersuchungen " of the Kommission zur

wissenschaftlichen Untersuchungen der deutschen Meere.

The concluding part (no. 3-6) of Vol. XVI of the Annales des

Sciences Naturelles, Botanique, issued in December, consists of an

extended discussion of the Ochnacez, by Van Tieghem.

A paper on Drosera, by Heinricher, is contained in the Zeitschrift

des Ferdinandeums für Tirol und Vorarlberg, far r9o2.

A note on Melocactus seedlings, by Suringar, is contained in the

Berichte der Deutschen Botanischen Gesellschaft of December 23.

An account of the *ayotli? or native gourds, of Mexico, by

Urbina, is in course of publication in the Anales del Museo Nacional

de México; beginning with the issue of December.

A habit-photogram of a fine American Elm is contained in

Country Life in America for January.

A largely illustrated paper on the morphology and ecology of the

Indian Podo-stemacez, by Dr. Willis, director of the garden, consti-

tutes part 4 of Vol. I of the Annals of the Royal Botanic Gardens,

Peradeniya.

An interesting account of the tuberiferous Coleus coppini, by

Heckel, is contained in fascicle 2 of the Annales de l'Institut Colonial

de Marseille, Vol. VIII.

An illustrated account of Carludovica jamaicensis, by Fawcett and

Harris, is published in the October number of the Bulletin of the

Botanical Department, Jamaica.

Notices of the Elephant’s trunk Banana, Musa wilsoni, with illus-

trations, appear in the Gardener's Chronicle for December 20 and 27.

Brodiea bicolor is described by Suksdorf in Zhe West American

Scientist for December.

“School Science for January contains an article on The cocoanut —

its composition and germination, by Kirkwood.

A phylogenetic study of Ginkgo, Cephalotaxus, and the Taxacez,

by von Spiess, is appearing in current numbers of the Gi sterreichische

Botanische Zeitschrift.

No. 435] NOTES AND LITERATURE. 219

A considerable number of illustrations of Platycerium occur in Zhe

Gardeners’ Chronicle of January ro.

The principal articles in the January number of Zhe Bryologist

are: — E. G. Britton, Sematophyllum recurvans ; Grout, Orthotri-

chum; and Holzinger, Karl Gustav Limpricht.

An important cytological paper on the Basidiomycetes by Maire,

is published as a supplement to the Bulletin dela Société Mycologique

de France, of December 31, which also contains a paper by Guéguen

on teratological specimens in the same group.

Spegazzini publishes a second series of “ Mycetes argentinensis "

in the opening fascicle of Vol. VIII, ser. 3, of the Anales del Museo

Nacional de Buenos Aires.

Mr. Massee notes the introduction of Cintractia patagonica into

England, in Zhe Gardeners’ Chronicle for January 3.

A study of yeast cells and their nuclear difference from unicellular

animal organisms, by Feinberg, is contained in the Bericht der

Deutschen Botanischen Gesellschafr issued on December 23d.

Traverso publishes a note on the Sclerospora of grasses, in fascicle

5-7 of Malpighia for 1902.

Fascicle 2 of the supplement to the Zzudex Kewensis, has appeared

and covers the phanerogamic genera Cymbidium to Iriha.

As one of the “Twentieth Century Text Books” issued by the

Appletons, an “Analytical key to some of the common flowering

plants of the Rocky Mountain Region,” by Professor Aven Nelson,

was published in June last.

The laws of plant distribution in the alpine zone are discussed by |

Jaccard in No. 144 of the Bulletin de la Société Vandoise des Sciences

Naturelles.

Several papers on antarctic plants are contained in the concluding

number of Nyt Magazin for Naturvidens Kaberne, for 1902.

The double Lieferung 22-3 of Ascherson and Grebner's " Synop-

sis der Mitteleurpüischen Flora" completes the grasses and enters

on Cyperacez.

A sand-hill flora study in Lancashire, by Bailey, is contained in

the Memoirs and Proceedings of the Manchester Literary and Philo-

sophical Society, issued December 15.

220 THE AMERICAN NATURALIST. [Vor. XXXVII.

Reiche's * Flora de Chile," published at Santiago, has reached the

third volume, devoted to Cunoniacez to Compositz-in-part.

* Flora of New Providence and Andros (Bahama Islands)" is the

title of a paper by Mrs. A. R. Northrop, wbich forms the opening

number of Vol. XII of the Memoirs of the Torrey Botanical Club,

issued December roth.

In a thesis on “Die Stelar-Theorie,” published at Groningen in

1902, Schonte reaches the conclusion that but a single stelar type

* Monostelie" occurs in the stem and root of vascular plants.

A study of style and stigma in the Cruciferz, by Villani, is con-

tained in Malpighia, Vol. XVI fascicle 5-7.

Recent numbers of the Bulletin du Muséum d'Histoire Naturelle,

of Paris, contain a considerable number of morphologico-systematic

papers by Van Tieghem.

Plant physiology in the high school is the subject of a paper by

Professor Barnes in School Science for December.

The use of bacteria in studying photosynthesis is discussed by

Beijerinck in Vol. IV of the Proceedings of the Section of Sciences

of the K. Akademie van Wetenschaffen te Amsterdam, in which Burck

writes on the irritable stigmas of Torenia and Mimulus, and on

means to prevent the germination of foreign pollen ‘on the stigma.

Guignard contributes a paper on the double fertilization of Cruci-

ferz to the Journal de Botanique for November, which also contains

the results of an anatomical study of Menispermacez, by Maheu.

A paper on neo-Lamarchian views as to organic evolution, by

von Wettstein, is separately issued from the 1902 Verbandlungen der

Gesellschaft deutscher Naturferscherund Ärzte.

C. A. White contributes a discussion of the mutation theory of De

Vries to the Smithsonian Report for 19or.

Mendel's law is discussed by Professor Spillman in the Popular

Science Monthly for January.

Mendel’s laws and their probable relations to intra-racial heredity

are discussed by Yule in recent numbers of Zhe New Phytologist.

An illustrated article on the Missouri Botanical Garden, by its

Director, is published in the Popular Science Monthly for January.

No. 435.] NOTES AND LITERATURE. 221

A Plea for hardy plants, with suggestions for their effective

arrangement, by J. Wilkinson Elliott, has been reprinted from the

Transactions of the Massachusetts Horticultural Society for 1895, with

additional plans and copious and exquisite illustrations by J. Horace

McFarland and others, and issued from the press of Doubleday, Page

& Uo. :

A delightfully entertaining little book on Children’s Gardens, by

the Honorable Mrs. Evelyn Cecil, better known under her maiden

name of Alicia Amherst, has been published by the Macmillans.

A sketch of the life and labors of Sir William Hooker, with por-

trait, is begun by his son, Sir Joseph Hooker, in the Annals of

Botany for December.

A portrait of Schumann forms the frontispiece to the December

number of the Monatsschrift für Kakteenkunde.

Commenced January, 1383 Twe volumes per year —

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9 Vor. XXXVII, No. 436 APRIL, 1903

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

Page

On tho Appendages of Tremataspis . . Frofessor W. PATTEN 223

Studies of Localized Stages in Some Plants of the Botanic Gardens of Harvard

II.

Unwin o e eae . JOSEPE A. CUSHMAN 243

III. Notes on the Reproduction of Certain Reptiles . C.S. BRINLEY 261

IV. Synopsis of North American Invertebrates. XVIII, The Amphipoda,

8.J. HOLMES 267

293

V. Publications Recoived

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WILLIAM M. DAVIS, M.E., Harvard University, Cambridge.

ALES HRDLICKA, M.D., Mew York Ci

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THE

AMERICAN NATURALIST.

VoL. XXXVII. April, 1903. No. 436.

ON THE APPENDAGES OF TREMATASPIS.

W. PATTEN.

IN a recent paper in the American Naturalist’ on the structure

of the Tremataspidza, I maintained that the marginal openings

(so-called gill openings) on the under surface of the head of

Tremataspis might be interpreted as the points of attachment

of metamerically arranged appendages. Whether all these

appendages projected freely from the body like the jointed

pectorals of Bothriolepis, or presented any one of the innumer-

able variations in structure and function to which such append-

ages might be subject, there was no means of determining. If

one admits the extremely doubtful systematic position of the

- Ostracoderms, there is no satisfactory reason for calling the mar-

ginal openings of Tremataspis gill openings unless one assumes

the whole point at issue by asserting that the Tremataspidze

are true vertebrates. But if the Tremataspide are provided

with oar-like appendages, similar in structure and mode of attach-

ment to those in other members of the Ostracoderms, such as

Pterichthys and Bothriolepis, then these appendages must have

! On the Structure and Classification of the Tremataspidwe. Vol. xxvii, No.

425, pp. 386 and 388.

224 THE AMERICAN NATURALIST. (VOL; XXXVII.

been attached to the head, near the anterior end of the series of

so-called gill openings. The anterior pair of incisions, or open-

ings (of which I have found at least two additional pairs in front

of the six pairs described by Rohon) are found on the lateral

margin of the anterior part of the head, in precisely the position

where the hypothetical oar-like appendages should: be attached

provided any were present. Moreover in Tolypaspis, Cyathaspis .

and Pteraspis, there is a very conspicuous rounded incision on

the margin of the dorsal shield that has usually been regarded as

the site of the lateral eyes, apparently because there was no

other interpretation for these openings at hand, and because

otherwise the lateral eyes. would appear to be absent. But,

these incisions also occur at points corresponding to the ones

where the oar-like appendages in Pterichthys and Bothriolepis,

and the anterior pair of incisions in Tremataspis, are located.

The conclusion to be drawn from these facts is that Tremataspis,

Cyathaspis, and Pteraspis were probably provided with jointed,

oar-like swimming appendages similar in character and location

to those of Pterichthys and Bothriolepis.

Moreover, and this is the important point, if Tremataspis has

a pair of jointed, armored appendages attached to the anterior

margin of the ventral surface, they must have been attached to

one of the anterior pair of marginal incisions. If this is the

case, there is no escape from the conclusion that be%ind the oar-

like appendages there are as many more pairs of serially homol-

ogous structures (but not necessarily similar to them in form or

function) as there are pairs of marginal incisions.

The assumption that Tremataspis is provided with jointed

pectorals is strengthened by the fact that Lindström! has

described what he considers to have been an appendage in

Cyathaspis. He states on p. 5: “ Along with some other frag-

ments was found, detached and broken into three pieces, what

may be regarded as the shelly covering of an extremity or limb

(Pl I, fig. 9-12). Itis plain that it is not a fragment of the

dorsal shield nor of one of the cornua. Its exterior ridged sur-

face is, to wit, inflected towards the interior surface along the

! Lindström, G.: On Remains of a Cyathaspis, etc.: Bihang F. K. Svenska

Vet. Akads Hand! Bd. 21, Afd. IV. No. 3.

No. 436.] APPENDAGES OF TREMATASPIS. 225

lateral edges and partly covers it. In all probability the muscu-

lar and other organs, which it has covered, were in the same way

sheltered on both sides by similar plates. It is narrow, elongated,

by degrees tapering and ending bluntly. The surface is of the

same glossy lustre as the other portions of the exoskeleton and

is covered with ridges which have a nearly parallel direction

and continue without interruption. On the interior surface (Pl.

I, Fig. 12) which is quite smooth, are seen the polygonal parti-

tion lines of the vacuole walls, and the openings of the minute

canals. There is no evidence that it has consisted of more than

one plate, nor that the supposed extremity has been covered by

several plates, as the limbs of Pterichthys." Lindstróm does

not attempt to justify his conclusion that the structure in ques-

tion is an appendage, or to indicate its point of attachment, or

the effect such a discovery, if confirmed, must have on the classi-

fication of the Ostracoderms. The importance of Lindstróm's

discovery is somewhat diminished by the fact that what he calls

an appendage is but a single plate. Hence it is uncertain

whether it covered a small part only or the whole appendage, or

whether it was part of a scale or spine paired or unpaired, belong-

ing either to the head or trunk.

An examination of some beautiful specimens of Pteraspis in

the British Museum indicates that the so-called lateral eye

openings are better explained as points for the attachment of

oar-like appendages, and that the projecting processes sometimes

attached to the openings should be regarded, not as a portion of

the matrix squeezed out of the orbits, but as either the remnants

of an appendage itself, or as the matrix that originally filled the

base of the appendage.

In view of the common bonds of relationship that unite the

various groups of the Ostracoderms, and in view of the presence

of at least one pair of jointed cephalic appendages in Bothrio-

lepis, Pterichthys, and perhaps in Cyathaspis and Pteraspis, the

presence of one or more appendages in Tremataspis appeared to

be within the range of possibility. It remained to discover some

traces of the appendages themselves.

Thus the matter stood at the time of writing in St. Peters-

burg, for the Imperial Academy of Sciences, my first paper on

the Tremataspidze.

226 THE AMERICAN NATURALIST. [Vor. XXXVII.

On my return to America, I began work anew on a few

fragments and isolated plates that had been obtained from Herr

A. Simonson, on my visit to the island of Oesel, and to which I

had previously failed to give adequate attention. One fragment

of special importance was almost concealed in the rock and its

true nature was not at first apparent; the other was similar to a

plate first described by Rohon. When it became evident that

these fragments were probably parts of appendages, it was too

late to do more in that paper than refer to the fact that what

appeared to be fragments of appendages of Tremataspis had

been discovered.

I shall now describe what I consider to be the distal portion

of an oar like appendage attached to the head of Tremataspis.

Fic. 1. Distal joint of cephalic appendage of l'remataspis seen from the ventral side. X ro.

Fic. 2. Same from median edge. X 1o

. rg

f. articular process. r. ridge. s. shoulder.

No. 436.] APPENDAGES OF TREMATASPIS. 227

The structure, Figs. 1-3, was at first deeply imbedded in

the matrix leaving only a small part of the dorsal surface

exposed, from which the outer layers has been worn off, reveal-

ing the characteristic polygonal meshwork of the middle layer.

After exposing the entire dorsal surface of the plate by

cautiously removing the matrix with fine needles, to insure

against accidents in the subsequent treatment, it was photo-

graphed and modeled in wax on a large scale. This proved to

be a wise precaution, as the extremely brittle and fragile plate,

in spite of the utmost precaution, was accidentally injured, in

one of the many occasions when it was being placed in position

for examination. After the dorsal surface had been thoroughly

studied, it was covered with hard balsam and fastened face

downwards to a glass slide. When the remaining matrix had

been removed and the balsam dissolved off, the completely iso-

lated plate could be examined from any desirable point of view.

The other plates to be described were treated in a similar

manner.

When seen from what I consider to be the dorsal side, Fig. 3,

the distal joint of the appendage appears something like the

blade of a knife. It is about 2.5 mm. wide by 7.5 mm. long

and of a light yellowish brown color. A cross section would be

triangular, with the thickest part on what I assume to be the

median side, or that side which when the appendage was in place

was nearest the body. The opposite or lateral margin, especially

toward its posterior end, is thinned down to a rather sharp edge,

ornamented by a series of glistening beadings or scallops. Fig.

36. The six beadings nearest the posterior end are the most

regular and they are readily seen from either surface. Toward

the anterior end of the lateral margin one can distinguish five

Or six more undulations, somewhat irregular in size and best

seen from the under surface, Fig. 1, 6. Towards its anterior

end, the lateral margin becomes thicker and more rounded so

that cross sections of the appendages at that point would be

more oval in outline. The dorsal surface is rather full and

rounded at the anterior end, flattening out posteriorly where,

near the apex, it takes a rather sharp downward curve, Figs.

2 and 3, as though the point were slightly bent. Near the

228 THE AMERICAN NATURALIST. [Vor. XXXVII.

posterior end, the median dorsal margin forms a rather promi-

nent shoulder, s, giving this portion of the appendage a charac-

teristic contour.

The anterior end, which slopes diagonally forward and out-

ward, is provided with a beautifully modeled articular condyle, c.

It is a smooth, rounded prominence, flattened dorso-ventrally,

slightly depressed on its anterior surface, and with a well

defined constriction around its base. Its median end is contin-

uous with a distinct ridge that extends along the middle of the

median surface of the appendage, Figs. 1—3, 7.

The entire dorsal surface of the joint was covered originally

with a polished enamel like layer, similar in color and texture to

that on the dorsal shield of Tremataspis. But a large patch of

this layer, on that part of the specimen that originally protruded

from the matrix, is worn off, thus exposing the hexagonal mesh-

work of vessels, filled with a reddish deposit, so characteristic of

the middle layer of the shell of Tremataspis.

The polished outer layer terminates abruptly at the anterior

end in a rounded edge that may be followed around the project-

ing anterior lateral angle, ^. On the dorsal surface, this polished

edge presents two well defined curves, a part of the posterior one

being destroyed. On the ventral surface, the whole of the enamel

layer is preserved, its sharply defined anterior margin forming

three well defined rounded incisions, Fig. 1. The resemblance

between what is left of the anterior enameled margin on the

dorsal side, and the intact margin on the ventral, indicates that

both were probably in the uninjured specimen quite similar.

The ventral surface of the appendage is perfect in every detail.

In contour and texture it differs somewhat from that of the dor-

sal surface. It is slightly depressed in the middle, there are

prominent thickenings on either margin, and the characteristic

shoulder of the dorsal surface is absent. It is covered with

coarse pits, the mouths of which look towards the distal end of

the appendage as though they had at one time been occupied by

coarse hairs directed backwards. This produces a surface texture

quite different from the smooth surface on the dorsal side of the

appendage.

When we look at the appendage from the median side, Fig. 2,

No. 436.] APPENDAGES OF TREMATASPIS. 229

its asymmetry is clearly seen. Unfortunately the characteristic

texture of the median surface is not well shown in the photo-

graph. This was due in part to the fact that the articulating

condyle was accidentally broken off and the shiny surface of the

cement used to repair it made it difficult to obtain good photo-

graphs. It shows, however, with sufficient distinctness that the

enamel layer which covers only the posterior end of the median

surface, abruptly terminates anteriorly in a thick fold, me’.

The remaining portion of the median surface is covered with

a peculiar layer entirely different in texture from that on the dor-

sal and ventral surfaces. It is much darker colored and its lus-

treless surface, which is irregularly folded, is devoid of pores,

except a few scattering ones of comparatively large size. It

has the appearance of having been a tough flexible membrane

that had become dried and wrinkled before fossilization. Nearer

the anterior end, the conspicuous median ridge and the well

moulded condyle indicate the presence of firmer bone-like

tissues.

I regard the whole structure as the distal joint of an oar-like

swimming appendage that served the same function for Trem-

ataspis that the “pectoral appendages” do in Bothriolepis and

Pterichthys. That we are dealing with a projecting appendage

of some kind and not a superficial plate is shown by the pres-

ence of the polished outer layers of the shield on both surfaces ;

and that it was freely movable, is shown by the prominent artic-

ular condyle at its anterior end.

We know with certainty that such a plate could not have

been a mere spine attached to either the ends of the cornua, or

to the lateral margins of the cephalic buckler back of the oral

region, or to any part of the dorsal surface of the head, for on

these parts, the structure of which is perfectly well known, there

are certainly no articulating surfaces that could serve for the

attachment of either spines or appendages of this character.

We may also assure ourselves that it could not be a median dor-

sal or ventral spine of the trunk, for the difference in texture

and contour of its two surfaces proves that it was a part, or the

whole, of a paired organ. We are, therefore, limited to the

230 THE AMERICAN NATURALIST. (Vor. XXXVII.

assumption that it formed a part, or the whole, of a paired organ

attached to the sides of either the trunk or to the anterior ventral

part of the head.

The probable mode of attachment of the joint, and its direc-

tion of movement, is indicated by the sharp slope of its anterior

end, the length of the process on its anterior lateral margin,

and by the shape and condition of the condyle. These condi-

tions indicate that the plate could not have been attached

directly to the sides of the body, for in that case it would have

to be directed diagonally backwards and the prominent anterior

fateral process would prevent that range of forward and back-

ward movement which the shape of the condyle clearly indicates

it did have.

The same argument would preclude the possibility of its being

attached directly to any of the marginal incisions in the oral

region of the ventral surface. In a word, there is no place

where there is any likelihood that such a structure could be

attached directly to the body, and there is no structure known

in the Ostracoderms with which it can be compared, if not with

the distal joint of the pectoral appendages of Bothriolepis or

Pterichthys. With this it corresponds fairly well except that it is

formed of a single plate, instead of several.

We therefore assume that the plate in question formed the

distal joint of an appendage something like that of Bothriolepis.

It now remains for us to find the plates that make up the armor

of the proximal portion of the appendages.

Basal joint: I regard the structure shown in Figs. 4 and 5

as one of these plates. Two specimens of this plate are in the

collections at Dartmouth College; a nearly perfect one from

which this photograph was made, and a fragment of another.

One or more of these plates are also in the collection of the

Imp. Acad. of St. Petersburg. The plate is 6 mm. long and

6 mm. wide. It is roughly diamond shaped, but not quite

symmetrical in outline, as the angle on the right side is higher and

broader than that on the left. There is a corresponding differ-

! Rohon makes his anterior median plate considerably smaller than this, namely

4 X 3mm. This difference I am inclined to believe is due to the fact that his

plate was imperfect, and not entirely exposed.

No. 436.]

APPENDAGES OF TREMATASPIS.

231

Fic,

Fic. 5. Sam

jas collar

Basal plate of Cephalic Rid se from dorsal surface. X 10.

e from inner, or visceral, aspect. X 10.

fold of y xclished layer on the ventral side. e.

cut into the polished n layer. m. m. & m. m? scle markings.

Dr, & e. g»

groove

232 THE AMERICAN NATURALIST. [Vor. XXXVII.

ence in the inner surface, showing clearly that the plate zs not

bilaterally symmetrical. What I shall call the distal portion of

the dorsal side (Fig. 4) is strongly arched and has a very smooth,

beautifully polished surface dotted with minute pores. It is a

slaty gray color, becoming a dark chocolate on the left, in that

part that was originally deeply imbedded in the matrix. The

texture changes rapidly at the anterior end, where there is a

broad band of coarse dimples in the polished floor of a shallow

groove, 6. 5. In front of this groove the plate narrows to form

a thick collar, e, very different in appearance from the rest of

the plate. It is lighter colored and has a smooth, but irregular

surface that lacks the brilliant polish and regularly distributed

minute pores of the posterior part. It is provided with a few

large openings irregularly distributed. The whole appearance of

the collar suggests that it was made to fit into a nearly circular

opening and that it was held in place there by stout ligaments

attached to its roughened surface.

A remarkably clear cut, semicircular groove is seen on the

beautifully polished and rounded surface of the plate a little way

below the band of pores, e. g. It is seen on all the specimens

examined (four in all) and must have been produced during the

lifetime of the animal by a rotatory movement of the plate that

brought it in contact with some hard object. The meaning of

this groove will be considered farther on.

By the exercise of some care, the matrix was successfully

removed from the inner surface of the plate, thus displaying a

somewhat remarkable structure, Fig. 5. A small fragment of

the edge of the plate was broken off in excavating the matrix,

the outline of the missing portion being indicated by a dotted

line. The plate is now seen to be of a uniform dark chocolate

color and with a concave inner surface. The posterior half of

the inner surface has a coarse rough texture different from that

seen anywhere else in Tremataspis, but otherwise it shows no

particular structure worthy of mention. In about the middle of

either side, the margin is thickened and here the enamel layer of

the outer surface extends a short distance on to the inner sur-

fare v. f.

There are two large oblong depressions on either side (m. m.)

No. 436.] APPENDAGES OF TREMATASPIS. 233

apparently made by the attachment of powerful muscles. The

floor of each depression is irregularly ridged, and has a dull,

almost shining, surface. „At its posterior end is a still deeper

rounded depression, m. m.,* that probably served for the attach-

ment of a special group of muscle fibres. The rounded depres-

sion is considerably deeper and more clearly defined on the right

side of the plate.

Fic. 6. Basal plate, A, from the side, B, from the anterior end. X 10.

The lateral margins of the collar that was seen on the outer

surface, Fig. 4, c, extend about half way down the inner surface

of the plate, forming a conspicuous ridge on either side of the

muscle markings. | When seen in profile from the anterior

end, Fig. 6, B, the collar looks like a small concave plate

adhering to the inner surface of the larger one. The concave

inner side of the collar is smooth and undulating, but with a dull

surface. It is provided with several large grooves leading to

conspicuous apertures that evidently served for the passage of

blood vessels to the interior of the plate. These grooves are

instructive as they indicate that, as it stands in the figure, the

blood vessels come to the plate from its upper end.

Rohon! has described a plate like the one we are now con-

! Zur Kenntniss der Tremataspiden.

Mélanges Géologiques et Palaontologiques. Aull. de l'Académie Imp. des

Sciences de St. Petersbourg, T. I. '94

234 THE AMERICAN NATURALIST. [Vor. XXXVII

sidering. His specimen was only partly exposed and he failed

to see many important details. He looked on it as the anterior

median oral plate. I have examined Rohon’s specimen and

there is no doubt that it is identical with the one in the Dart-

mouth collections. Rohon states, p. 188,

« Die fur die erste Reihe von mir angenommenen zwei vorderen Seiten-

platten sind am Original nicht allein zerbrochen, sondern auch gänzlich

verschoben. Als die vordere Mittelplatte vermag ich nur vermuthungsweise

die in den Figuren 12 und 13 a, b abgebildeten und isoliert gefundenen

Platten ansprechen. Die Platten sind durch ihre bedeutende Wélbung

ausgezeichnet; hingegen stimmen sie der Form und mikroskopischen Struc-

tur nach mit den ubrigen Platten vollkommen uberein. Dass jedoch die

zwei vorderen Seitenplatten zu den nachfolgenden gehoren, dafur spricht

sowohl ihre Lage wie ihre Beschaffenheit; sie sind ebenso an der Ober-

fläche glänzend und flach wie die meisten der nachfolgenden, wahrend die

in deren Nähe sichtbaren Stücke von dem Umschlagsrande stark gewólbt

und dünner erscheinen. Im Übrigen betrachte ich, wie gesagt, das

Thatsiichliche in Bezug auf die beiden vorderen Seitenplatten als etwas

Unsicheres.”

We must take exception to Rohon’s statement that the so-called

median oral plate agrees completely in form and microscopic

structure with that of the remaining oral ones. In the Dart-

mouth specimens, the surface is very smooth and_ shining

whereas in the cast of that specimen in St. Petersburg which

shows nearly all the plates “in situ" the surface of the oral

plates is broken into very delicate ripple marks similar to those

usually seen on the dorsal and ventral surfaces of the buckler.

Moreover, the isolated so-called median plate has rounded out-

lines and a strongly convex surface so that it would appear quite -

out of place among the true oral plates which have nearly

straight line contours and flat surfaces.

The smooth surface texture of this plate may be a specific

difference, as I have seen some specimens of Tremataspis with

very thick, strongly arched bucklers and smooth surfaces. They

appeared to belong to a different species from any of those

already described. However, a small recently discovered iso-

lated plate belonging to the oral region shows a smooth polished

surface similar to that of our shield-shape plate, indicating that

the surface texture of the oral plates may have been slightly

different from that on the rest of the body.

No. 436.] APPENDAGES. OF TREMATASPIS. 235

Neither can I agree with Rohon’s view that the oral surface

of the St. Petersburg specimen which shows the oral plates iv

situ is so distorted as to force the median plate strongly to one

side. As I have pointed out elsewhere the true anterior median

oral plate is here almost exactly in its proper place, and it is very

different in size and shape from the isolated one Rohon wrongly

supposed to be a median oral plate.

The location of this shield shape plate in Tremataspis, for

there can be no reasonable doubt that it belongs to this genus,

has been a great puzzle. The objections to Rohon's view that it

is a median plate lying just behind the mouth are overwhelming.

It is (1) much too large. (2) It is asymmetrical and cannot be

regarded as a median plate. (3) Its beautifully preserved struc-

ture is totally unlike a jaw which we must assume it to be if

placed in that position. (4) The rounded collar at its anterior

end, the two powerful and counteracting muscles attached to its

inner side, and the sharply cut grooves on the polished outer

surface, that were unquestionably worn by contact with some

other hard part, shows conclusively that this plate was attached

by the collar on its anterior surface to some circular opening and

that it had a wide range of rotary movement from side to side,

an improbable movement in a plate thus located.

The resemblance of this plate to the hypostoma of Trilobites

in its shape, muscle markings and its probable mode of motion,

led me at first to consider it as a preoral plate attached by its

collar to the anterior ventral margin of the dorsal shield. But

its asymmetry, which at first did not impress me so forcibly as it

did later, its large size, and the lack of correspondence between

the shape of its anterior margin and the anterior margin of the

dorsal shield, gave little support to such an idea. If this plate

belongs to the cephalic shield of Tremataspis, there is but one

place remaining where it could have been attached and that is

to the large anterior semicircular incision on the margin of the

cephalic buckler, that is at the point where, on other grounds,

we have reason to suppose that a pair of oar-like appendages

were attached. Fig. 7. If we compare this plate, on this

assumption, with the corresponding plate at the base of the

appendage of Bothriolepis, as figured by Whiteaves, we see that

236 THE AMERICAN NATURALIST. [Vor. XXXVII.

they are indeed somewhat similar. Both plates have a strongly

convex outer surface, a pointed posterior extremity and an artic

ulating anterior margin. In Figs. 7 and 8 I have reconstructed

the appendages of Tremataspis on the assumption that the basal

portion was covered by several plates similar to those of Bothrio-

lepis. In these figures the basal and distal parts of this hypo-

thetical appendage and also, as far as possible, the marginal

incision of the cephalic shield, are drawn to the same scale. I

have described this incision in my first paper and have shown

Fic. 7. Reconstruction of the ventral side of the head of Tremataspis, indicating the mode

of attachment of the cephalic canada

that it lies on the down turned margin of the shield. I assume

that the collar of the basal plate with its convex side facing

dorsally, was attached by ligaments, or a flexible membrane, to

the margins of the incision, the membranes or ligaments being

long enough to allow great freedom of movement. It is not

likely that the collar of the basal plate actually fitted into the

incisions, for its surface is not smooth enough, and if it did there

could hardly be the requisite freedom of movement.

No. 436.] APPENDAGES OF TREMATASPIS. 237

It is clear that in the swimming movements of such an append-

age, Fig. 8, the blade-like distal part would probably be carried

upward and forward with the sharp cutting edge in advance,

pas

Cccé

P e

US A M

Fic. 8. Reconstruction of the head and trunk of Tremataspis, seen from

above, 2.

thus offering the least resistance. On the return stroke, which

impelled the animal forward, the broad under surface would be

238 THE AMERICAN NATURALIST. (VoL. XXXVII.

in advance the whole appendage swinging backwards and down-

wards. In performing a complete stroke, the apex of the

appendage would describe either an ellipse or a fig. œ, while at

the beginning and end of each forward and backward stroke the

appendage as a whole would rotate on its own long axis, through

an angle of something like 45 degrees. These movements would

cause the dorsal surface of the basal plate to strike, and rub

against the folded margin of the dorsal shield, and would pro-

duce just such a series of grooves as are shown so clearly on

the plate. These grooves are well worth careful consideration.

There are three grooves arranged in the arc of a circle just

back of the band of coarse pores. On the left side of the plate,

Fig. 4, e. g.' the groove has a symmetrically rounded floor, is per-

haps a fifth of a millimeter deep, and narrows at either end till

it disappears. The anterior boundary of the groove is more,

sharply curved than the posterior. The groove on the opposite

side of the plate is less sharply marked, shallower than the first,

and its anterior and posterior boundaries are alike. A very nar-

row hardly visible groove continues the lines of the first two so

as to complete the arc of a circle, but it does not open directly

into either of the two marginal grooves. These three grooves

are apparently formed by friction against some hard object dur-

ing the same movement, except that the contact seems to have

been more forcible at the beginning and at the close of the move-

ment. It will also be observed that these grooves do not lie on

the most protruding part of the surface but in a kind of recess

between the most prominent portion and the pitted anterior

border, Fig. 6 A. It is clear therefore that the grooves must

have been formed by contact with some narrow projecting rim

and not by a broad flat surface. This is in harmony with our

supposition that the grooves were formed by the rotation of the

basal plate of the appendage against either the ventral margin

of the cephalic shield or against the adjacent oral plates on the

opposite side of the opening to which we have assumed the

appendage was attached.

If one constructs a rough working model of an appendage

such as I have described, it will be clear that as the appendage

approaches the beginning or end of a stroke, it becomes more

No. 436.] APPENDAGES OF TREMATASPIS. 239

and more nearly parallel with the long axis of the head, the tend-

ency being to draw the basal plate more and more diagonally

across the margin of the shield, instead of straight across it as

during the middle of the stroke. This factor and the sudden

change in the intensity of muscular contraction at the beginning

and end of each stroke furnish a very satisfactory explanation of

the increased depth and width of the ends of the eroded groove.

To return to the basal plate shown in Fig. 4. A close exam-

ination of its convex surface shows a second belt, not shown by

the photograph, farther back than the first where the surface

enamel has been very slightly eroded. It extends about two

thirds of the way across the lower portion and is coextensive

with the line of greatest elevation of the surface. It is hard to

see, on any supposition as to the location of the plate how this

abrasion could have been produced by contact with any part of

the animal’s body to which the plate was attached. I assume

that the abrasion, if made during the lifetime of the animal,

must have been produced by contact with the bottom over

which the animal was moving.

Fic. 9. Distal ends of two differe

of T: Pese ca seen from the Md pastas aad drawn

1 cale. B shows the same plate shown in

nig eae des

240 THE AMERICAN NATURALIST, | [Vor. XXXVII.

Since writing the above description of the pectoral append-

ages of Tremataspis, a new specimen of the distal joint has come

into my possession, Fig. 6 A. It is unquestionably of the same

character as that shown in Fig. 3. It has the same general form

and texture but differs from it in certain details. It is consider-

ably smaller, measuring only 5.66 mm. in greatest length by

I.65 mm. in greatest width. . There are six or seven very prom-

inent marginal serrations or spines. The curvature of the dorso-

median surface indicates the presence of the peculiar shoulder

seen in Fig. 3, but the surface is broken along the edge where

such a shoulder should be. The smooth anterior margin of the

shining surface layer appears to have been straight instead of

scalloped. There is the same peculiar wrinkled and lustreless

surface on its median side, extending not quite to the distal end.

At the proximal end, this layer forms an irregular prominent

surface with two or three round openings for the passage of

blood vessels, but without any trace of an articular condyle, or

articulating surface, such as was. seen in the first specimen.

The surface at this point is intact, showing that an articulating

surface could not originally have been present.

While there can be no doubt that the two structures in ques-

tion are of the same nature, and that both belong to Tremataspis,

the differences between them, in the above mentioned details,

are so great that we can hardly regard them as identical. We

are left then to the conclusions that (1) they are either homolo-

gous structures from different species of Tremataspis, against

which it may be urged that the surface texture is the same in

both and like that known in T. schmidtii only. (2) or they

may be serially homologous structures from the same species.

This conclusion harmonizes with the fact that both structures

agree in surface texture but not in form, and with the fact that

there is a series of marginal openings on the ventral surface

serially homologous with the anterior ones to which the pectoral

appendages are attached. I have already suggested that these

openings indicated the presence of other appendages of a smaller

size than the pectorals, although there was no evidence at that

time available that furnished any hint as to what the character

of these appendages might be. This fact we have always borne

No. 436.] APPENDAGES OF TREMATASPIS. 24I

in mind in view of the great range in structure and function

presented by such organs in other animals whose structure is

better known than that of Tremataspis, and also in view of the

fact that there is often such a great range of variation in a series

of these organs in the same animal. Nevertheless in view of the

above named consideration and while well aware of the uncer-

tainties involved, it seems to me we must not lose sight of the

possibility that there may have been more than one pair of

jointed appendages in Tremataspis, similar to those described as

pectorals, and that the fragment in question may have belonged

to one of these additional appendages.

In conclusion, therefore, I believe that we are justified in the

statement that Tremataspis is provided with a pair of oar-like

swimming appendages similar to those of Bothriolepis and Pteri-

chthys, and that similar appendages were probably present in

Pteraspis, Cyathaspis and Tolypaspis.

In Tremataspis, these appendages were probably attached to

the largest pair of incisions on the anterior ventral margin of

the head. Appendages attached there would agree (1) in point

of attachment with those in Bothriolepis and Pterichthys. (2)

they would be attached to a part of the head best fitted, so, far

the shape of the head and the location of its centre of gravity

is concerned, to serve as the support for a pair of swimming

appendages. (3) There is no other part of the head to which

such appendages could be attached. (4) They could not have

been attached to the trunk, because the trunk of Tremataspis

must have been of relatively small size, judging from the way

the shield tapers posteriorly, from the small opening at the

posterior end, and from what we know of the size of thè trunk

scales and of the size of the trunk in Thyestis and other Ostra-

coderms. It is therefore extremely improbable that large

appendages could have been attached toa body so small that

the centre of gravity must have been situated in the head, some

distance in front of the anterior end of the trunk.

That by far the greater share of the work of locomotion in the

Ostracoderms must have been performed by the oar-like cephalic

appendages is indicated by their anterior position and great size

242 THE AMERICAN NATURALIST. | [Vor. XXXVII.

in Bothriolepis and Pterichthys, and by the relatively small size

of dorsal and caudal fins in the latter genus. Such a condition

is in marked contrast to that in many of the most primitive of

the true fishes where the pectorals are much smaller rela-

tively, serving rather as balancers, the principal work of locomo-

tion being performed by the tail and caudal fin.

Among true vertebrates, the only structures suggestive of

the cephalic appendages of the Ostracoderms are the external

gills during their early embryonic stages, including among these

structures the “balancers of Amphibian larva. The large

size and anterior position of the latter appendages make them

especially suggestive of the oar-like appendages of Tremataspis.

HANOVER, N. H.,

April 19.

STUDIES OF LOCALIZED STAGES IN SOME

PLANTS OF THE BOTANIC GARDENS

OF HARVARD UNIVERSITY.

JOSEPH A. CUSHMAN.

In animals and plants interesting stages occur at certain parts

throughout the life of the individual. These are expressed in

Jackson’s law, “Throughout the life of the individual, stages

may be found in localized parts, which are similar to stages

found in the young, and the equivalents. of which are to be

sought in the adults of ancestral groups." In a preceding paper

the author has described such localized stages in some New Eng-

land plants. In the present paper the same line of investigation

has been extended to certain exotic plants which have been

recently studied. Jackson showed that such localized stages

occur in organisms at very definite areas and under definite con-

ditions of growth. This view is corroborated by the present

study.

My thanks are due to Prof. G. L. Goodale and Dr. E. W. Olive

for supervising my work, to Mr. Cameron for opportunities to

study a great number of seedlings in the greenhouses and for the

care of those under observation, and to Dr. Jackson for reading

this paper and for many helpful suggestions.

The following examples show the various

ideas here intended to be brought out in the

line of localized stages in development.

.Astragalus adsurgens Pall. Figs. 1-5.

This herbaceous plant, a native of Siberia,

shows in its early spring growth localized

stages in development. The leaves are com-

pound and so the differential characters are

more easily and definitely seen than in many

plants. Seedlings (Fig. 1) were obtained

beneath the adult plants which were at that — leaves, the

rst with 3, the second

time — April — just appearing. The cotyle- with teaftets.

243

244 THE AMERICAN NATURALIST. (VoL. XXXVII.

dons are comparatively large, green and fleshy, with a peculiar

one-sided appearance. The leaves are alternate, the first nep-

ionic leaf having three leaflets, the second five. In some cases

the third leaf had five leaflets like the second leaf and in other

cases seven, the greater number having five. This shows as

noted in the previous paper (Studies of Localized Stages of

Growth in some Common New England Plants," Amer. Nat.,

Vol. 36, No. 431.) a difference in acceleration of development in

individual seedlings. An interesting feature of the leaves is the

fact, as was mentioned in the preceding paper, that the tip of the

later leaves repeats the form of the tip of the first leaf. In look-

ing over the figures of this plant it will be noticed that the tips

of all the leaves do not correspond as shown especially in Fig. 4.

This seems at first sight an exception to the principle of local-

ized repetition, but on careful study was seen to be due to the

following growth. The first

few leaves seem to havea

further development after

unfolding. At first the tip

is as given in Figs. 2, 3,

and 5. Then later the tip

broadens and becomes dis-

tally notched. This is seen

also in the seedling, when

the leaf unfolds the tips

being pointed but after-

ward becoming reéntrant

as noted in Fig. 4. The

first leaves of seedling and

spring growth then are re-

tarded even in taking on

such a character as the

shape of the tip.

In early spring. growth

FiGs.2-5. Spring growths of Astragalus adsurgens,

figs. 2, 3, 5 weak growths, fig. 4 stouter more vigor- the first leaves of weaker

T gem shoots are trifoliolate as in

Figs. 2 and 3. Where this is the case the second leaf is

usually of five leaflets. These two leaves then correspond

No. 436.] LOCALIZED STAGES IN PLANTS. 245

exactly with the characters seen in the seedling. In stronger

growths, however, the first leaves have five leaflets or even more.

The second in such cases usually have seven.

Fig. 4 shows exceptionally well a series of stages in early

spring growth. The first leaf has five leaflets, the second seven.

The third leaf which is in the axil of number 1 has eleven leaf-

lets; the fourth, in the axil of number 2 has thirteen. Number

5 farther up the stem has fifteen, number 6 twenty-one. Num-

ber 7 which is not wholly unfolded shows that it will have many

more than the leaf preceding it. The specimen represented by

Fig. 4 was taken from one of the strongest growths found and

does not revert to the very first condition of the leaflets found

in seedlings or feeble growths, as that in Figs. 1, 2, and 3. In

Fig. 4 there would naturally be a stage between leaves 2 and 3

with nine leaflets. In the strong growths these intermediate

steps are very apt to be skipped by acceleration of development.

Such a stage however is found in specimens such as represented

by Figs. 2, 3 and 5. The similarity of the early spring growths

representing localized stages to the seedling representing direct

development is strikingly shown in such plants as this where the

number of leaflets may be actually counted.

Potentilla tridentata Ait. Figs. 6-9.

Seedlings of this plant were found in the spring about the

adult plant and were alike in

having the first nepionic leaf

trifoliolate, but differed in the

characters of the tip of suc-

ceeding leaves. The first nep-

ionic leaf has no distal notches

and is almost exactly like the ,

first leaf of Figs. 6 and 7. Fies. 6-8. Apes i sb. toed apa

The second leaf is very varia- TO Md: ee

ble, it may repeat the characters of the first leaf exactly or the

tip may be tridentate on all three leaflets or on the terminal one

only. When the terminal leaflet is tridentate the lateral leaflets

may be simple like leaf 2 of Figs. 6 and 7, or may have a single

246 THE AMERICAN NATURALIST. (Vor. XXXVII.

notch as in leaf 3 of Fig. 6. The typical leaf of the adult plant

has all three lobes tridentate at the tip — Fig. 8.

This species in its direct development is more accelerated

than others in the genus. Turning to Lubbock’s “Seedlings ”

’92, there are figured Potentilla fulgens Wall, Vol. 1, p. 488,

Fig. 315 and P. repetens L., p. 487, Fig. 314, in which the first

two nepionic leaves are simple, the third trifoliolate. In P. ezser-

ina L. and P. bifurca L. the first leaf only is simple, the second

trifoliolate. Here then is a distinct difference in the rate of accel-

eration, our species noted here having the first leaf trifoliolate.

A large majority of the first leaves in spring growth show

excellent localized stages. Usually the first

leaf is trifoliolate with simple tips on all the

lobes as in Figs. 6 and 7. This is exactly

like the first nepionic leaf of the seedling.

The most common condition for the second

leaf of spring growth, on weak shoots espe-

cially, is a tridentate tip on the terminal

lobe and a simple one for the lateral lobes.

The third leaf may take on the full specific

characters or may have the lateral lobes

with a single notch as in leaf number 3 of

Fig. 6. We have then in Fig. 6 a shoot

which repeats all the stages seen in the

various seedlings mentioned. It is a sort

of ideal condition, repeating all the stages

of the seedling of the same plant, in the

proper sequence without repetition.

The stages beneath the flower or as

shown here beneath the fruit, Fig. 9, repeat

in the reverse order the stages noted in the

straight development of the seedling and in

the localized development of the early spring growth. Above

the highest typical leaf is one which is like the first leaf of the

seedling and first leaf of spring growth, being a trifoliolate leaf

with tips which are simple. After one or two leaves of this type

the leaf drops the lateral lobes and becomes a simple unifoliolate

leaf to the base of the highest flower cluster. This last condi-

flowering and fruiting.

No. 436.] LOCALIZED STAGES IN PLANTS. 247

tion is a simpler and therefore more reversionary character than

any found either in the seedling or the early spring growth.

This may be compared to what is seen in simpler species as

figured by Lubbock, P. repetans L. and others as mentioned

above in which the first nepionic leaf is simple although dentate

in outline. Potentilla tridentata may therefore be considered

accelerated in development in that it has skipped the simple leaf

stage in early development. It is an important fact that in the

ontogeny of the individual the simplest stage found occurs not

in the young but as a definite feature of localized senescence.

Geum virginianum L. Figs. ro, rr.

Of this species seedlings were not obtained, but the early

leaves in other species as figured by Lubbock (Geum freemontit

Hort. and G. coccineum Sibth. and Sin. Vol. 1, p. 486, fig. 313)

are orbicular with varied dentations, more or less deeply cut.

In early spring growth of adults the first leaves are of this

same character. In the specimen, Fig. 10, the first three leaves

62.

Fics. 10-11. gini ing g

had dropped off before it was drawn. The fourth leaf has the

orbicular form like the seedlings as noted but begins to show a

three-lobed character by the deeper notches on the sides. The

fifth is deeply cut and the trilobate character much stronger.

The next leaf number 6 is not only trilobate but has begun to

248 THE AMERICAN NATURALIST. (VoL. XXXVII.

develop the next pair of leaflets proximally. Fig. 11 shows a

stronger growth with the same main features. Leaf number 5

although it has begun to develop the second pair of leaflets still

holds to the rounded form in the terminal leaflet. The next leaf,

number 6, however has become more split up and successive

developments of this character are shown in numbers 7 and 8.

The proximal part of the leaf here shows a failure to develop

full characters as seen in the more simple leaflets toward this

Beneath the flower there is a gradual reduction of these charac-

ters and a return to leaves much like those of earlier growth as

shown both in the seedling and in the early spring growth, the

stages being in the reverse order of sequence.

Lamium sp? (Dead Nettle). Figs. 12-18.

In the seedlings raised, Fig. 12, the cotyledons were foliaceous,

glabrous, petiolate; lamina oblong-reniform, deeply cordate at

the base with a slight distal median notch; petiole nearly terete,

slightly flattened above. The petioles become

much elongated and grow upward for some time

with their bases nearly touching, petioles in speci-

mens raised were an inch long two weeks after

germination. The first nepionic leaves are

Fis. 12. Seedling of Opposite and at this time seem apparently sessile

yledons, first two as the strength seems to be used in the length-

rg Sia es ening of the first intermode; later they become

petiolate. This is a progressive development of the proximal

portion during growth. These first leaves have an orbicular

form, crenate at the margin, a terminal rounded tooth and from

two to four lateral teeth on a side. Following leaves increase

the number of the lateral teeth and the broadest part of the leaf

moves toward the base.

In spring growth, Fig. 13, the first leaf was very similar to the

first nepionic leaf figured, Fig. 12. The number of teeth was the

same and the general orbicular form of the leaf. The second

leaf, Fig. 14, has another tooth added and the broadest part of

the leaf is nearer the base than in the first leaf. The remaining

No. 436.] LOCALIZED STAGES IN PLANTS. 249

leaves, Fig. 15—18, show the increase in the number of the teeth,

the migration of the greatest breadth to near the base of the

leaf, the cordate form of the base and the change in the tip as

Fics. 13-18. Progressive stages in spring growth of Lamium showing increase in number of

notches and gradual auricled form of the base with a pointed form of the whole. Compare

with leaf 1 of fig. 13, the nepionic leaves of fig. 12.

well as of the whole leaf in becoming more pointed.

At the upper part of the flowering stalk the leaves decrease

in the number of teeth until the highest leaves are simple and

lanceolate, usually with a clasping base repeating in reverse

order of sequence stages seen in the seedling and young growth

of early spring, becoming even more simple.

Artemisia stelleriana Bess. Fig. 19.

Although the seedling of this species was not obtained, other

species which have few-lobed leaves as figured by Lubbock (Vol.

Il, p. 135.) Artemisia anuua L. and A. mutellina Vill. have a

development much like what is represented

in Fig. 19. In the early spring growth

which is figured, the first four leaves are

simple becoming progressively spathulate

in succeeding leaves. The next four are

three-lobed and after that the five-lobed

character appears, representing the great- 19 [22

est complexity reached in the species. The fis. 19. Spring growth of

lobes in later leaves are much separated but N APENI mags

do not increase in number. reaching the typical five-

In weak and late growths there is a ma

return to earlier conditions, it being true localized senescence,

appearing late in the season at the end of the season's growth.

250 THE AMERICAN NATURALIST. [Vor. XXXVII.

The lobes are frequently progressively reduced to three and in

rarer instances to the simple form shown at the base of spring

growth. Here then where there is no flower to take the strength

of the plant from the leaves, the diminutions of growing strength

near the end of the season is sufficient to bring out regressive

characters.

Arabis albida. Figs 20-37.

The seedlings of this plant have after the cotyledons a simple

Fics. 20-24. Stages in sity of Arabis albida and may represent -

the stages seen in shoots from the e ground i in spring growth as the

are so nearly identical.

rounded-oval leaf rounded at the base, Fig. 20. The second leaf

is broadly ovate with the widest part at about the middle, four

slight notches making two teeth on each side, very slightly

raised from the general outline and an auricled base, Fig. 21.

| |

25 96 W

Wo om

Fics. 25-32. Stagesin spring growth of Arabis a/bid t from stems of preceding years growth.

These early leaves do not have the auricled base of the early growth from underground.

The third leaf is broadest nearer the base, more pointed and

with three teeth on each side, the base auricled, Fig. 22. The

No. 436.] LOCALIZED STAGES IN PLANTS. 251

fourth leaf has a still more pointed form, the widest portion still

nearer the base which is now deeply auricled, Fig. 23. The fifth

leaf has still more lateral teeth, Fig. 24, and so on.

The spring growth of this plant is of two distinct kinds. In

one the shoots come from underground and are succulent, grow-

ing rapidly with long internodes, in the other the growths come

from the part of the plant above ground which survived from

the previous year. This growth is not so rank and the inter-

nodes are much shorter; a decided difference in the leaves of

the two forms is noted.

In the growth from dd the stages passed through

may be represented by the figures of leaves of seedlings, Figs.

20-24, the stages being exactly comparable to those of the seed-

| 37

33 94 ^ 35 6

Fics. 33-37. Continuation of stages on same stalk as preceding es but nearing the —

x d tee the last leaf is d the flower. These all r revert to the auricled base

ling. In the growth from above ground, however, there is a

decided deviation from this form. The earliest leaves are taper-

ing to the petiole and with but one tooth on a side, Fig. 25.

The second leaf has two teeth on a side but decidedly different

from those of leaves from underground shoots. In both how-

ever the widest portion is near the distal end. The growth con-

tinues, Figs. 26-29, with the number of teeth increasing and

becoming doubly serrate, Figs. 30-32. The greatest width con-

tinues to be at the top.

Beneath the flower however the leaf changes in its characters.

The base becomes auricled, Fig. 33, like the leaves in the seed-

ling and in the growth from the ground. Also the greatest

252 THE AMERICAN NATURALIST. [Vor. XXXVII.

width moves toward the proximal end, the number of dentations

decreases until in Fig. 36 there are but three on each side and

in Fig. 37 there are but two, in form very much like Fig. 21,

the second leaf of early growth from the ground. Its base is

auricled like the seedling and early growth from the ground and

the upper part is like the growth from old shoots.

Poterium canadensis A. Gray. Figs. 38-41.

Early in the spring seedlings of this plant were obtained about

the adult. They were taken up and grown in

pots in order to study variation in the first

leaves. The seedling, Fig. 38, develops as

follows: the cotyledons are broadly oval,

Fic. 38. Seedling of

Poterium ca nse

showing cotyledons c.

c. and first nepionic

leaf

nearly orbicular,

9 obtuse, deeply cor-

date at the base,

> glabrous, sub-

j fleshy, deep green

[^ with petiole sub-

Wes terete, slightly

| channeled abov e.

The first nepionic

leaf is pinnately

trifoliolate, leaflets

petiolate, rotund,

incise-serrate, lat-

eral ones sub-cor-

Fics. 39, 40. Early leaves of Poterium camadense. The leaflets date at base, tip

lettered from a to f show!localized stages in separate leaflets and A

eere inim a 20 Eamon ee e with amie qui

No. 436.] | LOCALIZED STAGES IN PLANTS. 253

shorter serrature than the others. Usually the lateral leaflets

were like the terminal but in some cases were more compli-

cated, Fig. 38, having five teeth in the lateral and three in the

terminal leaflets.

In early spring growth from mature plants, localized stages

are seen especially well-marked in weaker lateral shoots from

the side of the clump. One such is figured, Fig. 39. The first

leaf was like the first leaf of the seedling except that it had a

greater number of teeth on each of the leaflets. The second

leaf was mainly trifoliolate with a trace of the second pair of

leaflets at a, which is noteworthy as it is like the leaflet of the

seedling in having three teeth. The third leaf has five leaflets

and the fourth seven. The later leaves increase the number of

leaflets until there are as many as twenty three or even more in

a single leaf. In the developing leaf the proximal part is the

last formed and oftener shows characters which, being a failure

to develop the full characters, may be compared to the young

and considered as localized senescence, in that individual leaf.

Similar localized reversion of the proximal part of leaves were

shown in the previous paper mentioned (Amer. Nat, Vol. 36, No.

361) in Tansy and here also in Geum. They are shown in

Gymnocladus disicus by Dr. Jackson (Mem. Bost. Soc. Nat.

Hist., Vol. 5, No. 4. * Localized Stages in Development in Plants

and Animals"). These proximal reversionary leaflets repeat

various stages in the development of the typical leaflet of the

plant as shown in leaves lettered a to f in Figs. 39 and 4o.

Leaflets in this part of the leaf may be found representing

various steps from the tridentate character at a which is typical

of the first nepionic leaf to one of thirteen teeth at f comparable

to a much later stage in seedling growth. These leaves also

approach nearer to the earlier leaflets in having less auricled

bases, a character which is especially noted in the seedling.

The leaflets in the adult show an overlapping in the distal portion

and distinct separation in the proximal.

The stages beneath the flower are definite and easily made

out. From a typical leaf having a great number of leaflets, in

succeeding leaves the number becomes gradually reduced until

at some distance below the flower leaves are found having nine

254 THE AMERICAN NATURALIST. (VoL: XXXVII.

leaflets or fewer. In the specimen figured the reduction has

produced at the base a leaf of seven leaflets. From this stage

the reductions following the figure, are as follows, Fig. 41.

At the base no. 1 is a leaf of seven leaflets. The lower pair of

leaflets in this leaf is somewhat changed so that they have almost

the character of stipules. This shows that the reduction should

be thought of as a

failure to develop

characters, even the

distal half of these

two leaflets being

much nearer the usu-

al form than the prox-

imal half. The teeth

on the distal side are

well developed and

the lateral secondary

veins show well, while

on the proximal side

the teeth are few and

poorly developed and

the veins are not dis-

tinguishable. By the

shortening of the

proximal side by its

weaker development,

the whole shape of the

leaflet is changed and

Fic. 41. Flowering stalk of Poterivm canadense showing greatly aborted from

localized senescence below the flower. Breaks in the stem . l f h

are simply to allow its ie put into smaller space. Leaves its usua orm. The

1-5 enlarged to nat same modification of

proximal leaflets is characteristic of leaves below the flower in

other plants. To continue with the development toward the

flower, number 2 will be seen to have lost this third pair and the

second pair has assumed a like abortive character. Here the

whole proximal half is without teeth. This simple part of the

lowest pair of proximal leaflets has varying degrees of reduction

in different cases. The leaf number 3 has been reduced to three

No. 436.] LOCALIZED STAGES IN PLANTS. 255

leaflets, the lower two of which have assumed the same abortive

form as described. Leaf number 4 has simply the terminal leaf-

let and that greatly reduced in number of dentations. Number

5 has but three dentations characteristic of the terminal leaflet of

the first nepionic leaf but the lateral proximal border expanded

in a manner suggestive of the proximal modification of preceding

leaves.

These stages are exactly comparable to the stages of the

seedling and early spring growth in number of leaflets. In this

reduction as noted before in other plants, stages are developed

as in numbers 4 and 5 which are simpler than any shown in the

direct development of the seedling or in the localized develop-

ment shown in early spring growth. This is again a case of

repeating stages in the reverse order from that of the seedling

and early spring growth.

Ribes aureum Pursh. Figs. 42-45.

This plant which is grown quite extensively as a garden shrub

shows excellent localized

stages in its spring growth

on all parts of the plant as

well as below the flower.

The flowering season is early

and the very earliest leaves

are persistent enough so that

they are still on the plant and

can be compared with stages

beneath the flower. In some

cases among the lowest shoots

which bear flowers the series

can be traced entire in the

growth from one bud, from

the simplest early leaf to the

complex and back through pes. ura. Spring growth of Rides aureum.

the simple again to the flower. Fig. 42 most complicated leaf attained by the

In early spring growth the — "^^ :

simplest leaf usually found is like leaf 1 of Fig. 43, a leaf with

256 THE AMERICAN NATURALIST. [Vor. XXXVII.

three lobes and tri-nerved. The next leaf is more decidedly tri-

lobed but with a division into teeth. The next leaf has the lower

lobes divided and this division becomes somewhat more marked

in later leaves until the most complicated leaf is developed as in

leaf 7, Fig. 42. In Fig. 44 is shown a leaf intermediate between

leaves 1 and 2 of Fig. 43. By obtaining more specimens a more

complete series may be secured between many of thestages. In

Fig. 43 a significant thing is shown in the size of leaf 2. The

first leaf grew under unfavorable conditions and its development

in size went but a little way. The weather changed however

and became warmer while leaf 2 was developing and its unusual

size is the result. Next it was colder and leaves 3 and 4 felt

the effect. This was noticeable in many cases at

this particular time.

Turning to the flower Fig. 45, the leaves are

reduced as shown in the lowest leaf figured to

FA. leaves corresponding to Fig. 43, leaf 1. But as in

e oo the cases mentioned before, the reduction is carried

reum below Still farther as leaf 2 is simple in outline with no

ee traces of the teeth or lobes of the simplest leaf of

spring growth. This simpler form is found in a great number

of cases in the last leaf below the flower. As in cases previously

mentioned this stage is simpler than in its own young.

Chrysanthemum speciosum. Figs. 46, 47.

The seedling of this species repeats the essential steps shown

in Fig. 47 of the spring growth, but less completely. In the

early spring growth more stages are represented. A very weak

growth is shown in Fig. 46. In this the first leaf was nearly

simple having a lobe on one side only. The next leaf was three-

lobed and the following also. The fourth was five-lobed. In a

stronger growth, Fig. 47, the first leaf was three-toothed, the

lobes being simply teeth and nothing more. The next three

were simply three-lobed and the fifth and sixth instead of having

five lobes produced a complication of the lower pair of lobes.

These have two teeth in leaf five and in leaf six one has the

characteristic three-toothed form. In the remaining leaves the

No. 436.] LOCALIZED STAGES IN PLANTS. 257

five-lobed character is shown and the lobes usually each assume

a three-obed character in

which each lobe is compar-

able to the first leaves of

Fig. 46.

Beneath the flower the

plant repeats, in the reverse

order, these stages with great

definiteness. Beginning with

the typical complicated leaf at

the base of the flowering t

branch, stages are repre- 46

sented gradually losing their ri. Pd Early spring growths of Chrysanthe-

Characters in the reverse """séeciomm. *

order to the development figured, until simple five-lobed leaves

are reached, then three-lobed ones and finally beneath the flower

are simple leaves which are even simpler than the first leaves of

the seedling. The reduction is complete and exactly retraces

the stages of the seedling and early spring growth as localized

senescence.

Artemisia abrotamum (Tourn. Neck. Figs. 48, 49.

In this species of the genus before given, Fig. 19, the leaves

become much more complex in their lobing and the plant fre-

quently blossoms. Seedlings were raised, the first nepionic

leaves being three-lobed, the

next five-obed and so on.

z Inthe spring growth figured,

Figs. 48, 49, the first leaves

49 o are three-lobed, Fig. 48,

48 : leaves 1-5, Fig. 49, leaf 1.

Fras. 48, 49. Two examples of early spring The next leaves are five-

E in Ave. abrotamum showing dif- lobed, Fig. 48, leaf 6, Fig.

49, leaves 2, 3. Then comes

à splitting up and complication of lobes until the adult leaf is

much dissected. Below the flower the return in the reverse

Order to simple condition is perhaps as well marked as in any

258 THE AMERICAN NATURALIST, | [Vor. XXXVII,

case yet given. The return to a five-lobed leaf, then a three-

lobed leaf, and finally to a simple leaf beneath the flower, a con-

dition simpler than any seen in the seedling or spring growth as

in so many other cases. This species is a good one for studying

localized stages as the leaves are much cut up and pass through

many, but nevertheless definite, stages.

Bryophyllum tubiflorum Harv. Figs. 50, 51.

This plant is known to many by its power of reproducing by

buds from the notches of the sides of the leaf. When a leaf is

placed on damp sand or earth a bud soon starts out from each

of these lateral notches, and young plants result, some showing

greater strength than others. The leaf shrivels and finally dries

up leaving a ring of new plants. These young

plants show localized stages comparable to the

seedling. One of the growths from such a

leaf bud is shown in Fig. 51. The first pair

of leaves are entire, the second have a single

notch on each side, the third two on a side

n and so on. This repetition of characters is

leaf in n Bryophyllum rom as strong as any seen in early spring growth.

on In the study of localized stages in develop-

ment, as also noted in the preceding paper, in the study of spring

growth the leaves of weaker shoots seem to be better than more

vigorous ones for study as they repeat the greatest number of -

stages and revert to more primitive characters. Not only is this

tendency to a repetition of characters found in spring growth

from the ground, but also from the stems of the preceding

year’s growth and even as in the last case noted in growths from

the leaves themselves.

In the leaves of flowering shoots or weak terminal growths at

the end of the season the same stages are passed through as in

the seedling or spring growth but always in the reverse order of

sequence. This which is considered localized senescence is

directly comparable to senescence of the whole individual as

shown by Prof. Hyatt in his studies of cephalopods. In several

cases described it is shown that beneath the flower the leaves

No. 436] LOCALIZED STAGES IN PLANTS. 259

are much simpler than in the seedling as shown in Potentilla

tridentata, Ribes aureum Poterium canadensis and others.

Such extreme reversion may be compared to the young of other

species in the genus in which the first leaves do have more primi-

tive simplicity.

This life history of a plant marked by stages in the leaves can

be made out in many, perhaps all plants where the leaves are

compound, notched or lobed and even sometimes where the

leaves are entire, by changes in venation and outline.

CAMBRIDGE, Mass.

January.

NOTES ON THE REPRODUCTION OF CERTAIN

REPTILES. !

C. S. BRIMLEY.

For several years I have handled a number of reptilian eggs

in order to secure the embryos, and the following observations

are based mainly on those collected during the past three sum-

mers (1900, 1901 and 1902). The eggs of several species of

snake and lizard are found in summer (from about the middle of

May to the middle of August) by farm hands who plough them

up, when breaking up land that has been in clover or wheat or

some other early crop; from this it can be understood that the

eggs are deposited at no great depth in the ground. Eggs of

the Painted and Mud Turtles (Chrysemys picta and Kinosternon

pensylvanicum) are often ploughed up in the low grounds.

The eggs of the Black Snake (Bascanion constrictor) are more

frequently found than those of any other species of snake, these

are a short oblong in shape, often lumpy and one sided or irregu-

lar, and covered with a thick skin with a rough surface to which

dirt readily adheres so that the color is usually a dirty white.

The eggs are free, not adherent to one another in clusters as is

the case with some other species; in size they vary from about

26 to 40 mm. in length, by 21 to 28 in width, and are found in

lots of from 5 to 22 in number, the larger lots usually consisting

of larger eggs than the smaller lots, from which one would natu-

rally infer that the smaller lots of smaller eggs were probably

laid by smaller and younger individuals, and the larger lots of

larger eggs by larger and older specimens.

The eggs must take at least a month to hatch, possibly much

longer. A lot of 13 eggs were brought in June 28, 1900,

and four of them were kept until they hatched on July 25,

twenty-seven days later, the young snakes emerging through a

1 Read before the first annual meeting of the North Carolina Academy of

Science, Nov. 29, 30. 1902.

261

262 THE AMERICAN NATURALIST. | [Vor. XXXVII.

longitudinal slit in the egg, sometimes there are several of these

slits; in this lot the markings of the young snakes began to

show on those preserved on July 6, nineteen days before hatch-

ing. Another lot of 21 eggs of this species brought in July 11,

1902, in which the spotted color pattern had begun to appear on

the embryos were kept till some of them hatched on July 28,

seventeen days later, three young snakes which hatched,

measuring respectively 285, 300 and 303 mm. in length.

Another lot, obtained in 1900, contained among others, one egg

which, though entirely normal in external appearance, was very

abnormal internally, inasmuch as it contained two embryos, and

one of these was a two-headed monster.

Occasionally an egg of this species is pyriform in shape, much

like a killdeer's egg, as was the case with one egg of the lot of

thirteen mentioned previously, and with one of a lot of five

brought in June 28, 19or.

Another species whose eggs are sometimes brought in is the

Spreading Adder (Heterodon platyrhinus). The eggs of this

species are about the same size and shape as those of the Black

Snake, but the skin of the egg is smooth and very thin, much

thinner than in any other species of snake whose eggs I have

handled. Like those of the Black Snake, the eggs of this

species are free, not adherent to one another in clusters.

A lot of 13 eggs was brought in June 23, 1900, from which

embryos were put up from time to time, till a young Heterodon

was hatched from the last egg on Aug. 14, fifty-two days later,

the young snake 200 mm. in length, emerging from a longitudi-

nal slit in the egg ; the only egg in this lot measured was 39

mm. long by 29 in breadth and oblong in shape.

Another lot of 26 eggs was brought in June 27, 1902 (ten

other eggs were said to have been broken when the lot was

ploughed up) and were about 33 mm. long, a short oblong in

shape, and were kept till a young snake 185 mm. long was

hatched from the last egg on Aug. 5, thirty-nine days later.

On Aug. 1, 2 and 3, 1901, a Spreading Adder in captivity

laid sixteen eggs, similar to the two lots previously mentioned

but whiter in color, owing to their not having been in contact

with the soil. These eggs although only just laid, contained

No. 436.] REPRODUCTION OF REPTILES. 263

small embryos and in this connection I may state that I never

yet opened a snake’s egg which did not contain an embryo large

enough to be recognized as a snake. Lizard's eggs, however,

not infrequently have the embryo small enough to escape notice,

and a large majority of all turtle eggs brought in are quite fresh.

The yolk of a snake's egg too is whitish in color and thicker in

consistency than the yellowish yolk of a lizard's egg, while the

white and yolk of turtle's eggs resemble those of bird's eggs very

closely in general appearance. These remarks, however, are

only meant to apply to the different species I have myself

examined.

The eggs of the King Snake (Ophibolus getulus) are long,

oblong in shape, with a smooth tough skin and are more or less

adherent to one another in clusters. A lot was brought in July

II, 1900, some of which were put up from time to time till

three young snakes, 275 mm. long, were hatched from the last

eggs on Aug. 14, thirty-four days later. These eggs were about

40 to 43 mm. long by 24 to 26 wide.

In July, 1900, a King Snake in my possession laid 17 eggs in

confinement ; these eggs were like the foregoing lot, but smaller,

and were also adherent in clusters. One egg of this lot con-

tained an embryo with two heads, and two bodies, the bodies

separate for the anterior one third of their length.

Another lot of 10 eggs laid in confinement in July, 1901,

were stuck together in two clusters, four in one lot and six in

the other, and measured 31 to 35 mm. in length.

On Aug. 17, 1901, a Striped Chicken Snake (Co/uber quad-

rivittatus) from Georgia, laid 9 eggs in confinement, two lots

of two each were adherent, the rest free. The eggs were long

and narrow with a tough white skin and measured 37 to 43 mm.

long by 17 to 20 wide.

On July 25, 1901, I found 17 eggs in a box in which I was

keeping two species of Coluber (guttatus and guadrivittatus);

these eggs were much like those of the King Snake but longer

in proportion and contained very small embryos. Nine eggs

were stuck together in a cluster, six in another and there were

two free eggs.

A black Chicken Snake (Coluber obsoletus) laid a number of

264 THE AMERICAN NATURALIST. [Vou. XXXVII.

eggs in confinement in the summer of 1899 from which young

snakes were finally hatched, but I am sorry to say I did not take

any notes. A large specimen of the western Bull Snake (Pzty-

ophis sayi) also laid eggs in captivity the same summer, but I

kept no record of these and can only say the eggs were consid-

erably larger than those of the Coluber obsoletus and that young

snakes finally hatched out from ten or twelve of them. I have

no record and cannot remember certainly whether these eggs or

those of the Chicken Snake were free or adherent, but I think

both lots were in the latter condition.

On July 5, 1902, a Milk Snake (Ophibolus doliatus triangulus)

from Michigan, in my possession, laid 15 eggs all adherent in one

cluster; there were about 30 mm. long, with smooth, tough,

white skin and short oblong in shape. On the same day a dead

Green Snake (Cyclophis estivus) was brought to me, which con-

tained in its oviducts four elongate, thin skinned eggs (20 to 22

mm. long) which contained small embryos.

On July 12, 1902, two lots of snake's eggs were brought to

me, different from any I had previously obtained. One lot con-

sisted of two elongate, smooth, whitish eggs, 23 and 25 mm. long.

One was put up and the other kept until Aug. 14 (thirty-three

days later) when a young Carphophiops amænus within a day

or two of hatching was taken from the egg. The other lot con-

sisted of eight eggs, short, oblong in shape, just about the size

and shape of the eggs of the lizard, Sceloporus undulatus, but

smooth skinned and one-sided and about 16 to 17 mm. long.

These were kept until Aug. 8, when two young snakes 185 mm.

long, also Carphophiops amanus were hatched from the last eggs.

Practically all the lizards' eggs brought in belong to two species,

Sceloporus undulatus (the Fence Lizard) and Cnemidophorus

sexlineatus (the Sand Lizard locally known as Sand Trotter,

Sand Skeeter and Sand Sister), more than an hundred eggs of

each of these species are brought in every year, those of Scelo-

porus predominating in the early part of the season and Cnemi-

dophorus in the latter part.

The eggs of Sceloporus undulatus are short, oblong in shape,

measuring from 14 to 18 mm. long and about 10 to 13 mm.

wide, and the skin is roughened causing dirt to readily adhere to

No. 436.] REPRODUCTION OF REPTILES. 265

it. The embryos show the characteristic dark cross bars across

the back, when only about half hatched. The number of eggs

laid in one lot is from ten to fifteen. One rather curious fact in

connection with this species and the next is that the eggs

increase in size after obtrusion ; when first laid the eggs are rela-

tively long and slender, but as the embryo develops the eggs

increase in thickness if not in length. On June 28, 1902, four

very slim and narrow eggs of this species were brought to me; I

preserved one and kept the other three in dry earth in a corked

bottle ; on July 23, the eggs had materially increased in size and

a second egg was preserved, on Aug. 11 a third egg was put up;

on Aug. 23, the remaining egg was still unhatched, but the skin

had become loose and flabby, on Sept. 3, the next time I looked,

a young lizard had hatched out, 67 days after the eggs were

brought to me (the egg, however, probably hatched a day or two

after Aug. 23).

The eggs of Cnemidophorus sexlineatus are similar in general

appearance to those of S. undulatus, but somewhat larger (about

17 to 22 mm. long by r1 to 14 mm. broad) with a smooth skin,

to which dirt does not adhere so readily as to the rough skin of

the Sceloporus eggs, and hence they usually look much whiter.

The markings on the embryo do not show until a week or so

before hatching. On July 8, r9or, three small white eggs

which from their size (93 mm. long) could only belong to the

Ground Lizard (Leiolepisma laterale) and which contained lacer-

tilian embryos were brought to me, and another lot of three sim-

ilar eggs on July 2, 1902, which were said to have been found in

a rotten stump. :

Of the Testudinata, the eggs most frequently brought m are

those of the little Mud Turtle (Kinosternon pennsylvanicum) ;

these are hard shelled usually rather blunt at the ends and vary-

ing from a rather short to a rather long oblong in shape, varying

from 23 to 29 mm. in length; the eggs are sometimes ploughed

up, but are often said to have been found in “holes in the bank

of Walnut Creek. Some of the larger eggs attributed to this

species may be those of Aromochelys odoratus the Musk Turtle.

In preparing the shell of an adult Aromochelys tristycha (a

smaller species than A. odoratus or K. pensylvanicum) from

266 THE AMERICAN NATURALIST. (VoL. XXXVII.

Texas, two hard shelled eggs were taken from the oviducts,

Aug. 20, 1902. These eggs were about 26 mm. long and

resemble the smaller eggs of Kinosternon.

The eggs of the Painted Turtle (Chrysemys picta) which are

not infrequently brought in, are larger (32 to 34 mm. long) and

more elongate than those of the Mud Turtle, and are covered

with a smooth, thin, crisp skin instead of a hard shell.

The eggs of the Chicken Turtle (Detrochelys reticulata), a

number of which were taken from the oviducts of two or three

dead females from Georgia, in Nov., 1900, are very similar to

those of Chrysemys, but larger, measuring from 37 to 40 mm.

long.

On June 9, 1901, 26 spherical white eggs about 26 or 27 mm.

in diameter and said to have been found buried in the sand near

a pond, were brought to me; these were said to be “turtle

eggs " i. e, those of Chelydra serpentina and I have no reason to

suppose the identification was incorrect. The eggs are covered

with a thin, crisp skin as those of the Emydoid turtles are.

Of the viviparous snakes of the Natricine group a few of my

observations deserve to go on record. On Aug. 2 and 3, 1894,

a Natrix leberis gave birth in captivity to thirteen living young.

On July 28, 1900, a Virginia elegans from Mississippi gave birth

to five young, and in August, 1899, a Liodytes alleni from

Florida to six.

RALEIGH, N. C.

SYNOPSES OF NORTH-AMERICAN

INVERTEBRATES.

XVIII. THE AMPHIPODA.

S. J. HOLMES.

THE present key is restricted to the species of amphipod

Crustacea of the Atlantic coast of North America. The amphi-

pod fauna of the Pacific coast is very imperfectly known and a

key to the forms already described would include only a small

proportion of the numerous species of that region. The species

from the Arctic regions of the American continent are also not

included. Most of these are circumpolar in their distribution

and only those are described which range into the region

covered. The majority of the known species of eastern North

America are, however, representatives of this circumpolar fauna

and are found also on the northern coast of Europe and Asia.

The amphipod fauna of Labrador is very similar to that of

Norway, the differences naturally becoming greater as we pass

southward along the shores of the two continents. Nevertheless

there are not a few species common to the Mediterranean and

the southern coast of New England. The tendency of some

writers to describe a species as new when met with for the first

time in North America has therefore resulted in the production

of many synonyms.

While the labors of several English, German, Danish, and

Norwegian naturalists have made the amphipod fauna of Arctic

America fairly well known, we have almost no information con-

cerning the Amphipoda of the southern portion of our own coast.

The veteran American naturalist, Thomas Say, has described a

few species from the shores of the southern states, but no

successor has followed in his footsteps. Prof. Smith in the

important Report upon the Invertebrate Animals of Vineyard

Sound by Smith and Verrill made a list of the Amphipoda of

southern New England and described several new species, and

1 From the Zoölogical Laboratory of the University of Michigan, Ann Arbor,

Mich.

267

268 THE AMERICAN NATURALIST. [Vor. XXXVII.

subsequently he added considerably to our knowledge of the

amphipods of northern New England and Labrador. Since the

publication of Prof. Smith's papers only a few forms have been

made known from any portion of the eastern coast of America.

Stimpson in his Marine Invertebrata of Grand Manan instituted

several supposed new species, described after the fashion of

that versatile zoólogist, with very unequal degrees of fulness ;

some of the species are very well characterized while the

description of others is so meagre their recognition is practically

impossible. Many of Stimpson's species are identical with forms

previously described from Europe or Arctic America.

Most of the species included in this key will be found more

fully described and figured in a paper by the writer on the Amphi-

poda of southern New England, which is now in course of publi-

cation. The specimens I have secured at Wood's Hole, Mass.,

and the collections of the U. S. National Museum, the U. S.

Fish Commission, the Boston Society of Natural History and

several smaller collections which were kindly sent me for

examination have afforded several new species and many others

which were heretofore not known to occur on the New England

coast. While it may be unfortunate that a specific name at its

first introduction to the public is unaccompanied by full descrip-

tion and figures, such a procedure is unavoidable in a key of this

kind and it will not be long, I trust, before more extended

descriptions and figures of the new species here mentioned will

appear.

Many of the numerous families of the Gammaridea which have

been proposed appear to me to be based upon very inadequate

foundations, and I find it practically impossible to separate them

all by an analytical key. As it would not be feasible to present

the needed changes in classification in this paper, even were I

prepared to venture upon such a task, I have disregarded the

so-called family ties in dealing with this group and have prepared

a key leading directly to the genera. Following previous con-

tributors to this series of papers on North American Inverte-

brates I have adopted the following symbols to. designate

geographical regions.

A. Arctic America to Cape Cod.

No. 436.] NORTH AMERICAN INVERTEBRATES. 269

M. Cape Cod to Hatteras.

S. Hatteras to Florida.

The Amphipoda may be defined as malacostracous Crustacea

with a well-defined head, no carapace, sessile and usually com-

pound eyes, a thorax of seven segments, and an abdomen

consisting typically of six segments and a telson. The gills are

borne on the inner side of the basal joints of the thoracic legs.

The first three abdominal appendages are fitted for swimming ;

the last three pairs are very different from the preceding ones

and are directed backward and fitted for springing. The eggs

are carried in a marsupial pouch under the thorax of the female.

The term perzeopod is here used to designate the thoracic legs

behind the first two pairs, or gnathopods.

The Amphipoda fall into three main divisions which may be

separated as follows:

Abdomen well developed.

Head generally large with very large eyes. Maxillipeds without palps.

eriidea.

Head and eyes generally not of unusual size. Maxillipeds with palps.

ammaridea.

Abdomen rudimentary. First thoracic segment fused with the head.

Amphipoda of aberrant structure. . . . . . . . Caprellidea.

Tribe HYPERIIDEA.

Head generally large and tumid with very large eyes. Maxillipeds devoid

of palps, the basal segments fused together in the middle line. Coxal plates

small. Last two segments of the abdomen coalesced. Uropods usually

with laminate rami.

The Hyperiidea are exclusively pelagic.

habit of living within the bodies of marine animals such as medusz,

ctenophores and salpæ. Like most pelagic organisms the species of this

group have usually a wide range. To insure including all the species liable

to be found off the Atlantic coast of North America would involve an

extensive treatment of this division of the Amphipoda; only a few of the

more common species, therefore, are here described. Full descriptions and

figures of most of the known species of this group may be found in

Bovallius’ excellent Monograph of the Amphipoda Hyperiidea. All the

species here mentioned belonging to the family Hyperiidae are described

and figured in Sars’ Crustacea of Norway. The other species in the key

are treated of by Dr. Stebbing in the Amphipoda of the Voyage of the

Challenger.

Many species have the peculiar

270

THE AMERICAN NATURALIST. (VoL. XXXVII.

KEY TO THE SPECIES.

A. Antenne retractile in depressions on the under side of the head.

AA.

Head produced in front into a long rostrum. Basal joints of the

fourth and fifth perzopods thin and weak; fifth perzopods small.

Oxycephalide.

Second antennz in the male five-jointed, wanting in the female. A dor-

sal carina from the rostrum to the tip of the telson. Proximal part

of head tumid, almost covered by the large eyes; outer margin of

the head serrate. A lateral carina on the rostrum, thorax, and first

three segments of the abdomen. First three abdominal segments

with a pair of acute projections at the lateral angles, the margins

of the upper projection finely serrate. Basal joints of the third and

fourth peræopods broad, coarsely serrate in front and finely serrate

on the very convex posterior margin.

Oxycephalus clausi Bovallius. M.

Antenne carried on the anterior side of the head. Head not

attenuate in front.

B. Head not large; eyes covering only a small part of the head; first

antennz with the first joint of the flagellum very T and com-

pressed, the terminal joints minute. . . Vibilia.

The species of this genus are icmesda: difficult to determine

without full descriptions and figures and the reader had best

consult the monograph of Bovallius above referred to. Several

species occur off the Atlantic coast of North America.

BB. Head large and tumid, the large eyes covering a large portion

of the surface.

C. Eyes divided into an upper and a lower portion. No mandib-

ular palp. Fifth peraopods normal. . . (Phronimidae).

Head about as deep as the length of the first six segments

of the thorax. Lateral angles of the first three abdominal

segments acuminate. Gnathopods small simple, the carpus

distally produced. Third perzopods enlarged and chelate

or subchelate, the hand very variable in form and differing

with age and sex. Often found in the tests of Salpa and

Pyrosoma. . . Phronima sedentaria (Forsk.) A. M.

CU per not divided. Fifth comnts reduced or transformed.

Uropods devoid of rami, the peduncles laminate (Phros-

inidz).

First and second gnathopods simple. First three pairs

of perzopods subchelate, the third very large and stout,

the carpus very broad with the lower margin nearly

straight and armed with 6 or 7 teeth ; propodus slender,

slightly curved, closing against the toothed edge of the

carpus ; a slender, scarcely half the length of the

propodus. . Anchylomera blossevillii Edw. M.

No. 436] WORTH AMERICAN INVERTEBRATES. 271

CCC. Eyes not divided. Uropods with rami.

Inner ramus of the uropods fused with the peduncle.

DD.

No mandibular palp . . (Cystisomida).

Head large, the upper half. or more covered by the

large eyes, the lower margin dentate. First two tho-

racic segments fused. Body with a median dorsal

carina; posterior margins of most of the segments

dentate. ^ Gnathopods small, complexly subchelate,

the margins of the carpal process and propodus acutely

serrate. Attains a length of four inches. Transpar-

ent. . Cystisoma spinosum (Fabr.) A. M. S.

dete ramus ot the uropods free. Head very large,

tumid, the sides entirely covered by the large eyes.

Flagellum of both pairs of antennz long and multi-

articulate in the male, but short and unjointed in

the female. Mandibles with a palp (Hyperiidz).

Carpal joint of both gnathopods produced distally

as far as the tip of the propodus . . Hyperoche.

Second antenna in the female much smaller than

the first. Gnathopods almost naked. Carpus of

the first two perzopods narrow, the posterior

margin acute and serrate. ` Carpal process of both

gnathopods narrowly triangular, somewhat

exceeding the propodi, and serrulate above.

Hyperoche tauriformis (Bate), A

KE. Carpal joint of both gnathopods not produced or

not so much so as in Hyperoche.

F. Third pereopods much elongated. Second

and third paio om with the carpus much

dilated. . . . Euthemisto.

Body compressed, called; the last two

thoracic and first two abdominal segments

with a marin upturned tooth. Carp

joints of the first two peræopods oblong-oval,

widest near the middle, furnished posteriorly

with short, unequal sete. Third peraopods

not greatly exceeding the others. Usual

length 12 mm. . Æ. compressa (Goes), A. M.

Body carinated above, the last two thoracic

segments with posterior recurved teeth.

Carpus of the first perzopods most dilated

near the base where the posterior margin is

strongly convex. Third pereopods greatly

elongated, the carpus very narrow, much

longer than the propodus, and pectinated in

272

THE AMERICAN NATURALIST. [Vor. XXXVII.

AF.

GG.

front with spines which increase in length

toward the distal end where they may equal

the diameter of the joint. Usual length,

15 mm. . . . JA. spinosa (Boeck), A. M.

Back not carinated, with no dorsal spines.

Carpi of first perzeopods not widest near the

base and furnished posteriorly with elongated

sete. Dactyl of the elongated third pere-

opods with a tuft of spinules near the base.

fmm... .-. .Jbellula (Mandi), A

Last three pereopods of subequal length.

Carpus of the first and second, perzopods

of the usual form

G. Propodi of the last three peraopods

greatly elongated. Antenne subeqaul

in the female. . Parathemisto.

Body rounded above. Gnathopods hir-

sute, the first not distally produced ;

carpus of the second pair produced

beyond the middle of the propodus.

P. oblivia (Kroyer), A

Propodi of the three posterior perzeopods

not greatly elongated. Antenne very

small in the female. Head much

deeper than long, flattened in front.

Hyp

Gnathopods sparingly setose, the

carpus of the first pair produced into a

prominent lobe; carpal lobe of the

second pair reaching beyond the middle

of the propodus. Often found in

Aurelia. . H. galba (Mont), A. M.

Gnathopods thickly setose. Carpus of

the first pair scarcely produced distally,

that of the second not produced nearly

to the middle of the propodus. Often

found in Cyanea.

. medusarum (Müll), A. M.

Tribe GAMMARIDEA.

Head and eyes usually not greatly enlarged. Maxillipeds with palps, the

basal lobes not fused together in the middle. Coxal plates generally well

developed.

rare exceptions free.

bdomen not reduced in size, the last two segments with

No. 346.] NORTH AMERICAN INVERTEBRATES. 273

KEY TO THE SPECIES.

A. Eyes four, sometimes apparently only two (Haploops), or rarely absent,

each with a simple lens. Last two segments of the abdomen fused

together. Gnathopods slender. Terminal pereopods quite different

from the preceding ones, the basal joint enlarged and distally pro-

duced behind into a ciliated lobe. Telson cleft. . (Ampeliscida).

Lower eyes apparently absent. Basal joint of posterior perzeopods

not greatly expanded. Telson deeply cleft. . . . Haploops.

C. No long dorsal sete. Basal joint of posterior peraeopods

distally narrowed. . . #. tubicola Lillg. A.

CC. Back with fascicles of bas sete. Basal joint of posterior

perzeopods not distally narrowed. No corneal lenses.

D. Distal lobe of the posterior perzopods reaching about

to the middle of the merus. Antenna in the female

n and much over half the length of the body.

. AH. selosa Boeck, A.

DD. Distal tebe of the basal joint of the posterior perxo-

pods scarcely reaching beyond the ischium. First

antenne in the female markedly shorter than the

suit and less than half the length of the body.

19 m š . H. robusta Sars, A.

BB. Lower eyes plainly visible.

C. Telson elongated and cleft nearly to the base. . Ampelisca.

D. Posterolateral angle of the third abdominal segment

produced into an acuminate process.

E. Head about as long as the first three segments of

the thorax. Posterior perzopods with the basal

lobe nearly transverse below ; the carpus not strongly

produced anteriorly. 4. macrocephala Lillg., A. M.

Posterior perzopods with the basal lobe obliquely

truncated below, the carpus produced anteriorly.

Head much shorter than the first three segments

of the thorax.

A. eschrichti Kroyer, A. (= A. ingens Bate).

DD. Postero-lateral angle of the third abdominal segment

not produced.

E. Merus of the posterior pereopods distally produced

posteriorly nearly to the middle of the carpus.

Body much compressed, telson narrow.

A. compressa sp. nov., M.

Body not unusually compressed. Telson broad.

A. agassizi (Judd), M

EE. Merus of posterior pereopods not produced

274

THE AMERICAN NATURALIST. [Vor. XXXVII.

posteriorly, basal joint of the usual width,

obliquely truncated below.

A. spinipes Boeck, A. M.

CC. Telson short, seldom cleft to the middle. . . . Byblis.

Lower margins of the anterior coxal plates serrated. Body

with stellate pigment cells. . B. serrata Smith, M.

Lower margin of the anterior pem plates not serrated.

B. gaimardi Króyer, A.

AA. Two compound eyes, or rarely the eyes rudimentary or absent.

B. First antennz shorter than the second. Mandibles devoid of a

Terminal uropods with a single uniarticulate ramus. Telson

short and thick. Body compressed. . (Orchestiidz.).

First antennz exceeding the tip of the abun of the second

pair; aquatic forms. . . . Allorchestes.

First antenne nearly i T as lotig as the second which

are scarcely a third the length of the body.

A. littoralis St., A. M.

CC. First antennz much shorter than the peduncle of the second ;

terrestrial forms.

D. First gnathopods in both sexes subchelate. . Orchestia.

First antennz not quite reaching the tip of the penulti-

mate segment of the second. Hand of the second

gnathopods of the male with a notch near the posterior

end of the palm. Carpus of the posterior perzeopods

in the adult male much swollen. Common under

masses of seaweed near the shore. Very active.

O. agilis Smith, A. M. S.

First antennz reaching slightly beyond tip of the

penultimate joint of the second. Hand of the second

gnathopods of the male oval, the palm regularly curved,

with a slight prominence at the posterior end but not

notched. Common around salt marshes.

O. palustris Smith, A. M. S.

DD. First gnathopods simple in the female; large species.

Talorchestia.

Second antennz in the male about as long as the

body. Hand of the second gnathopods in the male

oblong, the palm with a large lobe near the middle

and a large prominence at the posterior end. Very

common on sandy beaches

T. longicornis. (Say), A. M.

Second gnathopods in the male about a third as long

as the body. Hand of the second gnathopods of the

male subovate, distally widened, the palm evenly con-

No. 436.] - NORTH AMERICAN INVERTEBRATES. 275

vex with no lobe near the middle, but defined poste-

riorly with a prominence. Habitat similar to that of

the preceding species.

T. megalophthalma (White), M

BB. Without the combination of characters of B.

C. First two pairs of perzopods devoid of spinning glands.

D. Last pair of pereopods much longer than the preceding:

ones, with the dactyl very long and styliform. Eyes

nearly contiguous above near the end of the projecting

ront A (CEdiceridz).

E. Carpus of as anterior Pod devoid of a

prominent posterior lobe. . . Parediceros.

Rostrum horizontally pródurad and abruptly

deflexed at the tip which forms only a slight

angle inferiorly. Second gnatho narrowly

oval, the palm longer than the upper part of the

posterior margin of the hand ; posterior process of

the carpus reaching the end of the palm. Telson

oblong, truncated atthe tip. Attains 22 mm.

P. lynceus (Sars), A

EE. Carpus of the anterior perzopods prolonged into

a long lobe which extends behind the hand.

Monoculodes.

F. Eyes near the base of the deflexed rostrum.

M. demissus St., A.

FF. Eyes in front of the base of the rostrum

Second gnathopods with the carpal process

scarcely extending beyond middle of palm ;

palm about as long as the upper part of the

posterior margin of the hand.

M. edwardsi, sp. nov. M.

Second gnathopods with the carpal process

extending much beyond middle of palm;

palm shorter than the upper part of the

posterior margin of the hand.

M. borealis Boeck, A.

DD. Without all the characters of D.

Rostrum produced into a hood over the antenna.

erimus perzopods much longer than the last

pai (Phoxocephalidz).

F. Pal * the first andi two-jointed.

Harpinia.

Lower margins of coxal plates fringed with

plumose seta. Eyes wanting. Infero-lateral

angle of the third abdominal segment pro-

276

THE AMERICAN NATURALIST. (VoL XXXVII.

duced into a long upturned spine. Basal

joint of the last pereopods with about

five more or less distinct serrations. Gnatho-

Wee of nearly equal size and of similar

H. plumosa (Kr.) A. M.

FF. ig of the fist maxillz one-jointed.

G. Second gnathopods dd larger than

the fret. ... . Phoxocephalus.

Eyes UMEN ON developed. Coxal

plates with simple marginal sete.

Infero-lateral angle of the third abdom-

inal segment rounded.

P. holbolii (Kr.), A. M.

GG. First and second PEREN of equal

sie... . Paraphoxus.

Eyes ideo Basal joint of

posterior perzeopods oval, not coarsely

serrate. Legs spiny; merus of third

pereopods broader than long.

P. spinosus sp. nov. M.

EE. Without all the characters of E.

Body compact, with well developed coxal plates.

i h

Mandibles not denticulated, and furnis

with a three-jointed palp. First antenna

with a short, thick base and a secondary

flagellum. Second gnathopods elongated,

slender, flexible, with the ischium elongated

and the hand small and furnished with dense

patches of short sete; dactyl rudimentary

(Lysianassidz).

G. Telson entire. . . Lysianopsis.

First Bahenol ae Infero-lateral

angle of the third abdominal segment

rounded. Color white.

L. alba sp. nov., M.

GG. Telson more or less deeply cleft.

f/f. Infero-lateral angle of the third abdom-

inal segment not produced.

No. 436.] NORTH AMERICAN INVERTEBRATES. 277

Z. Anterior coxal plates much less than twice as deep as their segments.

Euryporeia.

Eyes expanding below into two diverging lobes. Telson long, deeply

cleft, conically tapering. Attains 62 mm. . Æ. gryllus (Mandt), A

ZI. Anterior coxal plates twice as deep as their segments.

J. Anterior gnathopods slender; carpus elongated ; propodus narrow.

i ryphosa.

Eyes narrowly reniform. Hand of first mern tapering

distally, palm transverse. L. 7 mm. Abundan

T. pinguis ecd, A. M.

J/. Anterior gnathopods rather stout. Carpus not elongated.

Tryphosella.

Eyes very large, somewhat widened below. Fourth abdominal

segment with a deep dorsal depression behind which is an

angular prominence. Infero-lateral angle of third abdominal

segment nearly a right angle. Propodus of first gnathopods

shorter than the carpus. . T. horingit (Boeck), A.

H. Inferoiéietal angle of the third

abdominal segment produced into

an acute projection.

Z. Telson short, not cleft to the middle.

7. Anterior gnathopods not subchelate. . . . Menigrates.

Posterior perzopods very short, the bed Joints very large,

ovate, merus much expanded. Rami of terminal uropods naked.

M. DEES Boeck, A.

J/. Anterior gnathopods subchelate. - . Onesimus.

Lateral lobes of head obtuse. ‘Hand of anterior gnathopods

much longer than the carpus, slightly curved ; palm nearly trans-

erse and finely denticulated. . . . O.edwardsii (Kr.). A.

J7. Telson Eus to beyond the middle. i

J. Infero-lateral angle of the third abdominal segment gums into

a small tooth. . . Hoplonyx.

Hand of first t puto distally penes pus transverse.

Lateral lobes of head only slightly projecting and rounded.

Upper part of eyes narrow. Attains 18 mm.

H. cicada (Fabr.), A. M.

JJ. Infero-lateral angle of the third abdominal segment produced into

a large upturned tooth above which is a jns marginal sinus.

K. Eyes imperfect or wanting. . . . Centromedon.

Lateral lobes of the head dd out into a narrow sharp

process. Last pair of peræopods shorter than the pre-

inii. pair, the basal joint subquadrate and longer than

Antennz of subequal length. L.

the succeeding ones.

C. pumilus (Lillg), A

fie 1014

KK. Eyes well dod.

278 THE AMERICAN NATURALIST. [Vor. XXXVII.

L. Basal joints of the first antennz distally produced above ;

secondary flagellum small. ippomedon.

Tooth of infero-lateral inae i the third abdominal

segment broad, with a broadly rounded sinus above.

Basal joint of the — perzeopods Mug serrate

behind. .... . H. serratus sp. nov. M.

LL. Basal joints of ih first antennæ not distally ee

above ; secondary flagellum well develope

Anonyx.

Eyes large, elongated and expanded below. Hand of

first gnathopods scarcely tapering distally, the palm

transverse. Telson nearly rectangular, cleft nearly to

base. A. nugax (Phipps), A. M.

FF. "Without al the TERMI of F.

Terminal uropods uniramous or wanting.

amus of terminal uropods two-

Hen: . « + + (Stenothoidz).

Mandibles with a palp. . etopa.

Antennz of subequal bp poem slates very 3 Bm. the fourth

pair longer than deep. Hand of second gnathopods in the male

with a large sinus near the posterior end of the palm, the cor-

responding sinus in the hand of the female much smaller.

M. grenlandica Hansen, A. M.

ZI. Mandibles without a palp. . . . Stenothee.

J. Length exceeding 5 mm. Cpu of ist Puaibopoda nearly twice

as long as broad, the margins parallel. . .S. peltata Smith, A.

J/. Less than 5 mm in length.

Fourth pair of coxal plates — Dass and more or

less ovate in form ‘ . cypris sp. nov., M.

Fourth pair of cxi lids not ihesüsibe jm for the genus, not

GU. . a . S. minuta sp. nov., M.

HH. Ramus of terminal uropods one-

ed.

joint

Body slender, elongated. Ramus of terminal uropods much longer

e peduncle. . Neohela.

Eyes well developed. Both anina rer thas the body. Per-

æopods very long and slender. . N. phasma Smith, A.

4I. Body depressed, moderately stout. -Goia plates s small. Antenne

shorter than the body. . . Unciola.

Peduncle of terminal uropoda SARPUB distally into a lobe which

extends about to the tip of the short ramus.

U. irrorata Say, A. M.

HHH, Terminal uropods wanting.

Body slender ; coxal plates very small.

Dulichia.

No. 436] NORTH AMERICAN INVERTEBRATES. 279

First antenne a litde longer than the

second and about as long as the body,

secondary flagellum minute,three-jointed.

Hand of second gnathopod of the male

with a long thumb-like process above

the middle and a spine at the end of

the palm.. . . WD. porrecta, Bate, A.

GG. Terminal uropods biramous.

H. Anterior gnathopods with the carpus

and propodus forming a chela.

Leucotho.

Antenne of subequal length.

Carpus of the large second gnatho-

pods extending behind the hand to

the upper end of the palm. Telson

narrow and conically taperi

L. spinicarpa (Abildg.) A

HH. Notas in H.

Z. Carpus of the e ries in front of the proximal end of the

propodus. Eusirus.

Last thoracic aid first i two E aileni: iones sedes posteriorly

into a dorsal spine. Telson cleft about to the middle.

E. cuspidatus Kr. A.

ZI. Carpus joined in the usual manner.

J. Peraopods devoid of dactyls and peculiarly modified for digging.

Haustorius.

Three posterior pereopods dissimilar with the joints above the

propodus much expanded. Carpus of the first two perzeopods

with a large rounded posterior lobe; propodus widened distally

and rounded. Telson much wider than long and bilobed.

H. arenarius (Slabber) A. M.

JJ. Perzopods with dactyls.

K. First antenna with an accessory flagellum.

L. "Terminal uropods flattened and projecting beyond the

others. Gnathopods subchelate, larger in the male

than in the female, the second pair usually larger than

the first. Mouth parts normal Telson small

flattened, cleft or emarginate. . . (Gammaride).

M. Inner ramus of terminal odi scale-like, rudi-

mentary. First antennæ longer than the second.

Melita.

Carpus of first gnathopods as broad as long;

hand short and stout.

, parvimana sp. nov., M.

Carpus of first gnathopods much longer than

THE AMERICAN NATURALIST. [Vor. XXXVII.

broad; hand narrow. Posterior margins of the

abdominal segments produced into teeth. Ter-

minal uropods much elongated.

M. dentata (Kr.), A. M.

Carpus of first gnathopods much longer than

broad; hand narrower and shorter than the car-

pus.

Posterior margins of the abdominal seg-

ments not produced into teeth.

M. nitida Smith, M.

MM. Inner ramus of terminal uropods not rudiment-

ary, ee often considerably smaller than

the o

N. tss. ge slightly emarginate. Thorax and

NN.

abdomen dorsally carinated. . Gammarellus.

Dorsal carina high and produced posteriorly

on the segments along the middle of the body

into compressed spinous processes. Tip of

telson narrow, the cleft pes sma

i (F abr.), A

Dorsal carina not produced arias at the

ends of the segments. Antenne stout, sub-

equal. Telson broad at the tip which has a

rounded - ge

G. angulosa ADAE A. M.

Telson deeply cleft.

O. Last three segments of the abdomen with

fascicles of spines

bdomen dorsally carinated.

Carinogammarus.

First three abdominal segments

produced behind into acute teeth.

C. mucronatus (Say), M

PP. Abdomen not dorsally carinated, the

first three segments not produced

behind into teeth. . Gammarus.

Q. Inner ramus of terminal uropods

much less than half the length

of the outer. Eyes long and

arro Infero-lateral angle of

ile third abdominal segment

nearly right angled.

G. marinus Leach, A. M.

QQ. Inner ramus of terminal uropods

over half as long as the outer.

Fourth abdominal segment

No. 436.] NORTH AMERICAN INVERTEBRATES. 281

with median and lateral fasci-

cles of spines. Terminal

clusters of strong spines, the

inner ramus reaching nearly

gle of the third abdominal

segment pr (| VM

common.

terminal ods narrowly

abundantly at the surface.

G. annulatus Smith, M.

(— G. natator Smith).

OO. Last three segments of the abdomen

PF.

without fascicles of spines, although

there may be spiniform projections

from the posterior margins o e

segments.

Posterior margins of the abdomi-

nal segments armed with teeth

or spines. Gnathopods feeble.

First three abdominal segments

produced posteriorly into three

strong spines with smaller denticles

between and below them. Legs

very long and slender, the posterior

pairs with the basal joints not

expanded. Telson cleft about to

the middle.

M. spinosa (Goes), A

Body smooth above

Q. Terminal irdpads short and stout.

Elasmopus.

THE AMERICAN NATURALIST. [Vor. XXXVII.

Second antenna about as long

pods with the basal joints ex-

panded, the merus and carpus

expanded, especially in the

male. Terminal uropods pro-

jecting but little beyond the

others, the rami short and broad.

E. levis (Smith), M

QQ. Terminal uropods elongated.

Mera.

Coxal plates small. Basal

joints of the posterior three

pereopods long and narrow.

Rami of terminal uropods

narrow, elongated, and sub-

equal.

M. dane (St) A

ZL. Not with all the characters of L.

AM. Head with a prominent deflexed rostrum separated

from the sides by a deep lateral sinus. Fou

coxal plates usually smaller than the preceding

ones. Gnathopods feeble, the carpus elongated.

Telson much elongated . . . . (Syrrhoide).

N. Eyes large and coalescent above. . Syrrhoé.

Rostrum perpendicularly deflexed.. Poste-

posterior three pairs of perzopods deeply

serrated behind. Telson cleft nearly to the

ase. . S. crenulata Goes, A.

NN. Eyes round i scarcely coalescent above.

iron.

A small accessory eye below each large

eye. Last three abdominal segments pro-

duced posteriorly into a large median spine

but not serrated on the posterior margin.

T. acanthurus Lillg., A.

MM. Not with all the characters of M.

N. Body spiny; secondary flagellum minute.

achotropis.

Last two thoracic and first four abdomina

segments with a posterior median dorsal

spine, the first four abdominal segments hav-

No.436] NORTH AMERICAN INVERTEBRATES. 283

NN.

ing a smaller additional spine near the mid-

dle. A lateral marginal spine on the last

two thoracic and first three abdominal seg-

ments. Rostrum long. Hands of a similar

oval form. Telson cleft in its posterior

third. . . . . A. aculeata (Lepechin), A

Not as in N

O. Terminal uropods enormously enlarged,

the inner ramus rudimentary. Second

uropods with the peduncle expanded into

a lobe much larger than the quadrate,

distally serrated rami. Body depressed.

Chelura terebrans Phil. M.

OO. Notas in O.

P. Coxal plates enormously developed ;

tumid. i

body mandibular

palp. . . . . Stegocephalus.

Margins of e first five

coxal plates presenting an evenly

rounded contour. Rostrum

prominent, deflexed, with a deep

sinus below. Infero-posterior

angle of the basal joint of the

last perzopods produced and

cute. Telson cleft beyond the

middle, — ae in distal

half. S.

ws Kr, A. M.

PP. Coxal gins a ade size.

Mandibles with a palp. Last

joints of preceding pairs narrow.

ontoporeia.

Fourth abdominal segment with a

peculiar, upturned, bifurcated

process. P. femorata Kr., A.

PPP. Coxal plates rather small. Mandi-

bles with a palp. Last three

pairs of peræopods of similar

form and increasing in length

posteriorly. First few segments

antennz fused together in the

284 THE AMERICAN NATURALIST. bogs XXXVII.

male. . . Pardalisca.

Third E et aidoniial seg-

ments with a pair of posterior

dorsal projections ; fifth segment

with a single median spine-like

process. Eyes very large.

P. abyssi Boeck, A.

KK. First antennz with no accessory flagellum.

axillipeds with the palp small and two-jointed. Para-

ax mid.

Fourth coxal plates pointed below. Perzopods of

subequal length and furnished with strong curved

dactyls. Found on several species of fish.

L. sturionis Kx., A. M.

LL. .Not as in

M. Telson dei

N. First gnathopods rudimentary. . . Batea

Anterior coxal plates large. Telson cleft

nearly to the middle. First gnathopods

represented only by the coxal plate and basal

joint A minute species found among

hydroids. B. secunda sp. nov. M.

NN. First gnathopods not rudimentary.

. First three pairs of coxal plates pointed

elow. Body with a median dors

crest. Head with a very prominent

um.

P. Body with prominent tubercles on

either side _of the dorsal crest.

pimeria.

Dorsal crest extending from the

first segment of the thorax to the

fourth segment of the abdomen.

E. loricata Sars, A. M.

PP. Body devoid of tubercles. Anterior

gnathopods very slender and sub-

chelate. Telson not deeply cleft.

segments produced posteriorly into

spines. Postero-inferior angle of

third abdominal segment divided

into two prominent lobes the

No. 436.]

NORTH AMERICAN INVERTEBRATES. 285

margin of the lower of which and

sometimes also the upper is ser-

rated. . A. serratum(Fabr.),A

Dorsal crest not produced posteri-

orly into spines.

A. inflatum(Kr.), A

OO. First three pairs of coxal plates not

PP.

pointed below

No iendithar palp. Gnathopods

similar. Two last segments of the

abdomen completely fused together.

Dexamine.

First four aidondasd segments with

a prominent pointed dorsal projec-

tion at the posterior end.

D. thea Boeck, A. M.

Mandibles with a palp. Body

devoid of dorsal spinous processes.

Gnathopods slender; hands nar-

row. Pon dicm

Telon cleft t to beyond the mid

Terminal uropods with datae

lanceolate rami of subequal length.

Eyes large. Both antennz elon-

gated and furnished with calceola

in the male.

P. inermis (Kr), A. M.

MM. Telson not cleft.

jV. Body dorsally carinated.

O. Abdomen with tubercles or spines on

either side of the dorsal carina.

P. Body thickly armed with very long and

PP.

with long spines.

narrow spines . Acanthozone.

Thorax with five rows of long spines.

First thoracic segment with a large

horizontal spine projecting over the

head. Posterior margins of abdom-

inal segments armed with several

spines. Telson bue: to a nar-

row, veut

papse Pt (Lepechin), A

Body tuberculated but not armed

Pleustes.

Dorsal carina adig fons first thoracic

to fifth abdominal segment. Rostrum

THE AMERICAN NATURALIST. [Vor. XXXVII.

large. Hands oval, almost alike in form.

Sides of the abdomen tuberculated.

P. panoplus (Kr.), A.

OO. Body devoid of tubercles or spines except

in the mid-dorsal line and at the infero-

lateral angles of the abdominal seg-

ments. Antenne long. :

Paramphithe.

Anterior abdominal segments and a

variable number of the posterior thor-

acic segments produced into a large

compressed spinous projection in the

mid-dorsal line.

P. pulchella (Kr.), A.

Only the first two abdominal segments

produced posteriorly into a spin-

ous process. Telson oblong, distally

rounded. . P. dicuspis (Kr); A. M.

NN. Body without a prominent dorsal carina.

O. Antennz with calceole. Last peduncular

joint of the first antennz with a terminal

lobe.

P. Dorsal spines on some of the segments

ofthe body. . . . . Halirages.

Eyes large, oval. Last thoracic

and first two abdominal segments

with a dorsal posteriorly directed

spine. AH. fulvocinctus (Sars), A.

PP. No dorsal spines. . . Calliopius.

First and second gnathopods

nearly equal; hands ovate. Eyes

large. Telson oblong, distally

C. leviusculus (Kr.), A. M.

OO. Antenne without calceola. No terminal

lobe of the last peduncular joint of the

first antenne.

P. First antenne longer than the second.

Sympleustes.

First three abdominal segments

with a posterior dorsal prominence.

Second gnathopods stout, hand

distally widened, armed with several

short stout spines around the pos-

terior end of the oblique, somewhat

No. 436.]

NORTH AMERICAN INVERTEBRATES. 287

excavated palm. Perzopods short,

stout and subequal. Telson oblong,

distally rounded.

S. latipes (Sars), A

Abdominal segments not elevated

posteriorly. First three coxal

tooth. Second gnathopods much

less stout than in ZzZ pes, the hand

narrower. Peropods slender. |

S. glaber (Boeck), A

PP. First antenne shorter than the

second. Gnathopods small.

Coxal plates not large.

Apherusa.

Infero-lateral margin of the third

abdomin segment serrated.

Hand of second gnathopods oval.

A. gracilis sp. nov., M.

Infero-lateral margin of third

abdominal segment serrated.

Eyes very large. Second gnatho-

pods oblong, the palm oblique, the

osterior margin nearly straight.

Infero-lateral angle of the hea

produced into a prominent spine.

A. megalops Sars, A.

CC. First two pairs of perzopods with spinning glands.

D. "Terminal uropods uniramous.

EE.

Mandibular palp one-jointed. . . Siphonacetes.

Head with a long acute rostrum. Eye situated

on the lateral lobe of the head which is contracted

at the base. Second antennz longer than the

body. Length 6 mm

S. cuspidatus Smith, M.

Mandibular palp two-jointed. . . Corophium.

Second antenne enormously developed in the

male, the penultimate joint armed with an up-

turned tooth at the antero-inferior angle. Second

gnathopods simple, the merus joined along the

entire posterior margin of the carpus and

fringed with very long hairs. Last peraeopods

288 THE AMERICAN NATURALIST. | [Vor. XXXVII.

very much longer than the preceding pairs.

Length 4mm. . . C.cyZndricum G (Say), M

EEE. Mandibular palp three-jointed. Second gnatho-

pods of the male greatly enlarged, the carpus

produced below the propodus with which it

forms a sort of chela. Body depressed ; coxal

plates small.

F. Last two pairs of uropods uniramous.

Cerapus.

Ramus of last two uropods de shorter

than the peduncle. Telson short, bilobed.

Lives in tubes which are carried about by the

C. tubularis Say, M.

ima

FF. Only the cml uropods uniramous.

DD. Terminal

Erichthonius.

Carpus of second gnathopods in the male

nearly three times as long as broad across

the middle, the lower process armed with a

tooth near the middle; propodus about half

as long as the carpus; dactyl much shorter

than the propodus. Length about 4 mm.

E. minax (Smith), M

Carpus of second gnathopods in the male

much less than three times as long as broad

across the middle. Propodus much over

half the length of the carpus. Length 7 mm.

E. rubricornis (St), A. M.

uropods biramous.

E. Propodus of second gnathopods not subchelate.

tilocheirus.

Hand of first gnathopods widest across the

distal end, the palm nearly transverse.

Second gnathopods slender, the anterior

margin fringed with very long plumose setze.

15mm. Common. . P. pinguis St, A. M.

EE. Propodus of second gnathopods chelate or sub-

chelate.

F. Terminal uropods with short, hooked rami.

First antenne with an accessory flagellum.

and of the second gnathopods of

the male very large and having a

thumbdike process arising from

near the base of the lower side.

Coxal plates small. . Jassa

Hand of second piahia of the

No. 436.) WORTH AMERICAN INVERTEBRATES. 289

male oblong, longer than the rest of

the appendage; thumb long; a

tooth at lower end of the palm.

First two peraopods with the

merus much widened and produced

at the antero-distal angle into a

large rounded lobe. 1;

J. marmorata sp. nov., A. M.

HH. Hand of the second gnathopods of

male not as in Jassa.

Z. Antenne rather stout, i setiferous a flagella with few

segments. . Ischyrocerus.

Second gathop in the Tem with a T slender, basal joint;

hand long, narrow, curved, of nearly the same width throughout, the

concave posterior margin thickly setose and terminating inferiorly in

a stout tooth. Attains 10 mm I. anguipes Kr., A. M.

Antennæ slender, with mliia dapella. rubia.

Coxal plates large. First antenna as long as ; the body: the slender

flagellum nearly three times as long as the peduncle; secondary

flagellum small, not longer than the first two joints of the primary

one. Gnathopods in the male inp and fringed with long

plumose hairs. Length 13 m G. compta (Smith), M

GG. Firs irst antennæ devoid of a secondary

flagellum . Amphithæ.

Second antennæ e subpodiiam ; flagel-

lum often shorter than the last joint of

the peduncle. Anterior gnathopods

with the hand short and stout. Length

18 mm. . A.rubricata (Mont.), A.M.

(=A. manculata St. and A. valida Smith).

Second antenne with the flagellum

generally longer than the last joint of

the peduncle. First gnathopods with

the carpus and hand narrow and very

much elongated; palm very short,

transverse; dactyl when closed pro-

jecting far beyond the end of the palm.

Length 9 mm. Common. Color vari-

FF. Terminal Go with narrow rami devoid of

terminal hooks.

G. Second gnathopods much larger than the

rst. First antennæ with no accessory

flagellum. . Podoceropsis.

Antenna sibégusl. Fi irst gnathopods

290

THE AMERICAN NATURALIST. (VoL. XXXVII.

with the hand narrowly oval, a little

shorter and narrower than the carpus;

dactyl long, closing against nearly the

entire posterior margin of the hand.

Hand of the second gnathopods stout ;

palm excavated.

P. nitida (St.) A. M.

(= Xenoclea megacheir Smith).

GG. First gnathopods much larger than

the second. First antennze with

an accessory flagellum.

H. Second gnathopods in the male

very stout, the inferior lobe

armed with three or four stout

teeth; propodus about half as

ong as the carpus, the lower

margin furnished with two

blunt teeth.

M. gryllotalpa Costa. A. M.

inferior lobe narrowly triangu-

lar, acute, with a small tooth

near the base of the outer mar-

gin; propodus with a low

prominence along the distal

portion of the lower margin.

M. danmonensis (Bate), A. M.

HH. Second gnathopods of the males

simply subchelate. Autonoe.

than broad, shorter than

carpus. Last pair of peraeo-

pods much longer than the

preceding ones. Length 7

mm. A. smithi sp. nov., M.

No.436] WORTH AMERICAN INVERTEBRATES. 291

Tribe Caprellidea.

Head fused with the first segment of the thorax. Abdomen rudimentary.

Second gnathopods larger than the first. Usually gills are present only on

the third and fourth segments of the thorax. Anterior pairs of peræopods

usually wanting. No pleopods. Uropods rudimentary or wanting.

The Caprellidea comprise two families, the Caprellidae and the Cyamidz,

or whale lice.

Family Caprellide.

Body elongated, cylindrical. First thoracic segment separated from the

head by a dorsal depression. Antenne elongated, the first generally

much longer than the second. Posterior perzeopods prehensile.

KEY TO THE SPECIES.

A. Mandibles with a palp. ZEginella.

Second gnathopods at the idle MONEY the haril pradas into a

tooth in front of the dactyl; a tooth at the upper end of the palm; a

narrow tooth below the middle of the palm which is separated from

a broad prominence below by a narrow sinus. Second antennæ

scarcely half the length of the first pair. A. longicornis (Kr.), A. M.

The typical form of this species has the body smooth. In the

variety spinossissima (Aegina spinossissima Stimpson) the body is

thickly armed with slender spines. Intermediate forms occur which

connect the two varieties.

AA. Mandibles devoid of apalp. . . . . : + - + + « Caprella.

B. Body smooth or nearly so.

C. Body with short and thick €—À Head with a horizon-

' tally directed spine. . . . C.geometrica Say, M

CC. Segments of the body more or di elongated. Head devoid

of a horizontally directed spine.

D. Second gnathopods of the male with the basal joint

short (very much shorter than the hand).

Second gnathopods of the male joined nearly at the

posterior end of the segment. A ventral spine

between the bases of the second gnathopods. Two

first segments of the thorax much elongated in the

adult male. . C. equilibra Say, A. M.

Second guátbepéds at the male joined not so far back

although sometimes situated behind the middle of the

segment. No ventral spine between the gnathopods.

C. septentrionalis Kr., A

292 THE AMERICAN NATURALIST. | [Vor. XXXVII.

DD. Second gnathopods of the male with the basal joint

elongated and narrow. Last thoracic segments

sometimes with small dorsal elevations. Body

slender. à C. linearis (Linn.), A.

BB. Body tuberculated or spiny.

Body stout, armed above with large pointed tubercles which

in some specimens may become obsolescent. Antenna short

and stout. Second gnathopods of the male with short and

broad basal joint; hand ovate or suboval, armed with a large

tooth at upper end of the palm, a narrow tooth below the mid-

dle separated by a narrow sinus from a prominence below.

C. stimpsoni Bate (C. robusta St.), A.

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ANGLAS, J. Les phenoménes des métamorphoses internes. Paris, C. Naud, 1902.

Svo, 84 pp., 16 figs. “Scientia” Serie Biologique No. 17, 2 francs. — ENDERLEIN,

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text figs. 41-75. Cambridge, University Press, 1902.— HACKER, V. Ueber d

Schicksal der elterlichen und grosselterlichen Koristele. cic gc pak

träge zum Ausbau der Vorerbungslehre. Jena, Fischer, 1902. 8vo, 104 pp., 4 pls.,

16 text figs. 4 Marks.— Herrwic, R. A Manual of Zoologie. Translated from

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1902. 8vo, xi-704 pp., 672 figs. $3.00.— HoorPER, W. L. and WELLS, R. T.

Electrical Prokio for Engineering Students. “Boston, Ginn & Co. 8vo,

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Naud, 1902. vins 159 pp.. 9 e “Scientia,” Serie Biologique. Nos. 15-16. 4

Francs.— MENAGAUX, A.— La Vie des Animaux illustrée. I. TE Lémuriens.

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ORT, P. Traité de sylviculture. Principales essences forestieres. Paris, Félix

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Boston, Ginn & Co, 1902. 8vo, x-291 pp., 98 figs. $0.75.— Witt, W.H.

The A. B. C. of Photo-Micrography. A Practical fuas k for Beginners.

New York, Tennant & Ward, 1902. Svo, viii-155 pp. EBSTER, R.

W. Absorption of Liquids by Animal Tissues. Univ. EAR Decenuial Publ.,

Vol x, pp. 105-134.— WETTSTEIN, R. von. Der Neo-Lamarckismus und

seine Beziehungen zum Darwinismus. Jena, Fischer, 1902. Bie 30 pp. ark.

CLARKE, F. W. A New Law in Thermochemistry. Proc. Wash. Acad. Sci.

Le V, pp. 1-37.— Coss, J. N. epe Fisheries of the OFE spes

. U. S. Fish Com. for 190r. Pp. 383-499, pls. 21-27.— RB. The

nobis California expen Mai Myths. Audi. Os e Nat. "

Vol. xvii, Pt. II, pp. 33-118.— GRABAU, A. W. The Traverse Group of Michigan,

Am. Rept. Geol. Surv. Mich. pom zgor. Pp. 161-211.— GRAVES, S. The

Woodsman's Handbook. O. S. Dept. Agr., Bureau of Forestry Bull. No. 36.

148 pp., rs figs. Map.— Hinps, W. E. Contribution to a Monograph of the

Insects of the Order Thysanoptera inhabiting North America. Proc. U. S. Na,

Mus. Vol. xxvi, pp. 79-242, pls. 1-11.—Jorpan, D. S. and EvERMAN, B. W.

The Fishes and Fisheries of the Hawaian Islands. Rept. U. S. Fsh Com. for

zgor. Pp. 353-380.— JoRDAN, D. S. and FowLeR, H. W. A Review of the

Berycoid Fishes of Japan. Proc. U. S. Nat. Mus. Vol. xxvi, pp. I-21, 4 figs.

KNIGHT, W. C. The Birds of Wyoniig Buil. o. Agr. Exp. Sta., No. 55.

174 pp., 42 pls. and text-figs.— KRAEMER, H. On ie SAA of Protoplasm.

293

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v Am. Phil. Soc Volk xli, pp. 174-180, pls. 21-22.— KRAEMER, H. The

a of the Starch Grain. Botonical Gazette, Vol. xxxiv, pp, 341-354, pl. 11.

— Macoun, J. Catalogue e Canadian Plants. Pl. VII. Lichenes and Hepaticæ.

Geol. ea Canada. v—318-xix pp.— MAYER, A. G. Effect of Natural Selection

and Race Tendency upon ims Il Patterns of Lepidoptera. Se. Bull. Mus.

Brooklyn Inst. Arts and Sci. l. I, No. 2, pp. 31-86, 2 pls.— McCLATCHIE, A.

J. Eucalypts cultivated in Me lisi States. U. S. Dept. Agr., Bureau

Forestry, Bull. No. 55. 106 pp., 9t — NECOMBE, F. C. Sach’s angebliche

thigmotropische Kurven and adiens waren traumatisch. Sot. Centrallblatt.

Bd. xii, pp. 243-247.— NECOMBE, F. C. The me Zone of Roots. Ann.

Botany. Vol. xvi, pp. 429-447.— OGILVIE, S. H. “Glacial Phenomena in the

Adirondacks and Champlain Valley. Jour. Geol. Vol. x, pp. 397-412, 1 pl.—

OGILVIE, S. H. An Analcite-Bearing Camptonite from New Mexico. Jour.

Geol. Vol. x, No. 5, pp. 500-507, 4 figs.— PREBLE, E. A. A Biological Investi-

gation of the Hudson Bay Region. U. S. Deft. Agr. N. A. Fauna, No. 22, 140

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Pp. 289-351, pls. 6-20.— REID, J. A. The Igneus Rocks near Pajaro. Univ.

Cal. Publ., Bull. Dept. Geol. Vol. iii, pp. 173-190, pl. 18.— RITTER, W. E. The

Movements of the Enteropneusta and the Mechanism x- which they are accom-

plished. Biol. Bull. Vol. iii, pp. 255-261.— SHIMER, H. W. and GRABAU, A.

The Hamilton Group of Thedford, Ontario. Pa Geol. Soc. Amer. Vol.

xiii, pp. 149-186.— TowNSEND, C. H. Statistics of the Fisheries of the Great

La sh Com. for rgoo. Pp. 575-657.— TOWNSEND, C. H.

Statistics of the Fisheries of Fé sabe in River gs Tributaries. eft. .

Fish Com. for rgoo. Pp. 659-740.— WaLcorr, C. Cambrian telis

Acrotreta ; Linnarssonella ; bina With Description ie New Species. Proc.

LU. s. ol. xxv, pp. 577-612.— WILcox, W. A. Notes on the Fish-

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Argulide with a i ie of the Group and a Systematic Review of all

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text figs.

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de la Sociedad Aragonesa de Ciencias Naturaies. Tom. I, No. 8, Oct.— Cage of

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Jahrg. 19, No. 38.— New aa The. Vol. vi, Nos. 2-6.— Revu y deri

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VoL. XXXVII, No. 437

'The American Naturalist.

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VOLS XXXVII. May, 1903. No. 437.

ALBINISM, PARTIAL ALBINISM AND POLY-

.CHROMISM IN HAG-FISHES.

BASHFORD DEAN.

Ir hag-fishes are the most primitive of living ‘ fishes,’ it is an

interesting circumstance that they have not been known to have

differentiated a wide range in coloration. In other groups

mottlings and bright colors are the rule rather than the excep-

tion, but in hag-fishes blotches of color as well as colorless

conditions have not hitherto been described. If, however, we

consider that these fishes are, as a group, from relatively deep

water, it is not to be wondered at that they have run the

gamut of coloration common to deep sea forms— thus, in a

range of species they pass from black (Myxine circifrons Gar-

man), into dark purples, thence to violets and lavenders, then

into “meaningless " grays, sometimes uniformly colored, some-

times shaded. In the latter event the dorsal region is the dark-

est, the ventral the lightest. In some instances lack of pigmen-

tation in definite regions becomes a rather conspicuous feature :

thus the tips of the barbels are often, indeed generally, white,

and the mid-ventral as well as the mid-dorsal line in several spe-

cies is unpigmented. In one case at least (Homea stoutt) the

295

296 THE AMERICAN NATURAL/ST. (VOL. XXXVII.

lack of pigment in the mid-dorsal line becomes a prominent

character of the newly hatched young (larval coloration), Aside

from this we have had no detailed knowledge of the coloration

of hag-fishes, and we could not, therefore, answer the question

whether albinos, common in many and widely separate lines,

occur also in this phylum of lowly chordates and whether mottled

colors had already been evolved. In the event of mottled colors

occurring in a single species one might justly infer that the

blacks and violets and grays of this group are in reality but

symptoms of a deep-water, or possibly of a nocturnal habit.

In regard to the first of these questions we may now, however,

state definitely that true albinos occur among hag-fishes, and that

partial albinos are not rare. A perfect albino of 77. burgeri, Fig.

1, was collected at Misaki, Japan, but it was the only one observed

in upward of 800 examples. A specimen of H. stouti in part

albino, white from snout to gill region, somewhat mottled where

the white passed into the purple body color, Fig. 2, had formerly

been observed by me at Monterey, Cal. (1899), together with

several less perfect cases of the pigmentless condition,— these

out of many hundred specimens collected. So one can justly

conclude that in the myxinoid line albinism already plays its

usual róle among chordates.

The matter of motley coloring in hag-fishes is also elucidated

rather strikingly in the case of a specimen of H. polytrema

Girard, in excellent condition, which the writer recently received

from the neighborhood of Valparaiso, Chile? Fig. 3. The col-

oration of this species as far as one can determine from a single

specimen, is brilliantly mottled with black, ashen, umber and

ochre, to a degree which at once causes one to wonder whether

this form has not been actually mimicking a murray. Indeed we

learn from Dr. Delfin's interesting paper that the name of the

fisherman for this hag-fish is the “bearded murzena," indicating

! My friend, Mr. Naohidé Yatsu, to whom I am indebted for the figures

accompanying the present note, examined this specimen while it was still living

and tells me that its color was white, but tinted slightly with yellow, instead of

with pink as one would naturally suppos

? For this specimen he is greatly Shad to the kindness of the Chilean

ichthyologist, Dr. Federico Delfin, whose observations on the habits of this

myxinoid are in many regards the most complete hitherto recorded.

‘IYA *osreied[e A. vutco4270g v2uto ‘£ ^51 ‘rey *&219100]N. 774075 vao '£C»I 'uede( yesi ‘74aydang vamo Y 101

IJ L [PA 7 H A [kD W : H A IHESI 7420. ? j; A

“SHHSIA-OVH DINOUHOATOd ANV ONIJIYV

SS Ees

298 THE AMERICAN NAIURALIST. [Vor. XXXVII.

that the coloration of the present specimen is not abnormal.*

In this regard, we can, however, derive no light from Girard's

description published (1854) in the Chilean expedition report,

for he states that his material of Bdellostoma was too poorly

preserved to warrant any reference to coloration.

~ In the matter, accordingly, of the general color of hag-fishes,

we can now, I think, reasonably conclude that if one species is

provided with definite colors the entire group can hardly be

different in -this regard from ` other piscine groups, sharks,

chimæroids, teleosts, in which deep. sea forms are characteris-

tically monochromic, and shallow water ones. motley. And if we

accept this conclusion, and it seems to me a sound one, we have

‘still another ground for the belief — by analogy .with other

groups — that myxinoids are represented at the present. day by

but a small number of forms, in contrast with their maximum

development i in species and genea in early times.

lina TE RA to Wolnitzky (Baldómero), *Coast Fishery of the Province

Aconcagua," Buffalo Exposition (1901). Handbook, B. polytrema is referred to as

the *black Congrio," although this writer. remarks that the name is a misnomer .

No note as to coloration is, however, given.

COLUMBIA UNIVERSITY.

VARIATION IN LITHOBIUS FORFICATUS.

STEPHEN R. WILLIAMS.

As long ago as 1865, Dr. H. C. Wood in his “ Myriapoda of

North America" called attention to the great variation in that

group and tabulated many variations. I have chosen Lithodius

Jerficatus, the most common of the Chilopoda in the eastern

part of our country, for a quantitative variation study.

Lithobius forficatus is a cosmopolitan species, found in Europe

as well as America. This paper will furnish a place-mode for

the species at Cold Spring Harbor, Long Island. Comparisons

with place-modes from distant locations will be instructive.

Since the variations found are in specific characters such as

prosternal teeth, coxal pores, antennal joints and spines we

might hope for suggestions as to whether selection in Lithobius

is tending in any definite direction? Are the polygons skew

in any special way? Is Lithobius forficatus a stable or unstable

species? Any satisfactory answer to such questions would help

our knowledge of the method of origin of species, and in so far

advance this, the chief aim of modern biology.

Cold Spring Harbor has a moist climate with abundance of

vegetation, a corresponding wealth of insect life and, correlated

with these conditions, an abundance of the carnivorous Lithobius.

The prevailing species is Z. forficatus, although in looking over

my material I find three specimens of Z. multidentatus which

must have been taken at Cold Spring Harbor. Myriapods of

other genera are also common, Scolopocryptops sexspinosus and

a Geophilus, probably G. mordax, represent the Chilopods and

Polydesmus, Polyxenus, and some of the Iulidze the Chilognaths.

The animals were collected during the summers of 1899 and

1900. The greater portion came about equally from a “trap”

of mowed yard grass on the upland 300 feet above tide level and

from a shaded moist region down within a few feet of highest

tide. Here some logs and planks served as traps. The Litho-

299

300 THE AMERICAN NATURALIST, [Vor. XXXVII.

bius breed in this latter location, since great numbers of very

small white individuals 3 to 8 mm. long are

40 to be found in the hiding places during the

| months of July and August.

^s | I took the length of 200 individuals,

rH 100 males and 100 females (Fig. 1). The

h length was measured from the head be-

‘ tween the bases of the antennz to the end

of the anal segment — not to the ends of

the posterior legs. The polygons were

also drawn for the 100 males alone (Fig. 2),

for the 100 females (Fig. 3), and

for 100 mixed specimens, 51

males and 49 females, which

Fic. 1. The length polygon

of 200 individuals.

12 al 5

had been selected entirely at ‘+44 H

random and so with no regard d SE

to sex (Fig. 4). On this last Fig. 2. Length poly- Fic. 3. Length

100 all the other determina- pe ee Mies uA

females

tions were made. The com-

plete data from this last lot are given in Table 9

Because Lithobius grows by moults, only

e those specimens 16 mm. long or more were

" measured in order to have a somewhat homo-

geneous group. They are probably all adults

sonst. ^ or nearly so. The longest male was 28 mm.

FiG.4. Length polygon of

100 specimens, 15: males long and the longest female was 25 mm. long.

rain pavo There are two apices for the length polygon,

the greatest number of individuals (37) falling in the 20 mm.

class and the next greatest (29) in the median class, 22 mm.

It would take but three individuals to level up the depression at

21 mm. so that the polygon might be considered unimodal. It

is skew to the right, due to the rejection of the smaller, younger

individuals which would have fallen to the left side of the poly-

gon. The same two classes are the modal classes in the mixed

polygon. (Fig. 4) and the polygon for the 100 females, (Fig.

3). The polygon for the 100 males (Fig. 2,) has but one

mode at 20 mm.

No. 437.| VARIATION IN LITHOBIUS FORFICATUS. 301

TABLE 1

Lengthinmm. 16 17 18 19 20 21 22 2 2 45. 20. 27 35

200 indiv. COEMTO O47. 439. 431. 34 29 49. I5. 11 3 LE d

100 males 5 8 7 9 12 hb HM 9 4 653 14 1

100 females T5 5 12, 14 20 AE 0 0 5 |

100 mixed I 5 5 ri iy B 6 lo 6 2 1 í

These data are represented graphically in Figs. 1 to 4.

TABLE 2

Length Polygons Type Mean. P. E. M. St. Dev. P. E. St. Dev. Coef. Var.

200 indiv. I 2043 +4.12 2.446 + .083 11.97

100 males 4 20.87 + .15 2.29 : 10.97

100 females I 19.96 + .15 2.245 + -107 ti

100 mixed I 21.41 --.15 2.35 4.11 10.99

For the 100 mixed individuals, 49 females and 51 males, counts

were made of :

I. The number of prosternal teeth.

2. The number of joints in the antenne.

3. The number of coxal glands, pits or pores which are

found on the coxz of the last four pairs of legs (the 12th, 13th,

I4th, and 15th). These data are given in their entirety in

Appendix A.

According to the key for the different species of Lithobius in

Bollman's “ Myriapods of North America" the number and

arrangement of the coxal pores, the spines on the legs, the

number of joints of the antennz and the prolongations of the

posterior angles of certain of the dorsal plates are the decisive

specific characters. I append an abbreviated key taken from

Bollman ('87) for the two species Z. forficatus and Z. multi-

dentatus. I intended to include in the data counts of the ocelli

also but it was impossible. As Bollman says “the ocelli are

distinct or not" and in many cases the fusion was nearly as

complete as in Scutigera where there is a close approach to the

facetted compound eye.

Posterior angles of the oth, 11th and 13th dorsal plates produced. Anal

feet with a single spine, the penultimate with two. Coxe unarmed. Coxal

pores in a single series. Antenne more than 3o jointed. Claw of the

302 THE AMERICAN NATURALIST. [Vor. XXXVII.

female genitalia Apres Coxal pores transverse, on 12th coxæ 6-9, on

13th 6-10, on 14th 6-9, on 15th (anal pair) 4-6 pores. Joints of antennz

33-43. Prosternal teeth 8-12 . . . . Lithobius forficatus.

(Coxal pores round in younger Nene:

Posterior angles of the 6th, 7th, oth, 11th, and 13th dorsal plates pro-

duced. Anal feet with a single spine. Coxe armed. Coxal pores multi-

seriate. Joints of antenna 19-23. Prosternal teeth 14-18. Coxal pores

arranged in 3 to 5 series. . . . Lithobius multidentatus -

(Specimens 12 mm. long ee Gaii pores in 2-3 series, those 10 mm.

long in 1-2 series and those 5 mm. long have round pores in a single series.)

PROsTERNAL TEETH.

TABLE 4.

Number of teeth 8.39. 30. 11 ; 12: - 13 14

Number of individuals 4 4 40 18 25 5 4

This distribution is represented graphically in Figure 5.

The tendency to bilateral symmetry here in

the number of teeth is very strong and so gives

a bimodal curve with apices at 10 and at 12.

But at the ends of the series the bilateral ten-

dency is overcome by the tendency to adhere to

the more typical numbers. The tendency to

variation, even though it be towards bilaterality,

is not so strong as the adherence to the more

usual number. There are more individuals

with 13 prosternal teeth than with 14 and as

many with 9 as with 8.

Figure 6 shows the ventral side of the head of a specimen of

L. multidentaius with fewer prosternal teeth than Z. forficatus

(Figure 7) has. The number of prosternal teeth is not a good

specific criterion as they overlap a great deal in the two species.

shown in Table 4

JOINTS oF ANTENNA.

TABLE ;5.

No, joints, 34. 35 36 y $ » 40 41 42 43 44 e 5 s 5 " "i

jg 8 $

3 2.4 B 13 I5

Left SD dak oet Mou. WE il

No. 437] VARIATION IN LITHOBIUS FORFICATUS. 303

There were 89 individuals possessing antennz (either one

or both) with as many as 34 joints. This minimum was chosen

arbitrarily because in Bollman's key Z. forficatus is said to have

FiG.6. Ventral view of the head of L. multidentatus. X 17.

Fic.7. Ventral view of the head of Z. forficatus. X 17-

304 THE AMERICAN NATURALIST. [Vor. XXXVII.

33-43 joints in the antenna. In the table of data (Table 9)

it is shown by underlining that two of the right antennae

counted ended abruptly (Numbers 8 and 50). On the left 33,

50, 57 and 86 had broken ends. The presence of a rounded tip

does not necessarily indicate perfectness but possibly merely

that regeneration took place at the last moult. Since the

antennz are so liable to injury not much stress can be laid on

the polygons (Fig. 8), derived from them. The mode lies

at 41 in the right with an average of 41.23. There are equal

numbers in classes 39 and

41 on the left side and the

average is 40.77.

The number of joints in

the antenna make a good

“quick” distinction be-

tween L. forficatus and L. multidentatus. The latter has fewer

joints (about as 21 to 43) but the individual joints are longer.

This can be seen on the left side of Figures 6 and 7.

34. 36.38.40.42.44. 46. bu. 34.36. 38.40. 42, 44. 46.

Fic. 8. Polygons of the antennz, Right and Left.

CoxaL FORES.

In the very young individual the hinder pairs of legs are not

yet budded out. The 13th, 14th, and 15th pairs of legs grow

in rapid succession, they may possibly all be indicated at the

same moult. This increase in the number of legs takes place

when the animal is less than 10 mm. long. The 12th pair of

legs is the first pair to bear the pits on the coxz so that these

are the oldest pits of the series ontogenetically. The youngest

specimens that showed coxal pores at all had two pores, one on

each 12th coxa. These were round. They must of course

increase in number at the times of moulting until the adult con-

dition is reached. For the condition of the coxal pores in the

three hinder pairs of legs in both species under consideration,

see Figs. 9 and 10:

1Specimens of Lithobius of about this length are often violet in color while

those shorter are always white.

? Because of the magnification necessary to bring out the pores and the conse-

quent reduced size of field, the 12th pair of coxe could not be included.

No. 437-.| VARIATION IN LITHOBIUS FORFICATUS. 305

What is the function of these pits on the four posterior pairs

of coxa? Coxal pores

are found on many ar-

thropods and are consi-

dered to be homologous

either with the setiger-

ous glands or with the

nephridial openings of

Chaetopods. In Peripa-

tus, according to Sedg-

wick ('95) page 19, a

series of pairs of glands

lie in lateral compart-

ments of the body cavity

with ducts opening on

the lower surface of the

legs. Peripatus has no

malpighian tubules but

has nephridia like those

of the Annulata, which

also open at the base of

FIG. 10.

multidentatus. X 17

1G. ọ. Ventral view of the last three segments of Z.

forficatus showing coxal pores and female genitalia.

X 17.

thelegs. The slime glands

at the base of the oral pa-

pille may be coxal glands

modifed for defence.

Closed coxal glands occur

in adult scorpions, scorpion

spiders and many spiders.

They are found at the base

of one or more pairs of

legs. In recently hatched

individuals the duct can be

traced to the exterior.

These animals all possess

one or more pairs of mal-

pighian tubules. The

«brick-red ” gland found in

Ventral view of the last three segments of Z. Limulus, whose duct.in the

306 THE AMERICAN NATURALIST. [Vor. XXXVII.

adult was demonstrated by R. W. Tower ('95, page 471), may

correspond to these glands and if it does they are certainly renal

in function.

Lithobius possesses malpighian tubules and sections have not

demonstrated any connection or passage-way from these coxal

glands to the body-cavity. The cup-like depression is lined with

columnar gland cells which appear to be in condition to secrete

actively. The glands may be secondary characters connected

with reproduction for they increase rapidly in number up to adult

life and then remain in a condition of comparative equilibrium.

There is also the possibility that they secrete a recognition

substance.

The number of pores is certainly very Vitale as only r9 out

of the roo examined had them arranged in a bilaterally sym-

metrical fashion.

TABLE 6

Number of pores 4 5 6 " 8 9 IO

12th leg left = I 9 As 34 XdL

I2th leg right - 1 i 120 —39 8$ I

13th leg left -= - 2 2 do —O^ 2

I3th leg right - 4 209/542 3186 2

14th leg left - 1 2 PROMOS Ti I

14th leg right E I 9 2d 4I . I2 —

15th leg left 2 Spo 44 1h 3 2 -

15th leg right 2 AT o4 13 Mo o 4

"a LE LI Ll.

ú | L. hi Hi

wid mI | til |i |

TE BIBBERI PCC IEEE i

22 Eid —|

" I TENNE

24 LETT E: E

20 LL z

» it} | 5

(s In it iul]

" | it} | Im ] m im!

AIL | Hl I'm | LL Im |

| EFLALLILA Í i

6. 7. 9. 5. 7. 9. 6. 8.10. 6. B. 10. 5. 7. 9. 6. 7. 9. 4. 6. .8 . 6. .8

iA deg act n leg 1 e leg hi xd bet d id 1 94 iting

Fic. 11. Polygons of the coxal pores on the 12th-r5th pairs of legs of L. lalis

These distributions of frequencies are illustrated mro

in Figure 11.

No. 437.] VARIATION IN LITHOBIUS FORFICA TUS.

Coxal pores

12th leg left

I2th leg right

13th leg left

13th leg right

15th leg right

Type

TABLE 7.

Mean. P. E.

Fact.

++ |

Skew. St. Dev. P, E.

o04— .84 -L.o4

002— .865 + .041

07 79 + .038

397

Coef. Var.

11.29

11.7

10.56

9.9

11.88

11.3

15.4

12.94

What the significance of the fact that the average number of

pores is greater on the left side than on the right is I have been

unable to determine.

also tends to be more to the right even where (14th leg) the

curve is skew to the left.

The factor of skewness of the left curves

There is a similar instance recorded

by Bateson, ('94, p. 283), where the abnormality in the number

of nipples in the human is higher on the left side than on the

right.

CORRELATIONS.

TABLE 8.

Coeff. Corr. or p

Anal pair legs

12th pair legs

P. E. Coeff. Corr. or P . E.

-575

.686

.58

Coxal pores of the legs correlated.

039

+ .021

+ .029

+ -039

II. Coxal pores of different legs correlated.

Anal R. & 12th L.

Anal R. & 14th R.

14th R. & 13th R.

14th L. & 13th L.

13th R. & 12th R.

308 THE AMERICAN NATURALIST. (VoL. XXXVII.

III. Length correlated with different characters.

With cox. pores

Anal leg R. 222 + .062

14th leg R. .308 4L .659

I3th leg R. 298 + -059

I2th leg R. 205 063

b. With number of joints in antennae —.013 -E.067

c. With number of prosternal teeth X3tf + .066

CONCLUSIONS.

1. From the latter part of Table 8 it will be seen that

length has little to do with the number of joints in the antennz.

If the — sign were significant it would mean an inverse correla-

tion, the longer the animal the fewer antennal joints. But the

probable error is + .067 so that p may as likely as not lie any-

where from —.08 to + .054. There is thus essentially no corre-

lation. You can say, a priori, that the antennz of the larger,

presumably older specimens are more likely to have been broken

and to be found regenerating. There is no way of telling a

regenerated terminal segment from an original termination.

2. Length of body and number of prosternal teeth have

little to do with each other, the coefficient of correlation varying

between .065 and .195. I picked out from the data the four

individuals with 14 prosternal teeth, the maximum number.

Their lengths were 24, 24, 22, and 21 mm. _ In the four indi-

viduals with 8 teeth, the minimum number measured, the lengths

were 20, 22, 23 and 23 mm. This tells roughly what the coeffi-

cient of correlation tells precisely. In the case of the curve for

the prosternal teeth, which is strongly bimodal, the bimodality is

due to the tendency towards bilateral symmetry. At the ends

of the series this tendency is overcome by the tendency of varia-

tions to revert toward or group around the mode. Hence the

larger number of individuals with 9 and 13 teeth compared to

those with 8 and 14 teeth.

3. The length bears a more decided relation to the number

of coxal pores. The number of coxal pores on each of the right

legs was correlated in succession with the length. The coeffi-

No. 437.] VARIATION IN LITHOBIUS FORFICATUS. 309

cients of correlation of the pores of the right side were, according

to Table 8: Anal, .227; 14th, .308; 13th, .298 and 12th, .205.

That is, the anal and 12th leg coxal pores are more independent

of the length of the animal than are those of the 13th and 14th

legs. In other words the correlation is less at the ends ofa

linear series. I had expected the anal pores, the youngest onto-

genetically, to vary quite closely with the length of the animal,

the fewer pores on the shorter animal and vice-versa. This is

found to be true if a group of shorter animals, 6-15 mm. in

length be measured. The coefficient of correlation of the pores

of the right anal legs with the lengths in a group of 49 young

individuals is .88.

4. In the first part of Table 8 are correlated the coxal pores

of the pairs of legs, right with left. Here again the correlation

is smaller at the ends of the series. The correlation is much

closer than it was with the length but the pores of the anal pair

have a coefficient of .575 and the 12th of .58 against .686 for

the 13th and .69 for the 14th. The order is the same as in the

length correlations except that now it is the anal pair of legs

which shows the least correlation whereas it was the 12th which

corresponded least closely to the length.

I tried also one pair of diagonal cross correlations and some

serial correlations with very interesting results. The coefficient

of correlation of the coxal pores of the right anal legs and the

pores of the r2th legs is .43. That of the anal pores R. with

the r2th pores R. is .44. Consequently, diagonal correlation of

the ends of the series is nearly as close as the correlation of the

terminal members of the series on one side of the animal.

Calculating the correlation of the pores of each leg with the

one next it on the right side of the animal there is shown again

the difference in closeness of correlation between the ends and

the middle of the series. But the closeness of correlation of

the r4th R. pores with those of the 13th R. (.722) was so

high that I tried the opposite side, the 14th L. with the r3th L.

and found not quite as close a relationship but yet one higher

than any previous correlation obtained. There is here an

unusual case. In a bilaterally symmetrical animal the relation

existing between two adjacent segmentally arranged groups of

310 THE AMERICAN NATURALIST. [Vor. XXXVII.

organs is greater than that existing between the two symmet-

rical groups of one segment. That is, the morphological kinship

between successive segments ts greater than the likeness between

the two sides of a segment.

In trying to trace the ancestral history of any species, resem- `

blances which point toward a related species are valuable. We

have these in Lzthobius forficatus and L. multidentatus. The

drawings of the two types of coxal pores, Figures 9 and 10,

show how the normal condition in Z. forficatus can be suggested

by variations in £L. multidentatus and vice versa. We know that

the first condition of the pores in both species is the same, a

single row of small round pores. This is probably the ancestral

condition. A fusion of two pore rudiments in £L. multidentatus

would give the oblong shape natural in Z. forficatus and to be

seen twice in the drawing of L. multidentatus. On the other

hand a further constriction of the middle of the long narrow

pores of L. forficatus (left hand upper coxa) would result in a

two rowed condition. How the many rowed condition arises

from the single row in L. multidentatus and the phylogeny of

coxal pore patterns in general is a subject for further study.

That abnormalities in one species may indicate the normal con-

dition in a related species has been shown by Davenport (’00).

SUMMARY.

I. ‘A place-mode is furnished for Lithobins forficatus, for the

years 1899 and 1900, at Cold Spring Harbor, Long Island.

2. Length of body has essentially nothing to do with the

number of antennal joints in specimens 15 mm. long or more.

3. Length has very little to do with the number of proster-

nal teeth.

4. Length has. some bearing on the number of coxal pores

in the adult, the correlation being closer on the 13th and 14th

legs than on the 12th or 15th legs.

5. Coxal pores show a greater segmental or serial correlation

in the case of the 13th and 14th legs than bilateral symmetry.

6. Variations in the one species of Lithobius point toward

the normal condition in the other species under consideration.

No. 347.) VARIATION IN LITHOBIUS FORFICATUS, 311

In conclusion I wish to express my thanks to Dr. C. B.

Davenport for his many kindnesses in directing the work and

for criticising the paper.

LITERATURE CITED.

BATESON.

'94. Materials for the Study of Variation.

—

2. BOLLMAN, C. H.

'93. Myriapoda of North America. Edited by L. M. Underwood.

Bull. U..S. National Museum. No. 46.

3. DAVENPORT, C.

'99. Statistical Methods.

DUROS

'00. On the Variation of the Statoblasts of Pectinatella. Am. Nat.

Vol. 34, No. 408.

5. SEDGEWICK, A

'95. Peripatus. Ca»b. Nat. Hist. Vol. 5.

6. Tower, R. W. -

'95. The external opening of the “ Brick-red " gland in Limulus.

Zool. Anzeiger Dec. 1, p. 471.

7^. Woon, H: G, Jr

'65. On the Myriapoda of North America.

Trans. Am. Phil. Society Vol. 33 N.S. Part2. 137-248. 3 Pls.

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ON THE MORPHOLOGICAL AND PHYSIOLOGICAL

CLASSIFICATION OF THE CUTANEOUS

SENSE ORGANS OF FISHES.

C. JUDSON HERRICK.

In determining the rank and meaning of any given sense

organ there are three criteria to which we may appeal (1)

Doubtless the most important is direct introspective knowledge,

the psychological criterion. This criterion obviously is not avail-

able to us in the study of the senses of the lower animals, where

we are shut up to the remaining two. (2) Of these the most

important is direct physiological experimentation. The sense

organs under consideration are subjected to various types of

stimuli under experimental conditions and the reactions noted —

the physiological criterion. (3) The structure of the organs

frequently permits of inference as to probable function, a

method of small value except as controlled by other data — the

anatomical criterion.

For over two centuries it has been known that the fishes

possess various highly specialized sense organs in the skin, and

for over half a century it has been recognized that these belong

to two distinct morphological types. The problem of the mor-

phological and physiological significance of these organs has exer-

cised some of the ablest zoólogists during the whole of these

periods without, however, any agreement having been reached.

The reason for this unsatisfactory condition is not far to seek.

Most of these authors have been content with inferences as to

function based on studies of the structure of the organs— a

perilous course at the best—and few carefully wrought out

physiological experiments have been made. By the coóperation

of a number of students this condition is now largely remedied

and positive conclusions become possible. We shall summarize

these conclusions without in this place undertaking to give the

evidence for them in detail or to cite authorities, taking up first

3!3

314 THE AMERICAN NATURALIST. [Vor. XXXVII.

the anatomical findings which have laid the foundation historic-

ally for the physiological differentiation of these sense organs.

In the skin of fishes there are three types of sensory nerve

endings belonging to the cerebro-spinal system, aside from

sympathetic nerves of uncertain physiological significance. (1)

there are free endings everywhere in the skin, but especially on

the lips, barblets and exposed surfaces generally. (2) Sense

organs in canals or pits obviously belonging to the lateral line

system and termed canal organs and pit organs, or collectively

nerve hillocks or neuromasts. (3) Sense organs resembling

taste buds of the mouth, lying freely exposed on the surface of

the skin, never sunk below the surface, and variously termed

flask-shaped organs, end-buds, terminal buds, etc.

The nerves which end free, those of the first type, usually

lose their medullary sheaths some distance below the skin and

have sometimes been overlooked. Merkel, for instance, in his

great monograph denies the presence of these endings in the

skin of fishes. Recent students of nerve components have

shown that the nerves of this type are anatomically distinct from

those for both the other types of sense organs for their entire

extent, in all cases being provided with separate ganglia and

entering the brain by distinct roots. In the trunk region these

* general cutaneous" nerves make up the greater part of the

dorsal spinal roots and terminate in the dorsal horns; in the

head they enter by the V, X and sometimes by the IX pairs of

cranial nerves and all terminate in the funicular nucleus or the

gray matter associated with the spinal V tract, all morphological

equivalents of the dorsal horns of the spinal cord. Morpholog-

ically the general cutaneous system of nerves is a well defined

unit. Physiologically its function is unquestionably in the main

tactile. Practically all parts of the body are sensitive to touch

and are reached by these nerves, whether they are supplied by

other types of sensory nerves or not. The more acute the tac-

tile sensibility, the more rich the innervation by fibers of this

system.

The differentiation of the sense organs of the second and third

types has proven very difficult, for the reason that rows of

undoubted lateral line organs which are certainly homologous

No. 437.] SENSE ORGANS OF FISHES. 315

appear in one species as canal organs, in another as pit organs

and in still another as naked sense organs, and the separation of

the latter from the terminal buds of the third type is often a

matter of difficulty. This has led many of the students of these

organs to deny the validity of the distinction between neuro-

masts and terminal buds, classing both as variants of one type.

In 1870 Schulze discovered that the neuromasts, whether

enclosed within canals or exposed upon the surface, are charac-

terized by the presence of specific sensory cells, the pear cells or

hair cells, which extend only part way through the sensory

epithelium, while the specific sensory cells of the terminal buds,

like those of taste buds, extend from the external to internal

limiting membrane of the epithelium. This has been generally

confirmed and receives further support and interpretation from

the recent demonstration that neuromasts and terminal buds

receive distinct and strictly characteristic innervation.

All neuromasts, whether canal organs, pit organs or naked

organs, are innervated by fibers which are separable from all

other types of nerve fibers, with separate ganglia and roots, and

all of which terminate in the tuberculum acusticum or in the

cerebellum. The sense organs of the internal ear have the same

general structure as the neuromasts of the skin and are likewise

innervated from the tuberculum acusticum, so that the whole

system is termed the acustico-lateral system of nerves and sense

organs. The tuberculum acusticum and its derivative, the cere-

bellum, are morphologically intimately related to the general

cutaneous centers of the dorsal horn, and the whole acustico-lat-

eral system is in all probability phylogenetically derived from the

general cutaneous system.

This probability is strengthened by the results of recent

physiological experiments upon this system. The lateral line

system of fishes as a whole is undoubtedly concerned in the

maintenance of bodily equilibrium, and the method of stimulation

here is closely similar to that of ordinary tactile nerves, and

doubtless derived from it. The function of orientation in space

is especially localized in the semicircular canals of the internal

ear, of like phylogenetic origin with the lateral line canals, and

this part of the system persists in all terrestrial vertebrates.

316 THE AMERICAN NATURALIST. | [Vor. XXXVII.

Parker has also shown that the lateral line organs of fishes are

sensitive to mechanical jars of low frequency. This again is

closely related to the tactile function and has doubtless given

rise phylogenetically to the power of perceiving rhythmic vibra-

tions of higher frequency, viz, hearing, a part of the sense

organs of the acustico-lateral system within the internal ear

having been set apart for this function in the course of verte-

brate evolution.

Having now a morphological criterion for defining the cutane-

ous sense organs belonging to the lateral line system, it may be

stated briefly that all other specialized cutaneous sense organs of

fishes at present known may be grouped with the taste buds of

the buccal cavity both on the basis of their structure and of their

innervation. The nerve supply of all of these organs is now

known to be from a system of nerves distinct for their entire

extent from those previously considered, but intimately related

centrally to the sensory nerves from visceral surfaces in general.

This is known to students of nerve components as the communis

system of nerves and sense organs, because its fibers all end

centrally in the gray matter connected with the fasciculus

communis (=f. solitarius of human anatomy).

This system of nerves, like the acustico-lateralis, is well

developed in certain cranial nerves only, and, as the latter system

is supposed to have evolved from the general cutaneous system,

so the communis system has probably been differentiated from

the general visceral sensory nerves of the trunk. Peripherally

it is easy to distinguish the unspecialized component of this

system from that which is distributed to special sense organs ;

but centrally this is much more difficult. This, however, I think

I have accomplished in part at least in the case of Ameiurus.

At any rate, the chief ascending gustatory path in these fishes

is clearly separable from all other reflex paths from the primary

communis centers.

No important distinction can be drawn either in structure or

in innervation between the terminal buds of the outer skin and

the taste buds of the mouth, and to complete our argument it

remains to show whether these sense organs are similar in

function also. This has now been accomplished. In July, 1902,

No. 437.] SENSE ORGANS IN FISHES. 317

I presented before the American Association for the Advance-

ment of Science at the Pittsburg meeting the report of certain

experiments made upon the common cat fish, Ameiurus, which

go to show that this animal actually tastes with the terminal

buds known to be freely distributed over the body surfaces and

especially on the barblets. Since that report I have extended

these observations upon a number of marine fishes, particularly

the gadoids, and the report upon this work is now in press in

the Bulletin of the U. S. Fish Commission.

It may be regarded as established that fishes which possess

terminal buds in the outer skin taste by means of these organs

and habitually find their food by their means, while fishes which

lack these organs in the skin have the sense of taste confined to

the mouth. The delicacy of the sense of taste in the skin is

directly proportional to the number of terminal buds in the

areas in question. Numerous unrelated types of bony fishes

from the siluroids to the gadoids which possess terminal buds

have developed specially modified organs to carry the buds and

increase their efficiency. These organs may take the form of

barblets or of free filiform fin rays and the free rays of the pelvic

and dorsal fins of some gadoid fishes are thus explained.

The results of this examination may be summarized in the

following form :

I. Organs of the general cutaneous system. Free nerve endings distrib-

uted over the general body surface in all vertebrates. Innervation

by general cutaneous nerves ; primary centers, dorsal horns of spinal

cord and homologous centers of oblongata. Function, touch.

II. Organs of the acustico-lateral system. Peripheral organs, neuromasts,

or special sense organs with hair cells among indifferent support-

ing cells, the former extending only part way through the sensory

epithelium. Typically arranged in lines on various parts of the

body according to a tolerably definite pattern whose details are,

however, exceedingly variable. The internal ear is a specialized

part of this system. Innervation by the acustico-lateralis nerves ;

primary centers tuberculum acusticum and cerebellum. Neuromasts

may assume one or several of the following forms in a given fish:

1. Canal organs, regularly arranged in canals lying in the dermis or

dermal bones and communicating by means of pores at frequent

intervals with the surrounding water. Function, perception of

mechanical jars or slow irregular vibrations and the maintenance

of bodily orientation. Occur in nearly all fishes.

318

a de

Sei

III.

N =

THE AMERICAN NATURALIST. [Vor. XXXVII.

Pit organs. Similar to the last, but each organ sunk in a separate

^ pit which opens.to the surface by a pore. Usually regularly

arranged in lines which are closely related to the canals and

which may replace the canals. Function unknown, but probably

similar to the last. The pits in which these organs are placed

may become very shallow or disappear altogether, the organ

becoming superficial, with no apparent change otherwise. Occur

in most ganoids and teleosts and sometimes (perhaps generally)

in elasmobranchs.

Small pit organs. Similar to the last, but smaller and always in

deep pits. Not arranged in definite patterns, but irregularly

distributed over the skin. Known only in siluroid fishes.

Ampulle. Organs similar. in plan to pit organs, lying at the bot-

tom of long slender tubes opening by pores at the surface of the

skin, the pores scattered over the head, but the inner ends of

the ampulla grouped in definite clusters. Occur only in elasmo-

branchs. Function unknown.

Vesicles of Savi. Closed vesicles, found only in the torpedoes.

Cristze acustica. Found in the semicircular canals of all verte-

brates. Function equilibration (reaction to rotary movements).

Macule acustica. Found in the sacculus and utriculus of all

vertebrates. Function equilibration (reaction to translatory move-

ments, and static sense ?) and hearing (?).

Papilla acustica basilaris. The essential nervous part of the

organ of Corti. Function hearing. Found only in vertebrates

above the fishes.

Organs of the communis system. Peripheral organs are special

sense organs with the specific sensory cells extending through the

whole thickness of the sensory epithelium ; organ generally rest-

ing on a raised papilla of the dermis. Present in the mouth of

most vertebrates and in the outer skin of some ganoid and tele-

ostean fishes. Innervation by communis nerves; primary centers

gray matter associated with the fasciculus communis (f. solitarius),

in fishes the vagal and facial lobes. Function taste. There are

recognized two forms with no important differences other than

position.

Taste buds, within the mouth.

Terminal buds, in the outer skin, often on barblets or other spe-

cialized organs for their reception.

THE COMPOUND EYES OF MACHILIS.!

FRANCES SEATON.

Tus study was made from the eyes of Machilis variabilis?

This insect is found in great numbers on the under surface

of stones which lie near the water's edge at the bottom of Fall

Creek gorge, Ithaca, N. Y.

During the last of June, when the first trips for collecting

these insects were made, they were found to be of two different

sizes ; but in August only a few of the small ones were met with.

FiG. 1. A vertical section through the two compound eyes. showing their appearance before the

pigment is removed.

Whether this difference in size corresponds to a difference in sex,

or whether the larger ones were some that were hatched during

the previous autumn and had survived the winter, was not ascer-

tained.

In June and July, the exuvie were abundant on the stones,

but in August so few of these were found that it would seem as

if these insects must reach the adult stage during the latter part

of the summer. At this time also, the largest insects measured

I cm. in length, excluding the antennae and caudal filaments.

Early morning proved the best time for collecting Machilis.

! Contribution from the Entomological Laboratory of Cornell University.

? Identified by Mr. A. D. MacGillivary.

319

320 THE AMERICAN NATURALIST. [Vor. XXXVII.

Before ten o'clock, particularly if the mornings were damp or

cloudy, they were so plentiful that as many as thirty have been

captured within an hour. Toward noon as the sun neared the

zenith so that its rays reached the bottom of the gorge, I have

seldom found more than half a dozen specimens, and that too,

in the same place where, perhaps on the following morning they:

have appeared as abundant as ever. This has happened so

often that it has led me to believe that those creatures do not

enjoy much heat and to escape it, either go farther from the sur-

face or seek stones in a more sheltered place.

Machilis were generally found with the body, antennae and

Fic, 2. A vertical section through one compound eye, depigmented and stained.

caudal filaments lying flat against the under surface of the stone.

Occasionally the posterior end of the body was raised so that the

appendages of the last two and sometimes of the last three seg-

ments were raised above the ground, giving the animal an alert

appearance. Unless touched, they seldom offered to move, even

after the vial used in capturing them, was placed over them.

They appear to be sluggish, depending upon their protective

coloring as a means of escape from their enemies.

During the summer they have been kept in the laboratory in

bottles containing pieces of softened, partially decayed wood upon

which they have been seen feeding. Those kept in this way

No. 437.] COMPOUND EYES OF MACHILIS. 321

moulted every seven days though no one insect lived longer than

three weeks.

Methods.—This study was made entirely from serial sections.

Some of the sections were cut five microns thick but owing to

the difficulty of obtaining such thin sections through the chitin,

the greater part of the material was cut ten microns thick.

In preparing the eyes, the animals were first killed by being

dropped into hot water after which the heads were placed in

the fixing reagent. Before the heads were cut off, the antennz,

palpi and as much as possible of the pronotum were removed

since it was found that these parts, if left on the head, interferred

with the process of imbedding and cutting. The removal of

these parts had to be done with great care for the pressure of

the hand in holding the animal often caused a distortion of the

parts of the eye thus ruining the material for study.

Of the many different fixing reagents tried the best results

were obtained with Flemming's fluid, Picro-sulphuric acid, and

platino-aceto formaldehyde. The eyes were cut in paraffin after

which they were depigmented and stained on the slide. For

removing the pigment, nitric acid, caustic potash, and peroxide of

hydrogen were each tried, but the last proved to be the most

satisfactory.

The stains used were Heidenhain's iron haematoxylin, borax

carmine and Delafield’s haematoxylin. The first was often fol-

lowed by orange G. For staining nerve fibres, methelyn blue

was used by injection and immersion. The method used was

that given by Huber in the Journal of Applied Microscopy,

April,'98. The results were not at all satisfactory, owing to the

great amount of pigment in the eyes, which concealed all trace of

the nerves. In addition to methelyn blue, Kenyon's method for

the brain of the bee (Journal of Comparative Neurology, '96) was

also tried and although this brought out beautifully the structure

of the optic ganglia and the nerves proximad of the basement

membrane, the eye was so uniformly stained that a definite state-

ment as to which cells are the nerve-end cells can not be made.

The external appearance. — The two compound eyes occupy

the entire cephalo-dorsal part of the head, coalescing on the

epicranial suture for a distance of 2 mm. Each eye is almost

322 THE AMERICAN NATURALIST. (Vor. XXXVII.

circular in outline, the diameter from the anterior to the posterior

margin being slightly greater than that from the dorsal to the

ventral margin.

The eyes appear light green in the centre with a peripheral

band of reddish brown.

The corneal cuticle. — The corneal cuticle, a transparent con-

tinuation of the body cuticle, is divided into facets which are,

with a few exceptions on the periphery of the eye, uniformly

hexagonal and of the same size (Fig. 3 a). Each eye contains

between 450 and 525 facets. The facets have a maximum width

of 234 microns between their parallel faces and 25% microns

between opposite angles (Fig. 3 a).

The corneal cuticle differs somewhat in thickness in different

individuals, reaching a maximum in those insects about ready to

moult. The line of separation between the old and the newly

formed cuticle is very distinct. This, in individuals about to

moult is caused by the fact that the former is more deeply

staining than the latter (Fig. 3). The maximum thickness of

the cuticle at the centre of each facet is nine and one third

microns, while at the periphery it is but seven microns.

The corneal hypodermis.— Proximad of the corneal cuticle

and separating it from the cone-cells is the corneal hypodermis

—a layer of well differentiated cells and one of considerable

thickness (Fig. 3). There are two hypodermal cells beneath

each facet. The distal surface of each pair of hypodermal cells

is flat and hexagonal in outline (Fig. 3 b). This outline coin-

cides exactly with that of the corresponding facet.

Extending across the distal surface of each pair of cells and

perpendicular to two opposite sides of their hexagonal outline is

a straight deeply staining line which marks the separation of the

two cells. (Fig. 3 b). These lines are not all strictly parallel.

Proximad, the hypodermal cells become considerably smaller,

having a width at their proximal end equal to that of the under-

lying cone which is 6 microns less than the width of the distal

surface of the hypodermal cells. The Nene surface of each

pair of cells is concave. (Fig. 3.)

The nuclei of the hypodermal cells are NS and deeply stain-

ing. In transverse sections through the distal end of these cells

No. 437] COMPOUND EYES OF MACHILIS. 323

p: XD J- azr Ye.

D AS

Fic. 3c. Fic. 3g.

PLATE I.

Fic. 3. A eins em Jess three ommatidia et are bë of an insect about ready to

moult.

Fic. 3a. A transverse section through a, Figure 3.

i through

Fic. 3f. A transverse section through /, Figure 3-

Fic. 3g. A transverse section through g, Figure 3-

324 THE AMERICAN NATURALIST. [Vor. XXXVII.

the nuclei are semi-elliptical, lying with their long axes parallel

to the line separating the two cells. (Fig. 3 b.)

In Machilis maritima, Oudemans (87) describes four hypoder-

mal cells, but I am sure that there are but two in Machilis vari-

abilis. As Oudemans figures only a longitudinal section of the

ommatidia, it may be that he has made an error in observation

on this point, for in such a section, two cells present the same

appearance as four.

The cone-cells, —In each ommatidium, proximad of the two

hypodermal cells, lie four long cone-cells (Fig. 3, c. c.) These

cells are in close contact along their entire axial surface, forming

a cone forty-five microns long with a diameter of seventeen

microns at its base or distal end.

At its proximal end the cone is about two and one third

microns wide. Although these four cone-cells are closely

applied along their axial surfaces, yet their intercellular walls

are distinct in all transverse sections of the cone from its base to

its proximal end (Fig. 3 d and 3 e).

At the extreme distal end of the cone, lie the granular and

deeply staining nuclei of its four cells, (Fig. 3, ». c. c) In -

transverse sections through these nuclei, each is seen to have the

same triangular form and size as that of the cells to which it

belongs, thus entirely filling its distal end. (Fig. 3 c) In

longitudinal sections, these cone-cell nuclei appear to have their

greatest thickness over the centre of the cone and gradually

diminish in thickness toward the periphery. (Fig. 3, n. c. c.)

The convex distal surface of these cone-cell nuclei fits into the

depression on the proximal surface of the two hypodermal cells.

(Fig. 3.)

Whatever the substance of the cone cells may be, it is but

slightly affected by dyes. In most preparations it appears

absolutely unaffected by them, remaining a perfectly hyaline

structure.

The distal pigment. — Surrounding the posterior two thirds of

each cone, is a sheath of pigment which appears black in masses,

but whose separate large round granules are maroon in color.

(Figs. 3 e, and 7.) Outside of this thin sheath of black pigment

and separating the cones of the different ommatidia, is a brown-

No. 437.] COMPOUND EYES OF MACHILIS. 325

ish yellow pigment. The cells containing this last pigment :

extend from a short distance distad of the proximal end of the

cones up between the cones and hypodermal cells tó the corneal

cuticle to which they appear to be attached. (Fig. 3. The

nuclei of these cells lie between the outer ends of the cone-cells

(Fig. 3, z. y. c.) In a transverse section through the cone,

just proximad of the cone-cell nuclei, the nuclei of these yellow

pigment cells appear arranged in a circle around each cone (Fig.

3d). Asthese nuclei lie at different levels in the eye, their exact

number is difficult to determine. Not less than eight have been

counted and in some sections, as many as ten or twelve, so that

each cone is surrounded by at least three yellow cells and pos-

sibly as many as six.

It may be possible that these yellow cells constitute an iris

tapetum as in the cabbage butterfly ; or they may contain a pig-

ment that acts as such, as in the Dragon fly. (Exner ’91.)

In transverse sections through the proximal half of the cones,

the black pigment mentioned before occupies a narrow ring

around each cone from which it is separated by the peripheral

cell wall. (Fig. 3 e) Outside of this ring of black pigment

and filling up the interstices between the cones, lies the brownish

yellow pigment. (Fig. 3 e.) It seems probable from both

transverse and longitudinal sections that the two kinds of pig-

ment which surround each cone represent two distinct circles of

cells, although no nuclei have been found in the narrow area

occupied by the black pigment.

Although eyes have been examined from insects which have

been kept for three hours in the dark previous to killing, as well

as eyes from other insects which have been kept in the light, for

an equa length of time, no change in the position of the black or

* iris " (Exner) pigment was noticed.

The rhabdoms.—The rhabdoms are the long, rod-like struc-

tures which lie between the proximal ends of the cones and the

basement membrane (Fig. 3). They occupy about two thirds

the entire depth of the ommatidia. At its distal end each rhab-

dom is equal in width to the proximal end of the cone against

which it presses (Figs. 2, 3, and 5). Proximad, the rhabdoms

taper slightly until at the basement membrane they are only one

326 THE AMERICAN NATURALIST. [Vor. XXXVII.

micron in width. The rhabdoms are fluted, having in trans-

verse sections the form of a seven-pointed star (Figs. 3 g, and 7).

In sections from which the pigment has not been removed,

the rhabdoms appear as perfectly hyaline structures, while in

depigmented and stained sections, with the exception of a small

central core, they invariably appear non-granular and deeply

staining. (Fig. 3, and 2). Thus the rhabdoms present a

decided contrast to their cones which are but slightly affected

by dyes, if at all.

There has been no indication in any of the sections that the

rhabdoms consist of seven parts or rhabdomeres, except that in

cross section they appeared as seven pointed stars. (Figs. 7 and

-8.)

In many longitudinal sections they often have a peculiar

beaded, or sometimes a corkscrew appearance. (Fig. 9.) This

peculiarity has been noticed in sections so differently treated

that, at first, it seemed as if it must be due to the presence of

nerve fibres in the rhabdoms but it now seems that it is prob-

ably an artifact.

That the cones and rhabdoms are in Machilis distinct and

separate structures, is evident for three reasons :— first, the

rhabdoms always appear as deeply staining structures, while the

cones do not; second, in transverse sections the cones appear

circular in outline and divided into four parts or cells while the

rhabdoms invariably have the form of a seven-pointed star;

third, in longitudinal sections there is always a distinct trans-

verse line where the cone-cells end and the rhabdoms begin.

(Figs. 2 and 3.) Then, too, in many sections, the cones appear

partly broken away from the rhabdoms at this point and when-

ever the cones were entirely separated from the rhabdoms, the

break invariably occurred at this particular point. (Fig. 5.)

This is a very different condition from that which exists in the

lobster’s eyes as described by Parker (’90), where the four cone-

cells are continued as fibers outside of the rhabdom to the base-

ment membrane. It also differs materially from the condition

found in the eyes of Mantis (Patten '86), where the cone-cells

extend to the basement membrane through the centre of each

ommatidium, there being no distinction between cone-cells and

rhabdom.

No. 437.] COMPOUND EVES OF MACHILIS. 327

The retinule.—In specimens which have not been depig-

mented the rhabdoms are surrounded throughout their entire

length, by a thin sheath of reddish brown pigment. (Fig. 9.)

In cross sections, this pigment is seen to extend up close to each

rhabdom, filling the spaces between its seven points (Fig. 8.)

In longitudinal sections this narrow area of reddish brown pig-

Fic. 4. A vertical section c the two peer eyes before pigment was removed ;

amera lucida d

€ ò

A vertical section the cones and

por a three adjacent ommatidia ; a,

cone and rhabdom in close Sonaactioi at

pple : O d fro

the rhabdom; c, cone an bdom cane

separated, camera lucida pen

ment surrounding the rhabdoms, appears of the same width, and

as if it might be continuous with the thin sheath of black distal

pigment which surrounds the cone-cells. (Fig. 7.)

In transverse sections of eyes that have been depigmented

and stained, the narrow area immediately surrounding the rhab-

dom, which was before filled with pigment, now appears as a

clear non-staining area and across it extend many fine lines

from the rhabdom (Fig. 7). Outside of this, the area which

before was faintly granular, now appears as a granular, deeply

staining area whose outer edge is divided into seven distinct

(Figs. 3 g and 7.) Thus in transverse sections through

area surrounding the rhabdom.

parts.

328 THE AMERICAN NATURALIST. (Vor. XXXVII.

the proximal half of the eyes, the ommatidia appear as many

rosette-like figures which lie so close together that there can be

little space between

them in eyes. (Fig.

3 g). Each rosette-

like figure has in its

centre the star-like,

deeply staining rhab-

dom which is sur-

rounded by a narrow

Fic. 7. A transverse section, showing the rosette-like appear- hyaline area. Out-

om the rhabdomes and retinule, depigmented and žige of the latter is

the deeply-staining

granular area whose outer margin shows a distinct division into

seven parts. (Fig. 7.)

In many longitudinal sections of depigmented and stained

eyes, the narrow hyaline area surrounding the rhabdoms, can be

seen with a high power, to be crossed at regular intervals by

many very finelines. (Fig. 6.)

Whether the two areas surrounding each rhabdom represent

two distinct circles of cells, the

inner — the retinulae proper —

and the. outer,— accessory cells,

— or whether these two areas are

two parts of a single whorl of cells

is difficult to determine. It seems

probable, however, that since no

nuclei have been seen in the inner

area, we have in each ommatidium

a single whorl of seven cells, the !19-8- A transverse section through the

: 2 eye showing the rhabdomes and pigment.

retinula, surrounding each rhab-

dom. These seven retinulz are of uniform size and have the

pigment massed on their axial border. The number of retinulz

differs from that in Machilis maritima, where Oudemans ('87)

described six.

In longitudinal sections of the ommatidia, the seven retinulz

are seen to extend slightly distad of the proximal end of the

cone-cells, where each is considerably enlarged to accommodate

No. 437.] COMPOUND EVES OF MACHILIS. 329

its large nucleus. (Fig. 3.) The nuclei of the retinule do not

all lie in the same plane. (Fig. 2.)

In addition to the pigment along the axial border of the

retinule each contains con-

siderable pigment in its prox-

imal end so that in longitudi-

nal sections of the eyes, this

appears as a narrow band of

pigment distad of the base-

ment membrane. (Fig. 1.)

Since there is in Machilis

no shifting of the iris pigment

and since the rhabdoms are

of uniform width, the insect,

has according to Exner (91)

day eyes with apposed images.

The nerves — Proximad of

the basement membrane, ly-

ing berwein Itant e Fic. 9. A somewhat oblique section showing the

cell sheath of the optic gang- narrow area proximal of the basement membrane

across which the nerve fibers pass from the optis

ganglia to the ommatidia.

lia, is a narrow area contain-

ing hundreds of minute nerve

fibers which enter the eye through the basement membrane.

(Fig. 9.) Iam unable to say which cells of the ommatidia the

nerves enter.

Abbreviations employed in the figures.

€ and c'. corneal cuticle. n.rt. nuclei of retinule.

hyp. corneal hypodermis. rb. rhabdom.

n. hyp. c. nuclei of hypodermal cells. b. m. basement membrane.

d o cone-cells. d.p. distal pigment.

N. 6 €. nuclei of cone-cells. i. line of separation between

Ke Pls nuclei of yellow cells. hypodermal cells.

re. retinulz. Ja. facets.

SYNOPSIS OF NORTH AMERICAN INVERTE-

BRATES.

XIV. Part IV. Tue ScvPHOMEDUSE.

CHAS. W. HARGITT.

Tue following synopsis is a continuation of that upon Hydro-

medusze which appeared as XIV of the American Naturalist

series, during April, May and July, 1901.

As in the preceding parts, while depending largely upon my

own records of the Scyphomedusz, I have at the same time

drawn freely upon the literature wherever found, but chiefly

Heeckel’s * System der Medusen” and to a less extent Mayer's

numerous papers. (Bull. Mus. Comp. Zool.) Fewkes papers,

chiefly of the same series, including also L. Agassiz, “ Contr.

Nat. Hist. United States,” 1862, and A. Agassiz’ “Catalog

N. A. Acalephzs," 1865.

In only a few cases has any attempt been made to present

accounts of the synonymy of the several species, and then only

so much as might serve to obviate ambiguity.

In general form, habit, structure and distribution the Scypho-

medusze have much in common with the Hydromedusz and

probably sustain a more intimate relation to them than to any

other ccelenterate Class.

They may however be somewhat sharply distinguished by the

following characters :

I. Absence of a true velum. The velarium of the Cubo-

medusze has important structural differences, though doubtless

serving an identical function.

2. Sense organs when present are modified tentacles,

variously designated as tentaculocysts, rhopalia, etc.

3. Entodermic origin of sexual products.

In development there is general correspondence between this

and the preceding Class. In some the medusa arises by direct

33!

332 THE AMERICAN NATURALIST. [Vor. XXXVII

(hypogenic), development from the egg ; while in others, and by

far the larger number, development is indirect (metagenic),

exhibiting perfectly evident alternation of generations; in this

case however, involving a distinct metamorphosis, the polyp

giving rise to a free-swimming ephyra which is in turn trans-

formed directly into a medusa. It should also be noted that

asexual budding is, unlike that of the former class, by the

transverse fission of the polyp body into a series of disks which

become free as ephyre, as already noted. Direct asexual

budding from medusoid organs, common in many Hydromedusz,

is unknown among the Scyphomedusz.

SYNOPSIS OF THE ORDERS.

I. STAUROMEDUS&. Scyphomeduse with vasiform or sub-conical

umbrella. In some cases sedentary, attached by an aboral peduncle or

stalk. Wholly devoid of sensory organs, but with eight tentacles or

tentacular organs which serve as anchors. Stomach with four wide

gastric pouches which communicate with a marginal canal. Gonads in

four crescentic loops on the floor of the gastric pouches.

II. PEROMEDUSÆ. Scyphomeduse with bell more or less conical in shape

and with a usually well-developed horizontal constriction which divides it

into two regions ; an aboral, resembling quite remarkably the apical projec-

tion of the bell of certain Hydromeduse ; the marginal portion, which

is eight or sixteen lobed and bearing tentacles and rhopalia or tentacu-

locysts. Stomach capacious with four gastric pouches which are separated

by narrow septa, and extending into a circular sinus. Gonads much as

in the former order.

III. CUBOMEDUSÆ. Scyphomeduse with a distinctively quadrate

umbrella, ptovided with a well-defined velarium, which is supported at the

radial angles by thickenings or frenula. Marginal tentacles four,

interradially disposed, and with four perradial rhopalia. Bases of

tentacles often provided with wing-like expansions, pedalia

IV. Discomepus&. Scyphomedusz with shallow, or db shaped, eight

lobed umbrella. Marginal sense organs eight, per- and interradially dis-

osed about the margin. Tentacles often very numerous. Manubrium

often very large, pendulous and complexly frilled or plaited. Stomach

with four to eight or more gastric pouches, within which are borne

the gonads.

The medusz of this order are often of large size. Specimens of Cyanea

reaching a diameter of from four to six feet in some cases and with ten-

tacles having an extent of more than fifty feet when fully extended. The

average size however, even of this species, is very much smaller, as will

be noted later.

No. 437] NORTH AMERICAN INVERTEBRATES. 333

SYNOPSIS OF FAMILIES OF STAUROMEDUS;E.

I. TESSERIDÆ. Margin of umbrella devoid of definite lobes or anchors ;

the umbrella at-

tenuated at the

apex into a hol-

low stalk, which

in some genera

serves as

means of attach-

dial and four in-

terradial.

^no far as

known no rep-

resentatives of

this family come

within our

range.

Halichvsfu "uhr: Clark

41 Ly V

FIG. 1.

II. LUCERNARIDÆ. Margin of umbrella definitely lobed, each terminat-

ing in tufts of delicate knobbed tentacles ; exumbrella attenuated at the

apex as an organ of attachment; margin of umbrella with eight tentacles,

arranged as in previous family, but sometimes modified as anchors.

KEY TO THE GENERA.

A. Without gastrogenital pockets in the sub-umbrella wall of the radial

pouches. :

1. Umbrella with 8 marginal anchors - . Haliclystus.

2. Umbrella without marginal anchors . . Lucernaria

B. With four perradial gastrogenital pockets in the subumbrellar wall

of the four radial pouches.

3. Margin of umbrella with 8 anchors . . Halicyathus.

Haliclystus auricula Clark. 18635.

Fig. 1.

Haliclystus auricula, Clark, 1863, 1878.

“ primula, Heckel, 1877.

t 1865.

Lucennaria " x

Haliclystus auricula, " 1880.

334 THE AMERICAN NATURALIST. [Vor. XXXVII.

Umbrella octangular-pyramidal, umbrella stalk quadrate-prismatic, approx-

imately as long as the bell height. Eight arms, arranged in pairs; four

perradial sinuses broader and deeper than the four interradials; each arm

with from 100-120 tentacles; eight large marginal anchors.

Color.— Very variable, often -E almost every tint of the spectrum,

though generally having a single color

Size— Broad diameter 20-30 mm. Height, including stalk, 20-30 mm.

Distribution.— From Massachusetts Bay northward to Maine, etc.

Haliclystus salpinx Clark. 1863.

T salpinx Clark, 1863.

H. salpinx A. Ag. 1

Lucernaria salpinx Heckel 1865.

Haliclystus salpinx Heckel, 1880.

Umbrella octangular, stem quadrate, prismatic, with four interradial longi-

tudinal muscles; eight arms, symmetrically disposed, each with a tuft of

60-70 slender tentacles. Marginal anchors very large about as long as the

arms.

Distribution.— Chiefly Northeastern Atlantic coast.

Lucernaria quadricornis O. F. Müll. 1776.

Umbrella flat funnel-shaped or quadrate-pyramidal, approximately twice

as broad as high. Stem cylindrical, single-chambered, about as long as the

bell-height and with four interradial longitudinal muscles. Eight arms

arranged in pairs, the four perradial sinuses of the bell margin as broad and

deep as the four interradials. Each arm with from 100-120 tentacles.

olor.— Variable, gray, green, yellowish brown to red-brown.

Size — Umbrella 50-60 mm., height including stalk, 50-70 mm.

Distribution.— As in Haliclystus.

Halicyathus lagena Heckel. | 1880.

Lucernaria auricula Fabr. 1780.

L. typica Greene 1858.

L. fabricii L. Ag. 1862

L. lagena Heckel 1865.

Manania auricula Clark, 1863.

M. auricula A. Ag. 1865.

M. lagena Heck. 1877.

Halicyathus lagena Heck. 1880.

Bell deep flask-shaped, about twice as high as broad ; stalk slender cylin-

No. 437] WORTH AMERICAN INVERTEBRATES. 133

drical, single chambered, much longer than height of bell. Arms eight,

arranged in pairs, not longer than broad; each arm with 60-70 delicate

tentacles. Eight marginal anchors.

Color. — Black or dark brown, occasionally reddish- or yellowish-brown.

Size.— 5-7 mm., height including stem, 20-30

Distribution. — Eastport, Me. (Stimpson), seinni, (Ag.), Greenland.

SYNOPSIS OF THE FAMILIES AND GENERA OF PEROMEDUS;E.

Family PERIPHYLLIDÆ. Rhopalia 4, marginal lobes 16, tentacles 12.

Rhopalia 4, marginal lobes 8, tentacles 8.

Family Pericolpide.

A single genus only of the Peromedusze is represented within

the range of the present synopsis, namely, Periphylla and under

this three species have been recorded.

Generic characters : — Umbrella with four perradial, buccal pouches and

with four basal funnels; gastric

pouches with two rows of filaments.

Periphylla, hyacinthina Steenstrup.

1837.

Fig. 2.

Umbrella bell-shaped, about as

broad as high; marginal lobes nearly

rightangled truncated below ;

double the length of the bell-height.

Manubrium extending to the base of

the marginal een and about double

as broad as

Color. Enab reddish, pedalia

and marginal lobes red to violet, tenta-

cles bluish. (Hæckel.)

Distribution. — Greenland, Steen-

strup, Gulf Stream go-10o miles S. E.

off Martha's Vineyard (Fewkes). Fic. 2.

Periphylla peronii Heckel.

336 THE AMERICAN NATURALIST. (VOL. XXXVII.

Periphylla humilis Fewkes. 1884.

Bell low conical, diameter twice that of height. Rhopalia 4, provided

with protecting hood; marginal tentacles 12, of yellow color. Color of

exumbrella brown, rough and opaque; central disk and corona rather uni-

form brownish in color.

Distribution.— Off Martha’s Vineyard as for previous species.

Periphylla peronii Heckel, 1880.

Charibdea periphylla, Per. & Les. 1809.

C. periphylla, L. Ag. 1862, Cont. Nat. Hist. U. S.

Stomolophus periphylla, Fewkes ?

Umbrella low conical, about as broad as high. Marginal lappets 16,

eight tentacular and eight ocular. Tentacles long and stout, about as broad

at the base as the marginal lappets. Manubrium about as broad as high,

somewhat cubical.

Distribution.— Tropical Atlantic, (L. Agassiz), St. George's Bank (S. I.

Smith).

FAMILIES OF CUBOMEDUSÆ :

Of the Cubomedusæ only a single Family has been represented by spe-

cies within the range of this synopsis, namely Charibdeidæ, and under this

but a single genus and species.

Charybdea verrucosa Hargitt, 1902. Fig. 3.

Several specimens were taken at Woods Holl during the

summer of 1902 and have been

described by the present writer, Az.

Nat. July, 1902. Bell ovoid in out-

line, as seen in profile, cuboid viewed

from the aboral pole. Size from 2 to

3 mm. in short diameter by 4 to 5 mm.

in the height. Surface dotted irregu-

larly with light brownish, warty clus-

ters of nematocysts. Rhopalia 4, pre-

radially located, set in rather deep

pockets, and shielded by projecting

hoods. Tentacles short and spindle-

like, with deep annulations, interra-

dially situated. Velarium well-devel-

FiG. 3. Charybdea verrucosa Har-

gitt.

No. 437.] NORTH AMERICAN INVERTEBRATES. 337

oped, but without distinguishable canals, supported by frenulz

on the inner perradial corners of the sub-umbrella. Gonads

were undeveloped, and no distinguishable gastric filaments were

present.

In color the specimens were light amber being darker on the

tentacles.

As pointed out in the description before cited, the specimens

under consideration show many points of difference or contrast

as compared with typical Charybdea species. Mayer who has

described a similar species from the Tortugas ascribes this to

immaturity. This has seemed to me somewhat doubtful, and

some hesitancy was entertained as to whether they probably

come within the Charybdeidz ; but in the absence of specimens

in sufficient numbers or undoubted maturity it seems impossi-

ble to more definitely settle the problem.

Mayer has described two species from the Tortugas, namely,

C. aurifera and C. punctata. Both species were based on single

specimens and both seemed immature. Hence the same doubt

rests upon these as upon the previous species. A comparison

of Mayer's figures, Bull. Comp. Zool, XXXVII, No. 2, will

show many points of. similarity and suggests close relationships.

SYNOPSIS OF SUB-ORDERS AND FAMILIES OF DISCOMEDUS#.

Sub-order 1. CANNOSTOM.

Discomeduse with simple, quadrate mouth, devoid of oral lobes or tenta-

cles; marginal tentacles short, solid.

Family EPHYRIDÆ. Radial pouches usually 16, broad and simple; no

marginal canal. Chiefly deep-sea forms, occasionally taken at the surface.

Family LINERGIDÆ. Radial pouches broad, terminating in numerous

branching, blind distal canals.

Sub-order 2. SEMOSTOM;E.

Discomedusz with quadrate mouth, and with elongated, oral arms, or

lobes, which are often complexly folded and frilled: marginal tentacles

hollow, often very long. Marginal lobes usually 8.

Family ULMARID&. Radial canals of small size, but usually numerous

and branching, the branches often anastomosing into an intricate network

and finally uniting with a definite marginal canal.

338 THE AMERICAN NATURALIST. [Vor. XXXVII.

Family CyANEID#. Radial canals broad and pouchdike, and with

numerous ramifying, blind, lobular canals; no circular canal; 8-16, rarely

more, marginal lobes.

Family PELAGIDÆ. Radial canals rather broad but simple and without

ramifying branches; no marginal canal; usually 16 marginal lobes.

Sub-order 3. RHIZOSTOM.

Discomedusz in which the mouth early becomes more or less overgrown

and obliterated by the 8 rootlike oral arms; gastric cavity extending into

arms and opening by funnel-like mouths on the edges and sur-

faces. Devoid of marginal tentacles.

Family ToREUMIDa&. Radial canals 8-16, narrow and with anastomos-

ing branches; devoid of marginal canal; rhopalia 8-16. Suctorial fun-

nels on the outer (dorsal) surface of the oral arms.

Family PILEMIDÆ. Radial canale 8-16, occasionally more, variously

branching and anastomosing ; rhopalia 8. Suctorial funnels on both outer

and inner surfaces (dorsal and ventral), of the oral arms.

KEY TO THE GENERA.

EPHYRID2.

1. Gonads four, simple, horse-shoe-shaped ; devoid of Te lobes or

lobular pouches . : . Ephyra

2. Gonads 4, Lobitkr voltus i6 8 oiis, 8 biotidali Pihyhuch:

3. Gonads 8, symmetrically disposed; 16 lobular pouches, ocular,

Nausithoé.

4. Gonads 8, ewes — 32 lobular pouches, 16 ocular, 16

tentacular, . . Nauphanta.

5. Gonads 8, nec i in pairs ; Tebuk adole pat 28, press indefi-

nit ) Atolla.

6. Goa 4, twodobed. with interradial orda E E R T paa

ULMARIDÆ.

7. Rhopalia 8; tentacles numerous, short, borne on under margin of the

umbrella without the velar lappets; oral arms 4, simple but with the

margins fringed with nematocysts, . Aurelia.

8. Rhopalia 16; tentacles numerous, long, 'i in a6 alteri on | the lower mar-

gin within the velar lappets ... . . . . . . . . Phbhacellophora.

CYANEID.

g. Rhopalia 8; tentacles very numerous, long, arranged in 8 clusters,

each comprising several rows. Oral lobes four, but highly folded and

fringed, ov Ce ee ee. e es Cyanea

No. 437] NORTH AMERICAN INVERTEBRATES. 339

PELAGIDA.

10. Marginal tentacles 8; marginal lobes 16, . . . . . . Pelagia.

1I. Marginal tentacles 24; marginal lobes 32, . . Chrysaora.

12. Marginal tentacles 4o; immature specimens, din. js in younger

individuals ; gei qe lobes 48; . . 5. . . . Dactylometra.

Ephyroides rotaformis Fewkes. 1884.

Report U. S. Fish Commission, p. 949.

Among meduse of the Gulf Stream Fewkes has described what is con-

sidered by him a new genus and species of an Ephyra-like medusa.

The generic characters given are somewhat indefinite, no mention being

made as to gonads, radial pouches, sense organs, etc. The following brief

notes are condensed from the above cited report:

Umbrella flat discoid, and viewed from the aboral pole Sees three

zones :—‘ Discus centralis; Zona coronalis; Zona marginalis.” The

last named zone is marked by definite marginal lappets of large size with

rounded outlines twice as long as broad, and 16 in number. Interposed

between the lappets are a similar number of gelatinous elevations, * socles,"

ending a short distance from the deepest point of the marginal incision and

abutting the line of junction of the discus centralis and zona coronalis. The

marginal lappets are supported at their base by a pair of gelatinous * socles.”

Distribution. —

Nausithoé punctata Koll.

Bull. Mus. Comp. Zoól. Vol. XXXVII, p. 67.

Umbrella flat, 9-10 mm. broad. Marginal tentacles 8, stiff, about 7 mm.

long. Rhopalia 8, alternating with the tentacles. Marginal lappets 16,

long and flexible; gastric pouches 16, simple, and extending to the lappets.

Mouth simple, quadrate, devoid of lobes or tentacles.

Distribution. — Bahama and Tortugas Islands. (Mayer.)

Nauphantopsis diomedeg Fewkes. 1884.

Op. cit. p. 946.

From a fragmentary specimen collected by the Albatross in the Gulf

Stream Fewkes has proposed the new genus and species here mentioned.

The following very brief synopsis of characters are condensed from his

description. Report U. S. Fish Commission 1884.

Umbrella high disk-shaped, with marginal walls probably somewhat ver-

tical. Marginal lobes 32. Tentacles 24, rhopalia probably 8.

Distribution. — Lat. 38° N., long. 69? W.; depth 2.033 fathoms.

340 THE AMERICAN NATURALIST. (VoL. XXXVII.

Atolla bairdii Fewkes. 1884.

Report U. S. Fish Commission, p. 936.

Umbrella disk-like with aboral center convex. Marginal lappets 44.

Marginal tentacles 22, each supported by a gelatinous * socle." Rhopalia

22, situated in notches between the lappets. Manubrium large, with simple

mouth. Gastric pouches 22. ?

Color. — Slightly bluish, with rust-colored patches, especially on the border

of the coronal furrow.

Distribution — Gulf Stream, between N. lat. 35-38; W. long. 72-75.

One specimen from the depth of 991 fathoms, the other from surface.

Arolla verrilii, Fewkes. 1884.

OP. cit. p. 939.

Umbrella flat discoid, six to eight times broader than high.

Marginal tentacles 22 to 28, with same number of interposed rhopalia.

Marginal lappets same number as the combined number of tentacles

and rhopalia.

Distribution. — Between lat. 38-40; long. 68—71; from depth of from

373 to 2,369 fathoms.

Linerges mercurius Heckel. 1880. Fig. 4.

Op. cit. p. 950.

Umbrella mitre-shaped, with

arched crown and usually verti-

cal sides, diameter about twice

that of height. Lobular canals

bowed and rounded out. Ten-

tacles | cylindrical. Gonads

horseshoe-shaped. Size 12 to

16 mm. broad. 6 to Io mm.

high.

Distribution — Bahama and

Tortugas Islands (Mayer). Gulf

of Mexico. Straits of Florida

(Fewkes).

Bathyluca solaris Mayer. 1900.

Bull. Mus. Comp. Zool., XXXVII, p. 2.

Umbrella flat and rather thick, aboral surface dotted with batteries of

nematocysts. Marginal lappets 24; tentacles 16, long and hollow. Rho-

No.437] WORTH AMERICAN INVERTEBRATES. 341

palia 8. Manubrium cruciform, simple, devoid of arms or appendages.

Gonads 4, horseshoe-shaped, beneath which on the subumbral wall are four

open sub-genital pits. Stomach large and with 16 gastric pouches, eight of

which extend to the ocular lobes and eight to the tentacular lobes.

Color. Disk translucent, slightly bluish; clusters of nematocysts dull

yellowish-brown ; tentacles slightly greenish.

Aurelia favidula Per. & Les. Fig. 5.

Fic. 5. Aurelia favidula Per. & Les.

Umbrella flat and disk-like, somewhat arched above; margin normally

eight-lobed and with eight rhopalia located in the marginal sinuses. Many

variations from the normal octamerous form are found in some collections

reaching as high as 25%.

Marginal tentacles numerous, short, forming a delicate fringe about the

entire margin except at the rhopalial sinuses. Radial canals 16, of three

sorts, per- inter- and adradial ; the first two series branching and anastomos-

ing freely, the last usually straight and simple from its origin to its junction

with the marginal canal.

Manubrium cruciform in cross section, and with four long oral arms which

are more or less fimbriated and the margins bearing numerous batteries of

nematocysts. Gonads crescentic in form, borne upon the floor of the four

gastric pouches.

Color.— Aurelia is among the duller colored of the Scyphozoa, the bell

being quite transparent, but with a bluish opalescence. The gonads present

a pale pinkish hue, though the ova are almost clear white as examined singly.

Distribution —Aurelia is one of the commonest of the Atlantic coast

medusz and ranges from the coast of Maine to Florida. It is most abun-

dant during the early summer or spring along most of the New England

coast, though fairly abundant northward till late in summer. Its breeding

habits seem to be somewhat continuous during most of the summer. The

scyphistoma stage is a somewhat extended one, probably lasting over the

entire winter season. Kept for weeks in aquaria they showed no signs of

metamorphism. I have taken them in all stages of strobilization during

342 THE AMERICAN NATURALIST. [Vor. XXXVII.

April and early May, when ephyrz were being discharged in great numbers.

During the summer season the polyps bud and stolonize very freely, from a

single Mte: a colony of many individuals arising within a space of

ten days. Figur such a colony reared within a watch-glass aquarium.

Aurelia marginata L. Ag. 1862.

Cont. Nat. Hist. U. States, Vol. IV.

Umbrella flat dome-shaped to hemispherical, three times as broad as high.

Mouth-arms relatively small, considerably shorter than the umbrella radius.

Gonads very large.

A southern medusa, reported by Agassiz from Key West, Florida.

Callinema ornata, Verrill. 1869.

Umbrella flat and disk-shaped, rather thick and aborally rounded; the

exumbrella surface covered with wartlike papilla; walls transparent and

with prominent radial canals which are of two sorts, one branching and

anastomosing, the other simple and straight, each 16 in number. Margin

with 16 lobes deeply incised within which is located a conspicuous rhopalium.

Tentacles numerous and of varied size and length, arising from the under

surface of the margin beneath the marginal canal. Manubrium large and

pendulous and with prominent folded oral lobes, somewhat like those of

Cyanea. Gonads 8, in prominent pouches within the gastric cavi In

size specimens vary from 10—18 inches in diameter. Distribution, taken at

Eastport, Maine, by Verrill, and later by Fewkes, from whose account this

description is condensed. Cf. Bull. Mus. Comp. Zool. Vol. XIII, No. 7.

Cyanea arctica, Per. & Les.

Umbrella flat and disk-like, with a central aboral convexity, with 8 prin-

cipal lobes and 16 or more secondary lappets; ocular pouches small sub-

triangular, tentacular pouches two or three times as broad as the ocular.

o. adial pouches purplish to brownish ; oral lobes deep chocolate

brown ; gonads yellowish white ; tentacles variably colored, yellowish, orange,

purplish or brown.

Size. — From 100 to 500 mm. in diameter, though in many cases larger.

A. Agassiz notes one having a size of seven feet and with tentacles more

than 100 feet in length.

Distribution. — Almost the entire Northeast coast of the United States.

L. Agassiz has described two additional hom namely, C. fulva, and

C. versicolor. These are of doubtful distinctness, variation in size and

coloration being the chief differences Vis recognizable. Collections

No. 437.) NORTH AMERICAN INVERTEBRATES. 343

made from a wide range of New England coast waters show every feature

of intergradation between the several extremes and sufficiently establish the

fact that at most only C. versicolor, of the Carolina coast, has possibly a

varietal distinctness.

In this connection it is pertinent to refer to the morphological variation

in the common species. C. arctica, which is nearly as marked as in Aurelia,

including variation in the radial symmetry, number of gonads, oral arms,

etc. Variation in color is sufficiently indicated in the references just given.

Pelagia cyanella Per. & Les. Fig. 6.

Fic. 6. Pelagia cyanella Per. & Les.

Umbrella disk-like, with rather highly arched aboral surface; marginal

lobes 16, and with 8 rhopalia and 8 tentacles symmetrically and alternately

disposed at the lobular sinuses. Gonads 8, forming conspicuous pouch-like

masses within the gastric pouches of the tentacular radii. Manubrium

large and pendulous, with four frilled oral arms approximately as long as

the tentacles.

Color.— Disk translucent bluish tint, sprinkled with reddish-brown pig-

ment spots over the entire exumbrellar surface, the more numerous near

344 THE AMERICAN NATURALIST. [Vor. XXXVII.

the margin and forming crescents at the marginal lobes; manubrium

similarly mottled on the outer edges of the arms, inner edges and frills

delicate flesh-colored ; tentacles a dull, madder-ed ; gonads pale purplish.

Two specimens of this medusa have been taken in the Woods Holl region

recently, the last in July, 1902, some 65 miles south of Marthas Vineyard.

According to Agassiz, Contr. Nat. Hist. U. S., the development of this

medusa is direct, skipping the fixed polyp and strobila stages.

Dactylometra quinguicirri L. Ag.

Umbrella rather high and arched aborally much as in Pelagia, disk three

to four times as broad as high. Manubrium long and pendulous with

slender oral arms, which are more or less frilled as in P. cyaze//. Rhopalia

8, marginal tentacles 4o, marginal lobes 48. In arrangement five tentacles

are located between each pair of rhopalia in adult specimens, though in some

cases only three are present, particularly in small specimens. Gonads in

four masses within the gastric pouches, and beneath each gonad in the sub-

umbral wall is a prominent subgenital pit.

Color.— In general much like Pelagia, though less brilliant, the various hues

being paler and somewhat more delicate. Exumbrella delicate bluish,

mottled with reddish brown fading into yellowish; tentacles reddish to

orange; oral arms pale pinkish with bluish tint rd blended, making

this medusa one of the most beautiful among the Pelagidz

Distribution.— Is less extended than that of Aurelia or Cyanea. It is

quite common in Buzzard's Bay, Vineyard Sound, Nantucket.

Like several of the preceding Dactylometra exhibits considerable varia-

tion. According to Mayer, Bull. Mus. Comp. Zool. Vol. XXXII, No. 7,

the tertiary tentacles arise on either side of the ocular lappets. In several

specimens examined during the past summer this was not found to be the

case. On the contrary they sprang in every case examined between the

primary and secondary sets. Again according to the same observer the

after attaining a size of 130 mm. in diameter. On the contrary I found

them well developed in specimens having a size of only 40 mm. and where

no gonads were developed. There was also noted the same variation in

the marginal lobes and other organs which have been noted in connection

with species previously noted.

Dactylometra lactea L. Ag. 1862.

This is a southern medusa, no record of its occurrence north of Florida

having come to my notice. In general aspects it is much like the preced-

ing species, though of smaller size. Its color is milk-white with a purplish

iridescence, and with yellowish dots over the exumbrella. It has been

reported from the Bahama and Tortugas Islands, from the Gulf of Mexico,

and from the coast of South America.

No. 437] NORTH AMERICAN INVERTEBRATES. 345

Cassiopea frondosa Lamarck. 1817.

Polyclonia frondosa L. Ag. Cont. Nat. Hist. U. S. 1862.

Umbrella disk-like, arched, about three times as broad as high, with 12

distinct, broad, ocular radial stripes. Margin with 12 broad velar lobes.

Manubrium approximately as long as the bell-radius, very s pinnz of

mouth arms variously parted and distally plumose or frondos

Color.— Bluish to olive-green; arms greenish or sees with whitish

terminal filaments.

Distribution.— Coast of Florida, Tortugas Islands, etc.

pees meleagris L. Ag. 1862.

Nat. Hist. U S.

Umbrella high, arched, more than hemispherical, with 8 deep ocular

incisions, and with 96 marginal lappets.

. Color—Whitish-blue, the margins becoming yellowish-brown, margin

lappets dark-brown.

Szze.— About five inches broad by about three inches high.

Distribution.— Southern Atlantic coast, Savannah, Charleston, etc.

SYRACUSE UNIVERSITY,

The Zoólogical Laboratory,

Feb. 10, 1903.

NOTES AND LITERATURE

GENERAL BIOLOGY.

Development and Evolution.!— In this work, which is a collection

of short papers originally published in various magazines, the author

attempts to apply the biogenetic as contrasted with the psychogenetic

method to the facts of ontogeny and phylogeny with the purpose of

determining “which sort of a theory of biological evolution” is most

satisfactory. As a psychologist Professor Baldwin tends to empha-

size the importance of the psychic in evolution and development ;

his theory is consequently psychophysical, not vitalistic.

Of the three parts of the book the first deals with problems of

genesis, the second with the method of evolution, and the third with

bris psychological facts and philosophical problems.

It has been the psycho-physical, not the physical alone, which

has been the unit of selection in the main trend of evolution" says

the author. In support of this statement he presents facts of social

transmission, and individual intelligent accommodation to conditions.

Upon the plasticity of the organism, its imitativeness, and its ability

to make intelligent adjustments Professor Baldwin lays great stress.

In fact it sometimes seems as if he might explain everything by sim-

ple imitation,

The whole work, in so faras it can be unified for purposes of

brief description, is a statement of the author's theory of “ Ortho-

plasy " in connection with a marshaling of the evidences of organic

selection. In contrast with natural selection, which is usually thought

of as due to the destruction of the unfit, organic selection is essen-

tially due to the fact that individual accommodations keep certain

individuals alive, and thus permit of that accumulation of variations

which determines the direction of evolution in later generations. If

such a process as that of organic selection is occurring it is obvious

that the assumption of the inheritance of acquired characters, in the

usual sense, is unnecessary. That use-inheritance is not a factor in

Including Psychophy-

1 Baldwin, James Mark. Development and Evolution.

ory of Genetic Modes.

sical Evolution, Evolution by Orthoplasy, and the The

New York. Macmillan, 1902. 8vo. xvi+ 395 pp.

347

348 THE AMERICAN NATURALIST. [Vor. XXXVII.

the evolution process almost every paragraph of “ Development and

Evolution " serves to show.

Intelligence is represented as the highest form of the process of

“accommodation,” for with it comes adaptiveness, educability, and

the ability to profit by social tradition.

'The book, although bad in form, contains much valuable material.

One can but feel that the author might well have taken the trouble

to carefully rewrite it in a systematic and logical fashion instead of

merely throwing together a lot of fragmentary discussions, without

any attempt at the avoidance of repetition. Professor Baldwin

evidently likes to make his readers work. R. M. V.

Biological Laboratory Methods.'— According to the author's

introduction, Biological Laboratory Methods is a book intended to

meet a demand “for suitable text-books which will give full and clear

instructions concerning the use of the microscope and the other

instruments and methods required in these [biological] laboratories."

Furthermore, Dr. Mell says, such a book * should begin at the begin-

ning and treat of all matter relating to the subject in simple lan-

guage...." An examination of Dr. Mell's book reveals an unfor-

tunate discrepancy between aim and achievement. The book begins

anywhere but at the beginning, while fullness, clearness and simplic-

ity of language are qualities which are not everywhere evident. Ina

book of 321 pages intended for *the beginner " in biological work, 44

pages are given to the microscope and 66 pages are devoted to chap-

ters on “Preparation of the Tissue for Mounting," *Imbedding

Methods,” “ Stains, Their Preparation and Use," and “ Mounting

the Tissue for Preservation," while 84 pages are given to photogra-

phy and bacteriological methods. A description of Born's recon-

struction method and “ Methods for the Preservation of Marine

Organisms” occur under the heading “Maceration.” Essential

details, particularly in the description of the microscope, are lacking,

while the non-essential is everywhere present to the inevitable confu-

sion of an inexperienced student. Under *Imbedding Methods,"

directions for imbedding in paraffin having been given (with the

warning that *Prolonged heating at any time is injurious to many

forms of vegetation"), the student is told that the mass should be

“shaped into a rectangular form, so that when placed in the micro-

tome one face of the mass will be square with the knife and the

1Mell, P. H. Biological Laboratory Methods. New York. Macmillan, 1902.

8vo. xii + 321 pp., 127 figures.

No 437] NOTES AND LITERATURE. 349

opposite edge parallel with it." With these directions as an example

of clearness, the accounts of numerical aperture and polarized light

may best be imagined. Students will find the *too voluminous"

“works of reference” of Carpenter, Gage and Lee not rendered less

useful by the appearance of “ Biological Laboratory Methods."

LWR

ZOOLOGY.

Sexual Dimorphism.— Sexual dimorphism among animals and

the evolution of secondary sexual characters form the subject matter

of an interesting volume by Cunningham.’ The author points out

the inadequacy of natural selection asan explanation of the very con-

stant characters upon which animal classification is based in that it

must be admitted that many of these characters are of no obvious

advantage to their possessors. He next turns his attention to sec-

ondary sexual characters and claims that here too that special form

of selection called by Darwin sexual selection is ineffective because

again the differences are not of a kind to afford a basis for the

selective process. In his opinion the origin of these characters has

been due to Lamarckian factors. It must be admitted that the

influence of the environment profoundly changes animals. Those

changes that occur at the breeding season are dependent upon the

changes of habits characteristic of that period. They consequently

form the basis for the evolution of secondary sexual characters.

That these changes may be inherited is well known, hence we should

not deny that they are examples of the inheritance of acquired charac-

ters because we are ignorant of the method by which their inheritance

is accomplished. From this standpoint the author reviews a large

range of secondary sexual characters from the mammals to the low-

est metazoa in which such characters are known. While the line of

argument will probably not be convincing to even the milder Weis-

mannians, the wide range of illustrations brought together by the

author will afford interesting reading to every zoólogist.

1Cunningham, J. T. Sexual Dimorphism in the Animal Kingdom. London.

Black, 1900. Svo, xii + 317 pp., figs.

350 THE AMERICAN NATURALIST. [Vor. XXXVII.

Parasites and Geographical Distribution. — The value of para-

sites of different animals for the investigation of the geographical

distribution of the latter has been pointed out recently by H. von

Ihering.! The object of modern zoogeographical research is chiefly

to trace the origin of the different forms of life, and, with respect to

the fauna of a limited section of the earth’s surface, it is important

to settle the question whether the inhabitants originated there, or

whether they immigrated from other parts, and, in the latter case,

whence they came.

Von Ihering studies the present South American fauna from this

point of view, and points out that, among the fauna of this continent,

we can distinguish two chief elements: the one is peculiar to it, that

is to say, was present there before the second half of the Tertiary,

while the other immigrated from the North, after the Miocene.

Then he proceeds to demonstrate that the parasitic worms found in

these two groups of animals exhibit peculiar differences, so that it is

possible, under certain circumstances, to draw the opposite conclusion

that the parasites of a certain species of animal indicate, whether the

latter belongs originally to South America, or whether it immigrated

in the later Tertiary.

The instances quoted are taken chiefly from among Mammals and

Birds, but it is evident that also other groups may furnish examples.

In conclusion, von Ihering condenses his results in three funda-

mental “biological laws,” which we reproduce here, freely translated :

(1) Land animals, even if they migrate over a large extent of

territory do not lose the parasitic worms peculiar to them because

the lower animals which serve as intermediate hosts offer everywhere

analogous conditions, provided everything else remains unchanged.

Although, in new areas of distribution, some new parasites may be

added, the old conditions largely remain unchanged, which is very

evident in South America, where the parasitic worms of the holarctic

region are not found with the indigenous (autochthon) mammals or

birds, but only with the strangers (heterochthon) that immigrated at

a late period.

(2) Under these circumstances, helminthology becomes a valu-

able aid for the analytic method of zoogeography, and we may con-

fidently obtain by it important results as to the history of such groups

in which we do not possess satisfactory geological material, or in

which such material naturally cannot be expected.

hering, H. von. Die Helminthen als Hilfsmittel der zoogeographischen For-

schung, Zoolog. Anzeig. Bd. 26, 1902, pp. 42-51.

No 437] NOTES AND LITERATURE. 351

(3) Helminthology treated in this way, may also aid paleonto-

logical research, since the relations between parasites and hosts, and

their migrations and geological age, permit conclusions to be drawn

as to the age of the single larger groups (of the parasites) and even

of their genera and species. A. E.

Two papers on the Nautilus.!— These two works which appeared

nearly simultaneously form the most important contributions to our

knowledge of the tetrabranch cephalopods which have appeared for

years. Dr. Griffin had for his material numerous specimens collected

by the Menage expedition of the Minnesota Academy of Sciences,

while Dr. Willey collected his among the islands of the Eastern

Archipelago where he went in the hopes of obtaining the embryology

of this most interesting animal.

Neither of the papers — which extend over 95 and 91 pages and

are illustrated by several text figures and 17 and 9 plates respectively —

can be summarized here. The two, to a great extent, supplement each

other. Dr. Griffin has endeavored to give a connected account of the

anatomy, utilizing not only his own dissections but the accounts of

his predecessors and hence gives a wealth of detail. Willey on the

other hand describes rather what he himself has investigated and his

comparisons are those of the broader morphological treatment. A

single example will illustrate the different points of view. In treating

of the digital tentacles Willey gives comparatively little about the

anatomical structure but tries to work out a numerical nomenclature

of these parts, in which he comes to results widely at variance with the

previous studies of Vayssitre. Griffin, on the other hand describes

the anatomy in great detail, but says nothing regarding the arrange-

ment, although he knows of Vayssiere's work. He gives however a

plan of their position which differs in some respect from that of

Willey. Willey further enters with the question whether these tentacles

are to be compared to the arms of the dibranch cephalopods or to the

acetabula as has been suggested, inclining to the former view.

The sections relating to the foot in Dr. Willey's paper are of

interest. Accepting Grenacher division of the molluscan foot into a

median protopodium and lateral epipodia and discussing change of

function and its relations to change of organs and to topography he

argues for the conclusion that the siphon represents the protopodium

rifin, L. E. The Anatomy of Nautilus pompilius, Memoirs National Acad.

Science, viii, 1900 (1902). — Willey's Contribution to the Natural History of the

Pearly Nautilus, in his Zoi/ogzca/ Researches, part vi, August, 1902.

352 THE AMERICAN NATURALIST. [Vor. XXXVII.

of the gasteropod and the tentacles the epipodia. He also claims

that the post-anal papilla are osphradial in nature and are therefore

an additional evidence for metamerism in these forms, while on the

other hand he fails to find any metamerism in the ccelom.

Besides dealing with Nautilus Willey presents numerous other

facts in this number of his * Results" among them many details as

to the anthropology of the regions visited, and notes on tunicata,

Amphioxus and Enteropneusta. K

BOTANY.

Recent Literature on Seedlings.— In Zorreya (Vol. II, pp. 113-

117, August, 1902) Lloyd discusses “ Vivipary in Podocarpus." Dur-

ing the last winter a specimen of Podocarpus ma£ayibore an excellent

crop of fruit which germinated almost without exception and on the

parent plant. The hypocotyl, he finds, is, when developed, of that

club shape characteristic of certain other viviparous plants, as the

mangroves, and is rich in food material, especially starch, which

seems to be derived not only from the endosperm, but from its own

photosynthetic activity as well as may be inferred from its greater

weight, green color, and the presence of stomata. In most cases

the primary root does not develop, but its place is taken by one

or usually two lateral roots formed near the end of the hypocotl.

He calls attention to vivipary in Ae/ecamma bambusoides and its

possible existence in other grasses, in Zi//andsia albisiana and in

Quercus fusiformis as well as interesting similarities in other oaks,

and concludes that vivipary is by no means the unusual condition it

is supposed to be. F.W. Rane (“ How to grow a Forest from Seed.”

Bull. N. H. Ag. Exp. Sta. 95, November, 1902), figures a few tree

seedlings though the bulk of the paper is naturally of an economic

nature. In Aroc. Cambridge Philosophical Soc., Vol. XL, pp. 445-

457, Pl. 5, 1902, Gardiner and Hill consider the histology of the

Endosperm during the germination of Tamus communis and Galium

Tricorne. Chauveaud in Bull, Mus. d Hist. Nat., 1902, No. 1, pp. 52-

59, discusses the arrangement of the vascular system in the cotyledon

of the onion, A//ium cepa.

NO. 437.] NOTES AND LITERATURE. 353

Cyril Crossland (“ Note on the Dispersal of Mangrove Seedlings,”

Ann. of Bot., Vol. XVII, pp. 267—270, fig. January, 1903) observed

mangroves growing in large numbers in the crevices in the hard coral

limestone surface near high water mark on the east coast of Zanzibar,

but only occasionally found them growing in mud, where the well-

known method of planting may be observed. He frequently found

embryos planted in holes in the rock at a distance of a hundred yards,

and in a few cases some miles from the nearest parent tree. He

frequently found the embryos floating vertically in the sea with the

leaf bud just projecting above the water and concludes that the embryo

is planted in any softness or crevice of the bottom upon the falling of

the tide. The observation adds a quite distinct method of distribu-

tion for this interesting plant.

A contribution of importance to our knowledge of seedlings is that

of Willis in his paper on the * Morphology and Ecology of the Podo-

stemacez.”! [nan earlier number of the same publication Mr. Willis

treated the systematic relations of the forms found in India and

Ceylon and in the present paper for which the other was preparatory

he discusses the anatomy and ecology of the different forms and

when material was available the developmental stages as well The

peculiar ecological conditions under which the members of the

family are to be found are discussed, and in addition to the descrip-

tions and figures of the young stages of several forms given in the

systematically arranged portion of the paper some theoretical consid-

erations are taken up in his quite extensive general discussion and

summary.

The germination of the seed of Peperomia and Heckeria has been

studied by Johnson (Bot. Gaz., Vol. XXXIV, pp. 321—340, Pls. IX and

X. 1902). He finds that the swelling of the embryo and endosperm

bursts the seed coats and that the endosperm protrudes through the

rent as a sack, which surrounds the small, undifferentiated embryo

until cotylelons and root have been developed when the root breaks

through the endosperm which still surrounds the tips of the cotyle-

dons and remains imbedded in the seed till all the starch of the

perisperm is absorbed. While only careful chemical work can yield

a definite answer to the question the morphological features indicate

that the aleurone containing endosperm of these forms serves not as

a storage organ for food material, but as a digesting and absorbing

1 Willis, J. C. Studies in the Morphology and Ecology of the Podostemacee of

Ceylon and India. Ann. Roy. Bot. Gard. Peradeniya. Vol. i, pp. 267-465, Pls.

IV-XXXVII. 1902.

354 THE AMERICAN NATURALIST. [Vor. XXXVII.

apparatus for transferring the reserve starch of the perisperm to the

embryo. He suggests that in several genera of Cannacez, Poly-

gonacez, Phytolaccaocez, Caryophyllacez, and others, a thin layer

of endosperm separating perisperm and embryo may serve the same

function. J. A. Harris.

The Origin of Monocotyledons.— The importance of a study of

the seedling stages of plants in classification is being much empha-

sized of late. In a recent number of this Journal, Professor Campbell

(Am. Nat., Vol. XXXVI, pp. 7-12, January, 1902) touches on this

question. In a more recent number (dm. Nat., Vol. XXXVI, pp.

981-982, December, 1902) was reviewed the preliminary paper of

Miss Sargent in which she announced a theory of the origin of the

Monocotyledons from a dicotyledonous type. In the following

number of the Mew Phylologist, (Vol. I, pp. 131-133, June, 1902)

Tansley commends very highly in some ways the paper by Miss

Sargent, but calls attention to the freedom with which the hypothesis

of the derivation of a simple structure from a more complex one is

nowadays used in morphological work, and to the danger of the too

free use of this hypothesis in biological speculations, unless the

special conditions determining the reduction are to be ascertained,

since there seems to be no reason for believing that there is any

general cause leading to reduction as compared with the primary

tendency to increase in bulk and complexity of structure. He thinks

that, while of the greatest interest in many ways, the generalized

conclusions of Miss Sargent's paper should not be accepted without

careful consideration.

In the current number of the Annals of Botany, Miss Sargent!

presents in greater detail the theory recently announced, though she

states that the evidence she has accumulated will not be published in

detail until her monograph on the comparative anatomy of seedlings

of the Liliaceae is completed. In the opening pages she discusses

the nature of the evidence employed in formulating her theory,

This is followed by observations on the anatomy of seedlings,

occupying nearly sixty pages, and considering the tribes Scillez,

Tulipez, Asphodelez. Alliew, Dracanee, Asparagee and Aloi-

nez of the Liliacez, to which her work has been principally con-

fined, with fewer examples from the Amaryllidacee, Iridacez,

1 Sargent, E. A Theory of the Origin of the Monocotyledons, founded on the

Structure of their Seedlings, 4». of Bot., Vol. xvii, pp. 1-92, Pl. I-VII, Jan-

uary, 1903

No 437.] NOTES AND LITERATURE. 355

Aroidez, Palmez and Scitaminez, of the monocotyledons and the

Ranunculacez of the dicotyledons. In the third part she takes up

general considerations on the origin of the monocotyledons.

In the space of a review, it is necessarily impossible to state in

detail the evidence presented, but some of the main points of argu-

ment may be given. As in her previous paper she affirms her belief

in the real systematic value of some of the vascular characters of the

young seedling —at least of the Liliaceæ— and emphasizes the

structural similarity of some of the Ranunculaceous seedlings to those

of certain forms which she concludes represent the primative type of

vascular arrangement in the Liliaceæ. She expresses her belief in

the genetic connection of Eranthis and Anemarrhena, but even if

there be no historical connection, she maintains that the structure of

Eranthis may illustrate the double origin of the Anemarrhena coty-

ledon. Two tables are given, one listing dicotylodenous seedlings

with a well-marked cotyledonary tube and the others those in which

the union of the cotyledons occurs along one margin only. The

ecological relations of these seedlings are discussed in relation to

their bearing upon the development of a permanent monocotyled-

onous type, and it is found that almost all belong to plants of

geophilous habit and that in some cases the whole structure remains

under ground during the first year of their growth, developing under-

ground organs for the tiding over of the unfavorable vegetative period

to follow. Such considerations have led her to regard the monocoty-

ledon as an organism adapted primarily to a geophilous habit, and

she thinks that when considered from this point of view many

puzzling details of structure in the monocotyledons become compre-

hensible. A brief discussion of some of these points is given. A

bibliography of forty-five titles lists the literature to which reference

is made.

While the presentation of the theory is confident, it is not without

due reserve. “The evidence is obviously incomplete. The theory

itself cannot be considered as proved in any sense. It is brought

forward as a working hypothesis which I have found in practice to

be suggestive and illuminating.” The favorable tone of the present

review may be attributed to the desire of the reviewer to present the

theory from the point of view of the one who proposes it, but no one

will deny that the data presented in the paper is of the greatest value

and that the theoretical considerations will receive the careful attention

of botanists and be of importance in the elucidation of the great

problem, to the solution of which the paper is an important contri-

bution.

356 THE AMERICAN NATURALIST. (VoL: XXXVII.

A recent paper by Mr. Lyon,’ whose work on the embryogeny of

Nelumbium has been a stimulus to work of this nature, has apparently

not come to the notice of Miss Sargent. This paper considers

primarily not the origin of the monocotyledons and dicotyledons but

the origin of the cotyledon itself. His conclusions are of suc

interest, and part of them so directly connected with the questions

with which Miss Sargent’s paper is concerned that it seems well to

state them briefly. Of the three elements, cotyledon, stem and root

of the typical embryo of the pteridophytes and angiosperms, the

cotyledons do not represent modified leaves, but are primarily haus-

torial organs, originating phylogenetically as the nursing foot in the

Bryophytes and persisting in the higher plants. Thus the monocotyle-

donous condition occurring in the bryophytes, pteredophytes, and

Monocotyledons is to be regarded as the primitive one while the

two or sometimes more cotyledons of the dicotyledons are the

homologues of the single cotyledon of the monocotyledons.

J. A. HARRIS.

French Forestry.”

French and American foresters have to work, Professor Mouillefert’s

Traité de Sylviculture, the first part of which has just appeared, is well

calculated to be of much service even in this country. The remain-

ing three volumes of the series are in press and are promised for the

current year.

The bulk of the present volume is devoted to detailed accounts of

the principal species both native and foreign with which French for-

estry is concerned. Each is described and illustrated by remarkably

good figures showing commonly the winter habit, twigs with buds,

the leaves, inflorescence, flowers, fruit, seeds, germination and wood.

Reference is made to the geographical distribution, the tree's prefer-

ence as regards soil and climate, and the best methods of propagating

and tending. Considerable attention is given to the wood, especially

as regards distinctive structural features and economic uses. ‘The

uses of other parts is also given, as well asan account of the general

economic importance of the tree, exhibited so far as possible by sta-

tistics. Finally, its more important insect enemies and vegetable

parasites are mentioned and the nature and extent of the injuries they

inflict briefly indicated.

‘Lyon, H. E. The Phylogeny of the Cotyledon, Pos/e/sia, Vol. i, pp. 57-86,

y yogeny y 5

1902.

2 Mouillefert, P. Principales Essence Foresti?res préceedées de notions de statisque

forestières. Paris, Felix Alcan. 1903. 12mo, xii + 545 pp., 630 figs.

No. 437.] NOTES AND LITERATURE. 357

Near the end of the book is a synoptical table or key to the prin-

cipal woods distinguished by features visible to the naked eye or

under a magnification of about 20 diameters.

A preliminary part of about 40 pages states clearly and concisely

important conclusions drawn from French forestry statistics, modern

views of the influence of forests on water supply and climate, and

the influences of climate, soil, forest covering, commercial and eco-

nomic considerations, etc., on forest production. ‘The volume closes

with a full index, thus making this part of the treatise complete in

itself.

Kraemer's Course in Botany and Pharmacognosy ! presents in

convenient form a good general account of the minute and the gross

anatomy of vegetable drugs and their microchemistry. Following the

general morphological part are chapters devoted to the description

and discrimination of drugs in the crude state and in powder. In

this part excellent keys are included by means of which the student

is helped to recognize any official drug. A third part deals briefly

with the most useful reagents required in pharmacognosy, and with

simple methods of making microscopic preparations. There are 17

plates containing 128 figures, 6 of which are colored. All are clearly

drawn and well printed on clayed paper. Besides a full general

index there is a special index to powdered drugs.

The book is remarkably well calculated to give students of phar-

macy all the botany they need in preparing for their profession. The

style is unusually clear and direct, and an orderly comprehension of

the more difficult topics is much facilitated by the use of tables. In

preparing this work Professor Kraemer has done a good service to

many students and teachers.

F LS

Notes. — Zhe Journal of the New York Botanical Garden, for

February, contains information concerning the research scholarship

recently established at that Institution; an interesting account by

Mrs. Vail of Jonas Bronck and his Bouwery in New Amsterdam, and

some chemical studies of Sarracenia purpurea, by Gies.

“Why Popcorn pops” is the subject of an article by Wilbert in the

American Journal of Pharmacy for February.

1Kraemer, Henry. A Course in Botany and Pharmacognosy. Philadelphia,

1902. 12mo. 384 pp. 128 figs.

358 THE AMERICAN NATURALIST. [Vor. XXXVII.

Professor Spalding’s presidential address before the Society for

Plant Morphology and Physiology, dealing with the rise and progress

of ecology, is published in Sczence of February 6th.

An address before the Western Railway Club on Timber Preser-

vation, well illustrated with figures of botanical interest, has been

described by Dr. von Schrenk.

An account of silkworm food plants, well illustrated, has been pub-

lished by Oliver as Bulletin 34 of the Bureau of Plant Industry of the

Department of Agriculture.

An illustrated account of plants injurious to sheep, based on a

Bulletin of the Nevada Experiment Station, is contained in Zhe

Pacific Rural Press of February 7th.

An article on electromotive force in plants, by A. B. Plowman, is

published in the American Journal of Science for February.

No. 18 of Dr. Holm’s Studies in the Cyperaceae, dealing with Carex

Jusca and C. bipartita, is published in the American Journal of Science

for February.

Among other interesting botanical articles in Volume XXXV of

the Journal and Proceedings of the Royal Society of New South Wales

is one on the relation between leaf venation and the presence of cer-

tain chemical constituents in the oils of the Eucalypts, by R. T. Baker

and H. G. Smith.

A conspectus of the flora of Greece, by E. De Halacsy, — in the

form of a two-volume octavo, — has recently been issued from the

Engelmann press of Leipsic.

A good plate of detailed figures of Prunus besseyi is contained in

Volume III, Fascicle 7, of the /cones Selecte Horti Thenensis.

Under the editorship of Professor Sargent a new serial entitled

Trees and Shrubs, devoted to illustrations of new or little known

ligneous plants, prepared chiefly from material at the Arnold Arbore-

tum of Harvard University, has been launched from the press of

Houghton, Mifflin and Company. The first part, issued November

26, 1902, contains in addition to other things a considerable number

of new species of Crataegus not included in the Silva, and a new genus

of Scrophulariacee Faxonanthus, with a single species, F. pringlei

Greenm. In form and typography Zrees and Shrubs agrees with

Professor Sargent’s Siwa of North America, and the plates are as in

No. 437.] NOTES AND LITERATURE. 359

that work from drawings by Mr. Faxon, which, however, are photo-

mechanically reproduced.

The Plant World for February contains the following articles : —

Safford, “ Extracts from the Note-book of a Naturalist on the Island

of Guam,— III.” ; Hastings, “ Notes on the Flora of Central Chile” ;

Rowlee, “Conditions of Plant Growth on the Isle of Pines”; and

Maxon, “A Botanist's Mecca [Chittenango Falls, N. Y.]."

The Bulletin of the Southern California Academy of Sciences, of

January rst, contains the conclusion of the botanical portion of Dr.

Yates’ “ Prehistoric California,” an article by S. B. Parish on certain

California trees, and a second addition to the flora of Los Angeles

County, by Abrams.

The American Botanist, (a popular journal) for gees contains

the following articles: —W. W. Bailey, “ Bar Field,

“Cereus giganteus”; W. A. Terry, “ Partridge Berries and Winter-

green Berries”; and, W. N. Clute, * The Scouring Rush in Winter."

A revision of the described North American species of Leptochloa,

by Hitchcock, constitutes Bulletin 33 of the Bureau of Plant Industry

of the Department of Agriculture.

Data on the rapidity of growth of Populus grandidentata are given

in Forest Leaves for February.

The appearance of Volume II, Fascicle 3, of Coste’s Flore descrip-

tive et illustrée de la France, etc., gives occasion to once more call

attention to an admirably executed flora, with thumbnail habit and

detail illustrations of all of the species included.

CORRESPONDENCE.

To the Editor of the American Naturalist:

Str: Dr. T. D. A. Cockerell calls my attention to the fact that two

generic names of fishes used in Jordan & Evermann's Fishes of North

America have been earlier used in a generic sense.

The first is Falcula Jordan & Snyder, Zw. U. S. Fish Comm. for

1899, p. 124 (1900) type F. chapale. For this genus of Mexican

Cyprinidz, we suggest the new name of Falcularius Jordan & Snyder.

Falcula Conrad, Amer. Journ. Conch. VI, p. 77, is an earlier genus of

mollusks.

The second is Xenochirus Gilbert; Proc. U. S. Nat. Mus. XIII,

1890, p. 9o, a genus of Agonidze. The same name, Xenochirus,

was used by Gloger in 1842, for a genus of Mammals.

As a substitute for Xenochirus, Dr. Gilbert proposes the new name,

Xeneretmus Gilbert. The type of Xeneretmus is Xenochirus tria-

canthus Gilbert. There is also a genus Xenochira (Huswell, 1879)

but that name being spelled differently from Xenochirus, is suff-

ciently distinct.

D. S. JORDAN.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS AND DEATHS.

EDUCATIONAL GIFTS.

Atlantic City, N. J., $60,000 for a library from Andrew Carnegie.

Barnard College, $1,000,000, from an anonymous donor for the purchase of

additional land.

Bates College, $10,000 by the will of Ario Wentworth.

Boston Society of Natural History, $20,000 from the estate of the he:

R. C. Billings.

Brooklyn Institute of Arts and Sciences, $50,000 from Robert E. Woodward.

Brown University, $5000 from Edgar L. Marston for a scholarship.

Colby College (Maine), $5000 by the will of Robert O. Fuller.

olgate University, $100,000 from Jas. B. Colgate.

Columbia University, $10,000 for a scholarship by the will of Mrs. E. J.

Bowker; $100,000 from the Duke of Loubat for a chair of American

Archeology.

Cornell University, $150,000 from an anonymous donor for a pension fund.

College of Physicians in Philadelphia, a conditional gift of $50,000 from

Andrew Carnegie; $10,000 from F. W. Vanderbilt and $5000 from

Clement A. Griscom for library purpo

Dartmouth College, $5000 by the will of Poles Sylvester Waterhouse.

Davenport Academy of Science, the estate of the late Mrs. Chas. E. Putnam.

including $24,000 for a publication fund.

Denison College (Ohio), a conditional gift of $60,000 from John D. Rocke-

eller. .

Harvard University, $50,000 by the will of Rebecca C. Ames; $5000 by the

will of Professor Sylvester Waterhouse ; $ 5o09 from Mrs. John Markoe

for a scholarship. i

Hobart College, $5000 from Mrs. Vail.

Johns Hopkins University, $5000 from R. B. Keyser for plans for improv-

ing the new site of the Universi

Massachusetts Institute of Technology, $100,000 by the will of Ario Went-

worth.

Oberlin College, $50,000 from an anonymous donor.

Rochester Atheneum and Mechanics Institute, $50,000 by the will of Mrs.

Susan Brevies.

Rockefeller Institute for Medical Research, $1,000,000 from John D. Rocke-

feller for land and buildings.

361

362 THE AMERICAN NATURALIST. [Vor. XXXVII.

Vassar College $8,000 by the will of Dr. Elizabeth L. McMahon.

Washington University (St. Louis) $25,000 by the will of Professor Sylves-

ter Waterhouse.

Western Reserve University, $100,000 from Andrew Carnegie for a library

training school.

APPOINTMENTS.

‘Elmer D. Ball, professor of zoóbiology in the Utah Agricultural Col-

lege.— Dr. Joseph Barrell, assistant professor of geology in Yale Univer-

sity.— Dr. Richard Beck, professor extraordinary of economic entomology

in the veterinary school at Tharandt, Germany.— Dr. Maurice A. Bigelow,

adjunct professor of biology in Teachers College, New York City.— Dr.

Pierro Marcellin Boule, professor of paleontology in the Paris Museum of

Natural History.— Dr. Hermann Braun, professor of zoólogy in the veteri-

nary school at Tharandt, Germany.— Dr. Cavallie, professor of anatom

in the school of medicine. at Clermont-Ferrand. — Stewart Culin, curator

of ethnology in the museum of the Brooklyn Institute of Arts and Sciences.

— Dr. D. J. Cunningham of Dublin, professor of anatomy in the University

of Edinburgh.— Bruce Fink, professor of botany in Iowa College.— Dr.

Eugen von Daday, professor of zoólogy in the Hungarian Polytechnicium

at Budapest.— Joseph Burtt Davy of California, state agrostologist an

botanist to the Department of Agriculture in the Transvaal.— Dr. Giovanni

Battista DeToni, professor of botany in the University of Modena.— Dr.

James J. Dobbie, director of the Museum of Science and Art, Edinburgh.

— Dr. Livingston Farrand, professor of anthropology in Columbia Univer-

sity and assistant curator of ethnology in the American Museum of Natural

History, New York.— Dr. Otto Frank, professor extraordinary of physi-

ology in the University at Munich.— Dr. Friedrich Richard Fuchs, docent for

physiology in the University at Erlangen.— Dr. Frederick DeForest Heald,

adjunct professor of plant physiology and bacteriology in the Univer-

sity of Nebraska.— Dr. Hóflich, teacher of agricultural bacteriology and

anatomy of domestic animals in the agricultural school at Weihenstephan.

— Dr. F. von Huene, docent for geology and paleontology in the Univer-

sity at Tübingen.— C. W. Johnson, curator of the Boston Society of Natu-

ral History.— Prof. Franz Loewinson-Lessing, of Dorpat, professor of miner-

alogy and geology in the Polytechnic Institute at St. Petersburg.— Dr.

Lubosch, docent for anatomy in the University at Jena.— Dr. Otto Maas,

professor extraordinary of zoólogy in the University of Munich.— Margaret

tby, adjunct professor of botany in Teachers College, New York.—

Dr. Benjamin L. Miller, associate in geology in Bryn Mawr College.— Dr.

Muth, docent for botany in the Karlsruhe technical school.— Dr. Friedrich

Oltmauns, professor of botany in the University at Freiburg i. B.— Dr.

. Ortmann, of Princeton, curator of invertebrate zoólogy in the Carnegie

Mosea, Pittsburg— Dr. K. Alfred Osann, professor extraordinary of

No. 437.] GIFTS, APPOINTMENTS, RETIREMENTS. 363

mineralogy in the University at Freiburg i. B— Edmond Perrier, professor

of comparative anatomy in the Paris Natural History Museum.— Dr.

Porier, professor of anatomy in the medical faculty of the University of

Paris.— Dr. W. H. C. Redeke, director of the zoólogical station at The

Helder, Holland.— Mr. Gragg Richards, assistant in geology in the Massa-

chusetts Institute of Technology.— Dr. H. M. Richards, adjunct professor

of botany in Teachers College, N. Y.— Dr. Auguste Roude, assistant

professor of anatomy in the University at Lausanne.— E. Dwight Sanderson,

professor of entomology in the Texas Agricultural College.— M.

Saville, professor of American archeology in Columbia University.— Dr.

Arnold Spuler, professor extraordinary of anatomy in the University at

Erlangen.— Walter Stahlberg, custodian of the museum of Oceanography

in Berlin. — Dr. Fred. Wilhelm Richard Thomé of Jena, docent for anatomy

in the University at Strassburg.— Dr. George Tischler, docent for botany

in the University at Heidelberg.— Professor William Morton Wheeler,

curator of invertebrate zoólogy in the American Museum of Natural His-

tory, New York.

RETIREMENTS.

Sir Michael Foster, from the chair of physiology in the University of

Brunn Technical School.— Dr. Felix Kreutz, from the professorship of min-

eralogy in the University of Cracow.

DEATHS.

A. H. Chester, professor of chemistry and mineralogy in Rutgers

College, April 12, aged 60.— Francois Desbois, student of orchids, in

Brussels, Sept. 14, aged 75. — Dr. Hermann Dingler, professor of botany in

the forestry school at Aschaffenburg, aged 55.— Major Alfred Fichlin, ento-

mologist, in London.— Josef F. Freyn, student of plant geography at

Smichow, near Prag.— Herbert D. Geldart, botanist, at Thorpe Hamlet near

Norwich, England.— Dr. Franz Graeff, professor of mineralogy in the Uni-

versity at Freiburg i. B., aged 47.— Dr. Alexander W. M. van Hasselt,

entomologist, in Amsterdam aged 88.— Dr. Paul An bape of

mineralogy in the Faculty of Sciences in Paris.—

Schafer, botanist, at Ratisbon, January 21.— Charles 5 tpe sind

gist, in Ilkley, near Leeds, England.— E. A. Hudak, ioloptétulgist, in

Gólwicz-bayna, Hun .— Max Kossmann, coleopterologist, in December.—

Dr. Leonhard Landois, professor of physiology in the University at Greifs-

wald, aged 65.— Dr. Adrien Lemaire, student of diatoms and vegetable

anatomy, in Nancy, Oct. 23. Orga Lemberg, professor emeritus of min-

eralogy, in Dorpat Nov. 20, aged 6o.— Gustav Limpricht, bryologist, in

Breslau, Oct. 20, aged 68.— pa Lohde, student of coleoptera, in Ber-

364 THE AMERICAN NATURALIST. | [Vor. XXXVII.

lin, January 6, aged 21.— Dr. Ernest Mehnert, professor of anatomy at the

University at Halle, aged 65.— Louis Montillot, entomologist, in Montrouge,

France, in December.— Dr. Heinrich Nitsche, professor of zodlogy in the

Forestry School at Tharandt, Nov. 8.— Dr. Julius Pethó, chief of the Hun-

garian Geological Survey, at Budapest, Oct. 14, aged 55.— Dr. Wilhelm

Pfitzner, professor of anatomy at the University at Strassburg, January 1,

aged 49.— Luigi Pozzi, entomologist, in Modena, April 1, 1902.— Mrs.

Charles E. Putnam, president of the Davenport Academy of Science, Feb-

ruary 20, aged 70 years.— Gustav Raddi, director of the museum at Tiflis,

Caucasus.— Oscar Lamarche de Rossius, lepidopterologist, in Hamoir,

France, Sept. 7, aged 66.— Michael Schieferer, student of the leaf-mining

Lepidoptera, at Graz, Styria, Mar. 31, 1902, aged 74.— S. Sirodot, professor

of botany at Rennes.— Robert A. Sterndale, author of a work on Mammals

of India and Ceylon, in St. Helena, Oct. 3.— Dr. A. J. Stuxberg, zoólogist

and intendant of the museum at Góteborg, December 1.— Dr. Pierre Jules

Tosquinet, entomologist, near Brussels, October 28, aged 78.—Dr. Woronine,

professor of botany in the University at St. Petersburg, aged 79.— Dr. T.

Zaaijer, professor of anatomy in the University at Leiden.

(Wo. 436 was mailed May 22nd.)

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VOL. XXXVII, NO. 438 JUNE, 1903

THE

AMERICAN.

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

I. The Colors of Northern Gamopetalous Flowers . . - JOHN H. LOVELL 365 .

II. The Causes of Acceleration and Retardation in the Metamorphosis of Ambly-

stoma tigrinum: a Preliminary Report . . . . -JLE POWERS 385

III. The Originofthe Sporophyte . . - . Dr. BRADLEY MOORE DAVIS 411

IV. Notes and Literature: Zodgeography, The Atlantic Problem; Zoógeograph- 481

ical Development of the Indo-Australian Archipelago — Botany, Influ- - 434

ence of Light and Darkness on Growth and Development, A Popular

Introduction to Forestry, Notes

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ASSOCIATE EDITORS:

. ALLEN, PH.D., American Museum of Natural History, des York.

. ANDREWS, PH.D., ees ape Universi A Baltim

idge. ;

jose ides v Warkocion.

ee Univer.

PHD., Gainey. of California, Berkeley.

, PH. EE Lake Fi orest University.

IANN, Pu.D., Carnegie Museum, Pittsburg.

'W, S.B., F.R.M.S, “McGill University, Montreal.

, S.D., Columbia Univer. ersity, New York.

SSELL, Pu.D., Harvard University, Cambridge.

RU ISSELL, LL.D. — acs quss eui iones

SMITH, S.D., U. S. ce

NEGER, L

THE

AMERICAN NATURALIST.

Vor. XXXVII. June, 1903. No. 438.

THE COLORS OF NORTHERN GAMOPETALOUS

FLOWERS. !

JOHN H. LOVELL.

THe Gamopetalz are characterized by the more or less united

petals. This series is not derived from the Choripetale, but

represents an independent though parallel-line of development.

The union of the petals has not been induced by insects, and

“is not,” says Goebel, “ the result of a cohesion of originally free

parts, but is due to the excessive growth of the zone of the floral

axis in which the rudiments of the petals are inserted." Much

uncertainty prevails as to the phylogeny of the gamopetalous

Series, which can only be settled by further investigation.

There are two points of culmination. By common consent the

Compositze are placed at the head of this series and of the plant

kingdom. In this family it is the community of flowers rather

than the individual flower that is of chief importance. A

Secondary culmination occurs in the Labiatæ and Scrophular-

! The Colors of Northern Monocotyledonous Flowers, Amer. Wat., vol. xxxiii,

P- 493; The Colors of Northern Apetalous Flowers, Amer. Nat., vol. xxxv, p.

197; The Colors of Northern Polypetalous Flowers, Amer. Nat., vol. xxxvi,

P- 203.

365

366 THE AMERICAN NATURALIST. (Now. XXXVII.

iaceze, families in which the flowers are zygomorphous and often

of large size. It has been suggested that these two divisions

represent two distinct lines of development and are of very

different origin. The Composite are derived through the

Rubiaceze and Caprifoliacee from the Umbelliferae, while the

aroids are assigned as the remote ancestors of the labiate forms.

It seems better, however, to regard the series as homogeneous.

The gamopetalous families probably had a common starting

point and are derived from a group of primitive forms allied to

the progenitors of the Choripetalæ, but a later offshoot, and still

retaining affinities with the monocotyledons. The flowers are

characterized by definiteness in the number of parts and by their

frequent cohesion, characters, says Engler, which must have

become fixed at a very early period in their history. The Gamo-

petala are of later occurrence in the fossil state than the

Choripetalz.

The first of the gamopetalous orders, the Ericales, which

includes six families, is the most primitive in structure. The

flowers have the corolla lobed or pointed, or the petals may be

entirely distinct ; the ovary is superior except in the Vaccinia-

cec; there are two typical staminal whorls, and the carpels

equal the petals in number. In the genera Clethra and Pyrola

the petals are but slightly, or not at all united, and in Monotropa

and Ledum are regularly separate. Of the 81 flowers belonging

to these six families 47 are white and 24 are red; while only 1

is yellow, 6 purple, and 3 green.

The Pyrolaceze are low perennials with clustered evergreen

leaves, growing in woodlands and swamps. The flowers are

racemose and 7 of the species are white, 2 red, r purple and t

green. They vary but little in size. Several of the white

flowered species of Pyrola are tinged with green, while in

Moneses and Chimaphila the white flowers are often pinkish.

The purple Pyrola uliginosa is a bog plant. The flowers of this

family are visited by bees, flies and beetles. The primitive color

was doubtless greenish-white, which has varied into pink and

purple. Flowers of different sizes have been observed on 7.

minor.

The Monotropaceze are saprophytes without chlorophyll, and

No. 438.] NORTHERN GAMOPETALOUS FLOWERS. 367

with the leaves reduced to bracts. The plants which live in rich

. woods are white or colored with anthocyan. J/onotropa uniflora

(Indian pipe or ghost-flower) is usually pure white, and stands

out in strong contrast with the dark brown soil of its woodland

home. Occasionally the whole plant is pink or red. Specimens

in drying turn black. M. Aypopitys has the entire plant pale

yellow, with yellowish or pink flowers. On the island of Wollin

Knuth saw a bumblebee sucking the flowers. The plant of

Monotropsis is pale pinkish brown with fragrant pink flowers.

— The flowers of the Ericacez are produced in great profusion,

and often display bright masses of color, which are visible at a

long distance. This family reaches its highest development in

the temperate zones, the structure and form of the plants

enabling them to endure the severities of alpine summits. The

heaths proper are most abundant at the Cape of Good Hope,

where more than 400 species have been described. Of the 38

northern species 22 are white, 1 yellow, 10 red, and 5 purple.

Some of the handsomest of the wild flowers belong to the

genus Azalea. Of the five northern species two are white, one

orange and two rose color, and all have large flowers with slender

tubes. A. /ufea (A. calendulacea) in spring is covered with

orange flowers which turn to flame color. In the Alps on a hot

June day the hillocks projecting from the surrounding snow are

densely carpeted with the crimson flowers of the smaller and

more primitive A. procumbens over which flit numerous bees and

butterflies. Florists offer hybrids of A. pontica from Asia

Minor, which display a great range of yellow and red coloring.

Closely allied to the Azaleas and equally beautiful are the Rhodo-

dendrons. The flowers are red or purple, or in R. maximum

rose or sometimes white spotted with orange, and are fertilized

by bumblebees. Rhododendron maximum and Kalmia latifolia

have been called the two handsomest North American shrubs.

Bigelow remarks in his Medical Botany that “ these shrubs, in

luxuriant size and vigor, are seen to cover tracts of great extent,

at one season presenting an unbroken landscape of gorgeous

flowers.” Asa Gray in one of his letters speaking of the

glorious view from Roan Mountain, North Carolina, writes,

* The valleys and mountains all around, covered with rich forest,

368 THE AMERICAN NATURALIST. [Vor. XXXVII.

are adorned with Rhododendron maximum and Kalmia latifolia

in immense abundance and profuse blossoming, of every hue

from deep rose to white, and here and there among other shrubs

Azalea calendulacea, of every hue from light yellow to deepest

flame color.” ! The flowers of Kalmia are red or red purple and

are pollinated by bees. In northern New England the com-

monest species of Kalmia is A. angustifolia, which produces its

crimson flowers in late spring. The central flowers of the

corymb are smaller and imperfectly developed; the corolla is

white and without pouches, while the stamens stand erect and

are not exceeded in length by the pistil. Most of the northern

genera of the Ericacez, however, are shrubby plants with small,

white, racemose or clustered, drooping flowers attractive to the

honeybee and bumblebees. Such are Andromeda, Leucothoé,

and Gaultheria procumbens, or the checkerberry. The flowers

of this last species are partially concealed beneath the leaves and

in the grass and moss, amid which the plant grows. Yet I have

found the honeybee and three species of bumblebees very

frequent visitors. The aromatic scented flowers are white with

reddish calyx and peduncles. It would be difficult to find greater

extremes in conspicuousness than is presented by the magnificent

color masses of Rhododendron and Kalmia, and the small con-

cealed flowers of Gaultheria, yet these latter do not want for

pollinators. None of the true heaths (Erica) are endemic in

America, but two species (Æ. cinerea and E. tetralix, adventitious

from Europe) occur in small patches on the Island of Nantucket.

E. carnea unlike most red flowers of this family is adapted to

butterflies. Both the petals and sepals are pink or crimson, and

the peduncles are still more intensely colored. The inverted

flowers were doubtless first adapted to bees, as is usually the

case with campanulate flowers in this position. The entrance to

the corolla has now become so narrowed that only the slender

proboscis of the butterfly is admitted. Miiller saw a bumblebee

in vain striving to suck the honey, In the Alps he found it

frequently visited by Vanessa cardut.

Of the Vacciniacez, or huckleberry family, ro species are

! Letters of Asa Gray, vol. ii, p. 692.

No. 438.] WORTHERN GAMOPETALOUS FLOWERS. 369

white, 11 red, and 2 green. The primitive color of the corolla

seems to have been green, as it is still greenish, greenish-white,

greenish-pink, or yellowish-green in the genus Vaccinium. The

flowers are bell-shaped and the pollinators are bees. V. pennsyt-

vanicum (the common blueberry) is white often tinged with red

on the side exposed to direct sunlight. At the time of blooming

in spring the leaves are reddish, and patches of these bushes on

a hillside are visible at a long distance. The flowers of Gaylus-

sacia resinosa, or the huckleberry, are red, and the entire plant

exhibits a marked tendency to develop red coloration. The

floral bracts and stems are reddish, and the leaves when stung

by a gall-fly turn a deep red, while the brilliant crimson of the

foliage in autumn is unsurpassed. The nodding flowers are

visited, according to my observations, by nine species of bees

besides several butterflies. If the entrance to the corolla, which

is already quite small, should be further narrowed to the exclu-

sion of bees, as has been described in Erica carnea, this species

would become also adapted to butterflies. It might then readily

be assumed that the red coloration had been produced in response

to the visits of these insects, whereas it has arisen very early in

the history of this flower independently of any supposed prefer-

ence for this color which butterflies may possess. This the

writer believes to have actually happened in the case of not a few

red butterfly flowers. Conclusions as to the influence of insects

in developing particular colors, it is evident, must be made with

great caution. No other gamopetalous families contain so many

red flowers as the Ericaceze and Vacciniaceze. In the dev elop-

ment of red and the exclusion of blue coloring they resemble

the Rosacez. But in fruit blue and blue black are in the Vac-

ciniaceze of common occurrence.

In the Primulales there are also two staminal whorls, and

distinct petals still occur. The Primulacez contain 4 white, 11

yellow, and 7 red flowers. Trientalis, Lysimachia, Steironema,

and Anagallis are pollen flowers and contain no nectar. The

corolla is small and rotate. The white Trientalis americana,

(star-flower) grows in damp woodlands, and is rarely visited by

insects. Anagallis arvensis, or poor-man's weather-glass, has

scarlet flowers, which are sometimes white, and in the variety

370 THE AMERICAN NATURALIST. [Vor. XXXVII.

cerulea are blue. The closing of the flower in the afternoon

effects fertilization in the absence of insects. The species

of Lysimachia and Steironema are yellow sometimes with a red

center, or dark spotted in Lysimachia quadrifolia and L. terres-

tris. Of L. vulgaris there have been described three forms.

The smallest, which grows in the shade, has light yellow flowers

with greenish yellow filaments and is self-fertilized. The most

conspicuous and largest form grows in sunny localities, has dark

yellow petals with a red center and is rarely self-fertilized. The

third form is an intermediate one. The flowers of Lysimachia

are largely fertilized by bees of the genus Macropis. These

pollen flowers, which attract few visits, display white, yellow, red,

scarlet and blue colors, and the evidence is wholly insufficient

to prove that the coloring has been determined by the selective

agency of insects.

The species of Primula, well-known for the laborious investiga-

tions of Darwin, are largely dimorphous. The corolla is tubular

and the honey is deeply concealed. The flowers are yellow and

red. Bumblebees are frequent visitors to the yéllow-flowered

forms, and Müller found six red-flowered species in the Alps

fertilized by butterflies. On P. farinosa, which is red with a

yellow eye, he collected forty-eight species of Lepidoptera. In

Glaux the corolla is wanting and the calyx is pink and petal-like.

Sometimes the corolla is duplicated, as in the “hose in hose "

flowers of Primula acaulis.

The Ebenales contain four families, the Sapotaceæ, Ebena-

cea, Symplocaceze, and Styracee. There are but eight species

in the Northern States, two with yellow and six with white

flowers. In tropical regions these families are represented by a

large number of trees and shrubs of which the most important

is the ebony-tree, valued for its black heart wood.

The characters of the Gentianales and Polemoniales are so sim-

ilar that the two orders cannot be readily distinguished. They

differ from the preceding families, which are more closely allied

to the Choripetale, in having but a single whorl of alternate

stamens, and in the smaller number of carpels. Up to this

point white and red colors have been most common in the Gam-

opetalous series, but these two orders are remarkable for con-

taining a large number of purple and blue flowers.

No. 438.] NORTHERN GAMOPETALOUS FLOWERS. 371

The Gentianales include six families. The olive family, or

Oleaceze, contain two white, one purple and seven green flowers.

The green flowers belong to Fraxinus (ash) and are the result of

retrogression, as a part of the species have lost both calyx and

corolla and are wind-fertilized, while others still retain the peri-

anth. The common lilac (Syringa vulgaris) exhibits a great

variety of beautiful shades including white, red, purple and blue.

Showy yellow flowers occur in Forsythia.

The Loganiacez represent the stem-form of the order Genti-

anales. The presence of stipules distinguishes it from all of the

other families of this order. The genera are synthetic in char-

acter and exhibit affinities in many different directions. Here

are found the ancestral types of the Gentianacez, Apocynaceze

and Asclepiadacez, as well as of the Scrophulariaceze and of

the Rubiacez. The family is abundant in tropical regions but

in the Northern States is represented by only four species; one

of which is yellow, two white and one red. Our species are

related most closely to the Rubiacez. The bird flower Spigelia

marilandica (Maryland pinkroot) is scarlet outside, and yellow

within, with a corolla one and one half inches in length, resem-

bling in its coloring Lonicera sempervirens.

The Gentianacez are the first of the Gamopetalous families

in which blue flowers are common. There are 7 white, 1 yellow,

10 red, 4 purple, and 16 blue flowers. The gentians form the most

important genus. These plants many of which are characterized

by a brilliant blue color are widely distributed in temperate

regions, and are very abundant at alpine elevations extending

upward to altitudes of over 15,000 feet. The genus contains

white, yellow, red, purple and blue flowers. In the mountains

of Switzerland the traveller is delighted by the vivid masses of

blue coloring displayed by the gentians. Red-flowered species

occur in the Andes. Müller regards Gentiana lutea as repre-

senting the primitive form of this genus. The rotate flowers

are yellow with slender pointed petals, which are but slightly

united. The honey is fully exposed to beetles and flies as well

as to bees, but so few are the visitors that the plant is unable

to dispense with spontaneous self-fertilization. The blue bell-

shaped flowers have been evolved by bumblebees. The transi-

372 THE AMERICAN NATURALIST. (VoL. XXXVII.

tion from yellow to blue is shown in G. purpurea, which is yellow

inside and blue outside. A number of species by the lengthen-

ing and narrowing of the corolla have become adapted to Lepi-

doptera alone, or to bumblebees and Lepidoptera, but the blue

color it is noteworthy has been retained. In the genus Sabbatia

the handsome flowers are wheel-shaped, and the coloration seems

to have changed from yellowish to white or pink, or pink-purple.

S. lanceolata and S. paniculata are white turning yellowish in

fading; and S. augustifolia and S. campanulata are pink with a

yellowish eye. Several of the rose-colored species are sometimes

white. In Erythraea the slender funnel-form flowers are as the

name of the genus denotes usually red. In Frasera carolinensis

the rotate corolla is light greenish yellow with brown purple dots.

Bartonia which grows in swamps and meadows shows evidences

of retrogression in the thread-like stems and awl-shaped leaves.

The flowers are small and white or yellowish.

The Menyanthacez are marsh or aquatic plants, which in our

species have yellow or white flowers. The flowers of Limnan-

themum (floating-heart) are short-lived. As the petals wither

they become pulpy and exude a thin layer of liquid which is

attractive to flies. A similar change occurs in Tradescantia.

The Apocynaceze are confined chiefly to tropical regions.

There are only seven species in the northern states but they

exhibit a wide range of color, two are white, one yellow, one red,

one purple and two are blue. The dogbanes (Apocynum) have

small, bell-shaped, white, or pink flowers in cymes. - The pinkish

flowers of A. androsemifolium are attractive to butterflies. The

white species are visited by numerous bees. Vinca_minor, or

the myrtle, is blue, while V. rosea from the West Indies is rose

or white. Both species are fertilized by bees. The Oleander

(Nerium) cultivated from the Levant has large, scentless, showy

rose-colored flowers that are fertilized by the hawkmoth Sphinx

nerii. The caterpillar of this moth lives exclusively upon the

leaves of the oleander. The species of this family in the north

are too few to afford much information as to the relations of

insects to their coloring.

Like the preceding family the Asclepiadacez are most abun-

dant in the tropics. There are 11 white, 3 yellow, 5 red, 13

No. 438.] NORTHERN GAMOPETALOUS FL OWERS. 373

purple and 7 green flowers. The inflorescence is in umbels.

Pollenization in the genus Asclepias is affected in a curious and

remarkable manner, analogous in many respects to that of the

Orhidacez. The pollen coheres in waxy masses called pollinia,

which by means of an ingenious clip mechanism are clamped to

the legs or antennz of insects, and are thus transferred from one

flower to another. When the pollinia come in contact with the

stigma they adhere very firmly, and the insects obtain their free-

dom by snapping the connecting bands. Conspicuousness is

gained by two whorls of petaloid bodies, the corolla and the

corona. The corona is composed of five hood-shaped appendages

bearing horn-like processes, within which the scented honey is

plentifully secreted. The color of the corona is usually the

same as that of the corolla, white, orange-red, or purple, though

it is frequently of a different shade. But sometimes the flowers

are distinctly bicolored ; in A. quadrifolia the petals are pink

and the corona is white, while in A. /anceo/ata the corolla is red

and the corona orange. The visitors are Hymenoptera, butter-

flies and flies. Of the 22 northern species, 2 are greenish, 6

white, 3 yellow or orange, 4 red and 7 purple. The white and

greenish-white flowers are the smallest. The two species of

Asclepiodora have the corolla greenish and the hoods purplish.

In Acerates, or green milk weed, the corolla is greenish and the

hoods white, yellow, or purplish. In Vincetoxicum the corona

is reduced to an annular ring adnate to the corolla, and the

brown or red purple flowers possess a putrid odor attractive to

carrion flies. The Asclepiadacez is one of the five Gamopeta-

lous families, which contain seven or more greenish flowers.

The prevalence of greenish and white flowers and the absence

of blue would indicate that the coloring was but little developed

in this family. The corona is perhaps analogous in its origin

to the ligule of grasses. !

The fifth gamopetalous order the Polemoniales, or Tubiflorz, -

is the largest and most difficult of this ‘series. The families

differ but little from each other, and are closely allied to the pre-

ceding order. About goo species belong to the Convolvulacez,

or morning-glory family, but they are confined chiefly to the

tropics. In the Northern States this family contains 7 white,

374 THE AMERICAN NATURALIST. [Vor. XXXVII.

1 yellow, 7 red and 3 blue flowers. The corolla is campan-

ulate or funnelform. In Ipomæœa and Conyolvulus many of the

flowers are bicolored, or tricolored. [pomea_ purpurea is purple,

pink, variegated, or white, and a variety in my garden produced

a purple flower striped with red, the whole flower fading to red.

Both red and purple flowers may occur on the same plant. Z.

versicolor cultivated from Mexico has small, reddish flowers,

which change to orange and yellow. Convolvulus tricolor has a

blue corolla with a whitish throat and a yellow tube. The two

species of Quamoclit have scarlet, salverform corollas adapted to

humming-birds. The genus Cuscuta is a group of yellow, or

orange-colored, parasitic plants destitute of chorophyll, and of

world-wide distribution. The flowers are white or tinged with

rose. In Maine I have found that the visitors were small bees.

The Polemoniacez, or phlox family, are most abundant in the

Western States. In the Eastern States there are 7 white, 10

red, 3 purple, and 8 blue flowers. The corolla of Phlox is salver-

form, and several of the red species have been observed to be

fertilized by butterflies. — P//ox paniculata and P. drummondii

are favorites in cultivation for the brilliant effects they produce

when massed. They have yielded innumerable shades and com-

binations of white, yellow, red and purple. A large number of

the varieties are bicolored, in which the center may be white,

red, purple, or even blue. A curious color variation of a peren-

nial phlox of the variety Monsieur Maille is given in the Ameri-

can Garden for January, 1890. * In the morning the flowers

were of a clear blue, remaining of this color until nearly noon,

when they gradually changed to a delicate pink and by evening

were a beautiful deep rose. This was repeated every day while

the plant. was in bloom." : Many handsome red and blue flowers

occur in the genus Gilia; in G. tricolor the lobes of the corolla

are purple or white, and the throat brown purple, with a yellow

tube. The genus Polemonium possesses pure blue flowers. In

Cobea scandens from Mexico the flowers are at first green

changing to purple. Yellow is not common in this family. In

the Polemoniacez, as in the Convolvulacez, the flowers are

remarkable for the variety of their colors, for the number of

combinations of two or more colors in a single flower, and for

No. 438.] NORTHERN GAMOPETALOUS FLOWERS. 375

the changes of color in individual flowers. These color relations

will be again referred to under the Boraginacez.

Of the Hydrophyllaceze 8 species are white and 10 blue.

Frequently these latter species revert towhite. They are mostly

woodland flowers where blue contrasts with the yellowish or

brown background. This is a small family intermediate between

the preceding family and the Boraginacez.

The coloration of few families is so interesting as that of

the Boraginacez, since the individual flowers often undergo

several changes of color, and “seem to recapitulate to us the

evolution of their colors." There are some 1500 widely distrib-

uted species. The inflorescence is in one-sided scorpioid spikes,

racemes, or cymes. The flowers are small and regular, except

in Echium, but the corolla varies much in length. In Myosotis

and Asperugo the tube is short, and there is a great variety of

visitors; while in Borago and Symphytum the flowors are

inverted and the honey is accessible only to long-tongued

bees. There are 6 yellow, 19 white, 1 purple and 17 blue

flowers. Heliotropum polyphyllum is white but in variety /eaven-

worthii of Florida bright yellow, and in Æ. curassavicum (sea-

side heliotrope) the corolla is white with a yellow eye, changing

to blue. Most of the species are, however, white or blue. The

common heliotrope from Peru (Z7. peruvianum) has the tube

white with the lobes purple, which in fading grow whitish. In

this genus yellow and white appear more primitive than purple

or blue. Echinospernum has small bell-shaped flowers, which

are visited by flies, bees and moths. Æ. Jappula in the bud is

white, red before expanding, and afterwards bright blue. The

coloration of Myosotis, one of the simpler genera, is remarkably

variable. M. palustris and M. laxa are sky blue with a yellow

eye; M. arvensis in bud has a yellow tube with pale pink corolla

lobes, which change to blue and finally fade to white ; M. versi-

color is yellow changing to blue and violet, and M. aipestris

produces dark blue, bright blue, reddish, and even snow white

flowers. The flowers are visited by many species of flies.

Lithospernum purpureo-cceruleum is at first purple-red and later

blue. Pulmonaria officinalis is red and later blue-violet.

Echtum vulgare. (blueweed) has large, showy, irregular flowers,

376 THE AMERICAN NATURALIST. [Vor. XXXVII.

which change from red-purple to bright blue. Symphytum

officinale has a yellowish-white corolla, which is rarely purplish.

Borago officinalis has a bright blue corolla, with which the black

cone of anthers offers a marked contrast. The last three species

are fertilized chiefly by bees. To this family belongs Arnedza

cornuta. When the flowers open each of the five petals is

marked with a dark purple spot, which by the third day has

faded entirely away leaving the corolla bright yellow. In the

Boraginaceze, it is evident, that yellow and white are more prim-

itive than red, purple, or blue. A change in the cell sap from

an acid to an alkaline condition explains the coloration of flowers,

which are at first red but afterwards blue. The transition from

yellow to blue may be caused by the development of a soluble

blue pigment masking the yellow plastids, as in the purple pansy.

The species of Boraginaceze are very rich in anthocyan. The

roots and in some instances the leaves and stems, as in Echium

and Lithospermum, stain violet the herbarium paper on which

they are mounted. From the roots of Z. canescens (puccoon)

the Indians derive a red dye. Another species of this family

Alkanna tinctoria (alkanet) is extensively cultivated in Europe

for its carmine dye used in coloring silk and cotton fabrics, and

also oils, wax and wine. In the sheep pastures of New Mexico

there grows another species Plagiobothrys arizonicus, called

blood purslane, * when the sheep find a patch of it, it colors

their heads red clear to the ears." *

The flowers of the Verbenacez are two-lipped. Of the 12

northern species, two are white, two purple and eight blue. The

species present various interesting color changes and combina-

tions of colors. The native species of Verbena are white, pur-

ple, or blue. The cultivated varieties are remarkable for their

brilliant scarlet and crimson colors. Lantana mixta from Brazil

has flowers, which at first are white changing to yellow, orange

and finally to red. According to Fritz Miiller, the flowers when

yellow are sought by bees, and when red by butterflies. Z, nivea

has sweet-scented white flowers, which in the variety Z. “muta-

bilis change to bluish. The flowers of Clerodendron are

1 Norton. Report Missouri Botanical Garden, 1898, p. 149.

No. 438.] NORTHERN GAMOPETALOUS FLOWERS. 377

bicolored with the calyx and corolla strongly contrasted; C.

trichotomum from Japan has a white corolla and a red calyx,

while C. ¢homsone from tropical Africa has a snow white calyx

and a bright crimson corolla.

The Labiatze is a large family of about 3000 species of widely

distributed plants, which are especially abundant in the region of

the Mediterranean. The foliage usually contains an aromatic

volatile oil. The flowers are distinctly zygomorphous, and

highly specialized both in form and color. In this family and

the Scrophulariacez the gamopetalous plants come to a second-

ary culmination. But while these two families belong to the

same order they are in different subseries, and are more closely

united with other families than with each other. The Labiate

connect with the Verbenacez, both having one-ovuled carpels ;

and the Scrophulariaceze are related to the Solanaceze, both hav-

ing carpels ‘with numerous ovules. T

The 120 species of the Labiatze contain 24 white, 4 yellow,

12 red, 47 purple and 33 blue flowers. According to the

length of the corolla tube and the consequent limitation of the

insect visitors, the flowers may be arranged in three groups.

In Mentha and Lycopus the tube is short, and the visitors are

chiefly flies. In Thymus, Origanum and Betonica bees become

of increasing importance, though flies and other insects are also

numerous. While Stachys, Ajuga, Teucrium, Salvia, Lamium,

Galeopsis are almost exclusively fertilized by bees. Individually

the flowers are often inconspicuous and significance is gained by

massing. In Mentha and Lycopus the stamens are reduced to

four or two, the nearly regular corolla is 4-lobed, and the small,

pale purple or white flowers, which are fertilized chiefly by flies,

are in dense axillary whorls. These genera have evidently

retrograded and departed from the typical labiate form. The

flowers of Lycopus are the smallest of this family. The sma:l

flowers of Thymus and Origanum are in terminal clusters,

2-lipped with a five-cleft corolla. In both genera there occur

large hermaphrodite and small pistillate flowers. The color is

purple and the visitors are flies, bees and butterflies, but the

largest percentage is of flies. The flowers of Køællia (Pycnan-

themum), or basil, are more or less bilabiate. They are small,

378 THE AMERICAN NATURALIST. [Vor. XXXVII.

white, or purple-dotted, and for the most part are in terminal

clusters. The species, like the mints, are strongly aromatic-

scented. In many purple-flowered species of the Labiate family

the leaves and stems also develop a purple pigment, as in

Mentha and Origanum. The entire plant of the summer savory

(Satureia hortensis) turns purple in autumn, while on the other

hand the white-flowered water horehound (Lycopus) has pale

green foliage.

A great number of the larger labiate flowers are chiefly, or

almost exclusively, fertilized by bees. A number of species,

however, have scarlet flowers and are adapted to humming-birds.

The species of Stachys are white, yellow, red, or purple, and the

lower lip is frequently spotted or variegated. The flowers are

rather large and bees are the most important pollinators. The

flower of S. recta is yellowish-white with the border of the upper

lip marked with two purple stripes, and the lower lip purple

spotted. S. annua has the corolla tube whitish-yellow with the

lower lip red-spotted. The corolla of S. palustris is bright

purple with a white and dark red marking on the under lip, and

the flower of S. sz/vatica is red with a purple and white path-

finder on the lower lip. Our four northern species of Lamium

are pollenized for the most part by bumblebees. Three are

purple or purple-red and one is white. The under lip is varie-

gated. Z. album is large, white, or sometimes rose-colored,

with the under lip pale yellow, marked with olive-colored dots.

The pathfinders of the purple-red Z. maculatum are dark red

with white markings, and of the bright purple Z. purpureum, a

dark red flake and dark red lines The flowers of Galeopsis, or

hemp-nettle, display as great a variety of colors. G. tetrahit, so

common in waste places, is purple witha pathfinder on the lower

lip of a yellow spot crossed by a network of red lines ; while G.

versicolor is yellow with the lower lip in front dark violet but at

its base yellow. The species of Prunella are also bee flowers ;

P. vulgaris is blue-purple, white, or rose-colored. Salvia prat-

ensis is usually purple, but Müller mentions a pink variety grown

in his garden. Ajuga repans is a bumblebee flower, and also

Knuth, Paul. Handbuch der Blütenbiologie, bd. ii, teil 2, p. 259.

No. 438.] WORTHERN GAMOPETALOUS FLOWERS. 379

has blue, rose, or white flowers. The native species of Salvia

are blue, or blue and white, and are fertilized by bumblebees.

But in tropical America there occur scarlet and fire-red species,

which are visited by humming-birds. Further illustrations of

variegation may be observed in Teucrium, Scutillaria, Nepeta

and Monarda. It will be noted that the bee flowers are larger

and more variable in color than the species to which flies are the

most important visitors.. Bee fertilization is correlated with high

specialization in form and color. Pathfinders usually occur on

the lower lip and consist of two, three, or even four colors, one

of which is very frequently yellow. The Labiate have but four

yellow flowers, while the Scrophulariacez have thirty-three.

These four species are pale or greenish-yellow, as in Lophantus

nepetoides. The colors of the Labiatze will again be considered

under the Scrophulariaceze.

The Solanacez, or potato family, belong to the same sub-series

as the Scrophulariacece, or figwort family. The flowers of the

Solanacez arë regular or nearly so, and the corolla tube varies

greatly in length from rotate and campanulate to salverform.

There are five perfect stamens of equal length. This family

represents an earlier stage of the Scrophulariaceze. Petunia is

a transition form, which has the border unequal and four of the

stamens didynamous and the fifth aborted. Many species of

Linaria, a well known genus of the Scrophulariacez, have been

known to become regular! A peloric form of Antirrhinum

majus is also known. Moreover at an early stage in their

development the flowers of the Scrophulariacez are regular.

Nine species of the Solanacez are white, twenty-one yellow,

two purple, and eight blue. The genera Lycopersicum (tomato)

and Solanum (nightshade) contain pollen flowers, and are de-

void of honey. They are visited sparingly by both bees and flies.

The rotate flowers of Solanum are white, yellow, purple and

blue with a cone of yellow anthers in the center, or in: oS

heterodoxum the lowest anther is violet like the corolla. The

violet corolla of S. dulcamara (nightshade) is marked at the

base with ten green spots; which Müller observed Syrphidae

! Darwin. Animals and Plants under Domestication, vat i5 p.35

380 THE AMERICAN NATURALIST. [Vor. XXXVII.

examining in search of nectar. The fruits of this genus are

green, white, yellow and red. The small, open, campanulate

flowers of Physalis (ground-cherry) are nectariferous. The

seventeen northern species are greenish-yellow, bright yellow, or

sulphur-yellow, sometimes throughout, but often with a brown

or purple center, or with purplish spots. The yellow anthers

are frequently tinged with purple, and the calyx in some species

is purple-veined. The berry is also yellow or purple. In Leu-

cophysalis the flower is white sometimes tinged with purple

with a yellow center. The fruiting calyx of the strawberry

tomato, P. alkekengi from Southern Europe, contains an intense

scarlet pigment.

The flowers of Nicotiana and Datura are funnelform, or

tubular, and adapted to Lepidoptera. Nicotiana rustica and N.

longiflora are nocturnal species; in the former the corolla is

yellowish-green, and in the latter it has a white border and a

green tube four inches in length. The common tobacco, or JV.

tabacum, which is a diurnal species, has red-purple flowers. In

Datura the large nocturnal flowers are four inches long by two

broad. D. stramonium has green stems and white flowers,

while D. tatula has violet- purple flowers and purple stems.

Brunfelsia grandiffora has greenish flowers two inches in length ;

while B. /atiflora has lavender flowers with a white eye, which

fade to white. Both of these species come from South America.

Two cultivated exotic genera, which also have funnelformed

corollas, are Salpiglossis and Petunia. The large handsome

purple or rose-colored flowers of Salpiglossis are remarkable for

being netted-veined and laced with golden yellow. The numerous

hybrids of Petunia under cultivation are chiefly derived from two

South American species, one of which is white and the other

red-purple. They exhibit a wonderful variety of bicolored, tri-

colored, and variegated flowers often veined or mottled in

endless ways. A unique form is striped or margined with green.

The coloration of the Solanacez, which is an extensive tropical

family, is of more than usual interest because of the many green

flowers of large size which it contains. These green flowers, as

well as many large white and yellow colored species, are funnel-

form or tubular, most strongly sweet-scented in the evening and

No. 438] NORTHERN GAMOPETALOUS FLOWERS. 381

adapted to night-flying Lepidoptera. Nocturnal fertilization has

prevented the development of red and blue colors, as in Cestrum

nocturnum with yellowish green, C. fargui with dull yellow,

Datura metel with white, and D. fastuosa with the flowers

violet outside and white within. Cestrum elegans and Nicotiana

tabacum are day flowers and are rose purple. Of the bumblebee

flowers, Atropa belladonna is dull purple, and Hyoscyamus niger

is dull yellow with purple veins. The colors of the rotate

flowers of Solanum and Physalis are white, yellow, purple and

blue, and are sought by both bees and flies.

Unlike the Solanaceze the Scrophulariaceze are most abundant

in temperate and arctic regions. There are about 2500 species.

Of the 113 northern flowers, 13 are white, 33 yellow, 7 red, 32

purple, and 28 blue. As inthe Solanacez the number of yellow

flowers is much larger than in the Labiatae. In Verbascum and

Veronica the flowers are wheel- shaped. Three of the northern

species of Verbascum, or mullein, are yellow or sometimes white.

Gartner made many experiments in crossing the yellow and

white varieties, and actually raised one variety from the seed of

the other. V. phaniceum is purple, rose-colored, or white.

When the purple variety of V. phaniceum was crossed by the

yellow variety of V. d/attaria, the corolla of the hybird was pale

crimson? The leaves of Verbascum thapsus are yellowish

green, often spotted with yellow, or turning completely yellow.

The visitors are a miscellaneous group of insects, of which bees

are the most important. Miller devotes considerable space to

the consideration of the colors and manner of fertilization of the

speedwells, or the genus Veronica. Most species have light or

dark blue flowers often with darker stripes; but those of V.

urticifolia and V. perigrina are pink. The flowers, which are

small and rotate, with the sepals and petals reduced to four and

the stamens to two, have evidently departed widely from the

primitive type of the family. The visitors are chiefly flies,

especially Syrphidz, the honeybee and the smaller bees. Müller

finds it difficult to explain the origin of the blue coloring by the

lDarwin. Animals and Plants under Domestication. Vol. ii, p. 83.

*Kerner. Natural History of Plants, vol. ii, p. 567.

382 THE AMERICAN NATURALIST. (VoL. XXXVII,

selective influence of the pollenizers, and regards the question of

its evolution as still unsolved. It is surprising that he does not

give more consideration to the influence of physiological, as well

as of ecological causes, in determining the particular colors of

flowers. In this genus, as in many others, it is the character of

the pigment-forming function, which is of chief importance.

On the Alps Müller found 7ozzia alpina to be also a fly flower.

The corolla is'bright yellow except that the three lower petals

are spotted with dark purple. In view of the fact that fly-

flowers are white, yellow, red and blue, the evidence that their

particular coloration has been determined by the color sense of

Diptera must be regarded as unsatisfactory.

The species of Scrophularia are wasp flowers, green or

purplish without and brown purple and shining within. The

corolla is short and round with easily accessible honey. By far

the greater number of species of the Scrophulariacez are adapted

to bees and many to bumblebees alone. The species of Linaria

are for the most part bumblebee flowers. Z. vulgaris is bright

yellow with an orange-colored palate as a honey-guide, while Z.

canadensis has small blue flowers with a whitish palate.

Antirrhinum majus is bright purple or white with a yellow

palate. A. crontium, according to Knuth, is red, or rarely

white, with darker red lines. The under lip displays a pale

yellow pathfinder and a white zone besides other markings.

Chelone glabra (turtle head) is white with reddish lips. In

Maine this species is fertilized exclusively by bumblebees. C.

obliqua is bright rose color. Digitalis purpurea is purple with

dark purple spots within on the lower side. D. /ufea is yellow.

Melampyrum lineare is whitish with the lower lip yellow. Other

species of this genus have bright yellow flowers. Pedicularis

with the flowers yellow and red is also fertilized by bumblebees.

Rhinanthus crista-galli (yellow rattle-box) is bright yellow with

the lower lip purple-spotted.

In a preceding paper the capability of the calyx to develop

different colors was shown to equal that of the corolla. In con-

nection with the genus Castilleja the variety of coloring in tlie

floral bracts may be considered with advantage. In C. coccinea,

or the scarlet painted cup, the flower is greenish yellow and the

No. 438.] NORTHERN GAMOPETALOUS FLOWERS. 383

floral bracts bright scarlet, or occasionally both bracts and calyx

are yellow. In C. zzdivisa both bracts and calyx are bright red,

and in C. acuminata ‘yellowish or purplish, but in C. sessz/zffora

they are green similar to the leaves. In Monarda didyma of the

Labiatz the corolla is scarlet and the bracts red, in M. media

both are purple; but in M. clinopodia the bracts are pale or

white, and the flower is yellowish-pink. In Cornus canadensis

the involucre is white and white or red in C. florida. Bourgain-

villea glabra has three conspicuous purple-red bracts. Finally

in the Proteaceze Protea globosa has “the upper foliage leaves

grouped into a large outer envelope, which surrounds the spher-

ical golden yellow inflorescence, and these crowded leaves are

colored blue in contrast to the lower scantier foliage which has a

grass green color.” Other illustrations of colored floral bracts

occur in the aroids, spurges and Composite. The floral bracts

evidently may develop as wide a range of colors as the floral

leaves, though blue is much rarer.

Müller in his A/pend/umen has considered the relation of bees

to floral colors at considerable length. In many genera, when

the flowers are wheel-shaped and adapted to short-tongued visi-

tors, the species are colored alike, mostly yellow or white, as in

Ranunculus and Potentilla, the Alsinez, Cruciferae and Umbel-

liferee. Even when the honey is partially concealed this may

hold true, as in Mentha and many Cichoriacee and Composite.

The short-tongued insects visit these flowers indiscriminately

and not infrequently the bees. Genera adapted to bees on the

contrary usually display a variety of colors, as white, yellow, red,

violet, blue and brown, especially when they bloom in the same

locality at the same time. Among the examples given by Müiler

are Aconitum lycoctonum yellow, A. napellus blue; Lamium

album white, L. maculatum red, Galeobdolon luteum yellow ;

Salvia giutinosa yellow, S. pratensis blue; and Pedicularis

tuberosa whitish yellow and P. verticillata purple. An excep-

tion to this rule is offered by the PapjJionacez, which contain

a great number of yellow flowers. Here Müller thinks that the

color is so strongly transmitted that not even small variations

appear, and there is consequently no opportunity for the

development of a variety of colors. Asa rule, however, genera

384 THE AMERICAN NATURALIST. |. [Vor. XXXVII.

with rotate open flowers are monochromatic, while genera with

highly specialized corollas adapted to bees are polychromatic.

How have these color differences arisen, and how far are they

due to the selective influence of bees? That bees in collecting

nectar adhere closely in their visits to a single species is well

known. Even when the flowers are nearly alike in form and

color the honeybee often shows a remarkable power of discrim-

ination. If all of the flowers blooming at the same time in a

locality were of the same color, it is evident that bees could not

distinguish between them as readily as when there is a variety

of colors contrasting with each other. This is the reason of

the development, according to Müller, of the numerous color

differences in bee flowers. There can be no doubt that bees

can distinguish between different hues, and can make their visits

more quickly and easily because of the contrasts of flowers in

coloration. If, says Müller, we assume that one of two forms,

closely allied in structure and of the same color, should vary

in- color, bees would distinguish it at once, and follow: it more

easily in their visits. He found, moreover, that a majority of

bee flowers are red or blue. Of one hundred species of bee

flowers observed by him on the Alps thirty-four were white or

yellow, and sixty-six red or blue. In the German and Swiss

flora he records that 152 bee flowers were white or yellow, and

330 red, violet, or blue. The honeybee and bumblebees showed

a much larger percentage (about 20 %) of visits made to red

and blue flowers of all kinds, including not only flowers with

the honey partially or deeply concealed but also those with it

fully exposed, than to white and yellow flowers. To explain the

greater frequency of their visits to red and blue flowers, it does

not seem to the writer necessary to assume that they find

greater pleasure in these hues. The more specialized bees learn

from experience, and it is not difficult to understand that they

would soon come to associate with red and blue colors a more

abundant supply of nectar and greater freedom from competition.

(To be continued.)

THE CAUSES OF ACCELERATION AND RETARDA-

TION IN THE METAMORPHOSIS OF AMBLYSTOMA

TIGRINUM: A PRELIMINARY REPORT.

J. H. POWERS.

DuniwG the past six seasons the writer has been collecting

facts on the metamorphosis of Asmblystoma tigrinum, both

by observation and by experiment. It is hoped to publish an

extended account latter; but as relatively definite results have

already been reached, the interest of the subject seems to war-

rant the present publication of an outline, giving conclusions

and something of the evidence from which they have been drawn.

The extreme variability of our tiger salamander in regard to

the time at which it undergoes metamorphosis is well known.

Individuals may breed in the larval condition: witness the Mex-

ican Axolotl, which the concensus of scientific opinion has now

practically reduced to the rank of a sexually mature larva of our

common species. At least one, and probably two, specimens of

our common species, male as well as female, have come into the

writer's possession which showed every character of the Mexican

animal, even to sexual maturity. More striking than these facts,

however, are the anomalies of size which frequently occur. My

largest larva weighed, before the beginning of metamorphosis,

one hundred and thirty-six grams; my smallest adult weighed

but three grams. Thus a larva may outweigh an adult in the

ratio of forty-five to one. The above mentioned giant larva,

while kept in an aquarium, ate daily, for about two weeks, at

least one fair sized adult Amblystoma of its own species; some

days it ate two, or followed its cannibalistic meal by devouring a

piece of liver the size of a man's thumb.

What are the causes which produce these almost unparalleled

extremes of variation? Those familiar with the literature of the

subject know that many regard the question as already definitely

settled: favorable conditions for aquatic life are supposed to

385

386 THE AMERICAN NATURALIST. [VoL. XXXVII.

prolong the larval, gill-bearing stage; while unfavorable condi-

tions for aquatic life — e. g., the drying up of ponds, with forced

aerial respiration — are thought to be the regular causes of meta-

morphosis. Heat and light are regarded by many as subsidiary

causes.

This explanation of the metamorphosis of Amblystoma — and

of allied forms — as due to a direct response to changing condi-

tions of environment is traditional. It has received further sup-

port from casual observations of naturalists, who have seen these

animals undergoing metamorphosis on the mud of evaporating

ponds. But the final sanction which raised the hypothesis into

an almost universally accepted datum of science was Weissman's

great article, *On the Change of the Mexican Axolotl to

an Amblystoma."! This article was based upon Marie von

Chauvin's experiments with five larvae only. Later and much

more extensive experiments by the same person were much less

favorable to Weissman's conclusions. Indeed a careful study of

their methods and results seems to the writer rather to cast

doubt upon the entire conclusion that enforced air breathing

caused the metamorphosis of these supposed Axolotls. Buta

consideration of these experiments in detail is beyond the scope

of the present article. Brief reference will be made to them

latter. I will now pass at once to the results of my own obser-

vations and experiments, which have been made on Amblystomz

in the vicinity of Crete, Saline County, Nebraska.

First, metamorphosis, in the writer's vicinity, occurs rarely if

at all as the obvious result of enforced air breathing through the

drying up of ponds. Diligent search has been made; in one

summer over one hundred ponds and larger pools were examined.

Many contained larvze; but none were found in ponds or pools

that were less than one foot in depth. The last remnants of

large ponds, where larvae had been abundant a few weeks before,

showed no trace of them, although the water might be alive

with the tadpoles of the common frog. Experiment showed

that Amblystoma larvae could not usually withstand the temper-

ature of very shallow water exposed to Nebraska sunlight in

June, while the tadpoles of Rana were unharmed by it.

1 A translation of this article will be found in the Smithsonian Report for 1877.

No. 438.] AMBLYSTOMA TIGRINUM. 387

However, not death, but early metamorphosis is the probable

explanation of this early disappearance. Metamorphosis fre-

quently takes place early in the season, even in June. I have

taken small specimens in metamorphosis as early as June from

ponds with a depth of water from four to six feet. A careful

series of observations, in the case of a single pond, showed the

period of wholesale metamorphosis to occur, in this instance, in

the latter half of August. Yet in this case unusually heavy

August rains had raised the pond to its maximum height and

even caused considerable overflow. In spite of repeated search

at appropriate times and places, no Amblystoma have been

found in metamorphosis on the mud of drying ponds. That

they are so found in other localities the writer knows from

childhood experience, as well as from report and record. Our

ponds are usually too muddy for direct observation, but the con-

stant use of the dip net has shown me that these larvae undergo

metamorphosis in a considerable depth of water, preferably about

three feet. My field notes show. but few instances of single

specimens in metamorphosis taken from the shallow borders of

ponds; many more from deeper water, even up to six feet. In

but a single instance have I seen an Amblystoma leaving a pond

before the metamorphosis was quite complete.

The second point at which.I have failed to make my observa-

tions tally with the statements of others is in regard to the habit

of rising to the surface for air. It has been stated that rising to

the surface for air, or at least the more frequent rising, pre-

cedes or ushers in, metamorphosis. I have been favorably situ-

ated for the observation of this habit, and have followed its

beginnings, its sudden acceleration, its slow or still more sudden

cessation, rebeginning, etc.; but whatever the significance of

this strange and intricate phenomenon may be, it is certain that,

with our larvee, it stands in no immediate relation to metamorpho-

sis. It may begin and, within a day or two, become incessant

with larvae far too small for metamorphosis, under any condi-

tions. I have seen a group of about a hundred large larve, in

well aerated clear water, ten feet in depth, so incessantly playing

to the surface that the water seemed as if rained upon with large

drops at the beginning of a shower. Day after day I found them

388 THE AMERICAN NATURALIST. [Vor. XXXVII.

equally active in the same place; their gills were long and well

fringed; not one showed the labored upward ascent or the dead

after-sinking which characterize the larva in which metamorpho-

sis has begun. Suddenly, within twenty-four hours, this air-

taking at the surface entirely ceased; a half hour’s watching

showed hardly a swirl. Yet the larve were still there; fora few

days later, the water being let from the basin, I caught them,

not one showing a trace of gill shrinkage. Experiment with

larvae under controlled conditions conclusively confirmed these

general observations. Moreover, larvæ in metamorphosis do not

rise with especial frequency; and times of rapid surface play

have never coincided with times of wholesale metamorphosis.

It has been further said that, just as the bringing of Amblys-

toma into the air would force the change to the adult condition,

so enforced aquatic conditions would prolong the branchiate

condition,— larvae would not undergo metamorphosis if kept in

water and unable to crawl out, e. g., if kept in glass vessels with

perpendicular sides. Yet several hundred larve have undergone

metamorphosis in my laboratory under precisely these conditions ;

not one, in fact, has refused todoso. The form of the aquarium,

the accessibility or inaccessibility of a foothold out of the water

have exerted no noticeable accelerating or retarding influence

upon the duration of the larval state. Indeed, I have thus far

been unable to raise larve in the laboratory to their maximum

size. I have tried hundreds of them, have varied conditions in

many ways, and they thrive well, too well, as the sequel will

show; but my largest specimen was but 17.7 cm. in length,

while, in one instance, I have seen hundreds of larva in a pond

that exceeded this dimension. Out-of-door experiments, in tanks,

cisterns, etc., have, with one exception, to be mentioned later,

led to precisely similar results. Larvae undergo metamorphosis

where the possibilities of terrestrial life are not present; they

undergo metamorphosis under conditions where larve flourish,

but where adults slowly die of starvation.

The most crucial proof, it has been thought, of the power of

adaptive response in the young of Amblystoma has been the

bringing of the young into shallow water or out of the water

altogether, that enforced lung breathing might induce the transi-

No. 438.] AMBLYSTOMA TIGRINUM. 389

tion to the terrestrial form. That animals so treated have, in

some cases, metamorphosed is certain; but have sufficient con-

trol experiments been conducted to eliminate the possibility of

other causes than aerial respiration? The writer has repeated

these experiments, and does not find that air-breathing, taken by

itself alone, is the natural cause of metamorphosis, or even, in all

probability, an accelerating condition. I will instance one set

of typical experiments.

Five larva, 14 to 18 cm. long, placed on mud with but water

enough to partially cover them; refused to eat; kept quiet;

all began. metamorphosis three days later, and finished eight

days later.

Second: Fourteen larvae, same as above, placed in large

aquarium with five inches of water; larvae quiet; not fed.

Began metamorphosis four to five days later, and completed

same ten to twelve days later.

Third: Twenty larvz, same size as above, placed at same

time, in aquarium with water just sufficient to cover back fin.

Took little food; frequently excited and active. Eight showed

signs of metamorphosis in three days. Thirteen at five days.

All had completed metamorphosis in eleven days.

Fourth: Twenty larvze, carefully assorted, with one exception,

as to size and other characteristics to match the preceding, were

placed in an identical situation ; but with a depth of one foot of

water. They ate well from the start, the eating habit being

established before metamorphosis began. Yet three began

metamorphosis on the third day; many on the fifth day. And

all, with the exception of the one peculiar, broad-headed speci-

men, almost simultaneously with those in experiment three.

Fifth. Simultaneously with the above, more than twenty

larvee, not assorted as to size, and mostly smaller than the pre-

ceding were placed in a large aquarium supplied with running

water. They ate readily. Metamorphosis was more irregular ;

but on the sixth day many had begun the transition, and it was

completed in about the same period as with the foregoing.

Sixth. Above fifty small larve from the same source as the

above, deemed quite unsuitable for experiment, as they had not

reached the minimum size at which metamorphosis commonly

390 THE AMERICAN NATURALIST. [Vor. XXXVII.

takes place, were placed in the stone basin of a fountain. The

food supply was scanty or none; but there was more or less

shade and a continual spray of cool water. Here, it was thought

growth could be delayed and the larvae kept for later experiment.

Yet on the eighth day the astonishing fact was discovered that

many were in metamorphosis. And on the fourteenth day three

adults, 9.5, 10 and 11 cm. long respectively, were found.

Others had probably escaped ; as, despite the perpendicular stone

ascent of several inches, it was found they could do.

I have instanced this series of experiments, first, because of

all I have conducted, it is the most favorable to the received

hypothesis. Second, because it will serve to illustrate further

points as well. It will be noticed that there is a slight apparent

acceleration in the metamorphosis of the five larvae placedgwith

their backs out of the water; yet some of those in the aquaria

with abundance of water began quite as soon and finished but a

day or two later; all that were strictly assorted as to size and

other characters followed suit very soon, while still more astonish-

ing is the early metamorphosis of the undersized larvae in cool

well-sprayed water of the fountain. The well-nigh simultaneous

metamorphosis of these larvze so soon after the beginning of these

experiments indicates that the chief cause of metamorphosis was

alike operative in all cases ; what this case was will become obvi-

ous later in the discussion. In other instances I have had larvee

prove much more resistent to metamorphosis when kept partially

or entirely out of water. I will relate one, which is interesting

because it was a semi-natural experiment and, in part, on a large

scale.

Around the sloping cement sides. of a large artificial reservoir

frost had raised a strip of plastering, perhaps a yard in width,

leaving a crevice an inch or more in depth below it. Until mid-

summer this crevice had been several feet below water, and here

about two hundred larvae had formed the habit of concealment,

their abraded back fins telling plainly that their occupancy of the

crevice had not been temporary. When these had reached a

length of ten to thirteen centimetres the water in the reservoir

was slowly reduced in depth, during a week or more, until, at

first portions, and then the whole of the crevice was above the

No. 438.] AMBLYSTOMA TIGRINUM. 391

water level. The larva obstinately refused to leave their

accustomed shelter; they could obviously have swum out with

the receding water, and in most instances could have crawled or

even floundered down the incline into the water, as indeed they

did, if the plaster sheets were raised and the larvae poked a little.

I disturbed but few of them however; it seemed an admirable

opportunity to watch for enforced metamorphosis. Yet, despite

the favorable circumstances that the crevice remained moist and

that the larva were shielded from the direct rays of the sun,

within a few days most of them had died. In only two of the

survivors could I find any trace of gill shrinkage sufficient to

indicate probable beginnings of metamorphosis. Contrast with

this failure to respond to enforced air breathing, the case of ten

which, two years before, I had netted from this same reservoir ;

they had been accustomed to an unlimited supply of well aerated

clear water; their branchiz were unusually long; their whole

aspect promised (as I then thought; this being near the begin-

ning of my experience with Amblystoma) a long continuance of

the gill-bearing stage. The larvee were placed on Friday P. M.,

in a large aquarium with perpendicular glass sides, supplied with

abundance of the same tap water in which they had developed

from the egg. They were excessively refractory and restless ;

no attempt was made to feed them. On the following Monday

morning the astonishing fact disclosed itself that every larva had

undergone metamorphosis; every gillcleft was closed; the

remoulding of head and body shape were complete; only the

merest trace of tail fin in a part of the specimens, with gill stubs

less than two millimetres in length, were left. Cope, in his study

of museum specimens, would have reckoned every one of these

specimens as adults. Yet the transformation had taken place

from larvae of an unusually piscine type, within the space of sixty

hours; and entirely in water which was abundant, cool and well

oxygenated. To return to the case of the larvz left out of

water beneath the sheets of plaster.

Finding, about the fifth day, that most of them were dead, I

overturned the loose plaster and secured the last twelve surviv-

ors, two of which, as I have said, already showed barely percep-

tible signs of metamorphosis. All of them were placed on a

392 THE AMERICAN NATURALIST. [Vor. XXXVII.

moist surface, under large split sponges raised just enough to

give the larve sufficient room. The sponges were kept wet.

In this situation the two larvae which showed signs of metamor-

phosis when taken slowly completed the change, and one other,

one of the smallest in the lot, after weeks of hardihood in what

appeared like a most unnatural and unfortunate existence, under-

went still more slowly, an abnormal metamorphosis. The details

of this metamorphosis and its final results, both of which I have

occasionally had repeated under similar circumstances, are very

interesting ; but I will not discuss them here further than to say

that they appear to me to indicate that metamorphosis by early

enforced terrestrial life is wholly unnatural to these animals,

and, in a state of nature, would seldom produce an adult capable

of survival, even if the metamorphosis was itself successfully

passed, The other nine larve, taken from beneath the plaster

and placed beneath the wet sponges, all died, seven of them

without showing any signs of true metamorphosis, although

several withstood their terrestrial conditions for many days.

At this point and in connection with the last-mentioned larve,

I wish to protest against what seems to me a careless and well

nigh unpardonable misinterpretation of certain very simple facts

in connection with larve exposed to air. I refer to the

withering of the gill-tips and fringes and the like reduction of

the dorsi-ventral fin fold. Again and again in the literature of

this subject it is evident that observers and experimenters have

looked upon these changes as the natural beginnings of true

metamorphosis. That a novice should so consider them is nat-

ural enough, but how an observer, broadly and minutely. conver-

sant with the facts of amphibian metamorphosis could so

interpret them the writer is at a loss to know. The organs in

question do of course suffer when exposed to the air, even to

moist air; their extreme delicacy makes this inevitable. The

seven larvae, which, in the above trial, died before metamor-

phosis had begun did take on a somewhat dilapidated appearance

before they succumbed. The gills were more or less shrunken

at the tips and their fringes partially withered; the back fin,

too, had lopped to one side like a wilted plant, and its margin

was uneven and somewhat withered. But these changes, high

No. 438.] AMBLYSTOMA TIGRINUM. 393

authority to the contrary notwithstanding, bear no resemblance,

and apparently no relation, to the incipient changes of normal

metamorphosis, in which gill and fin-fold shrink, always, first at

the base, and never at the tip; and in which other fully as

important though less obvious changes are never wanting to tell

that a vital and inner crisis has arrived. As well interpret the

withering comb of the frost-bitten fowl as a sign of metamor-

phosis as the withering gill tips and fin margin of the air-

exposed larva.

It has been iiaitied: in connection with the hypothesis that

aerial respiration is the cause of metamorphosis, that a definite

relation exists between gill development and the disposition to

retain or discard the larval form. Large-gilled larvae resist

metamorphosis, it has been said; small-gilled larvae do not, but

are easily stimulated to change. Moreover, the claim has been

made, that the branchiz themselves develop more or less fully,

according to the amount of oxygen in the water in which they

live.

If there is any truth in these two corollaries of the received

hypothesis, it is certainly not the whole truth, and the observa-

tions of the writer have so far failed to substantiate it. The gills

of these animals vary much in size in a state of nature, without

obvious relation to the oxygen supply of the water. I estimate

that the surface for oxygen exchange varies, in larvae of equal

size, as one to twenty. The causes are by no means clear. In

general I find, in confinement at least, that rich nutrition and

quiet habits produce the largest branchiz. The finest specimens,

by far, that I have ever seen, in this regard, were two larvae

raised, almost from the egg, in a jar holding but two liters of

water which was changed rarely. The branchize of these two

specimens surpassed considerably, both in length of the rami

and in length and number of the fringes, the branchiz of the

finest specimens taken from the deep, clear water of the city

reservoir ; although this latter source does frequently yield long-

gilled specimens. Moreover, these two larva, despite their

superb branchiz, the wonder of all who saw them, were, none

the less, air breathers; they rose, not infrequently, to the sur-

face for air. And, still more surprising from the ordinary stand-

394 THE AMERICAN NATURALIST. [Vor. XXXVII.

point, these two larvze were, one after the other, striken by the

sudden impulse to change, the great gills withering almost in a

single night; they underwent metamorphosis at about fourteen

centimeters in length, in the very jar of water in which they had

been flourishing as larvz so surpassingly well. My aquaria have

furnished any number of essentially similar cases. Whether or

not it be true that oxygenated water develops the large gill, it

certainly cannot be too strongly emphasized that the large gill,

when developed, is no impediment to metamorphosis. I once

investigated a pond well stocked with thriving larvae which

struck me by the meagreness of gill development ; the rami were

short, and very thin, the fringes likewise, irregular, few and

short. A few weeks later the gills had considerably increased

in size and general development. Yet it was following this

period of gill expansion that rapid metamorphosis began. The

expansion of the gills was here doubtless in no sense the cause

of the metamorphosis. They may have been essentially unre-

lated phenomena. It is possible however that a connection

existed as follows: The smallgilled larvae, although having

reached considerable size, had still the delicate larval skin well

adapted in itself to subserve the purpose of aquatic respiration.

Metamorphosis seldom begins until this larval skin has given

place to the more or less thickened integument which is to

finally characterize the adult. This change in integument is not

here considered as a part of the metamorphosis proper; for the

larva may change its integument and yet long retain its aquatic

form. Yet this change is a necessary preparation for metamor-

phosis ; and it is quite possible that it should render the skin

less effective for respiratory purposes. The cutaneous circula-

lation might even be checked, and a corresponding increase of

blood flow to the gills, causing their growth. Certain it is that

larvee with the thickened dermis usually have at least well

developed gills, and such larvae, if the right stimulus comes,

are ripe for metamorphosis. I will mention in this connection

the surprising: fact, developed in the course of several experi-

ments, that the adaptation of these larva to entire aquatic res-

piration stands in no constant relation whatever to branchial

development. Many experiments were made to induce metamor-

No. 438.] AMBLYSTOMA TIGRINUM. 395

phosis in larvee confined strictly to aquatic respiration by nettings,

so arranged as to prevent the larve from rising to the surface

for air. But the surprising difficulty was at once encountered

that these larvæ are, all of them, air breathers from a very early

stage, and usually cannot, or will not, endure for any length of

time complete exclusion from air-taking at the surface of the

water. With larvæ of but two centimeters in length, when the

development of lungs is but slight, confinement below the surface

proved fatal. Even with but two specimens in a large aquarium

jar, where they had grown from the egg and thriven in the most

natural manner possible, the larvae repeatedly bored through the

netting at night to reach the surface of the water; upon the

netting being doubled, one larva still penetrated it, while the

other was dead beneath it. This experiment was repeated with

many variations in method, but little divergence in result.

Larve with very large branchiz, taken from deep, cool, clear

water, and introduced singly into a large aquarium freshly filled

with the very tap-water in which they had grown would some-

times die of asphyxia in a few hours if prevented from reaching

the surface. Such results are very striking. “ These animals,"

said an observer of my experiments, “have gills by the whole-

sale; but they seem to be mainly for ornament." Not only did

the largest gilled specimens succumb when confined beneath the

surface of standing water, and sometimes of running water, but

they showed no greater ability to live under water than the

specimens with the smallest branchiz that could be chosen.

Indeed, when, after many trials, we finally secured two specimens

that were indifferent or nearly indifferent to their confinement

under running water, neither of them were large-gilled forms,

and the one best adapted to this treatment was a very small-

gilled specimen. Even specimens in the earlier stages of meta-

morphosis show little less resistance under this treatment ; and

in one trial showed even more.

Hand in hand with this inability to live as gill-breathers goes _

the complete ability to live as lung-breathers, even at very early

stages. Seldom, if ever, have my larve died of bad or poorly

oxygenated water. Over three hundred larva, of perhaps two

to four centimetres, were, in one instance, transferred directly

396 THE AMERICAN NATURALIST. | [Vor. XXXVII.

from a reservoir where they had grown in pure, deep water, to'a

single aquarium. Here they lived and throve, although the

oxygen of the water was so exhausted that minnows died in a

few minutes of asphyxia. Yet the gills of these larvae remained

normal. (

If metamorphosis really were caused by enforced aerial res-

piration, and if loss of branchial surface were a chief factor in

the process, both of which are assumed by the common hypoth-

esis, then it would seem to follow as a natural or even necessary

conclusion, that cutting off the gills should stimulate to meta-

morphosis. True, European experimenters on the supposed

Axolotl did not find this to be the case; but their failure was

explained by the fact that the gills of the Axolotl were quickly

reproduced. Besides, the larvae experimented upon in Europe

were resistent to metamorphosis under most conditions. Gill-

amputation may have constituted a real stimulus to metamor-

phosis, and yet one insufficient to bring about the actual change.

It seemed to me that our larvee were much more favorable for

the experiment, for they are, most of them, predisposed to early

metamorphosis. And, moreover, I have not found that the gills

were reproduced with especial facility ; weeks may intervene

with but little growth, and no specimen in my aquaria has

reproduced a gill of normal type; they remain short and

truncated.

Yet here again experiment gave practically negative results.

At intervals during five days, I removed practically the entire

gills from twelve larvae, a few fringes that were situated on the

very gill arches alone remaining. The larve were kept in a fair

amount of water, which was changed once daily and they fed well.

Twelve similar larvae were treated in a similar manner in every

respect except that the gills were not removed. Individuals in

both sets soon began metamorphosis, two or three with the cut

gills showing a possible acceleration of about twenty-four hours

over the earliest specimens in the other lot. This, however, may

easily have been accidental, the metamorphosis of the majority

in the two lots running an almost exactly parallel course. At

the end of the twenty-fourth day, twenty-three out of the twenty-

four larvze, still in the aquaria, had completed the metamorphosis. ,

No. 438.] AMBLYSTOMA TIGRINUM. 397

One only had proved refractory, not having even begun the

change. This was one from which the gills had been removed.

Like all the specimens it had thriven and made good growth, yet

its gills had grown to barely one third natural size. At this

time the specimen was utilized for an experiment under running

water, and, despite its several weeks of life with practically no

gills, and the small size at which these organs still remained, it

withstood fairly well the confinement under water, dying at the

end of the second day just after metamorphosis had begun.

The foregoing facts, together with many more like them, have

led me to think that the acceleration or retardation of metamor-

phosis in our species of Amblystoma, is little, if at all, a question

of enforced air-breathing, of gill development, of oxygenated or

unoxygenated water. Is it then a question of temperature or of

light? Again the answer must be largely negative. Not that

the writer would deny to these important agencies all influence.

A very low temperature checks all life activities in Amphibia ;

metabolism sinks to the lowest ebb, and metamorphosis is

naturally excluded. Light, too, exerts important influences upon

the activities of these animals, and thus indirectly if not directly

affects growth; and growth and metamorphosis are intimately

connected. But abundant observations show that sudden and

early metamorphoses are not produced chiefly by excess of light

or heat; while délayed metamorphosis is certainly not alone the

result of darkness and low temperature. I will mention the fol-

lowing under light :

Larvae sometimes attain great size in Nebraska ponds despite

their complete exposure to the relatively constant sunlight of our

clear summer climate. In the laboratory, the largest larva I have

ever reared — 17.5 cm. — was kept in a rather small battery jar

exposed to the full light of an east window. Others in the same

window metamorphosed at very varying stages of growth. On

the other hand, larva in dark aquaria have frequently meta-

morphosed with the utmost readiness. In a long series of

experiments on the causes of color variation, larvae were exposed

for weeks before metamorphosis to all possible degrees of light

and darkness (not to lights of the different primary colors) and

no obvious retardation or acceleration of metamorphosis resulted.

398 THE AMERICAN NATURALIST. [Vor. XXXVII.

Many larvee have metamorphosed readily after introduction into

the almost total darkness of a closely covered, deep cistern.

As to temperature similar facts may be cited; indeed, many

of those already cited are obviously applicable. The instance of

prolonged larval growth in the east window is especially interest-

ing, for on several of the hottest days of the season other larve,

in like jars in the same window, died of the heat. On the other

hand scores of small larvae have undergone metamorphosis in

shaded aquaria supplied with currents of cool tap water. The

instance, already cited, of the phenomenally early metamorphosis

of numerous larvz in the cool water of the fountain basin may

be recalled. Even six small larvae. which I introduced into a

tank supplied with a stream of the coldest spring water, heavily

shaded with trees and covered with two-inch plank, all underwent

the change in the course of a few weeks. In short, these

Amblystoma larvae have, with the writer, proved singularly

indifferent to wide variations of temperature and luminosity.

What then does control the metamorphosis? But one cardinal

factor in the animal's economy is left, and observation and

experiment show it to be the dominant factor in question : nutri-

tion. Metamorphosis is a question of nutrition. Stated more

accurately, it may be said that metamorphosis is a matter of

metabolism, of anabolism passing into sudden katabolism, as the

result, usually, of checked nutrition. Other causes may and do

coóperate; buta check to nutrition, previously abundant, is by

far the most effective and the most frequently operative.

Liability to metamorphosis at any given time is great, in

direct proportion to the prevalence of anabolic change at that

time; the certainty of metamorphosis at any time is great, in

proportion to the suddeness with which anabolism is converted

into katabolism. The larva of Amblystoma is an organism,

physiologically, in an unstable equilibrium. Carnivorous, an

incessant feeder, capable, under the most favorable circumstances,

of growing to a length of eighteen centimetres in eight or ten

weeks, the overwhelming anabolic changes which are thus

maintained tend, when interrupted, to pass suddenly into their

opposite, into katabolism. Large larvæ, fed to the maximum,

and very fat, lose one quarter to one third of their weight during

No. 438.] AMBLYSTOMA TIGRINUM. 399

a metamorphosis that occupies but a few days. Some construc-

tive changes take place during metamorphosis ; but the changes,

as a whole, are plainly destructive. The beginnings of the

process, the resorption of fin-fold and gills, are plainly processes

of self-digestion of peripheral parts. I have seen cases of early

enforced metamorphosis, in which it seemed that this destructive

action attacked the periphery, not only in fin and gills, but in

the legs as well Now, without for a moment attempting to

explain, fundamentally, the nature of metamorphosis, it is thought,

none the less, that the description just given makes obvious its

intimate relation with nutrition. Looking at the matter thus

from the standpoint of physiological facts, and forgetting for the

nonce our teleology both “old” and “new,” it becomes probable,

a priori, that quick starvation will be more effective than enforced

air breathing in causing the flabby larva to digest its loose and

vascular tissues and take on the more compact form of the adult.

I may now summarize, briefly, sufficient evidence to support the

conclusion that, this is the case.

All of the numerous instances observed of extreme accelera-

tion of metamorphosis have been obviously the result of starva-

tion. The three-gram adult, spoken of at the beginning of this

paper, was the result of a small larva, accidentally overlooked,

and left for several weeks without food, in a jar of water. The

excessively early metamorphosis of nearly fifty specimens in the

cool water of the fountain is also a case in point, the three or

four larvæ that did not metamorphose made little if any growth

during the summer, thus showing the absence of available food

supply. Metamorphosis in the tank of cold spring water was

likewise attributable to this cause; several of the specimens

were under the size at which metamorphosis usually takes

place; but there was no visible food supply ; and the adults

that resulted showed very evident emaciation. Many essentially

similar instances have been observed.

But prolonged starvation is by no means necessary to stimulate

these unstable organisms to sudden change. With full fed larvee

that have reached the length of thirteen to fourteen centimeters

a failure to feed for one or two days is almost certain to be

followed by a number of cases of metamorphosis. Even over-

400 THE AMERICAN NATURALIST. [Vor. XXXVII.

eating, followed by indigestion and bloating, frequently upsets

the metabolic equilibrium sufficiently to induce the change, and

this even in quite small specimens; the accumulation of gas in

the digestive tract causes the animal to float for several days, no

food is taken, and by the time recovery has occurred the irre-

versible gill shrinkage, etc., has set in. A careless observer

might possibly interpret this floating at the surface, followed by

metamorphosis, as an instance where aerial respiration ushered

in the change. There is no occasion for such interpretation,

however: the animal floats on one side, with the mouth and

entire head under water.

Quite in accord with the explanation here given, and con-

stituting a minor confirmation of it, is the fact observed by

myself, and independently by an assistant in my laboratory, that

in any lot of larvae of approximately one age it is not the largest

specimens that metamorphose first; but an intermediate size.

The smallest larvee will not have finished certain developmental

changes which constitute the preparation for the metamorphosis ;

while the very largest are usually fat, phlegmatic, anabolic larvze,

which are less easily stimulated to sudden katabolic change.

Thus in seven experiments, in each of which three larvae were

taken varying somewhat in length, but all of them between

eleven and fourteen centimetres, there was but one instance in

which the largest of the three began metamorphosis noticeably

in advance of its companions, while in four instances the inter-

mediate or the smallest specimen was notably in advance.

Much more striking are the facts noticeable in aquaria containing

many larva, where a few extra large, anabolic individuals fre-

quently postpone metamorphosis for weeks after -— of the

smaller specimens have become adult.

In close connection with such facts as those just stated and

subject to the same explanation is the fact, which I have noted

again and again, that the tendency to sudden metamorphosis is

directly correlated with the disposition or temperament of these

larva. Variable as these animals are, in. no way do they show

greater differences than in their suceptibility to excitement.

One of the first ponds well stocked with Amblystoma that I

discovered was filled with larvae of the usual type, but which had

No. 438]. AMBLYSTOMA TIGRINUM. 40I

reached a size that is quite unusual in the writer’s vicinity.

These larvæ were excessively fat, and so tame that, despite the

clear water, they could almost be taken in the hand. The

specimens removed to the laboratory were equally phlegmatic

there, and underwent metamorphosis little if any sooner than

their relatives in the pond. The next larvæ discovered in the

same summer, likewise in clear water, were so wary as almost to

defy my best efforts with the dipnet, and the excitement of those

taken was incessant, even during the usually sluggish period of

metamorphosis, which, in this case, followed immediately upon

their transference to the laboratory. Restless larvæ have, indeed,

invariably, with me, undergone metamorphosis as the result of

capture and change of conditions. This exceptionally sudden

and rapid metamorphosis is no doubt due to the sudden katabolic

changes induced by the excessive activity. Even with larvae of -

an intermediate degree of excitability, a portion of the individuals

usually metamorphose as the result of removal to new quarters,

if they have attained the dimensions, etc., which makes meta-

morphosis easy. And the writer has been able to discover no

reason applicable to all such cases, other than that the shock of

new conditions, checking or changing food supply a little, always

upsets the accustomed rhythm of bodily functions and thereby

opens the way for change.

The facts thus far cited have, in the main, been instances of

acceleration of metamorphosis, but if their observation and inter-

pretation has been correct, it follows as a natural expectation

that facts of the opposite nature should be forthcoming. Ifa

rich food supply suddenly checked is the common cause of early

metamorphosis, then a moderate but constant food supply should

postpone metamorphosis, until a maximum larval size has been

attained, or perhaps postpone it indefinitely. The experimental

verification of this side of the proof has been far more difficult,

although final success has been reached in a few cases, and these

cases quite confirm the view of metamorphosis here adopted.

Moreover, the frequent failures are easily explained. As already

indicated, our ordinary larve grow rapidly, and tend, in the main,

to early metamorphosis, 7. e., at a length of fourteen centimeters

or less. Excepting occasional larve of a special type, to be

402 THE AMERICAN NATURALIST. [Vor. XXXVII.

mentioned later, the only instances of extra-large larvae which

the writer has met with in nature have been instances of larve. .

from late deposited ova, in ponds -formed late in the season.

Such larvz may live over winter and grow to a very large size

— twenty centimeters — by the next June, when metamorphosis

occurs. The common explanation would doubtless ascribe this

prolonged larval state to the lower temperature of the fall and

spring months. But until experiment has given fuller evidence

of the influence of temperature, the writer would lay stress on

the meagre but constant food supply. For ponds formed late in

the season are poorly supplied with Entomostraca and insect

larvae, which constitute the food of aquatic Amblystome.

But to return to experimental data. The larvæ worked with

thus far have always been rapidly grown specimens; and to

furnish them with an even, sparing food supply in the laboratory

has not, so far, been successfully accomplished. To keep a

constant supply of Entomostraca is not easy ; the substitution of

earthworms proves far too stimulating, if the supply be regular ;

and meat, although they thrive on it, is not better: if chopped

fine most specimens get none at all, so stupid are they in feeding,

and with larger pieces the result is again a too stimulating or a

too intermittent diet. The most prolonged larval growth that

has been reached, however, has resulted from regular feeding

with meat, usually of single specimens in jars. The great point

has been to secure the regular pong of food, and, irideed,

regular habits all round.

In one instance, two small larvae were kept the entire summer

and fall on Lemna minor, a plant which, curiously enough, these

carnivorous larvae will devour in great quantities, although they

obstinately refuse algze and the like. These two specimens (as

well as others similarly fed for a shorter time) made no growth

at all on their abundant vegetable diet. When my stock of

Lemna was exhausted, in the early winter, they refused other .

vegetable food, until one died of starvation. The other, supplied

with meat, grew rapidly, to a length of perhaps eleven or twelve

centimeters, when, after an interval in the feeding, it suddenly

underwent the same rapid metamorphosis which might, under

similar conditions, have befallen it the June before. I instance

No. 438.1 AMBLYSTOMA TIGRINUM. 403

this single case to show that mere delay, mere prolongation of

the larval period, does not in itself necessarily hinder metamor-

phosis. Stimulate the flagging life activities and suddenly check

the advancing anabolism, and metamorphosis will follow as easily

as had growth been rapid from the egg. The opposite opinion

has been held by Marie von Chauvin and others.

Laboratory experiments failing, success in rearing larvee of the

largest size was finally reached in cisterns. Although, here too,

success only followed after repeated failure. When larve of

twelve to fourteen centimeters were introduced into cisterns they

invariably underwent metamorphosis as a result of the shock

and check in food supply. Very small larve, introduced in

numbers, betook themselves to wholesale cannibalism, resulting

in irregular growth and early metamorphosis or death. But

larva of six or seven centimeters, introduced into cisterns in

small numbers, did, in some instances, establish themselves there

with interesting results. Several experiments of this kind have

been carried out or are now in progress. In one instance, three

larva withstood metamorphosis for one, two and three years,

respectively. The last was taken from the cistern on the fourth

summer, still in the complete branchiate condition, although,

from faint signs, it is judged that it would probably have become

adult during the summer. It was a male and rapidly approaching

sexual maturity, the testes showing the first division of the sper-

matids. These larve attained the size of large adults in about

sixteen months, growth, in the cistern, being thus only one

quarter to one fifth as rapid as the growth of feral or meat-fed

specimens. This slow growth indicates that the food supply

was light, and the steadiness of the growth, that it was rela-

tively constant. :

As a final proof that the metamorphosis of Amblystoma is due

to checked nutrition rather than to respiration of air, it seemed

advisable to experiment with these larve under water, 7. €., to

subject them to sudden starvation while quite preventing their

coming to the surface for air. As the adult Amblystoma fre- .

quently passes hours and perhaps days under water, and as

observation had shown metamorphosis to take place normally,

in the water, why should not the complete. change take place.

404 THE. AMERICAN NATURALIST. [Vor. XXXVII.

under water? Marie von Chauvin found that the larva of

Salamandra maculosa would not, at least contentedly, retain its

gills under water; but her experiments were not carried to their

full conclusion.

The attempts to produce the perfect adult Amblystoma under

water were not successful, although they fell little short of suc-

cess. Advanced stages of metamorphosis were produced; in

one instance only a trace of tail-fin and gill-stubs remaining.

As already indicated, the great difficulty encountered was the

unexpected fact that all of the larvae are air breathers, no mat-

ter how complete the development of the branchiz. No larva

could be found capable of living for more than twenty-four hours

below the surface of standing water, no matter how pure, or how

large the volume. "Their tenacity of life varied much. In a few

instances, by repeatedly reviving the semi asphyxiated animal in

air, or in running water, and replacing it in well aerated water,

under the netting, a partial adaptation seemed to be brought

about. Fewer movements were made and no distress or impulse

toward the surface would be shown for many hours. But grad-

ual or acute suffocation always occurred sooner or latter. When

a small stream of water was let fall directly into the aquarium —

in this case usually only a battery jar — the results were better ;

although many did not live long, when situated even thus, and

when the stream was checked, even for an hour, our very best

water-breathers always made efforts to reach the surface.

Although complete metamorphosis was not reached under

water, yet the tbeoretical importance of the partial metamor-

phosis which did result is hardly less than would have attached

to the completion of the process. The impulse to metamorphose

was plainly present. Over thirty larvae were tried in running

water, below netting; and not one was found, of suitable size,

that could there resist the tendency to starvation-metamorphosis.

All, which did not die of some other cause, began metamorphosis,

usually within six days. Sometimes, indeed, the metamorphosis

was begun almost immediately, as if the first shock of transfer-

ence and partial oxygen-starvation were enough, despite the low

temperature, to induce the change. Such individuals, did not,

however, survive long; those which carried metamorphosis to

No. 438.] AMBLYSTOMA TIGRINUM. 405

completer stages being such as began slowly, after at least

several days of contented life below water.

Before leaving these experiments, I will note that, in spite of

their fatal results and the convincing testimony they bear to the

inability of these organisms to make complete adaptive response

to even moderate environmental change, yet they did seem,

sometimes, to indicate, some attempt at adaptive response,— in

this case an attempt to retain the branchiate condition. For

when the fatal resorption of tissues began, the gills were not, as

is usually the case, one of the first organs to be attacked. In

some instances they showed no diminution in size or function

for several days after the back fin was greatly reduced. Such

instances also occur in larva in ordinary circumstances; but

they are not frequent, and may perhaps, in all cases indicate

that a high functional activity of the branchie is for a time

holding in check the katabolic changes which are elsewhere set-

ting in. Similar observations may be made on the occasional

overgrown, giant larvee, in which the branchial apparatus seems

frequently to resist metamorphic processes until other parts of

the organism are well under way in the change.

A fairly complete résumé has now been presented of the

classes of facts which seem to indicate that the metamorphosis

of Amblystoma, if not of all tailed amphibia, has heretofore

been, to some extent misinterpreted, and that the chief factors

in the process are always sudden shifts in metabolism, usually,

or at least most easily, induced by changed food supply. It

remains to answer one or two queries that naturally arise, and

to indicate one narrow group of cases that, at first Observation,

seem against the present hypothesis.

In the first place, it may very well be asked, if sudden destruc-

tive metabolism is the cause of metamorphosis, why should not

air breathing favor the change? It would seem to be favorable

to rapid oxidation of tissue. The answer is simple: theoretically

it should be favorable, and in very rare instances it may actually

beso. The writer has known one single instance of three larvae

which showed astonishing activity when accidentally removed

from the water, and they underwent the change rapidly. But

in all ordinary cases, larvae, when quite removed from water or

406 THE AMERICAN NATURALIST. [Vor. XXXVII.

left in water which but partially covers them, become at once

inactive. After a few efforts to crawl, and ineffectual turnings

about, as if in search of water again, they “settle down, and

wait for rain.” This inactivity partially compensates the first

effects of starvation; so that these air breathing larve may

metamorphose even less quickly than similar individuals sub-

jected to starvation, but kept in water which encourages move-

ment.

Second: What of the numerous observations on the meta-

morphosis of Amblystoma in the air, as the result of the drying

up of ponds? Do not these, after all, show that metamorphosis

frequently follows the enforced use of lungs? They do, indeed,

show that metamorphosis follows the enforced use of lungs; but

not that it is caused thereby. A moment’s thought will show

that this naive, natural interpretation may well be at fault. As

the water of a pond evaporates, what is the first result for these

larvae? Plainly a concentration of their food supply. Insect

larvee and entomostraca, moderately abundant before; become

now indefinitely easy of access. The larvae gorge themselves, for

a few days, to repletion. Suddenly the last stages of the

accelerating evaporation place them at a disadvantage ; freedom

|. of movement is checked, or they find themselves partially out of

the water. Experiment shows that under such circumstances

the larvae at once cease feeding altogether. The resulting

metamorphosis is obvious, and need be due to no other causes

than those which the writer has found effective under experimental

control.

It may seem more presumptuous to offer a similar explanation

for the results obtained by European experimenters on Ambly-

stoma and interpreted by them in so different a manner. Yet a

careful reading of a large part of the literature on the subject

seems to the writer to inevitably suggest that many of their

results are explicable as above suggested. This is especially the

case with the series of elaborate experiments made by Marie von

Chauvin. Fearing that her charges would die, as indeed they

sometimes did, she always prepared them for the trying ordeal

of metamorphosis by raising the temperature of the water in

which they were kept (to this she ascribes very great importance),

No. 438.] AMBLYSTOMA TIGRINUM. 407

and feeding to the maximum for several days (to which she

ascribes no other importance than giving the animals increased

strength). The animals were then brought immediately into

water sufficiently shallow to force them, for at least part of the

time to breathe air. In this latter condition the experimenter

complains again and again that it was next to impossible to induce

the Axolotls to take any food whatever. Thus in these experi-

ments, too, we have high feeding followed by practical starvation ;

and it seems that no control experiments were instituted to

determine what the effects of over and under nutrition might

have been with Axolotls still in an abundance of water. Yet

more interesting is it to note that even the varying degrees of

success and failure in inducing metamorphosis in these experiments

follow closely parallel to varying factors of nutrition, which factors

seem to have been wholly neglected in the final interpretation of

the results. Miz., as artificial methods of feeding were developed,

earthworms being made-to crawl down the throats of the refrac-

tory larvae exposed to the air, it became proportionately difficult

to induce metamorphosis by withdrawing them from the water.

Even for the space of thirteen months, larvae whose nutritive

equilibrium was thus maintained fairly constant, would sometimes

retain their aquatic organs despite their exposure to air. :

Before closing, brief reference should be made to one class of

exceptional cases of retarded metamorphosis which is of great .

interest, and which might at first thought, seem to contradict

some of the considerations set forth in this paper, while offering,

however, no support to the ordinary hypothesis. I refer to

certain cases of greatly overgrown larvae, differing in important

respects from the ordinary type. It was one of these that I

mentioned as a “giant larva," at the beginning of the paper. I

hope later to describe this interesting form in detail. At present

I will only mention that it is separated from the ordinary type

by its much broader and longer head, with parallel instead of

converging sides; its enormous gap of mouth ; its flattened or

usually concave profile, instead of the even curve of the common

larva and adult. It has, besides, unusually heavy limbs, with, in

most cases, much flattened toes and webbed feet. These larva .

exceed, by several centimetres in length, the size of even the

408 THE AMERICAN NATURALIST. [Vor. XXXVII.

largest larvae of the ordinary type which I have as yet seen.

The adults resulting from their metamorphosis are nearly as

distinct as are the young. They represent, in many features an

extreme development of what Cope has designated the ** Western

form" of Amblystoma tigrinum. But, in the writer's vicinity,

they seem to constitute only a very small minority of the species,

not more than one out of several hundred.

This form, as already indicated, seems to be very resistant to

metamorphosis; it has shown itself so under experiment; and,

in ponds where the ordinary form leaves the water at a very

small size, these occasional giants may continue in the larval state

until they have outgrown all but the very largest adult members

of the species. There is thus, in the case of these larvae, plainly

another factor present, in the retardation of metamorphosis.

And, as both the mature and the immature stages of this form

resemble, morphologically, the perennibranchiate types in several

respects, it seems, at present, natural to interpret them as rever-

sions toward a more primitive and perhaps perennibranchiate

ancestor.

But although these giant, flat-headed larvze resist metamorphosis

without the especial retarding conditions which usually prevent

the change, yet it is to be noted that they, too, undergo meta-

morphosis in the water, and that high feeding, followed by

partial feeding or by starvation will apparently invariably bring

about the result. I have taken several giants in the early stages

of metamorphosis from deep water ; and all the specimens in my

possession have finally undergone metamorphosis in aquaria.

The branchial apparatus seems more resistant than usual; the

whole process takes longer; and the order in which the several

component changes occur is different; but the physiological

aspect of the process, its causes, etc., remain the same, aside

from the matter of delay. :

A word, in conclusion, as to the significance and setting of

these facts in general theory. The writer does not, with

Weissmann, hold that the facts relative to the metamorphosis of

Amblystoma or the Axolotl may be sufficient to settle cardinal

differences of view in biological theory. The conduct of no

one species can be conclusive, or, indeed, more than merely

No. 438.] AMBLYSTOMA TIGRINUM. 409

suggestive, however completely it accords with one theory and

discredits another. But if the observations and interpretations

noted in this report prove correct for all Amblystomze, it is

evident their bearing on the theory of variation is not unimportant.

The variations in metamorphosis of the amphibia in general, and

especially of Amblystoma, have been looked upon by many as an

excellent example of the direct, purposive response of organisms

to their environment. Thus, in the Addenda to his work on the

Batrachia of North America, Cope quotes a writer who deems

these animals plastic and responsive in the highest degree, —

metamorphosis and remetamorphosis following upon the pre-

dominance of aquatic or terrestrial conditions as if the physi-

ological processes of these animals were gifted with both fore-

sight and free will. Marie von Chauvin, at the close of her last

article says: * Es sheint den Axolotln eine eminente Befáhigung

inne zu wohnen, sich den gerade gegebene Lebensbedingungen

anzupassen." The writer has no antipathy to the teleological

view of variation, providing facts can be found to demonstrate

it. Indeed, observations on Amblystoma were undertaken, with

the express purpose of giving, if possible, greater precision to the

interpretations suggested by Cope and ethers. But the facts, as

here outlined, all tend to support an opposite view. It may even

be said that, in the writer’s vicinity, the one representative

species of Amblystoma survives for no other reason than that, in

its larval condition it can appropriate large quantities of food

for which there are few competitors. Its other adjustments,

instinctive, physiological, and structural, seem very imperfect.

The species is indeed plastic ; but not purposive, or approximately

perfect in its reactions. And in no particular is this more true

than in respect to the various structures and structural changes

by which these animals are supposed to adapt themselves to

aquatic and terrestrial life.

After all, is not this view borne out by the natural history of

the Amphibia in general, with the exception, perhaps, of the more

progressive, modern type Anura? A half score of forms become

lungless in the adult condition, —a degenerative change of doubt-

ful adaptive value. Cryptobranchus and Amphiuma are perma-

nently aquatic, yet without branchie, despite their probable

410 THE AMERICAN NATURALIST. [Vor. XXXVII.

descent from branchiate forms. Siren and Pseudobranchus,

although especially aquatic types, exhibit the strangest anomalies

of development and retrogression, of use and disuse of gills.

Many forms of Amphibia develop gills in the egg, or in the

intrauterine state, which are destined to serve no purpose in

aquatic life, even when an aquatic larval state is soon to follow.

Of course there may be more adaptation than we know in these

anomolous conditions; but do they, in our present state of

knowledge, constitute a proof of adaptation or of direct response

to environing conditions ?

DoANE COLLEGE,

March, 1903.

THE ORIGIN OF THE SPOROPHYTE.

BRADLEY MOORE LAVIS.

ONE of the most interesting peculiarities of plant life is the

striking alternation of generations characteristic of all groups

above the alga and fungi (thallophytes). It is customary to

call these two generations by names that indicate the end of

their life activities. The gametophyte is the sexual plant, devel-

oping the sexual cells or gametes. The sporophyte is asexual,

producing spores. These functions of gametophyte and sporo-

phyte have very few exceptions among the higher plants, the

latter falling under the head of apospory and apogamy and

resulting, at least, in part from abnormalities of environment.

The fertilized egg always develops into a sporophyte and the

asexual spore, on germination always produces the gametophyte.

The life history becomes then a rhythmical alternation of

gametophyte and sporophyte only disturbed when a generation

introduces organs for vegetative reproduction and thus makes

possible a series of like individuals before the next generation.

Such methods of vegetative reproduction by brood organs

(gemmzae), buds, bulbs or various fragments are not uncommon

in the gametophytes of mosses and liverworts and are very com-

mon in the sporophyte generations of the seed bearing plants.

Since it is vegetative reproduction and the offspring are literally

chips from the parent block they in no way effect the underlying

principles governing the alternation of generations. .

These principles have become better understood with advances

in our knowledge of cell structure in plants, and certain very

interesting facts are now known which indicate that the pecul-

iarities of gametophyte and sporophyte are due to structural

conditions of the protoplasm that may in a measure be deter-

mined. So the problems have become largely an analysis of the

events in the cell activities preceding the formation of sexual

cells, also at fertilization, and immediately afterward with the

development of the fertilized egg, and similarly the period of

411

412 THE AMERICAN NATURALIST. [Vor. XXXVII.

spore formation and germination of the asexual spore has been

made the subject of careful study. It is evident that the criti-

cal periods of the plant’s life, when the changes are the greatest,

come with the passing of one generation over to the next or in

other words at the periods of gametogenesis and sporogenesis

with the early developments following each of these processes.

We have now an accumulation of studies upon this subject from

all three of the great groups of higher plants (bryophytes,

pteridophytes, and spermatophytes) and in certain regions the

investigations have been numerous. They have uniformly

yielded the same results in certain particulars that have estab- .

lished a foundation for some very interesting speculations on the

essential differences between the sporophyte and gametophyte.

It will be apparent that these differences must be very fun-

damental because they are the basis of extensive evolutionary

processes with the universal tendency to separate the gameto-

phyte and sporophyte further and further from one another in

structure and life activities. From the conditions among the

bryophytes in which the sporophyte is so closely associated with

the gametophyte as to have been called its fruit by the early

botanists, we pass through the pteridophytes to the spermato-

phytes where somewhat analogous conditions are found in which,

however, the relations. between the two generations are exactly

reversed. Among the seed bearing plants the gametophyte has

become so reduced as to live parasitically upon the sporophyte

passing its life in the interior of the asexual generation which is

thus made the phase that performs the vegetative activities that

we expect of plant life. There is no more interesting province

of plant morphology than that which traces the evolution of the

sporophyte and degeneration of the gametophyte as one passes

from the liverworts to the higher plants. There are involved in

these processes.a number of evolutionary principles of the

greatest significance but they do not fall within the range of the

present paper.

Can we establish a physical basis for the differences between

sporophyte and gametophyte in the cell and nuclear activities

during the life history. This has been attempted chiefly through

the study of the number of chromosomes present in the nucleus

No. 438.] ORIGIN OF THE SPOROPAYTE. 413

at the critical periods when one generation passes into the other.

The results have been remarkable. The nucleus of the game-

tophyte generally presents a fixed number of chromosomes as

shown in the figures of nuclear division in the various tissues.

We may let x stand for this number, which is generally not very

large (e. g., 4 in Anthoceros, 8 in Pelia, 12 in Osmunda, 12 in

the lily, etc.) The nuclei in the sexual cells (gametes) have this

gametophyte number (x). Their fusion, during the sexual act,

gives a sexually formed spore whose nucleus has 2x chromo-

somes, double the number of the gametophyte. This double

number (2x) prevails through the entire sporophyte generation

up to the time when the sporogenous tissue (archesporium)

appears. The events of sporogenesis reduce the number by one

half bringing the asexual spore back to the condition of the game-

tophyte with x chromosomes.

This cycle by which the number of chromosomes is doubled

and then reduced probably holds true for all plants above the

thallophytes, the conclusions ^being ‚based upon the study of

several liverworts and pteridophytes and a larger number of

spermatophytes. The essential facts of the history may be

expressed in an abbreviated form as follows. ;

d gamete

pesar ud en x chromosomes Fertilized HE sporophyte — Asexual — — Gametophyte.

X Chro s 9 gamete 2 X Chrom p = — dire —

Reduction of the chromosomes is a phenomenon that has been

much studied in connection with sexual, processes, especially

among animals. It is believed to result, from the supposed

necessity of keeping the chromosomes in all organisms relatively

fixed in number. Since the number of chromosomes is doubled

with every nuclear fusion in the sexual act the amount of chro-

matin would increase indefinitely and in geometrical progression

were there not a device for diminishing the quantity at some

time in the life history.

If the question is asked why are the chrombsomes so impor-

tant and why should their number be so significant, no answer

can be very satisfactory for our deep ignorance of the function

of the nucleus is exposed. However, the more detailed the

studies upon the structure and activities of the cell the greater

414 THE AMERICAN NATURALIST. [Vor. XXXVII.

has become the conviction that the chromosomes carry the keys.

to many and probably the most important problems of develop-

ment and heredity. We do not. know what the chromosome

does but its characteristic activities during nuclear division and

its behavior at critical periods in the life history are so remark-

able that the assumption of its importance in these events is

quite justified. The most attractive theory of reduction phe-

nomena assumes that specific characters are largely defined by

the amount and nature of the chromatin in the nucleus and that

a species, to keep true, must so provide that the chromatin con-

tent is relatively stable from generation to generation.

Reduction of the chromosomes at some period of the life his-

tory is almost universal among higher animals and plants but we

should note an important difference between the two groups in

the manner in which this is accomplished. Briefly stated for

animals, the reduction occurs just before the formation of the

sexual cells (gametes) which have in consequence one half the

number of chromosomes characteristic of the organism. The

fertilization of the animal egg by the sperm brings the male and

female nuclei together and as a result of their fusion the number

of chromosomes becomes again normal.

In plants above the thallophytes the history is very different

and in sharp contrast to that of the animal. There is no reduc-

tion at the time when the gametes are formed. The gametes

have consequently the same number of chromosomes as the sex-

ual plant (gametophyte). Their fusion gives to the sexually

formed spore double the number characteristic of the gameto-

phyte. This fact is believed to be largely responsible for the

peculiarities of the asexual generation that follows. The sporo-

phyte runs through its vegetative development, without any

change in the double number of chromosomes, to the time of

spore formation when the sporogenous tissue (archesporium) is

differentiated. There is then a period of growth during which

some or all of the archesporial cells become spore-mother-cells.

And during that preparation for spore formation (sporogenesis)

the number of chromosomes is reduced by half, becoming again

the number of the gametophyte. The reduced number first

appears in the nuclear divisions inside of the spore-mother-cell

No. 438.] ORIGIN OF THE SPOROPHYTE. 415

preparatory to the formation there of the characteristic group of

four spores.

It will thus be seen that there can be no genetic relationship

between the reduction phenomena of higher plants and animals.

They are not found at the same points in the life history and

there are also fundamental differences in the details of the pro-

cess that cannot be considered here.

With these points understood the sporophyte generation takes

on new interest in relation to sexual processes in plants for it

appears to be a development peculiar to this group of organisms

and probably related to a form of sexuality that differs from

that of animals in important respects. These differences con-

cern the maturation of the gametes which in animals takes place

with a very characteristic process of chromosome reduction

and in plants without this phenomenon.

So little is known about the origin of sex in animals that sat-

isfactory comparison with the much better understood history

for plants, is not possible. We have already discussed that topic !

and also sexual evolution.”

As there is no reduction phenomena in plants at the time

when sexual cells are formed perhaps we find in this fact a clue

to the reason of the sporophyte generation and its origin. It

must be clear that the most promising line of investigation

would deal with the doubling of the chromosomes at the sexual

act and the later reduction to the gametophyte number at the

end of the sporophyte generation.

The final explanation of the problems can only come through

the study of plants below the Bryophytes, that is among the

Thallophytes, and we have had as yet very little detailed research

on this point. So this paper must deal largely with speculations.

We know positively the main facts of sexual processes and chro-

mosome reduction in groups above the Thallophytes but for this

group almost nothing.

Nevertheless, we are justified in considering an hypothesis of

1 Davis, The Origin of Sex in Plants. Popular Science Monthly, Nov. 1901,

. 66.

2 Davis, The Evolution of the Sex in Plants. Popular Science Monthly. Feb.

1903, p. 300.

416 THE AMERICAN NATURALIST. [Vor. XXXVII.

the origin of the sporophyte and its relation to reduction phe-

nomena and this hypothesis may be tested among the thallo-

phytes and must stand this test if the suggestions are to become

a theory.

An explanation of the sporophyte upon a physical basis must

consider the problem in terms of protoplasmic organization. It

must attempt to isolate the structures and qualities given to the

egg by fertilization. It must determine the potentialities of the

sporophyte generation and also explain why this structure should

finally produce spores whose protoplasm returns to the condition

of the gametophyte. |

The sexual act in all plants above the thallophytes doubles the

number of chromosomes. This is the only morphological change

that we can observe in the structure of the protoplasm. It is

scarcely probable, however, that the doubling of the chromo-

somes introduces all of the changes that come over the fertilized

egg when it takes on the qualities that compel its development

into a sporophyte. There is of course the mingling of many

substances when the sperm fuses with the egg and we have good

reasons to believe that all regions of the protoplasm are effected.

But the nucleus gives us the most marked evidence of change in

structure and this is shown conspicuously in the number of

chromosomes. That the potentialities of the sporophyte are

chiefly bound up in the protoplasmic structure of the egg, there

can be little doubt. Fertilized eggs of plants above thallophytes

cannot be made to develop gametophytes. They are wound up,

to use a homely expression, to produce sporophytes and this

they will always do under normal conditions. Some peculiar

forms that omit certain stages in the alternation of generations

are believed to be the products of unusual environmental rela-

tions. They are apogamous or aposporous or examples of regen-

eration and are the exceptions that prove the rule.

The potentialities of the sporophyte generally remain indefi-

nitely long in all regions of the plant where the tissue is

embryonic in character, z. e., at all growing points or meris-

tematic areas. These tissues will reproduce the sporophyte or

add to its growth as long as they live and retain their undiffer-

entiated character. 7

No. 438.] ORIGIN OF THE SPOROPHYTE. 417

But more specialized and older parts of the plant show other

characteristics. Some of these regions become specialized for

various vegetative activities, assimilation, conduction of water,

storage, etc.; some become protective, as the epidermis and

cortical regions, and some strengthening. There comes however

sooner or later to certain of these older portions another activity,

that of spore formation or sporogenesis.

Sporogenesis in the simpler sporophytes generally involves

extensive regions of the plant, sometimes almost the entire

structure as in certain liverworts (Ricciales). However the

evolutionary tendency among higher forms is to develop more

extensively the purely vegetative tissues thereby reducing pro-

portionately the spore bearing areas. This very important prin-

ciple involves the sterilization of sporogenous tissue and may be

traced in a very interesting manner as the sporophyte increases

in complexity until the sporogenous tissue becomes confined to

special organs, termed sporangia. But we cannot consider

this topic at the present time. * ;

The significant result for us is the fact that at some period in

the history of every sporophyte certain tissues return to the

potentialities of the gametophyte and reproductive cells are

formed which can only grow into gametophytes. The mechan-

ism, wound up by the act of fertilization, runs down in a figura-

tive sense and the protoplasm, losing its sporophyte characters,

returns to the dead level of its ancestral plasm from the gameto-

phyte.

The only evidence of this reversion to gametophyte conditions,

as shown by the structure of the protoplasm, is in the reduction

of the chromosomes. This occurs just before the sporogenous

tissue (archesporium) is ready to form spores. The young cells

of the archesporium emerge from their last vegetative division

with the sporophyte number of chromosomes in their nuclei.

There is then a period of enlargement during which some or all

of the cells are stored with a rich supply of protoplasm becom-

ing spore mother cells. During this period of enlargement

the reduction phenomena takes place probably by the fusion of

the chromosomes in the resting nucleus. The resting nucleus

at this period is for some time in the state called synapsis when

418 THE AMERICAN NATURALIST. [Vor. XXXVII.

the chromatin network is very much contracted, a very con-

spicuous condition which is not well understood. The first

nuclear division in the spore mother cell presents the gametophyte

number of chromosomes and this is followed by a second divi-

sion so that four nuclei result, each of which is destined to

preside over a spore.

The fact that the number of spores formed in each mother

cell is four appears to have no important morphological signifi-

cance. Ithas no connection with reduction phenomena which, as

explained above, take place before these divisions. Among the

seed plants this division is very frequently omitted in the mega

spore mother cells which give rise directly to the gametophyte

(embryo sac) thus cutting out a portion of the usual history in

the development of spores.

However the fact that four spores are formed in each spore

mother cell is interesting because almost universal among the

bryophytes and pteridophytes and characteristic of pollen for-

mation in the Spermatophytes. We can see no reason why the

number should be four nor is any light thrown upon the prob-

lem from our knowledge of the thallophytes.

To summarize this explanation of the sporophyte; we assume

that the fusion of gametes (fertilization) gives to the sexually

formed spore a different mechanism from the parent gametophyte

and this mechanism, under normal conditions, runs a course, pro-

ducing the sporophyte. The result is a differentiation of the

cells through a constant tendency to develop vegetative regions

(somatic) in contrast to the reproductive tissue which is propor-

tionally reduced in quantity. The latter (archesporium) gener-

ally appears at certain periods of development and in definite

regions and represents the return of the sporophytic plasm to the

potentialities of the gametophyte. We do not mean to imply

that the entire development of the sporophyte is regulated from

within, which conception would be ultra preformation. There

must be numerous external factors (epigenetic) influencing the

vegetative regions and affecting the organography of the plant

and certainly the periods of spore formation. But there seems

to be the best of evidence that the initial stimulus to sporophyte

development comes from within. These are topics that may be

No. 438.] ORIGIN OF THE SPOROPHYTE. 419

better discussed in a consideration of the evolution of the

sporophyte than under the title of the present paper.

The origin of the sporophyte involves the examination of con-

ditions previous to the bryophytes, 7. e., among the groups of the

algze and fungi (thallophytes) and we will now consider these.

As is well known the algæ and fungi present organisms far

more various in the succession of forms comprising their life

histories than the higher plants. There is no general rule of

development in this assemblage of diverse groups but rather a

number of habits presented by the divergent lines in relation to

their various modes of life. In this respect the thallophytes

stand in sharp contrast to all plants above the bryophytes. The

antithesis of sporophyte and gametophyte, if present at all, is

greatly obscured by the intercalation of many and various means

of asexual reproduction which may give an indefinite and very

irregular succession of individuals. For many years botanists

have attempted to define among these lower plants life histories

comparable to the alternation of gametophyte and sporophyte.

Some of these suggestions appear probable, others have been

completely disproved by experimental studies on development.

The test of an alternation of generation involving a game-

tophyte and sporophyte must always lie with the activities of

the sexually formed spore. If this cell invariably produces a

phase different from its parent plant then we may properly

inquire whether such a structure does not have in itself potenti-

alities that separate it as a distinctly new form of development.

.It may then be called a sporophyte in contrast to the gamete

bearing individual and the succession of generations will be

antithetic at this point in the life history.

It does not matter how numerous are the successions of gen-

erations which depend on various methods of asexual reproduc-

tion. They may occur directly before or after the sexual act or

over a long intermediate period. Such generations are called

homologous since they all produce the same form of plant.

They do not affect the contrast of gametophyte and sporophyte

whenever that change occurs in a life history.

The establishment of alternation of generations among the

algæ and fungi depends on the determination of the various

420 THE AMERICAN NATURALIST. [Vor. XXXVII.

phases as either homologous or antithetic They are homologous

when they can be made to develop indefinitely the same form of

the plant successively, azzzhetic when the sexually formed spore

always gives rise to a distinctly new phase. This test of the

conditions in the life histories of the thallophytes is being gener-

ally applied throughout the group as opportunities are presented

and has led to some interesting results.

Some forms, that at one time were thought to present tenden-

cies towards an alternation of generation, are now known to give

merely a succession of homologous phases. The zygospore of

the moulds (Mucorales) and the eggs of Vaucheria, the Sapro-

legniales and probably the Volvocaceze may on germination pro-

duce a series of asexual generations or they may pass at once

back to the sexual plant according to the environmental condi-

tions. So there can be no antithesis of sporophyte and game-

tophyte among these types. These facts have been thoroughly

established experimentally by Klebs and illustrate clearly the

method by which the problems are attacked. However, the

details of the environmental conditions under which such plants

become sexual or asexual carinot be given here.

Again, certain thallophytes whose sexuality is highly devel-

oped present life histories with but one phase, a sexual plant,

and without the least tendency towards the development of a

sporophyte generation. Notable examples are found in the

Characez and Fucaceze where the eggs on germination produce

a sexual plant like the parent. The fact that the oospore of

Chara develops a small filamentous structure preliminary to the

establishment of the characteristic later growth does not affect

the general principle. Investigations on the nuclear history of

Fucus report a chromosome reduction after a peculiar manner

that cannot be reconciled with the conditions known for groups

above the bryophytes. The number of chromosomes is reduced

one half just before the differentiation of the oogonium so that

the gametes have half the number of chromosomes characteristic

of the parent plant. After fertilization the egg has again the

regular number and naturally develops into a plant like the par-

ent. The life history may be outlined in this manner.

gametes

Sexual plant <4 X chromosomes > Fertilized egg — Sexual plant.

X chromosomes 9" gametes X chromosomes

No. 438.] ORIGIN OF THE SPOROPHYTE. 421

A comparison of this formula with that given at the beginning

of this paper for all plants above the Thallophytes will make the

peculiarites of Fucus clear. "The reduction phenomena comes at

the same period as in animals, 7. e., previous to the formation of

the sexual cells, and the fertilization of the egg brings that struc-

ture back to the potentialities of the sexual plant. In contrast

to Fucus, the conditions among the Characez appear to be very

different. Investigations here have. failed to establish a reduc-

tion of the chromosomes at the time the gametes are developed.

We do not as yet know where the process occurs in this group.

These conditions in Fucus are unexpected and cannot be easily

brought into sympathy with the processes of gametogenesis as

illustrated in the higher plants. It is the only form among the

thallophytes in which the count of the chromosomes has been

made for the life history, and until we know the conditions in

some of the other types it must remain a puzzle. We are quite

in the dark as to its significance in the general problem of sexual

evolution among plants and in its relation to alternation of gen-

erations. It may present an exception to the usual history of

chromosome reduction among Thallophytes and this is to be

hoped as otherwise further studies may greatly complicate the

problems and require a reconsideration of the theories of the

evolution of sex in plants and the origin of the sporophyte.

There are left several groups of thallophytes which are gener-

ally supposed to present phases in their life history that are

either true sporophytes or indicate tendencies in that direction.

The most pronounced evidences of a sporophyte generation are

furnished by the red algze (Rhodophycez), Coleochæte and the

Ascomycetes. Tendencies in the direction of such alternation

of generations are probably shown in CEdogonium, Sphzroplea,

Ulothrix and the Conjugales.

The sexual reproduction of the red algze presents peculiarities

that have puzzled investigators for many years. The female

gamete (carpogonium), after fertilization, gives rise to a more

or less complex system of filaments that always remain attached

to the parent sexual plant and in many forms are undoubtedly

dependent upon it in part for nourishment and protection. Por-

tions of these filaments and sometimes almost the entire struc-

422 THE AMERICAN NATURALIST. [Vou. XXXVII.

ture become spores which on germination develop the sexual

plant. In 1898 Oltmanns presented the suggestion that the

filamentous growth from the female gamete was a true sporo-

phyte. Although we lack the confirmation required by the

close study of the details of nuclear activities, nevertheless the

theory is very satisfactory especially in its explanation of certain

peculiarities of this interesting process of development.

The simplest sporophytes among the red alga are illustrated

* gs f] : 4

Ficus

g f supp I phy he Rhodophycex ; shaded

structures sporophytic ; (x) auxiliary cells ; a, 2, c, Nemaiion ; d, e, f, g, A, Callithamnion.

by such forms as Nemalion (See Fig. 1, a, 6,.c) and Batracho-

spermum. Here we have a cluster of short filaments many or

most of whose cells become spores. But among the higher red

algee the conditions are greatly complicated by certain cell

fusions between the sporophytic growth and the filaments of

the sexual plant (gametophytic). Certain cells of the game-

tophyte are set apart as large richly nourished auxiliary cells

whose functions are to assist the developing sporophyte. The

fusion of the sporophytic filaments with the auxiliary cells stimu-

lates the growth of the former. It was at one time supposed

to be sexual in character but is now believed to be for nutritive

purposes alone. Apparently there are no nuclear fusions in

these unions of cells but only the mixing of cytoplasm with its

No. 438.] ORIGIN OF THE SPOROPHYTE. 423

opportunity for the extensive transfer of food material. The

main points in this interesting process are illustrated for Callith-

amnion in Fig. 1, Z, e, f, g, A

The sporophyte of the red algae holds a relation to the game-

tophyte somewhat similar to that in the bryophytes in as much

asit is always attached to the latter generation. The sporo-

phytic growth accompanied by developments, usually protective,

on the part of the gametophyte constitutes the so-called cysto-

carpic fruit of the Rhodophycea. It should be noted, however,

that this sporophyte does not exhibit the upward evolutionary

tendencies towards independence so characteristic of the higher

plants but rather a degree of dependence approaching parasi-

tism.

= LNT

a Ars E MA

am of ascogenous hyphz (shaded) in relation to

(Somewhat modified after Harper).

Fic. 2. Pyrone a, Group of gametes; b, diagr

the S duse: ad Lade rte of the inn plant.

The Ascomycetes furnish conditions somewhat similar to the

Rhodophycez but here also we lack precise knowledge of

nuclear structure at important periods of development. The

female gamete, called here the ascogonium or archicarp,

develops a system of filaments (ascogenous hyphae) so closely

associated with the parent organism that they can only be dis-

tinguished through special staining processes.

The ascogenous hyphz develop asci, a form of sporangium

424 THE AMERICAN NATURALIST. [Vor. XXXVII.

peculiar to the Ascomycetes, which are generally associated

together and surrounded by an envelope thus constituting a

fruit called the ascocarp. These conditions are illustrated

somewhat diagrammatically in Fig. 2 for Pyronema, the form

perhaps most completely studied.

Fic. 3. Coleochate: a, Group of antheridia ; 4, oógonium ; c, fertilized egg; d, odspore in cellu-

lar envelope; e, germinating odspore showing tissue in its interior (sporophytic). (After Olt-

ns.)

Coleochzete is a type upon which those who believe in a spor-

ophytic generation among the thallophytes lay great stress.

The principal reason for this emphasis is a certain resemblance

in form between a cellular body developed by the fertilized egg

and the simple sporophytes of the lower bryophytes. As is

shown in Fig. 3, 4 and e, the fertilized egg after a period of rest

develops in its interior a tissue each of whose cells gives rise to

a zoóspore which on germination produces a sexual plant. This

structure is very suggestive of the sporophyte of the simplest

liverworts (Ricciales), but we know nothing about the chromo-

some history during its development. It is not safe to relate

the liverworts to Coleochzete because the sexual organs are so

different, for the archegonium is an organ that can hardly be

derived from so simple a structure as the unicellular oógonium.

However we can readily conceive the Coleochztacez as a group

tending to develop a sporophyte generation along somewhat

parallel but totally independent lines from the Bryophytes.

Coleochzte is generally cited as the Thallophyte in which a

sporophyte generation is most evident but in reality the develop-

No. 438.) ORIGIN OF THE SPOROPHYTE. 425

ments from the female gametes in the red algæ and the Ascomy-

cetes are far more elaborate and show a much higher grade of

structural evolution. But the fact that the apparent sporophytic

tendencies in these latter groups are long divergent lines related

to the peculiarities of their modes of life and so very unlike that

of the bryophytes has not given them the degree of attention in

relation to the general problem that they deserve. If future

studies upon their nuclear conditions support the theory of

their sporophytic nature these structures should be cited as the

highest forms of sporophyte among the thallophytes but of

course in groups very far removed from the main line of ascent

to the bryophytes.

Fic. 4. Germination ot the sexualiy formed spores of a, Ulothrix ; 4, "scm cu €, d,

Oedogonium. (a, after Dodel-Port; 4, Cohn; c, Pringsheim; 4, Jura

There are left several algg somewhat similar to one another

in the behavior of their oóspores on germination, and the pecul-

iar group of the Conjugales. The fertilized eggs of Ulothrix,

Spharoplea and Oedogonium (see Fig. 4) generally give rise to

several zoóspores which develop into filaments like the parent

plant. There is thus a short period after the stimulus of the

sexual act when the protoplasm of the organism behaves differ-

ently from its usual vegetative activities. Some botanists con-

sider this behavior the beginning of a sporophyte generation

introduced after and the direct result of the sexual act. The

union of gametes is supposed to give a product (the sexually

formed spore) whose protoplasm has potentialities different from

the parent plant. There is in general a greater vigor and prob-

426 THE AMERICAN NATURALIST. [Vor. XXXVII.

ably a modified structure which demands some other form of

expression than the vegetative activities characteristic of the

species. Such a phase, intercalated after a sexual act, would

start its peculiar period of activity because the protoplasm had a

different chemical and physical composition from that of the par-

ent plant. Its fundamental characteristics are therefore per-

formed through the fusion of the gametes.

This view of the origin of the sporophyte as primarily the

result of sexuality carries with it. a comprehensive definition of

the phase. It becomes a generation, always intercalated after

the sexual plant, which is called a gametophyte by way of dis-

tinction. It is a generation always antithetic with the game-

tophyte because of potentialities within itself. When once

thoroughly established, the sporophytic generation would be

expected to have the power of developing the gametophyte only

after it had passed through its characteristic history. The

sporophyte could never be an homologous generation with the

gametophyte in the way that succeeding generations of Thallo-

phytes by asexual methods of reproduction are homologous with

one another.

We should not expect the sporophyte to have arisen with the

potentialities of immediate and extensive growth developments

but rather as a small beginning such as may well be illustrated

in Ulothrix, Spharoplea and Oedogonium. Here the peculiari-

ties of the sexually formed spore seem scarcely more than an

increased vigor that expresses itself in the formation of a number

of reproductive cells. However, we know nothing of the nuclear

activities in these forms and the assumption that the protoplasm

of the sexually formed spore is structurally different from that

of the gametophyte is speculation but seemingly reasonable. As

has been stated, there are several thallophytes whose sexually

formed spores develop directly into the sexual plant and indeed

the egg of Oedogonium has been known to grow directly into a

filament (Fig. 4 d). These forms must also be studied in com-

parison with the types in which the sporophyte generation seems

well established.

The Conjugales present some interesting conditions. The

union of the gametes gives a zygospore (see Fig. 5 a) that finally

No. 438.] ORIGIN OF THE SPOROPHYTE. 427

contains a single nucleus although the fusion of the gamete

nuclei is sometimes greatly delayed. The fusion nucleus divides

into four nuclei previous to the germination of the spore but

some of these degenerate. Thus in Spirogyra three nuclei break

down so that only one is left at the time when the new filament

ruptures the spore wall. In certain desmids (see Fig. 5) two of

the original four nuclei degenerate and the remaining two enter

into the formation of the pair of new desmids developed in each

zygospore. It may be suggested that the division of the fusion

nucleus apparently wasteful of material represents the expendi-

ture of energy infused by the sexual act and consequently a

sporophyte activity which is necessary to bring the protoplasm

Fic. 5. Germination of the zygospore of a desmid (Closterium). (After Klebahn.)

back to the potentialities of the parent organism. But again we

do not know the structural changes that take place in the nucleus

of the Conjugales at and after the time of fertilization. Certain

diatoms present activities resembling in some respects the Con-

jugales but the conditions are too complicated to be given here.

This attitude towards the significance of the nuclear divisions

following the sexual act is the keystone to the theory of the

antithetic origin of the sporophyte and its significance, a view

held by many prominent botanists including Bower, Strasburger,

and Klebs. The sexual act is assumed to give to the fused

gametes an organization that will always tend to differ from that

428 THE AMERICAN NATURALIST. [Vor. XXXVII.

of the parent plant. When the modification is sufficiently great

we should expect a new or modified morphology in the resulting

generation which would naturally react in a different manner to

its environment. The new generation becomes then a phase

intercalated after the sexual act and antithetic with the sexual

plant (gametophyte).

In its earliest beginnings the oe might differ so

slightly from the gametophyte in its potentialities for develop-

ment as to give no very decided results. Such expressions would

be an indefinite cellular body such as is illustrated by Coleochaete

or perhaps nothing more than a stimulus to develop several

reproductive cells as may be observed in Ulothrix, CEdogonium

and Spheeroplea.

An extensive elaboration of the sporophyte generation must

follow from any changes that lead to complexity either of exter-

nal conditions or internal structure through the mixing of pro-

toplasm in the sexual act. The character of the evolution must

depend largely upon the environmental relations, so that sporo-

phytes should be expected to diverge in structure from their first

inceptions. The most elaborate sporophytic developments in the

algze (e. g., probably the Rhodophyceze) cannot be like those of

land plants, and those of fungi must be expected to present

peculiarities of their own (Ascomycetes).

In closing we may summarize the principles that we have

discussed in relation to the sporophyte.

First: Its origin, a phase intercalated after the sexual act

because the fusion of gametes gives a protoplasm structually. dif-

ferent from that of the sexual plant (gametophyte).

Second: Its establishment as an azz?Aetic generation through

these peculiarities of protoplasmic structure that tend to express

themselves in morphological developments differént from the

gametophyte, which developments are at all time influenced by

environmental relations.

Third: Its end, the ‘production of asexual spores with the

potentialities of the gametophyte through a structural change in

the protoplasm by which the sporophytic characters disappear

and the ancestral qualities of the gametophyte again assert them-

selves.

No. 438.] ORIGIN OF THE SPOROPHYTE. 429

The sporophyte will vary in complexity with the environ-

mental conditions and the tendency must be always towards

structural divergence. There are several well: defined and very

important principles responsible for these complexities and they

will be considered in future papers dealing with the evolution

of the sporophyte.

UNIVERSITY OF CHICAGO.

NOTES AND LITERATURE.

ZOOGEOGRAPHY.

The Atlantis-problem has been re-investigated from the zoógeo-

graphical side by Dr. R. F. Scharff! Examining the land and

fresh-water faunas of the present Atlantic islands (Canary islands,

Madeira, Azores), he arrives at conclusions opposite to those of

Wallace. Although he does not deny that there are instances

which are apparently due to colonization of these islands by animals

transported by winds, currents, etc., he is of the opinion that a large

percentage of the fauna indicates that we do not have to deal with

truly oceanic islands, but that they once were connected with

present continents. He shows that there is reason to conclude that

* Madeira and the Azores, up to Miocene times, were connected

with Portugal; and that from Morocco to the Canary islands, and

from them to South America stretched a vast land which extended

southward certainly as far as St. Helena. This great continent may

have existed already in Secondary times, as Dr. Ihering suggested;

and it probably began to subside in early Tertiary times." Scharff

further believes that the northern portions of this latter continent

persisted until the Miocene, and that subsequently, in early Pleisto-

cene, there was again a connection of the Atlantic islands with the

Mediterranean countries (Africa and Europe).

These ideas admirably agree with those recently set forth by

Ortmann,? at least as far as it concerns the connection of West

Africa with South America, and it is quite interesting that both

authors arrived at the same conclusions, as did others before them,

namely: Suess (1888), Neumayr (1890), aud von Ihering (1891).

Zoügeographical Development of the Indo-Australian Archi-

pelago has been made the object of a special investigation by Pro-

fessor Max Weber’). He arrives at certain very important conclu-

1 Scharff, R. F. Some remarks on the Atlantis Problem, in Proc. R. Irish

Acad. vol. 24, sect. B, part 3, 1903, pp. 268-302.

? Proc, Americ. Philos. Soc. vol. 41, 1902, p. 348, ff.

3 Weber, Max. Der Jndo-australische Archipel und die Geschichte seiner Tier-

welt. Jena, Fischer, 1902. . 46 pp., map.

* 431

432 THE AMERICAN NATURALIST. (Vor. XXXVII.

sions, which seem well supported, at any rate they are better

supported than the previous very general theories concerning these

parts. We may summarize them here as follows.

1. We are to abandon Neumayr’s theory of a large Jurassic Sino-

Australian continent. Sufficient evidence has been brought forward

to show that at least a large part of the present Indo-Australian

region was covered by sea in Jurassic times, and, moreover, that this

sea was by no means a shallow one.

2. Zoólogical evidence supports the view of a continental connec-

tion of Asia and Australia during Cretaceous times. As to the size

and shape of this connection we possess hardly any indications: we

know that certain parts of the Archipelago were land during this

period, while marine deposits are known from other parts.

3. During the Cretaceous, and pre-eminently so during the Terti-

ary, this region was characterized by frequent changes, but during

all these times it was a region if not of continental character, certainly

of that of a shallow sea with many islands.

4. Wallace's line, a depression between Borneo and Celebes, and

said to be continued between Bali and Lombok, does not exist as a

zoógeographical boundary between Asia and Australia. —Between

Bali and Lombok, it does not exist at all, the sea being very shallow

there. There is indeed a depression between Borneo and Celebes,

but this does not separate Asiatic and Australian faunal development

Celebes possessing Asiatic elements in a high percentage. Never-

theless, Wallace's line holds good for a separation of Celebes from

Borneo, the former having received the oriental constituents of its

fauna not from Borneo, but from the Philippine islands.

s. The three larger Sunda islands (Sumatra, Java, Borneo) were

connected with one another and with the Asiatic continent in Plio-

cene times. The separation of these islands was not contemporane-

ous: Java was isolated first, and Sumatra last.

6. Borneo was also united, at about the same time, with the

Philippine islands and South China.

7. The smaller Sunda islands (Bali, Lombok, Sumbawa, Flores)

form a continuation of Java, possessing a fauna which in part is Ori-

ental, and in part Australian. The Australian elements possibly

arrived by way of Timor, which shows closer affinities to Australia,

and, no doubt, once was part of this continent.

8. Celebes was always separated from Borneo (beginning in the

Miocene), but it was connected with the Philippine islands on the

one side, and with Java and the smaller Sunda islands (Flores) on

No 438] NOTES AND LITERATURE. 433

the other. This accounts for its oriental elements. It possesses

also an eastern (Australian) faunal element, and this is explained by

a former connection by land with New Guinea by way of the Sula

islands, Buru and Ceram.

9. New Guinea, Aru, and Ki inae once formed part of the

Australian continent in Tertiary times. Temporarily connected with

this were the islands of Ceram, Misol, Waigeu, and Halmahera.

The Banda sea is a depression formed in Miocene times.

The general conclusions of M; Weber are given by himself in the

following words (translated) (pp. 41—43)

* [n Pre-Tertiary, possibly in Cretaceous times, Asia and Australia

were connected by a land mass. Eurasiatic types of animals popu-

ated this land, which subsided during the Eocene, and was dissolved,

into a southeastern portion: present Australia and New Guinea

where Monotremata, Marsupials, and ancient types of other groups

of animals were preserved, Cassowaries, Birds of Paradise developed,

while groups or universal distribution, as for instance Woodpeckers

and Sparrows, were not able to reach these parts. Toward the

North, a shallow coral sea extended, in which a few elevated parts

were present as islands, and possibly offered a refuge to a few older

forms of life like Cuscus, primitive Rodents, Insectivors, and others.

“Important changes took place in the Miocene: on the one hand

deep depressions were formed in the shallow sea, and on the other

hand stretches of land became dry, like Celebes, while in the western

parts continental conditions were formed, which offered opportunities

for the dispersal of Asiatic animals, such as we know from the

Siwalik beds of India. Meanwhile new changes of level took place

within this unstable part of the earth’s crust, which brought about,

by the end of the Pleistocene, the present configuration of the

archipelago. These processes caused the local disappearance of that

old Tertiary fauna, with the exception of a few relicts, while the

immigration of modern Asiatic types was favored. This immigration

was of longest duration for the greater Sunda islands, of which Java

was the first to become isolated. The fauna of these islands, con-

sequently, resembles most that of Asia. Toward the East, a mixed

fauna prevails, and the Asiatic types disappear gradually in this

direction, while Australian types become more frequent. This mixed

fauna forms a broad transitional zone between the greater Sunda

islands and the Philippines on the one side, and Australia, including

New Guinea, the Aru and Ki islands, on the other. The aunal

elements of this region are of different origin. Celebes preserved

434 7HE AMERICAN NATURALIST. [Vor. XXXVII.

some older Asiatic types, which came probably by way of the Philip-

pine islands. Younger Asiatic immigrants arrived in Celebes by

way of Java, and the smaller Sunda islands, but not directly from

Borneo, the old Strait of Makassar separating it from this island,

which also received immigrants from the Philippines, and sent others

in turn to them, from whence they were able to find their way to

Celebes.

“Tn addition, the fauna of Celebes received Australian types from

the Moluccas, where they were in part relicts, in part later immi-

grants from Australia. Possibly also the smaller Sunda islands

received a few immigrants from this latter continent.

A. E. O.

BOTANY.

Influence of Light and Darkness on growth and Develop-

ment.'— In this work is presented one of the most important, if not

the most important, single piece of research in plant physiology yet

published in this country. It is a record of a series of extensive

experiments covering several years (1895—1902), from which impor-

tant generalizations have been drawn. Heretofore too much of the

general principles laid down regarding the influence of light upon

plants has been based upon experiments with a comparatively lim-

ited number of forms. It was the author’s intention to avoid these

errors and for this reason he selected representative plants covering

a wide field in both relationship and form. No fewer than ninety-

seven forms were experimented with, and etiolated for an often long

period of time. These forms include very diverse orders of plants

and comprise practically every form of the plant body of importance.

Various typical and modified forms of stems, including those of suc-

culents and xerophytes, the different types of foliar organs and

various floral structures were investigated. A very careful and

detailed account of the histology of the different etiolated tissues, in

comparison with the normal, is also included.

! MacDougal, D. T. The influence of Light and Darkness upon Growth and

Development. Memoirs New York Botanical Garden. vol. ii. Jan. 20, 1903, pp.

i-xiii, —1-319, 176 figures.

No. 438.] NOTES AND LITERATURE. 435

As might be expected such an exhaustive investigation of the

relations of plants to light and darkness, leads to new and interesting

conceptions as to the action of illumination in determining the growth

of the shoots of plants. It is in this of course that the chief interest

of the work centers. As a record of careful and interesting observa-

tions the first and major part of the work is of great value, but with-

out entering too greatly into detail, it is impossible to give a fair idea

of its contents. For that the paper itself had best be consulted. In

passing, the following plants, unusual at least from the standpoint of

etiolation experiments, may be mentioned, Amorphophallus rivierti

Dur., Bowiea volubilis Harv., Cocos nucifera L., Lbervillea sonore

Grev.— and two sarracenias.

In discussing the theories, hitherto presented, the author points

out how inadequate and unsatisfactory the explanations of etiolation

phenomena have been. Under the heading *Morphogenic Influ-

ences of Light and Darkness” the author expresses his own under-

standing of the problem by a careful analysis of the facts, and the

following is intended to give the substance of his discussion.

Etiolation is not to be regarded as an adaptive modification on the

part of the plant, and the form that it assumes in darkness is not due

to an effort to reach the light. The various phenomena of etiolation

are of course, in the first instance, due to the absence of light, but it is .

only in some cases that a beneficial modification results. Aside from

the absence of chlorophyll, the basal fact connected with the condi-

tion of etiolated organs is that their tissues do not show the same

degree of morphological differentiation as do those of the normal.

This lack of differentiation, and the abnormal increase in size which

accompanies it, is most noticeable in the large parenchymatous tracts

of tissue. An incomplete deposition of aplastic material allows of a

much longer growth period. In this connection the author points

out that in organs where extension is possible by the formation of

additional parenchymatic cells we have the familiar elongation and

enlargement of etiolation, whereas in cases like those of dorsiventral

leaves, where the mechanical elements are prerequisite for the expan-

sion of the lamina, the organ remains small. A comparison of nor-

mal and etiolated plants shows that growth and differentiation are

not only independent phenomena, but are easily separable. ;

From this point of view then the phenomena of etiolation rests

upon the absence of the morphogenetic influence of light, or rather

as the author modifies it in considering the effect of continuous illu-

mination in producing partially elongated forms, we may suppose that

436 THE AMERICAN NATURALIST. [Vor. XXXVII.

the alternate action of light and darkness constitutes the stimulus

which finds expression in tissue differentiation. As is pointed out

we can very well imagine in the case of continuous illumination,

that the plant having adapted itself to this condition ceases to

respond, so readily to the stimulating action of light. It may be

narrowed down then to the fact that it is light which acts as the

stimulating influence in inducing morphological differentiation and in

its absence the tissues tend to remain in a more or less primitive con-

dition. The action of light, however, is not necessarily direct, since

the stimulative influence may be received by one portion of the body

and transmitted to another, as is shown by local aetiolation. The

author speaks of the enlargement of certain dorsiventral leaves

brought about in etiolated plants by the removal of concurrent

organs and mentions that even in their enlarged condition they do

not show much tissue differentiation, but he hardly explains the rea-

son for this behavior. Possibly this might be regarded as a reaction

to the untoward stimulus of wounding.

` The total amount of growth or increase in volume, that may be

accomplished by the shoot in the expansion of its imperfectly devel-

oped tissues during etiolation, is subject to great variation. In

many cases the total length, diameter, and volume may be actually

less than in a normally grown plant, and indeed the rate of growth

need not be so rapid. It may be said that light does not directly

affect the rate of growth, that it does not exert the retarding or para-

tonic influence commonly ascribed to it. This is shown by the action

of plants exposed to continuous illumination.

It will be seen that this is an important addition to our knowledge

of the influence of light on the growing shoot. Never before has

this point of view received such a comprehensive treatment or been

supported by such a wealth of facts. The author's insistence that

etiolation is not an adaptive reaction and the stress laid upon the

anatomical conditions presented by etiolated organs, leads to a far

clearer understanding of the actual influence of light as a morpho-

genetic stimulus. In this latter regard the evidence afforded is

especially convincing. It is not yet clear of course why light should

so affect tissues as to practically shorten the period of their meriste-

matic condition and to induce the formation of what is usually termed

"permanent" tissue. It is said that the aplastic material is not so

readily laid down in the absence of light, a statement which suggests

a chemical explanation. The exact nature of the chemical changes

which are supposed to take place in the differentiation of the usual

No. 438.] NOTES AND LITERATURE. 437

forms of tissue, must of necessity remain obscure for a long time to

come.

In conclusion the very copious index should be mentioned, in which

the novel and withal convenient practice has been adopted of enter-

ing author's names with a full bibliography.

H. M. R.

A Popular Introduction to Forestry. — Professor Filibert Roth,

formerly of Cornell University, and now, as chief of the Division of

Forestry, U. S. Department of the Interior, in charge of the work in

the Government Forest Reserves, has just published “A First Book

of Forestry." 1 ‘The little volume is intended for use in the public

schools and in country homes, and gives in non-technical language,

an exceptionally clear and readable account of some of the signifi-

cant aspects of forest life and growth, the most important principles

underlying the practice of forestry, and the methods now employed

in such common forestry processes as thinning the wood-lot; seed-

ing for succession; sowing cleared areas: harvesting the wood

crop; protecting the forest from fire, insects and fungus pests ; and

so on. He makes clear the forester’s aim so to harvest each wood

crop as to speed its successor, and discusses the six principal

methods of accomplishing this. Following these accounts of forest

life and forest reproduction and the description of the ways in

which forests should be protected and helped to make their best

growth, he also tells us some of the indirect benefits of forests, such

as the protection of soil and conservation of moisture, and gives us

a brief description of our own forests, and a history of forest culture

from the times of the Romans. Then the structural features and

physical and chemical properties of wood are taken up ; and finally,

a key is added for the determination of our common trees by charac-

ters of leaf, wood and bark, and the several valuable appendices in

addition to more technical tables include a convenient reference list

of the more important woods and trees of the United States. The

illustrations, which are excellent in themselves and admirably related

to the text, add greatly to the latter's value. As a popular educator

calculated to render more intelligent the growing sentiment in favor

of forest protection, this little book should have a wide circle of

readers, and its usefulness will hardly be less because of the con-

siderable number of other good books tener published by our

American foresters. J. S. P.

! Roth, Filibert. First Book of Forestry. Boston. Ginn and Co., 1902. 12mo.,

291 pp., illus

438 THE AMERICAN NATURALIST. (Vor. XXXVII.

Notes.— Zhe Botanical Gazette, for February, contains the following

articles : — Copeland, * Chemical Stimulation and the Evolution of

Carbon dioxide" ; Sargent, “The Genus Crategus in Newcastle

County, Delaware ” ; Long, * The Ravenelias of the United States and

Mexico” ; fittings. " PATE in Carya oliveformis”; and

Cotter, “Selected Notes.”

Rhodora, for February, contains the following articles: Harvey,

“An Ecological Excursion to Mt. Ktaadn”; Sargent, “ Recently

Recognized Species of Crategus”; Fernald, “ Andromeda polifolia

and A. glaucophylla” ; and Knight, “Some plants new to Maine."

Rhodora, for March, contains Bacon, *An Experiment with the

Red Baneberry "; Cushman, * Desmids of Bridgewater, Mass.” ;

O. Eaton, “Orchids of Chesterville, Me. "; Rand, * Observations on

Echinodorus parvulus"; Robinson, “Generic position of Echinodorus

parvulus ”; and Fernald, * A New Bidens from the Merrimac Valley."

The Bulletin of the Torrey Botanical Club, for February, contains

the following articles: De Vries, “On Atavistic Variation in Qinothera

cruciata” ; Clements, “Nova Ascomycetum genera speciesque ” ;

Peck, “New Species of Fungi”; Hollick, “A Fossil Petal and

a Fossil Fruit from the Cretaceous (Dakota Group) of Kansas”

Rowlee, “ Notes on Antillean Pines with description of a New Species

from the Isle of Pines ” ; and Murrill, * Polyporacez of North America,

II, The Genus Polyporus.”

Torreya, for February, contains the following articles : Underwood,

“Notes on Southern Ferns, I"; Seymour, * Trichomanes petersii

Found Anew”; Harper,“ A Unique Climbing Plant "; Britton, “ An

undescribed Eleocharis from Pennsylvania " ; and Earle, * Key to the

North American Species of Stropharia.”

Torreya, for March, contains Parish, “ Vital Persistency of Agave

Americana”; Earle, “Key to the North American Species of

Lentinus, I"; Ramaley, *Pubescence of Species of Astragalus ”

Cockerell, “Insect Visitors of Scrophularia "; and Haynes, * Some

interesting Hepaticz from Maine."

Vol. II of the Memoirs of the New York Botanical Garden, issued

January 20, consists of a treatise by Professor MacDougal on the

* Influence of Light and Darkness upon Growth and Development."

Minnesota Botanical Studies, 3d Series, Part 1, issued March 21,

1903, contains the following papers: Ramaley, ‘Observations on

Egregia menziesii” ; Butters, “ Observations on Trichoglcea lubrica?

No. 438.] NOTES AND LITERATURE. 439

Holtz, * Observations on Pelvetia” ; Hone, * Petalonema alatum in

Minn. ”; N. B. P. Nelson, “Observations upon some Algæ which

cause ‘Water Bloom'"; Hillesheim, “Some Observations on the

Staining of the Nuclei of Fresh-water Algæ ” ; Crosby, “ Observations

on Dictyospheria "; Brand, Stapfia cylindrica in Minn. "; Powell,

" Observations on some Calcareous Pebbles”; “ Lilley, Nitella batra-

chosperma in Minn." ; and Wheeler, “ Catalog of Minnesota Grasses.”

The Bulletin of the Southern California Academy of Sciences, of

February 1, contains the following botanical articles : Hasse, “ Con-

tributions to the Lichen-flora of the California Coast Islands";

Parish, “ Two New Plants from Southern California ” ; and Davidson,

“New Records for Los Angeles County.”

Professors Aven Nelson and Cockerell have described several new

plants from New Mexico in recent leaflets of the Proceedings of the

Biological Society of Washington, which also contain a revision of

Psilostrophe by the first-named author.

Bailey’s Queensland Flora has reached Part VI,— Alismacez to

Filices,— which concludes the work, making a volume of 2015 pages,

in addition to separately paged prefatory matter. It is illustrated by

88 plates. |

No. 8 of Koorder's and Valeton's Additamenta ad Cognitionem

Flore Arborez Javanice, issued as Mededeelingen uit i's Lands

Plantentuin, No. LIX, deals with Rubiacez and Oleacez.

An illustrated article on “Saving the Southern ichs d by Price,

is published in Ze Words Work for March.

F. L. Sargent has published— Cambridge, 1903 —a useful little

“Key to Common Deciduous Trees in Winter and Key to Common

Woods,”

Illustrated accounts of Sisal or henequen fiber, as produced in

Yucatan, are given in Advance Sheets of Consular Reports No. 1582,

the National Geographic Magazine for April, and The American

Znventor for May 1, all by Consul E. H. Thompson, of Progress.

Further studies on the Root-tubercles of Leguminosz and their

causation are published by Hiltner and Stórmer, as Heft 3 of the

third volume of Arbeiten aus der Biologischen Abtheilung für Land-

und Forst- Wirthschaft am Kaiserlichen Gesundheitsamte.

The John Crerar Library, of Chicago, has issued a list of bibli-

ographies of special subjects, among which botany finds place.

440 THE AMERICAN NATURALIST. [Vor. XXXVII.

The Botanical Gazette for November contains the following papers :

Johnson, “On the Development of Certain Piperacee”; Kraemer,

“The structure of the Starch Grain”; Aven Nelson, “ Contributions

from the Rocky Mountain Herbarium, IV ”; and Evans, ^A New

Hepatic from the Eastern United States."

Broteria, Revista de sciencias natures do Collegio de S. Fiel, is

the title of a new annual, issued at Lisbon. ‘The frontispiece of the

first volume is a picture of the statue of the botanist Broteri, in the

botanic garden at Coimbra, accompanied by a short account of his

life and work.

The Journal of Mycology, for December, contains the following

articles : Morgan, * A New Genus of Fungi [Sporocystis] " ; Sanders,

“Interesting variations in the appendages of Podosphera oxyacan-

the "; Holway, “ Notes on Uredinez, I ” ; Schaffner, “Ohio Stations

for Myriostoma”; Ellis and Bartholomew, “ New Species of Fungi

from Various Localities”: Morgan, “ The Discomycetes of the Miami

Valley, Ohio”; Kellerman, * Notes from Mycoiogical Literature,

III”; Kellerman, “Index to North American Mycology, continued.”

Meehan's Monthly, which has regularly included a plate with

commentary dealing with native plants, in each number, stopped with

the December number but is serially continued by a new journal,

floral Life.

Part 26 of Pittonia, the opening number of the fifth volume,

consists of a series of papers by Professor Greene, dealing with a

considerable number of genera of phanerogams.

The Plant World for October contains the following articles :

Safford, “ Extracts from the Note Book of a Naturalist on the Island

of Guam, II”; Fitzpatrick, “A Study of the Island Flora of the

Mississippi River near Sabula, Iowa”; Niles, “Origin of Plant

Names, II, the Lady’s Slippers and Moccasin Flowers”; Barrett,

“The West Indian Corkwood”; and Bates, * The Ke Longev-

ity of Certain Plants."

Rhodora for November contains the following articles: Evans,

“Notes on New England Hepaticz ”; Fernald, “Variations in Glaux

in America"; Jones, “ Pogonia aflinis in Vermont”; Holt, “Is

Cirsium palustre a Native of New Hampshire?” Morrell, * Lamium

album in Maine"; Fernald, *Lists of New England plants, X,

Carex"; and Beal, *Seed Throwing of Viola."

Torreya for November contains the following articles: Cannon,

No. 438] NOTES AND LITERATURE. 441

“Field Notes on Rhododendron catawbiense "; Earle, “Key to the

North American Species of Cortinarius, I"; Underwood, “Two

New Species of Selaginella in the Southern Flora”; Lloyd, “ Vaca-

tion Observations, I"; Howe, *Note on the Report of the Brown-

Harvard Expedition to Nachvak, Labrador"; and Hazen, * The

Habitat of the Slender Cliff Brake."

The Bulletin of the Torrey Botanical Club for November contains

the following articles: Underwood, * American Ferns, IV, the Genus

Gymnogramme of the Synopsis Filicum "; Knowlton, “ Notes on the

Fossil Fruits and Lignites of Brandon, Vt."; Piper, * New and Note-

worthy northwestern Plants”; and Salmon, Supplementary Notes on

the Erysiphacez."

Volume X of the Memoirs of the Torrey Botanical Club, issued in

November, is devoted to a history of pre-Clusian botany in its relation

to Aster, by Burgess.

In University Studies (of the University of Nebraska) for Decem-

ber, Dr. Clements analyzes the use of Greek and Latin in biological

nomenclature, lays down a series of rules, and gives a very large

number of correctly formed generic names which he proposes to

substitute for a like number of incorrectly formed names now in use.

The proper terminology of groups, in botany, is the subject ofa

short but incisive note by Professor Underwood in Science of No-

vember 28.

A flora of the West Indian island of St. Croix, by Millspaugh, is

printed as No. 7 of the current volume of botanical Publications of

the Field Columbian Museum.

In the Nuovo Giornale Botanico Italiano for October, is concluded

a critical study of the Italian species of Atriplex.

No. 17 of Dr. Holm’s Studies in the Cyperacez, in the American

Journal of Science for December, deals with segregates of Carex

tolmiei.

Curtis’s Botanical Magazine for November contains a figure of a

curious aquatic Amaryllid, Crinum natans, of Africa.

Out West, for December, contains an illustrated article on Pinus

torreyana, by Helen L. Jones.

A paper by Minnie Reed, on “Two New Ascomycetous Fungi

Parasitic on Marine Algæ,” is issued under date of November 20, as

a brochure of the first botanical volume of the University of California

442 THE AMERICAN NATURALIST. [Vor. XXXVII.

Publications. Both are referred to the genus Guignardia, and con-

sidered to be undescribed, as is also the species of Prasiola on which

one of them occurs.

An account of larch and spruce fir canker, and the fungi involved,

by Massee, is reprinted from the Journal of the Board of Agriculture

of London, for September.

Fascicle 2 of Sydow's Monographia uredinearum carries the

number of species of Puccinia up to 595.

An account of tea, its cultivation and preparation for the market,

by Williamson, is contained in part 4 of the fourth volume of Trans-

actions of the Edinburg Field Naturalists’ and Microscopical Society.

A study of the species of Eucalyptus cultivated in the United

States, by McClatchie, forming a large and well illustrated paper,

is published as Buletin no. 35 of the Bureau of Forestry of the

Department of Agriculture.

A practical little hand-book for the forester, dealing with measure-

ments of standing and cut timber and the like, is published by H. S.

Graves as Bulletin no. 36 of the Bureau of Forestry of the Depart-

ment of Agriculture.

An account of wild rice, Zizania aquatica, and its use by the

Indians, is published by A. E. Jenks in part 2 of the 19th Annual

Report of the Bureau of American Ethnology, and is illustrated by

numerous plates.

No. 437 was mailed May 26, 1903.

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I. The Colors of Northern Gamopetalous Flowers (continued), JOHN H. LOVELL 443

IL Rib Variation in Cardium . . > - = + FRANK COLLINS BAKER 461

. IXI. The Perforation of a Vein by an Artery in the Cat (FELIS DOMES TICA)

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AMERICAN NATURALIST.

VoL. XXXVII. July, 1903. No. 439.

THE COLORS OF NORTHERN GAMOPETALOUS

FLOWERS (continued).

JOHN H. LOVELL.

Tue Lentibulariacez, or bladderwort family, are mostly

aquatic or marsh plants. Only four genera and one hundred

and eighty species are known. The flowers are yellow, or vary

from yellow to purple and violet-blue. . Of the fourteen species

of Utricularia, or bladderwort, eleven species are yellow and

three purple. The rootless plants of Utricularia vulgaris float

near the surface during inflorescence. The deeply 2-lipped flow-

ers are bright yellow with the palate marked with reddish-brown

lines leading well down into the spur, which secretes the nectar.

According to Knuth, Heinsius found the flowers visited only by

long-tongued Syrphidz, the species Helophilus lineatus being

most numerous. This is certainly surprising as the closed flow-

ers appear adapted to bees. As the species is aquatic consider-

able patience is required to observe the visitors. After repeated

observations I have collected on the flowers in Maire only the

syrphid fly Helophilus conostomus. Like Utricularia the genus

Pinguicula is carnivorous, and the yellowish-green leaves are

thickly covered with sticky glands. The flowers are violet-

443

444 THE AMERICAN NATURALIST. [Vor. XXXVII.

blue with the palate covered with velvety white hairs. The

visitors are flies and bees.

The Orobanchacz are parasitic plants without chlorophyll,

usually colored yellowish or purplish. The flowers also are

frequently yellowish or purple. In variety luteum of Aphyllon

fasciculatum the whole plant is yellow. Sometimes the flowers

are bicolored, yellow or white, and purple.

The Bignoniaceae, or trumpet-creeper family, occur chiefly in

the tropics. Many of the species are bird flowers, one to two

inches in length, and crimson, orange or scarlet, as Bignonia

venusta and Tecoma radicans. Common examples of bird

flowers in North America are Lobelia cardinalis, Gossypium

herbaceum and Lonicera sempervirens. | The ruby-throated

humming-bird, however, visits many flowers fertilized by insects.

The Acanthacee, a large tropical family of some 1800 species,

also contains many scarlet bird flowers.

The order Plantaginales includes but a single family, the

Plantaginaceze, or plantain family. The inflorescence is in

spikes with small 4-merous flowers, which are mostly greenish

or purplish, and are wind-fertilized. They are of special interest

because they show the beginnings of adaptations to insect

visitors. In one or more species, *we have before us the

passage from anemophilous to entomophilous characters, the

evolution of an entomophilous from an anemophilous spe-

cies." Plantago media possesses a pleasant perfume and red-

dish filaments. Müller distinguishes an anemophilous and an

entomophilous form, which differ slightly in color, the stamens,

stigmas, and pollen. Twenty-four visitors have been collected

on the flowers. The limb of the corolla and sometimes the

border of the sepals of P. a/pizais red. Five insects in the

Alps have been collected on this species. According to Knuth,

the flowers of Plantago display a variety of colors; in P. major

the corolla is brownish, the filaments white, the anthers red,

brown, or sometimes yellow or even white, while in other species

yellow, red and violet appear.

The three orders, Rubiales, Valerianales, and Campanulales,

which terminate the Gamopetale, exhibit many affinities with

the families, which stand at the close of the Choripetalous

No. 439.] WORTHERN GAMOPETALOUS FLOWERS. 445

series. The individual flowers are usually small, and con-

spicuousness is gained by aggregation. The inflorescence is

cymose forming in the Dipsaceee and Composite dense in-

volucrate heads, and not infrequently contracted in the other

families belonging to this group into capitate clusters provided

with an involucre, as in Cephaélis tpecacuana of the Rubiacee.

Both actinomorphic and zygomorphic flowers occur, and the

sexes may be united or separated. By some writers the Rubia-

cec are derived from the Umbelliferze. While this derivation

is doubtful the terminal groups of the Choripetale and Gamo-

petalz certainly possess many points of resemblance, which

indicate a parallel development.

The Rubiales, which include the Rubiacez and Caprifoliaceze,

have opposite leaves, and usually the stipules are "present

in the first of the two families but rarely in the second.

Stipules occur elsewhere in the Gamopetalee only in the prim-

itive stem family of the Loganiacez. The corolla varies

greatly in length from rotate to funnelform and tubular, and is

in consequence adapted to a great variety of visitors.

The Rubiacez, or madder family, is of immense extent in the

tropics and contains about 5500 species. No other family con-

tains so many dimorphous flowers. The roots of several species,

as Rubia tinctorum and Galium boreale contain a red pigment

(madder red), which is widely used in dyeing. The flowers of

Galium, or bedstraw, are very small or minute, with the calyx

obsolete. In G. triflorum and G. circezans the flowers are green,

in G. boreale and G. mollugo white, in G. verum yellow, in

G. rubrum red, and in G. purpureum purple. The visitors are

chiefly flies, and the great variety of colors affords evidence that

they do not prefer one hue to another. Indeed the coloration

of the different species is probably determined by internal con-

ditions. Houstonia caerulea, or bluets, one of the common

spring flowers, is pale blue or nearly white with a yellow eye.

So abundant is this little plant that it often tinges the hillsides

and meadows. Other species are blue or purple.

The Caprifoliaceze, or honeysuckle family, are remarkable for

the variation in length of the corolla tube, and the consequent

adaptation of the flowers to a great variety of visitors. The

446 THE AMERICAN NATURALIST. (Vow. XXXVII.

white, wheel-shaped flowers of Sambucus contain no honey, and

are sparingly visited by flies and pollen-collecting bees. The

large, pyramidal or flat cymes are very numerous and conspicuous.

The small, rotate flowers of Viburnum are in large compound

cymes, which bloom in early spring and midsummer. They are

white, fragrant, and nectariferous. The most important visitors

are Andrenide, flies and beetles, to which the inflorescence with

its freely exposed honey is well adapted. I have found beetles

more abundant and in greater variety than upon any other

northern plants. The marginal flowers of V. a/nifolium and

V. opulus are sterile and greatly enlarged.

There are a few flowers adapted to wasps and to which these

insects are very frequent visitors. The most important wasp

flowers are Epipactis latifolia, Cotoneaster vulgaris, Scrophularia

nodosa, Symphoricarpos racemosa, and Lonicera alpigena, the last

two belonging to the Caprifoliacez. The flowers agree in

having abundant honey secreted in a short corolla, or pouch-like

receptacle, about the size of a wasp's head, and usually lurid

colors. In England Darwin found Æpzpactis latifolia visited by

swarms of wasps, but was astonished to observe that the sweet

nectar never proved attractive to any kind of bee or dipterous

insect. The small reddish flowers of Symphoricarpos racemosus

(snow berry) are campanulate and pendulous. Wasps thrust

their heads wholly into the flower to obtain the nectar. ' Loni-

cera alpigena is reddish-brown. Müller observed in the Alps

that it was visited by two species of wasps in great numbers.

The nodding blossoms of Linnea borealis are wine colored

with a yellow marking on the lower side, which serves as a honey-

guide, and exhale a sweet vanilla-like fragrance. It is a trailing

evergreen vine densely carpeting the ground in cold, open wood-

lands. I have collected on the flowers only the fly Empis rufes-

cens, which is rather common.

The large genus Lonicera is adapted to a variety of visitors.

The wasp flower Z. a/pigena is reddish-brown. The bee flower

L. tartarica is pink or white. The bumblebee flowers, Z. ciliata,

L. xylosteum and L. cærulea are yellow. The hawkmoth flower

L. periclymenum on the first evening it expands is white within,

changing to yellow on the second evening. The exterior of the

No.439.] WORTHERN GAMOPETALOUS FLOWERS. 447

flowers is purplish-red, and in fading they turn to a dingy orange-

brown. The bird flower L. sempervirens is scentless, scarlet out-

side and yellow within, or rarely throughout. The corolla of

Diervilla trifida, or bush honeysuckle, is light yellow with an

orange honey guide on the upper lobe. The older flowers turn

reddish, a color change which also occurs in Rides aureum and

in the genera Weigelia, Fuchsia, and Lantana. In Rides

aureum Müller states that the more intelligent insects immedi-

ately recognize by means of their red color those flowers which

no longer contain nectar, and consequently visit more blossoms

in the same time. Repeated observations by the writer failed

to show that the color change in Diervilla was of the same

significance. The honeybee was observed to visit the red

flowers both when solitary and when associated with yellow

flowers. Neither was there any preference manifested for yel-

low flowers, when flowers of both colors occurred in the same

cyme. An immense number of varieties of Weigelia have been

produced in cultivation by selection and hybridization, which are

remarkable for their wide range of coloring. There are white

and deep red forms with every intermediate shade ; white when

opening but changing to rose ; deep red in bud but rose-colored

in bloom ; flowers pale rose at first, changing to deep red ; yel-

low; light yellow, changing to white; pale yellow, changing to

pale rose ; and reddish-purple.

The herbaceous order Valerianales is intermediate between

the Rubiales and the Campanulales. The flowers of the Valeri-

anacez are in clustered cymes and are usually white or reddish.

The inflorescence of the Dipsaceze, or teasel family, is in invol-

ucrate, purplish heads, and is attractive to a great number and

Scabiosa atropurpurea of the garden is black-

variety of insects.

"The distinct anthers and hanging

purple, scarlet, or white.

ovule separate this family from the following order.

The Cucurbitacez, or gourd family, were formerly classed

with the Choripetalze, but are now placed in the order Campanu-

lales with the Campanulacez and Composite. The species are

herbaceous, tendril-bearing vines found chiefly in the tropics.

The petals are separate or united. The smaller flowers of

this family are white or greenish and the larger are yellow. The

448 THE AMERICAN NATURALIST. (Vor. XXXVII.

pollinators are bees. “The flowers of a species of Trianosper-

ma in South Brazil are visited, according to Fritz Müller, very

abundantly all day long by Apis mellifica and a species of Meli-

pona, although they are scentless, greenish, quite inconspicuous

and to a great extent hidden by the leaves." In this instance

as in some others the bees are probably guided by past expe-

rience in looking for the nectar. The large flowers of the culti-

vated Cucurbita are often wholly or partially concealed by the

leaves, yet are readily found by bees.

The stem-family, or line from which the other families of this

order are derived, is the Campanulacez, or bell-flower family.

Of the twenty-three northern species one is red and twenty-two

are blue. The flowers of Campanula are campanulate or rotate,

blue or white, and are visited by many Hymenoptera. Lobelia

has zygomorphic flowers which are usually blue or white. But

L. cardinalis, fugens, splendens and texenis, have fiery red corol-

las adapted to humming-birds. There is no more brilliant red

color in the northern flora than that of the corolla of Z. car-

dinalis. Phyteuma and Jasione are transition genera. s

At the head of the gamopetalous series stand the great

family of the Composite, which includes such familiar and

widely distributed plants as the thistle, aster, goldenrod, daisy

and dandelion. About 1000 genera and 12,000 species have

been described. Multitudes of these hardy weeds grow lux-

uriantly in our fields, and along our highways and hedgerows ;

and exhibit a remarkable vigor and ability to thrive under the

most untoward conditions. Many of the species tend to become

cosmopolitan, and have spread over both continents. The inflo-

rescence represents Nature’s greatest triumph in flower building.

Intercrossing by insects, economy of time and material, a large

number of seeds well adapted to germinate, and their wide dis-

tribution, have all been very perfectly attained. The individual

flower is often very small, and of little significance as compared

with the community. Conspicuousness is gained by massing a

large number of flowers in a head, an arrangement that also

permits insects to visit them very rapidly. In the goldenrod a

head consists of ten or fifteen florets, while in the white weed

the number may exceed five hundred. The capitulum with

No.439] NORTHERN GAMOPETALOUS FLOWERS. 449

its enfolding bracts often resembles a single flower, and was

termed by the older botanists a compound flower. The life his-

tory of the individual florets may be conveniently studied in the

garden sunflower, where they are of comparatively large size.

The Cichoriacez, or chicory family, are often treated as a

tribe of the Composite. There are 8 white, 53 yellow, 5 red,

2 purple and 5 blue species. All of the flowers of the head are

strap-shaped or ligulate, as in the dandelion. This species Tar-

axacum taraxacum (T. officinale) is gregarious, and in some local-

ities the plants are so numerous that the inflorescence covers

with a bright sheet of golden yellow entire hillsides. The visit-

ors are numerous ; in Low Germany Müller collected 67 Apidz,

7 Lepidoptera, 25 Diptera and 16 other insects. Most of the

genera of this family have yellow flowers as Hieracium (hawk-

weed), Latuca (lettuce) and Sonchus (sow thistle), but as a rule

they are much less conspicuous than the dandelion and have

fewer visitors. The great number of yellow flowers in this

family have already been referred to under the Scrophulariacez.

Cichorium intybus (chicory) has large bright blue flowers with

white and pink variations.

The Ambrosiacez, or ragweed family, are composed of small

greenish flowers, which in the absence of insects have reverted

to wind-fertilization. In Ambrosia the corolla has been lost.

At an earlier stage the flowers were homogamous or self-fer-

tilized, as is still the case in Senecio vulgaris which is visited

rarely by insects. The flowers ‘excellently illustrate the fact

that inconspicuousness is due to the absence of insects.

In the Composite the flowers are either tubular and all alike,

when the head is called discoid ; or the disk flowers are tubular

and the marginal flowers are ligulate, when the head is radiate.

There are 21 green, 126 white, 209 yellow, 4 red, 64 purple,

and 59 blue flowers.! When the heads are discoid the flowers

are all of the same color, but when they are radiate they are

frequently bicolored. In the garden daisy, or Bellis perennis,

the disk flowers are yellow, and the ray flowers are € white, pink,

or purple, with purple bracts. In Townsendia the disk flowers

' In classifying bicolored capitula preference is given to the color of the rays.

450 THE AMERICAN NATURALIST. (Vor. XXXVII.

are yellow, and the ray flowers are white, violet, or purple. In

Aster the rays are white, pink, purple, or blue, and the disk

flowers are yellow turning to red-purple or brown. In the

China asters (Callistephus) there is a great variety of colors,

and a single head is often tricolored, as a yellow center sur-

rounded by an inner white ring and an outer ring of purple.

The ray flowers of this genus may display almost every imagina-

ble shade of color, and individual flowers may change from white

to rosy red or lilac. But in Helianthus and in part of the spe-

cies of Coreopsis both ray and disk flowers are golden: yellow.

The original color of the genera, which was usually yellow, is

preserved by the central or disk flowers. The rays may vary

from yellow to white, red, purple or blue, and an innumerable

number of intermediate shades. Under cultivation Chrysanthe-

mum sinense has yielded a multitude of magnificent flowers.

The ray flowers have increased in number until they compose

the entire head, and there is scarce a tint or shade save blue that

is not known. The original colors. were a pale yellow, a white

and a very weak violet shade, and from these have been raised all

the colors and shades now seen in this flower. ‘This has been

accomplished by a very slow and persistent selection and cross-

fertilization. It is worthy of notice how intensified the yellows

have become, and how many shades of this color there now

are. The lilac has become pink of pure shading; then, as to

red, cullingfordii often presents us with a pure tone of red. The

most pronounced purple we have to-day is from the lightly tipped,

incurved Princess of Wales, being a sport named Violet Tomlin.

It is really purple. Now we cannot get purple without blue,

and to those who are at work in this field of development, a

blue chrysanthemum would not be such a great surprise.” !

Throughout the Composite the corolla has remained of small

size, and there is no reason to suppose it has ever been greatly

modified in form. The primitive colors have also been very

largely retained, for out of 483 northern species 209 are yellow

and 126 white. Fifty of our genera contain yellow flowers,

and some large genera consist wholly of flowers of this color,

! Thorpe, J. Amer. Garden, vol. xi, No. 1, p. 4.

No. 439.] WORTHERN GAMOPETALOUS FLOWERS. 451

as Chrysopsis, or golden aster, Solidago (with one exception),

and Senecio. The capitula are both discoid and radiate, and as

a rule both ray and disk flowers are yellow. But the disk

flowers in some genera have become brown or purple. In Rud-

beckia, or cone flower, the rays are yellow and the disk purple;

in Helianthus six species have the disk purple or brown, and in

sixteen. species the disk is yellow; and in Coreopsis both rays

and disk vary from yellow to brown.

One hundred and twenty-six species have white flowers. In

many instances where the ray flowers are white the disk flowers

are yellow. In these bicolored capitula there can be little doubt

that the white rays are derived from yellow-flowered progenitors.

In Verbesina (crownsbeard) all of the five species have yellow

disks, but one has white and four yellow rays. The white dis-

coid heads seem also to have been originally yellow. Of the

discoid heads of Hymenoppapus two species are yellow and

three are white. A number of genera, as Antennaria, Filago

and Gnaphalium, consist of white woolly herbs with yellowish

white often - inconspicuous flowers, which have undergone much

retrogression. The white-flowered species appear to be of later

origin than the yellow, and in numerous instances to be derived

from them.

There are only four red to sixty-four purple, and fifty-nine blue

flowers. The heads are both discoid and radiate. While the

rays may change directly from yellow to red, purple, or blue, in

many instances they have probably passed through an interme-

diate white stage. In Boltonia, which has the disk yellow, one

species has the rays white, and in two others they are blue or

violet. In Aster the rays are white in twenty-two species, purple

in six, and blue in forty-four. In Erigeron the white rayed

species frequently vary to pink or purple. In Coreopsis twelve

species have the rays yellow like the disk, but in one they are

pink, and in the variety Golden Wave they often change from

golden yellow to maroon. In some species of Aster the disk

flowers change from yellow to red or blue, as in A. roscidus, A.

carmesinus, and in A, curvescens. Whether the ‘purple discoid

flowers of Vernonia (iron-weed) have passed through a yellow

stage there is little evidence. The flowers of Artemisia (worm-

452 IHE AMERICAN NATURALIST, . [Vor. XXXVII.

wood) have reverted to wind-fertilization and are greenish or

yellowish.

The individual flower in the Compositze is small and of little

significance. Conspicuousness is gained by massing first the

flowers, then the capitula, and finally the plants themselves. If

the capitulum is large, as in Helianthus, it may be solitary, but

if small, as in Solidago, they may be aggregated into dense

flower-clusters. Many species are, moreover, gregarious, and so

abundant that they constitute important features in the floral

landscape. Such are the white weed, thistle, sunflower, golden-

rod, and aster. Kerner states that in New Zealand the small

white flowers of Haastia are so densely aggregated that they

form hemispherical mounds two feet high by three feet in

length. The plant is known as * vegetable sheep" since at a

distance it is frequently mistaken for that animal.

With the exception of the Umbelliferze, or carrot family, no

flowers are visited by so large and miscellaneous a company of

insects as the Composite. The guests of a single species may

exceed one hundred in number. The nectar is more deeply con-

cealed than in the Umbelliferee, and the percentage of long-

tongued visitors is consequently much greater. Throughout the

Compositae bright coloration is correlated with pollination by

insects; and when a genus reverts to wind-fertilization, the

inflorescence becomes inconspicuous. It is interesting to note

that the species, which attract the largest number of visitors,

display a variety of colors, as in the bright yellow goldenrods,

Chrysanthemum leucanthemum with white rays and a yellow disk,

Achillea millefolium white or tinged with red, the asters with a

yellow disk and white or blue rays, and the purple-flowered Can-

ada thistle. These differently colored species are visited by a

large company of Hymenoptera, Lepidoptera, Diptera and Cole-

optera, which are influenced by the length of the corolla tube

and the degree of conspicuousness obtained by a contrast of

colors and by massing; but there does not. seem to be any evi-

dence that they find greater pleasure in one hue than in another.

The white-flowered Eupatorium perfoliatum (thoroughwort) in

this locality is visited by a larger number of butterflies than any

other Composite plant. Bumblebees are also very common and

No. 439.) NORTHERN GAMOPETALOUS FLOWERS. 453

as pollinators far more important than the butterflies. No one,

however, would claim that the color of this Species was due to

the selective influence of either bees or butterflies. In a wood-

land pasture I found two large patches of the common elecam-

pane, or Zuu/a helenium, and the Canada thistle growing side by

side. The yellowish-red butterfly, Argynnis aphrodite was flitting

about upon the large yellow flowers of Inula, for which it

showed a decided preference, though occasionally it was observed

to fly over to the purple flowers of the thistle. The white cab-

bage butterfly on the contrary confined its visits almost exclu-

sively to the thistle blooms. As red has been supposed to be

the favorite color of butterflies, this singular behavior must have

been determined by other causes than the colors of the flowers.

Argynnis aphrodite also very frequently visits the small white

flowers of Aralia hispida, and Pieris rape delights in the white

or reddish flowers of the garden radish.

Bees not infrequently pass from one species to another in this

family, both when the flowers are closely allied and when they

are widely different. I have often seen bumblebees pass from

one species of goldenrod to another, and even back and forth

between goldenrods and asters. Occasionally I have seen them

pass between very different forms of flowers, as between sun-

flowers and the scarlet runner, or the goldenrod and the purple

vervain (Verbena hastata). On the other hand the honeybee

often displays a remarkable power of distinguishing between

closely allied species, even when they are of the same color.

One of the common golden-rods Se/idago lanceolata has its capit-

ula arranged in a crowded, flat-topped corymb. Another common

variety S, rugosa has the inflorescence panicled. In an upland

pasture these two species were found growing together, the pan-

icled form being much the more abundant. Honeybees, the

only insects present, showed a marked preference for S. /enceo-

lata, though they occasionally passed over to the other species.

They were repeatedly seen to leave S. /auceo/ata, and after flying

about but not resting on the flowers of S. /gosa return to the

plants they had left only a few moments before. In another in-

stance a bee was seen to wind its way among the plants of the

latter species until it found an isolated plant of S. /anceolata.

454 THE AMERICAN NATURALIST. [VoL. XXXVII.

A plant of each of the above species was bent over so that the

blossoms were intermingled, appearing as a single cluster; a

honeybee rested on S. /anceolata, and it seemed very probable

that it would pass over to the flowers of S. rugosa, but such

was not the case, for presently it flew away to another plant of

the former. The behavior of these bees in their endeavors to

adhere to a single species was thus attended both by loss of time

and repeated visits to the same blossoms. On another occasion

the whitish or cream-colored inflorescence of Solidago bicolor

was observed to be very frequently visited by the males of Bom-

bus bifarius, while the yellow-fiowered goldenrods in the vicinity

were entirely neglected. By holding yellow-flowered clusters

directly in their way, I repeatedly induced these bees to leave

S. bicolor; but they quickly perceived that they had passed to a

different flower, and invariably after a few seconds or sometimes

instantly returned to the cream-colored species. They were prob-

ably influenced by the greater supply of nectar in the flowers

of S. bicolor. The plants, which were growing on burnt land,

were of unusually large size, and secreted nectar very freely as

I ascertained by examination on my return home. These illus-

trations are sufficient to show that the influence of particular

colors in determining the visits of insects may be easily overes-

timated.

No. 439] NORTHERN GAMOPETALOUS FLOWERS.

455

THE COLORS OF NORTHERN GAMOPETALOUS FLOWERS.

: ; E d 5

"m S z e : T é 4

Orders. Families. 9 a 3 g 5 E 5

Go| + ood Geta lis ee ae e

Clethracae 2 2

| Pyrolace I 7 2 II

‘ Monotropacez I 4

Ericales < Erica ib 2 I P 10 5 38

Vacciniaceae zd abu II 23

E maet jud. E

iaaea a P lunfbaginaceze I I 2

(| Sapotaceze 2 2

Ebenales Ebghace : :

- Y 1 Symplocacez 1 I

Styrace 4 4

(| Oleaée 7 2 I 10

Loganiacez 2 I I 4

: Gentiánacez 7 I| 10 rr 38

Herpes 1 Menyanthacez 2 2 4

pocynaceze 2 I I il 7

Asclepiadacez TI 9 21: 51 39

Convolvulacee 7 ee, 3| 18

Cuschtacee 11 I 12

Poleihoniacez $ 10 31 8] 329

Hydrophyllacee 8 10 | 18

Boraginace: 48 1l12]| 4j

Verbefiaceze 2 $l 8-1

Labiatae 24 Pee Oe 47.| 331 120

Polemoniales — 4 | Solañaceæ 9| fi 2| 8| 40

he Scrophulariac e 13 | 33 "53238 113

ñtibulariaceæ 1I TL 2L m

Orobanch I 2 21 2 7

Bignoniacez 2 I I 4

Martyniacee I 1

[guten I E15 7

[| Phrymacee . I I

Plantaginales Prantaginaces 14 1 15

Xs Ru 4| 22 a TFSI X

Rubiales C aprifoliaces 22 | IE | 4 I 38

= Adoxac ; 1 be

í Valérianaceze 5 4 I

Valecunpies i Dipsacem . 4 4

Cucurbitac 4 I 5

Caffipanulaceze I BG

Campanulales Cichoriace wt $5813 2i 5i B

= Ambrosiaceze 15 15

Compositæ 21 | 126 | 209 | 4 | 64 | 50 | 483

. Total | 72 | 375 | 376 |106 | 198 |234 |1361

456 THE AMERICAN NATURALIST. (Vor. XXXVII.

SUMMARY AND CONCLUSIONS.

Numerical Summary.—ln the territory extending northward

from the parallel of the northern boundary of North Carolina

and Tennessee to the northern limits of Labrador and Manitoba,

and from the Atlantic Ocean westward to the ro2d meridian,

there are recognized in the Illustrated Flora of Britton and

Brown 4020 angiospermous plants. In the following table the

species belonging to the different series have been arranged

according to their predominant floral colors.

Series. Green. White. | Yellow. | Red. | Purple.| Blue. Total.

Monocotyledons 857 82 4I 22 22 34 1058

Dicotyledons

^ Choripetalze

Apetale er 175 89 51 45 24 384

Polypetale . 140 410 333 84 193 57 1217

Gamopetale . 72 375 376 106 198 | 234 1361

TM . 1244 956 801 257 4» 4 323 4020

In every 100 species there are 30.9 green, 23.8 white, 19.9

yellow, 06.4 red, 10.9 purple and o8. blue. The hydrophilous

and anemophilous species within this area, I place at about 1048,

of which 1021 are green, 1 white, 11 yellow, 3 red and 12

purple. A number of species vary between wind-fertilization

and insect-fertilization, and are differently classed by different

observers. Empetrum nigrum according to Warming is a wind-

flower, according to Lindman an insect flower, and according to

Knuth it is a wind-flower with occasional insect visits. There

are then in the district under consideration 2972 species, which

are fertilized by insects or are self-fertilized. Of this number

223 are green, 955 white, 790 yellow, 254 red, 425 purple, and

325 blue. In every 100 of these plants 07.5 are green, 32.1

white, 26.6 yellow, 08.5 red, 14.3 purple, and 10.9 blue. It is

evident that anemophily and small greenish flowers are corre-

lated, and that large bright colored flowers are due to insect

fertilization. The 1048 Anemophile and H ydrophilz are dis-

No. 439.] WORTHERN GAMOPETALOUS FLOWERS. 457

tributed as follows:— Monocotyledones 802 green; Apetalæ

134 green, 1 white, 11 yellow, 2 red, and 4 purple; Polypetalz

27 green, I red, and 8 purple; Gamopetale 58 green species.

The Pigments. — The colors of angiospermous plants are due

to three groups of pigments, occurring either singly or associated

together ; the green pigments or chlorophyll; the yellow pig-

ments which include carotin, xanthophyll and phyllofuscin ; and

the soluble red and blue pigments or anthocyan.

Chlorophyll. The characteristic green shades of foliage are

caused by chlorophyll, the most common of all plant pigments.

With the exception of the Fungi it is found in nearly all forms

of vegetation, though its presence is often partially masqued, as

in the Algae, by its association with other coloring substances.

Its wide distribution is explained by its activity in the synthesis

of carbohydrates. According to several late investigators there

is more than one kind of chlorophyll. This view was adopted

in 1895 by Gautier and Etard. Kohl in his recent work on

* Carotin" admits of two varieties, which he designates as a-

chlorophyll and f-chlorophyll.!. In a green leaf “the normal

chloroplasts contain much a-chlorophyll, little | 8-chlorophyll,

much carotin, little a-xanthophyll, and little 8-xanthophyll." ?

The a-chlorophyll is to be regarded as pure chlorophyll. Its

absorption bands lie in the red half of the spectrum. The

genetic relations of chlorophyll require further investigation.

Wiesner's theory that etiolin is the mother substance of chloro-

phyll has not been proven; and, according to Kohl, it can be

shown that in the greening of etiolated plants chlorophyll is not

formed at the expense of the etiolin. The different shades of

green observable in foliage are due partly to the quantity and

arrangement of the chloroplasts. The upper side of a leaf is

usually a darker green than the lower, because the palisade cells

contain three or four times as many chlorophyll granules as the

spongy parenchyma of the lower side? Ferns and mosses,

which habitually live in shady ravines, are a deeper green in

! Kohl, F. G. Untersuchungen über das Carotin und seine physiologische

Bedeutung in der Pflanze, p. 139

2 Ibid, p. 145.

3 Kerner. Natural History of Plants, vol. i, p. 374-

458 THE AMERICAN NATURALIST. [Vor. XXXVII.

such locations than when they grow in the open sunlight. The

color is also affected by a change in the position of the granules

under the action of intense light, as may be observed in Lemna

trisulca and many seaweeds.! Chlorophyll is readily soluble in

alcohol yielding a green solution, which is soon destroyed in

direct sunlight. There is a constant destruction and renewal of

chlorophyll in living leaves under the action of bright light, so

that on the same plant the leaves present different shades of

green. Green seaweeds, when left on the beach by the waves,

soon turn yellowish owing to the destruction of the chlorophyll.

Leaves and flowers may in some instances owe their particular

shade of color to the presence of chlorophyll mixed with some

other pigment. The dull purple of Scopolia atroipoides and

Atropq. belladonna, according to Hildebrand, are caused by green

grains mingled with violet-colored sap. In the gooseberry, says

Móbius, the brownish color of the flower is due to an upper layer

of cells containing red cell sap, and an under layer containing

chlorophyll. Many greenish yellow and purple flowers appear

to contain chlorophyll. The tints of autumn leaves are also

modified by its presence in greater or less quantities, while in

normal green leaves it is often accompanied by anthocyan.

= Yellow Pigments.— Chlorophyll is invariably accompanied in

the chloroplasts by carotin, the yellow pigment so common in

the root of the carrot. Tammes? and Kohl; found carotin to

be widely distributed in the blue, green, red, and brown Algee ;

in the Fungi, lichens, mosses, and ferns; in green, yellow,

etiolated and autumn leaves; and in flowers, fruits and seeds.

There is, however, no evidence of any genetic relation between

the two pigments; and carotin may exist independently in

organisms in which chlorophyll does not occur, as in Bacteria,

fungi, the root of Daycus carota and in yellow flowers and

leaves. Kohl finds that etiolin is identical with carotin, and

adds that the term etiolin in the sense used by Pringsheim

z gii Physiology of Plants, p. 618,

*'Tammes, Tine. eber die Verbreitung "e Carotins in Pflanzenreiche.

Flora wy Allg. bot. petens Bd. 87, H. 2, p. 24

? The distribution and properties of the yellow india are P at length

in Kohl’s exhaustive work on Carotin.

No. 439.1] NORTHERN GAMOPETALOUS FLOWERS. 459

should be stricken from the list of plant pigments. Etiolated

plant organs owe their coloring exclusively to carotin, with which

is often associated anthocyan. Also identical with carotin are

xanthopyll and anthoxanthin as these terms are commonly used.

Carotin (Cs, Hss) is easily dissolved by ether but is insoluble in

water. The melting point is 167.8C. Concentrated sulphuric

and nitric acid color it a dark blue. Its crystals are rhombic.

The functions of carotin, according to Kohl, are threefold. First

it aids in assimilation. Its absorption bands lie in the blue half

of the spectrum, and, together with those of chlorophyll, give

the absorption spectra of the crude leaf-green. “Both take an

important, though unlike part, in the assimilatory work of the

chloroplasts, both absorb supplementarily to each other a part of

the sunlight and assist in the deconiposition of the atmospheric

carbonic acid." Secondly, carotin may serve as a reserve product,

as in a number of Fungi and Algz and in the root of Daucus

carota, Thirdly, it is of biological importance because it renders

flowers, fruits and seeds conspicuous and attractive to insects and

birds, which aid in their fertilization and dissemination. Among

the flowers which owe their yellow color to carotin are, Abutilon

nervosum, Adonis vernalts, Cucurbita pepo, Eranthis h yemalis,

Forsythia viridissima, Geum montanum, Helianthus annuus,

Impatiens noli-tangere, Kerria Japonica, Enothera biennis, yel-

low flowered roses, Taraxacum officinale, and Tr ropeolum majus.

In the peel or pericarp of the lemon, in the flowers of the

yellow dahlia, in Linaria vulgaris, Corydalis lutea, the yellow

parts of Antirrhinum majus, and in all the yellow flowering

thistles, as well as in other flowers, the yellow pigment does

not occur in plastids, but dissolved in the cell sap. What is this

pigment? In a solution of crude leaf-green, in addition to caro-

tin, there are two other yellow pigments, one of which was

obtained by Tschirch in 1896 and the other by Schunck in

1899. Kohl proposes to designate the latter of these two pig-

ments as a-xanthophyll and the former as #-xanthophyll. They

differ both in their absorption spectra and chemical reactions.

The a-xanthophyll occurs in small quantities in normal chloro-

plasts and yellow autumn leaves. It is the -xanthophyll which

colors the peel of the lemon and the flowers with yellow cell

460 THE AMERICAN NATURALIST. [Vor. XXXVII.

sap. Both carotin and B-xanthophyll occur in species of Ranun-

culus, Verbascum, Caltha palustris and Ribes aureum. The

B-xanthophyll can be obtained in a yellow solution by boiling in

water the peel of the lemon. It becomes brown-colored with

sulphuric acid and with ammonia a deeper yellow. This pig-

ment was first isolated from the flowers of the dahlia nearly half

a century ago.

In the chloroplasts of golden yellow-leaved plants, as Sam-

bucus and Evonymus, Kohl finds yet another yellow pigment

largely soluble in water to which he gives the name of phyllo-

fuscin. In addition to this pigment he finds in yellow leaves

much carotin, and more or less 8-xanthophyll, but no a-xantho-

phyll or chlorophyll. Though they contain no chlorophyll such

plants grow and perform the work of assimilation, in which

process the chief part must be ascribed to carotin. Finally in *

yellow autumn leaves there is little or no chlorophyll, about

the same amount of carotin as in the green leaf, little a-xantho-

phyll and much £-xanthophyll.

The yellow plastids of flowers are usually round and small,

though sometimes angular as Tropzeolum. Several other modi-

fications also occur. In the tomato, asparagus, Crategus

coccinea, and in some species of Rosa and Physalis the plastids

of the fruit are spindle-formed or irregularly shaped, and are fire-

red, orange-red, or yellowish red. Tammes found that the red

plastids of the tomato gave the usual reaction for carotin. In

yellow leaves the plastids are round, but in autumnal yellow

leaves they occur in irregular masses. The scarlet poppy, tulip

and fire red canna owe their colors to a mixture of yellow plas-

tids and red cell sap. On the other hand dingy or dull colors

result from a combination of violet sap with yellow granules.

Anthocyan—vThe red and blue colors of leaves, fruits and

flowers are produced by a soluble pigment termed anthocyan.

The ecological significance of this coloring substance, which is

widely distributed in plants, is important and deserves further

study. It is of frequent occurrence on the stems, veins and

leaves of herbaceous plants, as well as on the under side of

aquatic leaves and of radical leaves growing in rosettes, as in

the Cruciferee. In early spring, in autumn, and at high ele-

No.439.] WORTHERN GAMOPETALOUS FLOWERS. 461

vation, it is particularly abundant. It probably serves to con-

vert light rays into heat, and at the same time protects and aids

in the translocation of the food materials. As in the previous

instances we have undoubtedly to deal with a group of pigments.

The formation of anthocyan has been studied by Overton with

the aid of cultures of aquatic and land plants. Experiments

with water cultures of Hydrocharis showed that light intensity

and low temperature were favorable to the development of red

cell sap. Plants of Hydrocharis were placed in a 2% solution of

invert sugar, and also in pure water. The conditions of light and

temperature were such that the water culture plants showed no

change in color, while in a few days the plants in the invert

sugar solution developed dark red coloring, especially in the new

leaves. Experiments with other aquatic plants gave similar

results. The red cell sap was contained chiefly in the palisade

cells, though extending also to other cells of the mesophyll.

Cut stems of Lilium martagon and other land plants placed

in a 2% invert sugar solution soon developed red color in

the palisade cells. The leaves of the control plants remained a

pure green. As the result of many observations Overton con-

cludes that a cell sap rich in sugar, low temperature, and intense

light are connected with the production of red color. During

the summer in the Alps the leaves of plants are much oftener

red-colored than in the lowlands, because the night temperature

is lower and the light intensity higher. "Winter leaves become

red-colored since the lower temperature causes the sugar con-

tent of the leaves to increase at the cost of the starch. In the

ripening of red and violet-colored fruits the appearance of the

coloring is also attended by the conversion of the starch into

sugar. A few experiments were made to determine whether

white flowering varieties of certain plants could artificially be

caused to vary into red flowering varieties,- but with negative

results. In the case, however, of the greater intensity of color

in Alpine flowers, and of white lowland flowers which become

red-colored in the Alps, and also of flowers which are brighter

colored in early spring than later in the season, it is probable

that the lower temperature causes the conversion of starch into

sugar.

462 THE AMERICAN NATURALIST. [Vor. XXXVII.

The red pigment is probably a glucoside, or a very closely

related compound, of which the constituents are a sugar and a

tannic acid. Since in many plants, the provision of the cells

with sugar increases the tendency to form red cell sap, there

can be little doubt that a sugar forms part of the raw material

out of which the pigment is built up. Tannins are also con-

tained in the cells in which the red color has been formed by the

artificial increase of sugar. The red color stuff is thrown down

by the tannin reagents coffein and antipyrin, and the precipitate

closely resembles those of the tannins. The behavior of the red

pigment indicates that it isa tannin compound. The supposition

that tannin is connected with the formation of the red and blue

pigments of flowers is not new, but was first suggested by Wig-

and in 1862. . It was observed that red color was formed only in

cells that contained tannin. “If we examine," says Overton,

*the reaction of the red color stuff upon different bases we

obtain support for the opinion, that this pigment represents a

weak bivalent or multivalent acid. For we find that its tinge is

almost unnoticeably changed by very weak bases as coffein,

antipyrin, etc, that with stronger bases, however, the color

turns first into violet and blue, and with a greater excess of a

strong base it finally changes into green. The most simple

explanation of these phenomena is that the free acid is only

little dissociated electrolytically and that the red color is peculiar

to the molecules of the acid that has not been dissociated, the

blue color would belong to the univalent, and the green color to

the bivalent ions of the acid. On account of the weakness of

the acid the bivalent ions would be found — in consequence

of hydrolytic dissociation — in larger quantities only when a

certain excess of a base is present." The capability of forming

red cell sap appears to belong chiefly to the phanerogams, for

the red color of mosses is confined to the cell membrane.!

Many of the pigments found in plants and used for coloring are

glucosides. The indigo blue of commerce is derived from the

glucoside indican, which occurs in the plants of the leguminous

genus Indigofera. Indigo red is also obtained from this gluco-

lOverton, E. Beobachtungen und Versuche über das Auftreten von rothem

Zellsaft bei Pflanzen. Jahrb. wissensch. Botanik, Bd. xxxiii, H. 2.

No. 439.] NORTHERN GAMOPETALOUS FLOWERS. 463

side. From indigo may readily be obtained aniline remarkable

for the great variety of dyes which it yields.

In darkness flowers differ greatly in the extent to which they

develop their colors. Sz/ene pendulata fails to show red color-

ing and Prunella grandiflora instead of developing dark violet

color remains a pure white; while 7 ultpa gesneriana forms its

red color and Crocus vernus its blue violet as perfectly in dark-

ness as in light. The explanation given by Sachs, where bulbous

plants produce normal flowers in darkness, is that the flower

forming substance was already collected in the bulb, and had

been stored up in a preceding period of vegetation in bright

sunlight. Leaves, flowers and fruits often display red coloration

only on the side exposed to direct sunlight. Kerner found that

the anthocyan in plants grown in an Alpine garden at an eleva-

tion of 2195 metres above the level of the sea was brighter

colored and more abundant than in the botanical garden at

Vienna. At a high elevation the glumes of grasses, the leaves

of stonecrops, and the pure white petals of some flowers become

red or purplish-red.

When a red flower or a solution of red cell sap is treated

with an alkali it changes to blue, but the red color is again

restored by an acid. Red color is more common in foliage

(where it is termed erythrophyll) than blue because an acid con-

dition usually prevails in the leaf cells. In the Boraginaceze

with a decrease in the acidity of the cell sap the flowers change

from red to blue; while an increase in the acidity of the cell sap

will cause a normally blue flower to vary into a pink variety.

* [n some rare instances the blue pigment occurs in a solid form

in flowers and also in fruit." In the fruit of the nightshade

Solanum americanum the coloration is due to intense violet-

colored crystalloids of rhombic form or in thin six-sided plates.'

Blue grained pigments also occur in Strelitzia regina, Tillandsia

amena, and in Delphinium elatum? The occurrence of blue

pigment in solid form is probably to be explained by the evap-

oration of the free water. It never occurs in chromoplasts.

Cells containing red and blue sap may occur indiscriminately

! Möbius, M. Die Farben in der Pflanzenwelt, p. 15.

2 Hildebrand, Friedrich. Die Farben der B/üthen, p. 45.

464 THE AMERICAN NATURALIST. [Vou. XXXVII.

near each other in the same flower, or the epidermis may con-

tain blue cells beneath which in the mesophyll is a layer of red

cells, as in Viola odorata. Yellow chromoplasts and anthocyan

occurring together give scarlet hues. The shades of flowers

depend upon the density of the chromoplasts, and the number of

layers of pigment cells, and the character of the epidermis.

Green Flowers.— Of the 223 green flowers classed as ento-

mophilous many have no petals, as fifteen species of the Poly-

gonacez and eight species belonging to the Caryophy llaceze,

also in several Rosaceze, in Acer saccharinum and Didiplis dian-

dra. Many are self -fertilized, as Triglochin and Scheuchzeria,

and the orchids Habenarea hyberborea and Epipactis viridiflora,

and the small green flowers of Lechea and Penthorum sedoides.

Some have the petals caduceous and depend upon | their scent to

attract insects, as the Vitaceæ. Many are visited by flies and

the smaller bees, as various Melanthacez, the Smilacez, the

Anacardiacez, and the green flowers of the Asclepiadaceae. But

the green flowers of Asparagus are visited by the honeybee. As

a whole, green flowers are small or even minute and attract few

insects. A transition stage is represented by the genus Ribes,

which contains species with greenish, white, reddish, and yellow

flowers. As is well known many flowers pass through a green

stage before their bright colors appear. Large green flowers,

which are chiefly fragrant and nocturnal, are found in exotic

Solanaceee. Other examples are exhibited by the orchids, as

several Brazilian species of Epidendrum. Green flowers, except

in some cases of retrogression, belong to an early stage of devel-

opment and their coloring requires no special explanation. The

petals are modified leaves, and their primitive color is green sim-

ilar to that of foliage. The larger green flowers may be

explained by the greater persistency of the chlorophyll, for some

species hold their colors much more strongly than others.

Yellow Flowers —The development of bright coloration in flow-

ers is an acquired habit. This is well illustrated by the sepals

of HelleZorus niger, which at first are large and white, but after

fertilization develop chlorophyll, become a fresh green color and

‘Mobius, M. Die Farben in der Pflanzenwelt, p. 3.

No. 439.] NORTHERN GAMOPETALOUS FLOWERS. 465

act as leaves. A similar change has been observed in many

orchids and liliaceous plants. Virescence, or the occurrence of

green flowers instead of those of the normal color, has been

observed in many Ranynculacez, Umbelliferee and Compositze.!

The formation of chlorophyll has but to cease, and under the

action of light the petals will quickly lose their green color, with

the result that in most instances the flower will change to yellow

or white. If the yellow pigments, which are invariably associ-

ated, as has been shown, with the chlorophyll in the chloroplasts

are persistent and continue to increase, the color of the flower

will be yellow. The quantity of yellow pigments, it will be remem-

bered, varies greatly in different plants. In some they are

scarcely perceptible, in others they are so abundant as to tinge

the whole plant yellow, while in a few golden yellow species they

exclude all other pigments even the chlorophyll. If, however,

the yellow color also vanishes we havea white flower. As would

be expected yellow and white flowers are the most common, and

are the earliest of the floral colors in their origin. A large num-

ber of yellow and white flowers with a mostly small, regular and

primitive perianth occur in widely separated families.

Melanthaceae 7 IO 5 2 24

Miacee . . 6 13 1 LI I 6 38

lygonacez 5 22 33 II 3 74

RÉnunculaceaé 38 26 6 3 13 " 97

iferz: .. 46 54 2 I IO I13

Saxifragacee 6 30 4 3 43

BHOCHE 24 Sin 39 35 4 13 4 95

Onagraces: . o . 24 14 3 rO 6 57

. Umbellifere . . 16 58 1 3 78

Primulacee . . II 4 7 22

Sol 21 9 2 8 40

Cichoriacem . . . 53 8 5 2 5 73

Composite . . . 209 126 21 4 64 59 483

Tet i v... 481 409 79 65 | 1n 92 1237

Many species of Composite, it will be noted, retain their prim-

itive colors. In a few families white flowers occur unaccom-

1 Masters, M. T. Vegetable Teratology, p. 339-

466 THE AMERICAN NATURALIST. [Vor. XXXVII.

panied by yellow. In the aquatic Alismacez the entire nineteen

species are white, and in the Caryophyllacez there are fifty-six

white flowers but no indigenous yellow species. The six species

of the Xyridacez on the other hand all produce yellow flowers.

In the anemophilous Betulacee there are eleven yellow species,

but flowers with a yellow calyx are rare in the Apetale. The

Hypericaceze are nearly monochromatic as twenty-two species

are yellow and only two red. The zygomorphic Orchidacez con-

tain ten yellow-flowered species, a larger number than any other

monocotyledonous family. A surprisingly large number of yel-

low flowers occur in the zygomorphic Papilionacez (33 species),

the Scrophulariaceze (33 species), and the Lentibulariacez (11

species). This fact Miiller attributes, and we think rightly, to

the persistence of the primitive yellow in certain genera, and its

little tendency to variation with the specialization of the flowers.

In many families of the Gamopetale yellow flowers are absent,

or are represented only by a single species, as in the orders Eri-

cales, Ebenales, and Gentianales, where the inflorescence is

chiefly white or red.

White Flowers.— White flowers are most abundant in the

American as well as in the European flora. A white inflores-

cence is evidently a less tax on the energies of a plant than one

containing pigments ; and trees and shrubs, which produce their

flowers in almost boundless profusion, as the Pomacez, Drup-

aceze, Ilicaceze, and the genus Viburnum, have almost exclusively

white blossoms. In the writer’s opinion white flowers are pri-

marily due to degeneration. In this connection the studies of

white leaves by Rodrique, Laurent and Timpe, which clearly

show evidences of degeneration, are of interest. According to

their investigations such leaves are thinner than normal green

leaves, and consist wholly of cellular tissue with the palisade

cells absent. It is desirable to consider very briefly some of the

conditions under which white flowers occur, and under which

they develop chromatism. They are derived both from primi-

tively green and from high colored flowers. Small, densely

clustered white flowers are common in the Cruciferz, Saxifrag-

aceæ, Umbelliferae, Cornaceze and Ericaceze. In these flowers

the stimulus to produce pigments is wanting and the leaf-green

No. 439.] NORTHERN GAMOPETALOUS FLOWERS. 467

colors, as may be observed in the Cornacez, fade away leaving

the petals white. A check in nutrition and growth will cause

bright colored flowers to become smaller and revert to white.

This may be caused by cultivation in an impoverished soil, by

transplanting, or by low temperature. In springtime white

flowers are noticeably common. In the Baltic flora the graphic

curve of white reaches its highest point in May, from which

it gradually sinks to its lowest point in late autumn. In the

arctic climate of Spitzbergen the flowers are chiefly white, and

there are few yellow and red, while blue appears to fail entirely.

In East Greenland the flowers are likewise chiefly white, and

among twenty-six species there is only one blue. Self-fertiliza-

tion also causes a diminution of the corolla in size and a paleness

or loss of color. Bright colored flowers fertilized artificially

with their own pollen in a few generations become paler; while

white flowers, as would be expected, and what is more surprising

white varieties of colored flowers adapted to insect-fertilization,

are both usually self-fertilizing. They may also exhibit evidences

of deterioration in their structure, as in Lepidium, Stellaria, and

Sagina, where the petals are usually present but sometimes are

wanting. In all of the instances cited there is a lack of vitality

in the corolla due to insufficient nutriment. Let the growth of

the plant now receive a stimulus and an increased brilliancy of

the flowers soon makes itself apparent, as when they are exposed

to clear sunlight or treated with nitrate of soda, and may also be

observed in the flushing of tulips, by which they lose their

variegated colors when treated with strong manure. The bright-

ness of the floral hues is also increased by crossing. When a

white flower is crossed by a yellow, red, or blue flower, a part of

the hybrid offspring contain pigments. When lowland white

flowers have been cultivated in the intense light of alpine heights,

they have in some species become red. Though the conditions

are abnormal a rapid development in size and color in an individ-

ual flower may be caused by the sting of a gall-fly; for example,

all of the organs of Crategus coccinea become bright red and

the change of coloring is accompanied by an increase in size.

; Hildebrand, F. Die Farben der Bliithen, p. 70.

2 Henslow, G. Oz the Self-Fertilisation of Plants, p. 327.

468 THE AMERICAN NATURALIST. (VoL. XXXVII.

The appearance of bright coloration is here marked by an

increased protoplasmic activity.

This view of the origin of white flowers explains why they are

commonest in Nature, accounts for their being most numerous

in families in which yellow flowers are likewise numerous, and

why they are most true to name under cultivation. We can

also understand that such flowers under forcing would be more

likely to develop a desired color than one already containing

pigments.

Red Flowers.— From its wide distribution among plants red

coloring naturally follows yellow and white in flowers. Light

which is destructive of chlorophyll stimulates the formation of

anthocyan. With the increase of white flowers in size and

vitality, accompanied by an increase of the sugar content,’ they

very frequently develop red coloration. In the Rosacez and

Pomacez a series of flowers illustrates every step of the transi-

tion from white to red. The species of Rubus and Cratzgus

are usually white or occasionally red, but Rubus odoratus is

purple red with a white form. In the familiar genus of Malus

the species are tinted or strongly shaded with rose, which in the

fragrant flowers of M. coronaria becomes the predominant color.

In Rosa the species are regularly rose or pink varying in several

species to white. Red flowers are derived often from white,

sometimes from yellow, and occasionally by reversion from blue.

They are the rarest in our flora. There are twenty-two species

in the Monocotyledons, forty-five in the Apetalz, eighty-four in

the Polypetale, and one hundred and six in the Gamopetale.

Red flowers occur both in the older and more recently evolved

families, while blue flowers are restricted to the latter. Red

coloration must be regarded of earlier origin in the sequence of

floral colors than blue ; and, as has been already pointed out, it is

also much more common in the vegetative organs of both the

angiosperms and cryptogams. In the following families red and

blue and blue-purple flowers are the most common :

! It is not unlikely that the higher intensity in color of Alpine flowers is due to

an increase of the sugar content, but, according to Overton, in most cases of

white-flowered varieties it is probably that some other compound rather than a

sugar is wanting.

No. 439) WORTHERN GAMOPETALOUS FLOWERS. 469

| A iments Flas a — — een ———

Lilaceg oo. onc i71 | I 13 38

Orchididacee . . 8. 5 ere 10 18 ti 61

Polygonacee . . LI | D ^g 5 22 s4 ] 74

Caryophyllaceæ — . 2$ | a | 56 8 88

sace i 13 | 39 1 3 4 9

Papiionacee . . 13 | € 88 | 33 39 197

al ji 1835 4 4 CREE. 26

Onagiacee . . . TON 6 24 14 3 57

Ene. 5 i8] & Tas I 38

Vacciniacee . . . II | 10 2 2

Gentianacee . . IO 16 (I 7 38

Pofemoniacee . . 10 8 3 | 7 28

Labiate . . . - 12 33 47 | 4 2 120

Tod: x. 154 87 181 | 149 | 250 62 | 883

It is evident that the families containing red flowers may be

separated into two series. In the first, which includes the

Polygonaceze, Caryophyllacez, Rosacez, Malvacez, Onagracez,

Ericacez, and Vacciniaceze, there are red flowers but no blue.

These families are primitive with regular flowers, which are

frequently of small size and but little modified. The Orchidacez

offer an exception in which, however, though there are no blue,

there are fourteen purple flowers. In the second series, which

includes the Liliaceze, Papilionacece, Gentianacez, Polemoniacez,

and Labiatz, there are both red and blue flowers, which are

highly specialized and often dependent on insects for fertilization.

Purple flowers belonging to the first series are chiefly red-purple,

while those of the second are blue-purple. The Rosacez and

Papilionacez are “sister families,” according to Engler; both

contain red flowers but there are no blue flowers in the Rosacez

though they are numerous in the Papilionaceee. The distribu-

tion of the red and blue coloration is probably to be explained by

the strong acidity of the cell sap in the first series, and its more

nearly neutral condition in the second, so that a comparatively

slight variation in the chemical conditions permits the develop.

ment of either a red or blue flower. A part of the hybrids

obtained by Darwin by crossing the red and blue species of

Anagallis were red and a part blue, while others were inter-

mediate in color. The same observer also records having seen

470 THE AMERICAN NATURALIST. (VoL. XXXVII.

a hyacinth with a truss of flowers perfectly blue on one side and

perfectly red on the other. Several of the flowers were also

striped longitudinally red and blue.

Anthophain. — n most instances the brown colors of flowers

are caused by a mixture of chlorophyll or carotin with anthocyan.

Among brown flowers containing two pigments are Calycanthus

floridus, Veratrum nigrum, Aristolochia glauca, Anona triloba,

Asarum, 4 donis vernalis, Ribes grossularia, and various species of

orchids. But in the black spots and brown markings on the flow-

ers of Vicia faba and of some species of Delphinium, Mobius

finds an olive brown pigment dissolved in the cell sap.! As its

chemical reactions and optical properties are sufficiently charac-

teristic to distinguish it from other plant coloring substances

he proposes tor it the name of anthophzin. The spots on the

wings of Vicia faba appear black largely because of the papilla-

formed structure of the epidermal cells, which become flatter

where the markings are brown. The properties of anthophzin

are very similar to those of phycophzein, the pigment peculiar to

the brown Algae ; but it differs from this substance in that it is

dissolved in the cell sap, while phycophaein, together with chlor-

ophyll, occurs in chromatophores. It is also less soluble in

water. Phycophzin is characteristic for an entire class of

plants, while flowers containing anthophzein are rare.

Purple Flowers. — There are twenty-two purple flowers in the

Monocotyledons, twenty-four in the Apetala, one hundred and

ninety-three in the Polypetale, and one hundred and ninety-eight

in the Gamopetalz. Purple flowers may be divided into green

or lurid purple, red purple, and blue purple. In the Melanthaceze

there are two small greenish-purple flowers adapted to Diptera.

In Trillium of the Convallariacee are four lurid purple flowers

visited by flies. In the Aristolochiacezee which are also adapted

to Diptera the calyx is lurid "purple. These flowers appear to

have been derived directly from the primitive green without pass-

ing through an intermediate stage. Greenish-purple flowers also

occur in the Polygalacezee and Asclepiadacez. Numerous other

families contain a few small purplish flowers, which evidently

! Móbius, M. Das on eta der braune Blüthenfarbstoff, Berichte sr

schen botan. Gesell. ess xviii, p. 341

No. 439] NORTHERN GAMOPETALOUS FLOWERS. 471

belong to a primitive stage of coloring. There are a few flowers

which are yellowish-purple. The petals of Asinima triloba are

at first greenish-yellow changing to a dull purple. In Geum

rivale the petals are purplish-orange and the calyx brown-purple.

Red-purple flowers belong toa higher stage of coloration. They

are common in the Orchidacez, Geraniaceze, Lythracez, and

Onagracez. Blue-purple are the most advanced of all, and are

common in the Papilionaceze, Labiate and Scrophulariacez,

families which contain numerous blue bee flowers, to which they

are akin in form and color. Many purple flowers also occur in

the Composite which are partly discoid and partly radiate.

Except in a few species where the color stuff is the rare olive

brown anthophzein, brown and brown-purple flowers usually con-

tain more than one pigment.

Blue Flowers. — There are only thirty-four blue flowers in the

monocotyledons of the Northern States, which belong chiefly to

the Commelinaceze, Liliaceae, and the Iridaceze. In the Apetalz

there are no blue flowers, and the purple flowers in this series

are primitive in their stage of coloring. The rarity of blue

flowers continues in the Polypetala. They are most common in

the Ranunculacez, Papilionaceze and Violacez. In the more

primitive “families of the Gamopetalz belonging to the orders

Ericales, Primulales and Ebenales blue flowers are again absent.

They belong chiefly to the three orders, Gentianales, Polemoni- .

ales and the Campanulales. It is, however, in the Polemoniales

that blue and blue purple species reach their maximum. There

are many bee flowers greatly modified both in form and color and

displaying a high degree of variegation. The culmination of

color specialization, as has been previously shown in detail, is

reached in this order. It will be observed that blue flowers

occur almost exclusively in the most specialized families, or when

they are present in a more primitive family, as in the Ranuncu-

lacez, it is in genera which have been highly modified, as in

Delphinium and Aconitum. These families and genera are the

most recent in their evolution, and blue is consequently the most

recent of the floral colors to be developed. Blue flowers are

usually derived from red or white forms, but in several families

they appear to have yellow-flowered ancestors. Müller believed

472 THE AMERICAN NATURALIST. (Vor. XXXVII.

this to be the case in the Violaceze and in Gentiana and in

Myosotis versicolor. The sequence of the floral colors has been

determined by the properties and distribution of the plant pig-

ments, rather than by the selective influence of insects.

Two Color Series. — The colors of flowers may be divided into

two series, a primitive series consisting of green, white and yel-

low, and a derivative series composed of red, purple and blue.

In the first the pigments are insoluble and are contained in

plastids or are absent. In the second they are dissolved in the

cell sap. Of the 4020 northern angiosperms 3001 belong to the

first series, while ror9 belong to the second. Of the 2972

entomophilous species 1968 belong to the first and 1004 to the

second series. The flowers of the second series are far more

numerous in the Polypetalae and Gamopetale than in the Mon-

ocotyledones and Apetalce. The pigments of the first series are

most common in primitive families, where the flowers are rotate

and but little modified. Very many flowers of the second series.

have the petals green, whitish, or yellowish in the bud or at the

base. In the color changes which takes place in individual

flowers green may be succeeded by every color, and red and blue

frequently pass through a white or yellow stage. In individual

flowers the tendency of green, white, and yellow to change to

red and blue is much stronger than the reverse.

Pigments not Induced by [nsects.—'The function of forming

pigments has not been induced by insects. It is a property

possessed by all plants from the lowest to the highest. Not

only chlorophyll but carotin and other pigments are widely distrib-

uted among the alge. This function is fully developed even

among minute unicellular plant organisms. The chromogenous

Bacteria are capable of producing colors of remarkable intensity,

as red, rose, yellow, orange, green, blue, violet, and black. Four

different pigments, as black, blue, green, and yellow, are pro-

duced by the Bacillus pyocyanus. The red of Micrococcus pro-

digiosus can be extracted by alcohol, discolored by alkalies and

restored by an acid. Intense light and acids in small doses

increase the production of the pigments, and the alkalies have

the reverse effect. ^ Bohn considers the study of Bacteria as

! Bohn, G. Z' Evolution du Pigment, p. 44.

No. 439.]] NORTHERN GAMOPETALOUS FLOWERS. 473

of much interest from their supposed similarity in origin and

composition to the pigment granules. According to this author

the chromoplastids have their origin in the nuclear chromatin,

and are designed to protect the organism from the chemical and

physical variations to which it is exposed. A remarkable

difference is exhibited by plants in their capability of forming

pigments. The four great divisions of the Algz are character-

ized by the presence of a green, blue, brown, or red pigment,

which in the last three classes is so abundant as to completely

mask the chlorophyll. The Fungi display many brilliant colors,

which in the Phalloideze become ecologically important, and pro-

phetic of their attractive office among the Phznogams. In

this family flesh flies are allured by the bright coloring, associ-

ated with a sweet substance and a nauseous scent, and aid in

disseminating the spores. In their form and vivid colors the

Balanophoracez show a marked resemblance to Fungi. Many

conifers and deciduous shrubs and trees display a bright yellow

foliage, from which chlorophyll is absent. There is also a great

variety of trees and shrubs and herbaceous plants both in tropi-

cal and temperate regions, which possess a red, purple, or varie-

gated foliage, which is highly ornamental. Conversely many pale

green species exhibit scarcely a trace of bright coloration. The

petals are only modified leaves and their colors are closely cor-

related with the coloration of the vegetative organs. It is often

possible from the inspection of the stem and leaves of a plant

to determine the color of the flowers it will produce.

Of no Physiological Significance.— With the exception of

chlorophyll the pigments are of no physiological significance in

the development of flowers. Their function is wholly ecological,

and any other effect they may produce is slight and incidental.

Negative evidence of this is furnished by the great number of

white flowers. Red and blue coloring frequently does not appear

until the flower is on the point of expanding. And even after

fertilization or in wilting the colors may brighten or change.

. Bright coloration in flowers, as in fruits, marks the approach of

maturity and decay. According to Massee many of the beauti-

ful colors of fungi are of no obvious use.'

1Massee, G. Evolution of. Plant Life, p. 145.

474 THE AMERICAN NATURALIST. [Vor. XXXVII.

Conspicuousness Due to Insects. — Bright coloring in flowers,

usually accompanied by an enlargement of the perianth, has been

evolved through the agency of insects. Wind-flowers are small

and green or dull colored. “In New Zealand where insects are

so strikingly deficient in variety, the flora is almost as strikingly

deficient in gaily-colored blossoms."! In many genera as the

flowers become more conspicuous, there is an increase in the

number of visitors and the power of self-fertilization is lost. A

colored perianth, which contrasts strongly with the surrounding

green foliage, can evidently be more easily seen by both insects

and birds. For the same reason a contrast in color between

different species in blossom at the same time is advantageous.

Insects would be likely to make their visits indiscriminately in a

monochromatic Flora, as now happens in the case of similarly

colored species of buttercups and goldenrods. In the Alps,

where owing to the shortness of the summer all of the species

blossom at the same time, there is the greatest variety of colors.

It is a well known principle of physics that when a red and yel-

low card are placed side by side each appears more brilliant than

when viewed alone, that is the effect of bringing two colors not

complimentary in competition is to move them farther apart.’

The utility of color contrast is sufficient to explain the evolution

of floral colors without recourse to the hypothesis that they afford

pleasure to insects.

Insects and Flowers — The influence of insects upon the evo-

lution of flowers has undoubtedly been greatly overestimated.

There is certainly no satisfactory evidence that the ancestors of

all angiospermous flowers were once entomophilous, and that the

wind-fertilized forms are the result of degeneration. In my

opinion not only the principal plant series but many families and

genera were developed before the habit of flower visiting became

established. The formation of this habit must have required a

considerable interval of time. Neither is there sufficient evidence

to support the claim that the color of every flower has been

determined by the pleasure it afforded to the pollinating insects.

| Thompson, George M. Fertilization of New uu ird e Plants, Zrans.

Proc. New Zeal. Inst. 1880. Opinion of A. R. Wallac

7 Rood, O. N. Zext-Boo£ of Coler, p. 246.

No. 439] WORTHERN GAMOPETALOUS FLOWERS. 475

Some of the adherents of this theory have not, however, hesi-

tated to cause a flower to undergo several changes of color in

order that its present hue may conform to their imaginary views

of its origin. Further observations are required to determine

how far a sense of color is developed among insects, but the

writer believes that the colors of flowers have determined the

color sense of insects rather than the converse. It is desirable

to review briefly the evidence of a preference for certain floral

colors in the four orders of insects, —the Coleoptera, Diptera,

Lepidoptera, and Hymenoptera,— which are important as flower

visitors.

Coleoptera. — There is no evidence that the Coleoptera have

exerted any influence on the particular coloration of flowers.

They are often poorly adapted for flower visiting, a habit which

they have acquired at a comparatively recent date. They prob-

ably give the preference to bright colors, but they do not avoid

dull yellow or green. None of our northern species are adapted

to Coleoptera, but they are very frequent visitors to rotate clus-

tered white flowers like Viburnum. No inference can be drawn

from the beautiful markings often displayed by beetles that they

take pleasure in the colors of flowers, for the most intelligent of

all flower visiting insects, the bees, wear the plainest dress.

Diptera. — The Diptera as fertilizers of flowers are of much

greater importance than the Coleoptera. The Syrphidz have

been thought to hover with delight over bright golden yellow

flowers ; while the carrion flies, it has been asserted, are attracted

by a lurid red or purple inflorescence. In number and impor-

tance as flower visitors the Syrphide, or drone flies, surpass all

other Diptera. The light blue Veronica chamedrys, the rose

pink V. urticifolia and the white Circea Lutetiana are adapted

to these flies, but they certainly furnish no evidence that their

colors have been evolved by their selective influence. Plateau

has recently shown that the Syrphidæ poise before many incon-

spicuous objects as green flowers, closed buds, green fruits, or

even the point of the finger, in the same manner as before yel-

low flowers. Poising upon the wing before a flower must, there-

fore, be regarded merely as a habit of flight, and not as evidence

that pleasure is experienced. It is, however, probable that as

476 THE AMERICAN NATURALIST, (Vor. XXXVII.

yellow is the color of honey and pollen the more acute insects

may from long experience, as in the case of yellow honey-guides,

associate this color with the presence of a supply of food.

Another group of flowers have nauseous or indoloid odors due

to the decomposition of some nitrogenous compound. They are

often flesh-colored, blood red, dull dark purple or red, and some-

times they are marked with livid stripes or spots. By some

authors they are regarded as resembling putrifying flesh or decay-

ing carcasses. In most instances the resemblance is not very

apparent. Malodorous flowers with other colors as yellowish

green or white also occur. These flowers are visited by carrion

and dung flies belonging to such genera as Musca, Lucilia,

Sarcophaga, and Scatophaga, which are believed to find the

supposed resemblance to putrid substances attractive. While

there is no improbability in this supposition, it is chiefly, if not

entirely, the nauseous odors which attract these insects. The

lurid coloring may often be explained by peculiarities of the

plants in the production of pigments, as in the Balanophoracee,

where not only the inflorescence but the whole plant is vividly

colored. There are also a large number of flowers with strong

scented rather than repulsive odors, which are attractive to flies,

as Anethum graveolens and some Umbelliferz.

Lepidoptera. — Narious birds and mammals, as is well known,

become greatly excited when a red object is held before them.

Humming-birds and honey-suckers are attracted by firered and

scarlet colors. Kerner has pointed out that flowers of these

colors are more abundant in the Tropics and in South Africa,

where these birds are most numerous; while they are rare in

Europe where there are no humming-birds. There would seem

to be no a priori reason why butterflies, as Müller believed, may

not be strongly influenced by red coloration. Of eight Alpine

butterfly lowers (Orchis globosa, Lilium martagon and L. bulbt-

ferum, Gymnadenia odoratissima, Dianthus superbus, D. silves-

tris, D. atroruber and Daphne striata), all were red colored.

Other red butterfly flowers are species of Silene, Lychnis and

Primula, Erica carnea, and species of Asclepias and Monarda.

On the other hand three species of Globularia with light blue

flowers are adapted to butterflies, **the only instance in the

No. 439.) NORTHERN GAMOPETALOUS FLOWERS. 477

German and Swiss flora of a blue color being produced by the

selective agency of Lepidoptera.” That butterflies visit very

frequently flowers of a great variety of colors is well known to

every observer. Of 1432 visits made by 100 species of Rhopalo-

cera, 44.8% were made to greenish-yellow, yellow and white

flowers ; and 55.2% to red, violet, and blue flowers. The per-

centage of visits to wasp and bee flowers was 16.7%, and to

lepidopterous flowers 13.8% ; but the greatest number of visits

was to flowers of the type of the Composite which was 43.2%.

The percentage of visits to flowers with the honey exposed or

not deeply concealed was small. Essentially the same results

were reached by the comparison of 2086 visits of 220 Lepidop-

tera. The above figures show that butterflies are influenced

more by the form of the flower than by its color. Red and blue

flowers are usually tubular and contain more honey than yellow

and white flowers, which are more often rotate and exposed to

pillagers of every sort. The flat, capitate inflorescence of the

Composite is especially well adapted to butterflies. It is also

noteworthy that in the families and genera, which contain red-

colored butterfly flowers, blue is very rare or wholly absent.

The evidence that red floral coloration is a source of pleasure to

butterflies cannot be regarded otherwise than unsatisfactory.

Nocturnal Lepidoptera are attracted by brightness, as white or

yellow and especially a bright light, rather than by hue.

Hymenoptera. — By putting different colored papers over the

entrance holes of ground wasps it has been proven that wasps

can quickly distinguish between colors? By a series of well-

known experiments Lubbock also showed that different colors

were readily recognized by the honeybee. Müller as the result

of numerous observations came to the conclusion that the honey-

bee prefers blue, violet, and various shades of purple and red, to

white and yellow and avoids scarlet and lurid colors. During

the past summer I repeatedly observed the honeybee collecting

pollen on the flowers of the scarlet poppy ; and am led to believe

that, if these flowers contained nectar, the color would not pre-

! Müller, H. Aldpenblumen, p. 523. :

* Morely, Margaret. Wasps and their Ways ; Peckham, G. W.and E. G. Some

observations on the Special Senses of Wasps, Proc. Nat. Hist. Soc. Wisc., 1887.

78 THE AMERICAN NATURALIST. [Vor. XXXVII.

vent the frequent visits of bees. Like butterflies bees are

greatly influenced by the form of the flower. The long-tongued

bees seldom visit butterfly flowers, pollen flowers, and flowers

with the honey fully exposed unless it is very abundant. They

are most frequently collected on wasp and bee flowers, and on

associations of flowers with the nectar deeply placed. The per-

centage of visits made by the long-tongued bees to yellow and

white flowers in Müller's observations was 36.6%, and to red

and blue 63.3% ; while the percentages of the short-tongued bees

were almost exactly the reverse, or 63.8% to yellow and white,

and 36.2% to red and blue flowers. This difference seems to be

chiefly due to the form of the flowers rather than to their color,

as the short-tongued bees are largely excluded from flowers with

the honey deeply concealed. The evolution of bee flowers and

that of the long-tongued bees has gone on pari passu. The pro-

genitors of the bee flowers were presumably regular, and mostly

white or yellow; while Apis, Bombus and the allied genera are

descended from forms resembling Prosopis. As the perianth

gradually became specialized a whole host of pillaging flies and

beetles were shut out, and a more abundant supply of honey

remained for the rightful visitors. If these partially developed

bee flowers displayed red or blue colors, they would be more

easily distinguished by the lawful guests from the great mass of

blossoms with the honey unprotected. As the result of long

experience the more intelligent bees would learn to associate

these colors with an ample supply of food and freedom from

unwelcome competitors. White and yellow flowers would tend

to disappear in these genera. A preference for blue coloration

shown by bees at the present time does not, therefore, necessarily

imply that blue affords them an zsthetic pleasure; but only that

they recognize the signal of flowers adapted to their visits.

Conclusion. — The colors of flowers both in general and partic-

ular have been determined by their utility rather than by an

aesthetic color sense in insects. Insects distinguish between

different colors, but they do not receive greater pleasure from

one hue than from another. Any preference they may manifest

has arisen from the association of the colors with the presence

of food substances. Conspicuousness, or contrast of the inflores-

No. 439.] NORTHERN GAMOPETALOUS FLOWERS. 479

cence with the foliage, has been induced by insects. It is of

advantage to insects since it enables them to find nectariferous

flowers quickly, and to plants because it aids in securing cross-

fertilization. Many colors are better than one since the flowers

are rendered more conspicuous by contrasts with each other as

well as with the foliage, and insects are less liable to visit them

indiscriminately. The sequence of colors, green, yellow, white,

red, purple, and blue depends apon physiological causes. Plants

vary greatly in their capability of forming the different kinds of

pigments, and the floral colors are correlated with the variability

of this function. The primitive colors green, yellow and white

have been determined by the nature of the chloroplast and its

pigment content; while red, purple and blue have arisen as the

result of various chemical and physical conditions.

Bibliography. — In the preparation of these papers constant

use has been: made of the works of Müller, Knuth, Kerner, and

Darwin; of the Manuals of Gray and Chapman; and of the

Illustrated Flora of Britton and Brown. Bibliographies of the

literature dealing with the mutual relations of flowers and insects

and with the colors of flowers will be found in Müller's Fertilisa-

tion of Flowers, translated by D'Arcy W. Thompson, and in

Knuth's Handbuch der Blütenbiologie (2871 titles). References

to 772 books and papers on plant pigments are given in Kohl's

Carotin und seine physiologische Bedeutung. In his paper

* Beobachtungen und Versuche über das Auftreten von rothem

Zellsaft bei Pflanzen" Overton briefly reviews the literature

relating to anthocyan. For the literature on the colors of ani-

mals Newbigin's Color in Nature may be consulted.

WALDOBORO, MAINE.

RIB VARIATION IN CARDIUM.

FRANK COLLINS BAKER.

CONTENTS.

I. Introduction. II. Material. III. Method of obtaining

quantitative data. IV. Discussion of data. Cardium robustum.

Cardium isocardia. Cardium muricatum. V. Comparison of

the three species.

I. INTRODUCTION.

Ar the present time biologists and zodlogists are paying

marked attention to the study of the laws of variation, such

studies having a strong bearing upon the origin of species. The

results of such studies will be of little value, however, unless

great care is taken in recording the exact locality from which

the material came. And likewise the results will be #/ unless

the material studied is of such a quantity as to form a good basis

for a quantitative examination. It frequently happens that a

careful study of the variation of a certain species is rendered

valueless because the working material has been too limited in

quantity.

That the time expended in such study is well spent is clearly

shown by the following statement made by the late Prof. Edward

D. Cope.! |

« So soon as sufficient material becomes available, the zoólo-

gist can make that kind of research into the permanency "and

variability of the characters of species which characterizes the

exact stage of the science. It is on such study that all useful

conclusions as to the origin of species depends. It is not the

orderly relation of species and genera to each other that demon-

strates the truth of the hypothesis of the derivation of the

1 As quoted by C. C. Adams, p. 208.

481

482 THE AMERICAN NATURALIST. (Vor. XXXVII

species, but the knowledge of their variations. Moreover, the

beginning of all investigation into the causes of those variations

is the knowledge of the direction which they take, whether they

are promiscuous or whether they bear some definite relation to

each other or to the environment."

The following paper is a contribution tothe study of variation

in the sculpture of the pelecypod Mollusca, as shown in the

genus Cardium.

II. MATERIAL.

The material upon which this paper is based consists of

three species of Cardium; C. robustum, C. tsocardia, and

C. muricatum. They were collected by Dr. J. W. Velie and

W. W. Calkins on the west coast of Florida, near Tampa.

Several hundred specimens of each species have been examined,

thus affording enough material for a wide range of variation.

To this material is added some data gathered by Dr. W. H.

Dall and ae c in his ** Contributions to the Tertiary Fauna

of Florida."

III. METHOD or OBTAINING QUANTITATIVE Dara.

The ribs on each valve were carefully counted and in order to

remove any possibility of error they were recounted several

times at intervals of two or three days. The size (length) of

each shell was determined by a pair of calipers spread from the

umbones to the ventral margin. These measurements are all in

millimeters.

In the curves plotted, the groups or classes having the same

number of ribs are indicated on the horizontal line, and the

number of specimens in these groups, the frequencies, are noted

on the vertical line. i

In the tables the number of ribs is indicated as a numerator,

and the individuals having the same number of ribs is noted as

a denominator.

No. 439.] RIB VARIATION IN CARDIUM. 483

IV. Discussion or Data.

Cardium robustum. ( — magnum.)

Figures 1 and 2

Table A.

_30 eis | a XM ee

! 33 7 128 81 39 3 2

Number of shells examined 364. The range of variation is

from 30 to 37.

The mode, or class with the largest number of individuals, is

33, with a frequency of 128. This curve

is remarkable for its regularity and for its

strong monomodal tendency. 2 soe See

In counting the ribs of this, as well as of #% 7 THE

other species, it was noted that the varia- 23 PH

tion was confined almost wholly to that &@

: i a

part of the shell anterior or in front of the 65} i

umbonal ridge, the latter being strongly A

indicated by a large, heavy rib extending — 42

from the umbones to the ventral margin, ime

and separating the flat posterior slope from jm

303132 3434 35 307;

the rounded anterior and lateral slopes.

There is one more rib on one valve than

on the other in this region, the numbers

being 7-8, or 8-9. The ribs of this area

were carefully counted and the following result obtained, the

figures being for the maximum number of ribs: m =a,

This gives the normal or mode at 7—8, with a frequency of iss.

If this were plotted on a diagram it would give a very sharp,

narrow curve, which always stands forstability. It is also note-

worthy that the curve for all the ribs and that for the ribs

posterior to the umbonal ridge are similar in form. The size of

the shell apparently does not change the result of these calcula-

tions. 41 specimens measuring 90 millimeters gave the result

shown in figure 2 and in the following table:

FiG. 1. Cardium robustum

Variation curve of “i

specimens.

484

THE AMERICAN NATURALIST. [Vor. XXXVII.

Table B.

Sues g 5 4380

I 14 15 8 2 I

As in table A the mode is at 33 (frequency 15) with an

7, ane

ENR ae a

ati itt

LIE I a

eters

species. It

additional minor mode at 32 (frequency 14).

This indicates a larger amount of variation for the

90 millimeter individuals than for the whole num-

ber of specimens, and illustrates the value of exam-

ining a large amount of material in order that

a false impression may not be given by the curve.

Dall! has examined a number of specimens of

this species and the results are interesting. He

found the range of variation to be from 30 to 35,

forty-five specimens being counted. Dall remarks

that there is a slight tendency to fewer ribs in the

southern than in the northern individuals of this

would be interesting to have a large number of

specimens from different localities examined and plotted, to

ascertain the exact amount of this variation.

Number of shells examined 222.

Cardium isocardia.

Figures 3 and 4.

Table C.

Eco ns D Uu 2» 5 s

22 73 79 13 3 I

The range of variation is

from 27 to 36. The mode is at 30 with a frequency of 33 and

a minor mode at 31 with a frequency of 70. The noteworthy

feature of the curve of zsocardia is the polygonal instead of the

triangular form and the marked symmetry of the two sides.

comparison of the curves obtained from an examination of

1 Dal

W. H. Contributions to The Tertiary Fauna of Florida.

1099.

Trans.

ice Znst. Sci., Vol. IIL, part V, p.

No. 439.] RIB VARIATION IN CARDIUM. 485

different sized shells is interesting, and is shown in figure 4 and

the following table :

Table D.

Size in millimeters. 28 29 320. 3f 32 33 34 35

30. 4 12 6 4 3 1 1

40. 6 3 16 16 5 3

50. 5 14 4 4

a 1

400 E li

oH = am 17

yi 16

aol 25 E

75 19 4

70! y. FTT

Las m SRERERE i —

pesans gegane SEHE

ARERERdE I

40) a oo oe oe m

J$F4—H1H-A HA 6 []

30} —1 + | `

25 = = $ p

25 m EN 2

70] LIT] 7

5 BERE 293031323] HIS

Fic. 4. Cardiu:

Fic. 3. M tso- tsocardia Varia-

cardia. riation curve tion curve aa size.

of 222 specimen:

The number of shells examined of the 30 mill. individuals was

31. The range of variation is from 29 to 35, with a mode at 30

and a frequency of 12. Of the 40 mill. individuals 49 species

were examined. The range is from 28 to 33, with a double

mode at 30 and 31 and a double frequency of 16. The 50 mill.

individuals included 27 specimens. The range is from 30 to 33,

with a strong mode at 14. The range of variation in the 30 and

50 mill. specimens is very uniform and the curves are almost

identical. In the 40 mill. individuals there is more variation as

shown by the broadness of the curve and its irregularity at the

lower part. This wider range of variation may account in a

measure for the polygonal form of the curve for the total

number of specimens.

The ribs of ¢socardia are much oodd on the anterior and

posterior slopes of the shell, where, also, the greatest spinosity

exists.

486 THE AMERICAN NATURALIST. [Vor. XXXVII.

Cardium muricatum.

Figure 5.

Table E.

4e. ue M oo d a D . 7 uo»

2 6 31 27 25 10 5 2 2

seer] Number of shells examined 110. The

E: range of variation is from 29 to 39, with a

36 .

34 break between 36 and 39. The mode isat

30l ; .

E L 31 with a frequency of 31, a strong minor

ZH mode at 32, with a frequency of 27 and a

yy . .

ott weaker minor mode at 33, with a frequency

/2- of 25. The width of this curve shows a con-

D siderable amount of variation. As in Zso-

Ooasuscsuns)- cardia, the ribs are crowded at the anterior

Fic. s. Cardium murica. 200 posterior ends and the side ribs are fre-

me Core ne quently quite smooth.

specimens. : P

Dall (oc. cit., p. 1090) has examined 55

individuals of this species, gathered from all parts of its habitat,

and the result is as follows :

Figure 6.

Table F.

e 7 m SM 5o SB LV. s » P 9

9 3 58 Ss 7 m 5 3 tu T

A comparison of the two figures is interest-

ing. The mode of curve No. 6 is at 31 with G44

a frequency of 9 and two minor modes at 33 1 AEH

(frequency 8) and 35 (frequency 8). The f "n H

breadth of this curve and its multimodal form ‘HHHH HH

shows a greater amount of variation in its

: ; Fic. 6. Cardium muri-

entire range than do the specimens from near ,4,, Curve of ss

Tampa, in which the number of the ribs is eh e pdt

i entire ra . .

more stable. The major mode, however,

remains the same, showing that the normal number of ribs for

this species is 31.

No. 439] RIB VARIATION IN CARDIUM. 487

Dall makes the following remarks concerning the rib vari-

ation of muricatum. ‘The only generalization that seemed

authorized is that the ribs are less numerous in specimens from

near the northern border of the range of the species, and also in

the fossils; the specimens with 37 to 41 ribs are nearly all

from the southern half of the area inhabited. There was no

diminution of ribs towards the southern extreme of the range

and no regularity in the variations of the murication which

could be correlated with difference of habitat."

Here again it would be interesting to know the results

obtained by a quantitative study of a large amount of material

from different localities.

V. COMPARISON OF THE THREE SPECIES.

Figure 7.

A comparison of the three species shows that magnum is the

least variable, while muricatum is the

most variable, as is shown by the width — 4L

of the curve. C. zsecardia seems to

stand midway between these two species.

In the study of quantitative variation

in the Mollusca the fact presents itself

that in each species there is a mode or

constant which remains unvaried and

from which certain individuals vary spo-

PES

r———À

LITI

gei

BH HHH

SORES eeu Sont

radically. These would seem to be

brought about by accidental variation ELE TTE S

rather than by natural selection. MUN. conde d da Mee

In figure 7 it will be seen that each species. 1, robustum. 2, iso-

: . : cardia. 3, muricatum. (Classes

species has a particular and different doubled.)

constant. C. magnum (1) has 33 as a

constant and shows a minimum of variation: zsocard?a (2) has

30 as a constant and muricatum (3) has 31 as a constant, but

shows a large amount of variation.

It is evident from the above study that the number of ribs is

not a safe character upon which to found a species. A small

488 THE AMERICAN NATURALIST. [Vor. XXXVII.

number of specimens, 20 or 30, might show an apparent break

in the numerical variation and seem to warrant the separation of

some individuals as species, but a larger number of specimens

shows that the number of ribs cannot be used in the separation

of species without other more important characters.

It would be interesting to know the results obtained by an

examination of a large number of specimens from different

localities, to ascertain the stability of the data recorded in this

paper.

My thanks are due to Professor C. B. Davenport of the Uni-

versity of Chicago for valuable suggestions in carrying on this

study.

CHICAGO, ILL.

THE PERFORATION OF A VEIN BY AN ARTERY

IN THE CAT (FELIS DOMESTICA).

ARTHUR W. WEYSSE.

In a cat which was being dissected in my laboratory a few

months ago, I noticed an artery passing through an opening in a

vein, and as further dissection showed the details of this abnor-

mality to be somewhat different from any similar condition which

I have found recorded it seemed well to publish the following

figure and description.

The cat was an adult female; the circulatory system had been

H "n 9H

H H 1

H E WE M

i I Ij

H * H

H H Hea

:d ae H

s , H

ioc: tat

CEES

e E RAE EE

injected with an ordinary starch injecting mass, the veins with

blue and the arteries with red. For the sake of added clearness

in the diagram the horns of the uterus were dissected free from

the body-wall and together with the urinary bladder were laid

back over the ventral side of the pubes; thus the umbilical

arteries and the uterine arteries and veins are directed backward

in the diagram instead of forward. It will be noticed that there

is a longitudinal slit about a centimetre in length in the right

489

490 THE AMERICAN NATURALIST. [Vor. XXXVII.

common iliac vein nearly opposite the point at which the

superior gluteal branch of the internal iliac artery in given off,.

and through this slit the superior gluteal artery passes. This

artery arises much farther cephalad than the left superior

gluteal ; the latter is in the normal position. The corresponding

veins, on the other hand, arise symmetrically from the dorsal

surface of the two common iliac veins. The opening does not

divide the vein into tubes of equal diameter, that on the median

side being much broader than the tube on the outside, but the

lumen is uninterrupted on both sides of the slit.

Abnormalities in the vessels in this region are very frequent

in the cat, and this same specimen shows one or two others.

Thus while the right umbilical artery leaves the internal iliac at

the usual point the left comes off much farther caudad than is

customary and is much smaller than the right. The median

sacral vein arises from the right common iliac instead of the left,

an abnormality which has frequently been observed. In other

respects, however, the vessels in this region of the cat in question

take the normal course.

Not a few cases are on record of veins which have been per-

forated by arteries. McClure (’00) mentions two instances in

the cat where lumbar arteries are found passing through a

foramen in the inferior vena cava, and the same writer ('00)

records four cases in the opossum where the spermatic arteries

pass through foramina in the same vessel, the inferior vena cava ;

Hochstetter ('96) describes an Echidna embryo in which the

inferior mesenteric artery passes through a foramen in the

inferior vena cava. All of these abnormalities may be explained,

however, if the inferior vena cava develops by a fusion of the

posterior portions of the cardinal veins as appears probable;

McClure (90) mentions a number of cases in which the

posterior cardinals persist in the adult cat, and if the fusion were

to take place after the formation of the lumbar arteries, they

would be left perforating the vein.

More nearly approaching the condition which I have found,

are the four cases recorded for the cat by McClure ('00) in

which the internal iliac artery passes through a foramen in the

common iliac vein very near the point at which it divides into

No. 439.] PERFORATION OF A VEIN. 49I

the external and internal iliacs. This condition is not so readily

explicable unless it should be shown that the internal iliac vein

develops as a branch of the external, which again seems probable.

If the internal iliac artery were first developed, the vein might

well develop on both sides of it and thus become perforated.

A case which resembles mine still more closely is that figured

by Treadwell (96), —a perforation of the right common iliac

vein by the right internal iliac artery; although this specimen `

was incomplete, the foramen appears to be much more cephalad

than the point at which the internal iliac vein is given off, but

here again it is the entire internal iliac artery which perforates

the vein, instead of merely the superior gluteal branch as in my

case.

Such abnormalities can only be explained by embryology, and

very little appears to have been published on the development

of the principal posterior branches of the dorsal aorta and the

inferior vena cava. As to the way in which the perforation in

question may have been brought about, the most reasonable

explanation that has suggested itself to me is that the internal

iliac artery grows out as a branch of the dorsal aorta before the

common iliac vein develops from the inferior vena cava. If this

should prove to be so, we can readily see that the anlage of the

vein on coming in contact with the artery, or in this case its

superior gluteal branch, might occasionally grow entirely around

it instead of passing to one side. The fact that in the case

figured the right superior gluteal artery arises from the internal

iliac much farther forward than is usual and so comes to lie

directly in the course of the right common iliac vein would seem

to bear out this suggestion.

Blood vessels in the pelvic region of a cat, ventral aspect; the veins are in outline, the

arteries shaded. The nomenclature adopted follows that used by Reigard and Jennings

in their Anatomy of the Cat.

A.a. Aorta abdominalis.— A. f. A. femoralis. — A. g. s. A. glutea superior.— A. 4. A. hypo-

gastrica (iliaca interna) — 4. i.e. A. iliaca externa.— A. ilio. A. iliolumbalis. S m.i. A

mesenterica i .£. A.spermatica interna. — 4. u. A. umbilicalis.— V. c. z. V.

cava inferior.— V., f. v femoralis. Fx v. prs superior — Vite. V. iliaca

munis.— V. ito. V. iliolumbalis.— V. s. V. tica interna.— r. A. prona

femoris.— 2. A. and V. glutea inferior.— 3 A. and v. a media.— 4. A.

V. uterina.— 5. 4. and v. sacralis media.

492 THE AMERICAN NATURALIST. (VoL. XXXVII.

BIBILOGRAPHY.

HOCHSTETTER, F.

'96. Beitráge zur Anatomie und Entwickelungsgeschichte des Blutge-

fássystems der Monotremen, in emon's Zoolgische Forschungs-

reisen in Australien und dem SUN ischen Archipel. Bd. 2, Lief. 3;

Monotremen und Marsupialier ; pp. 188-243, Taf. xvi-xix, 3 fig.

McCLunE, C. F. W

:00. On the frequency of Abnormalities in connection with the

Postcaval Vein and its Tributaries in the Domestic Cat (Felis

domestica) Amer. Nat., Vol. 34, pp. 185—198, 9 figs.

McCLung, C. F. W

:00a, The Variations of the Venous System in Didelphys virginiana.

Anat. Anzeiger, Bd. xviii, pp. 441—460, 21 figs.

TREADWELL, A. L.

'96, An abnormal iliac Vein in a Cat. Anat. Anzeiger, Bd. xi, pp.

717-718, 1 fig.

MASSACHUSETTS INSTITUTE OF TECHNOLOGY,

Boston, Mass.

A PECULIAR MODIFICATION AMONGST PERMIAN

DIPNOANS.

C. R. EASTMAN.

Tne genus Sagenodus is represented in this country by less

than a dozen species, of which only three have been described

from the Permian of Texas. These are S. dialophus, S. periprion

and S. porrectus, all founded on detached dental plates of small

size. The presence of a fourth species, different from any

hitherto described, and displaying quite unusual modifications

amongst ceratodonts, is indicated by several well preserved

mandibular and palatine dental plates which have recently been

brought to light by Dr. E. C. Case, of the State Normal School

at Milwaukee, and kindly placed by him in the hands of the

writer for description.

The present species occupies a unique position amongst fossil

dipnoans in having a dentition adapted for cutting instead of

crushing, thus paralleling the conditions found in certain

Palzeozoic sharks and in recent Gymnodonta. This divergence

is the more striking in view of the singularly uniform type of

. dental system pervading lung-fishes throughout their entire

geological history. Whether so extreme a variation is to be

correlated with the change from marine to brackish-water condi-

tions that took place during the Permian, with its very pronounced

effects upon the environment and food-supply, may perhaps be

plausibly conjectured.

In the new form, which may be named Sagenodus pertenuis

in allusion to its chief peculiarity, the coronal grinding surface

has become reduced to practically »/ in the lower jaw, owing to

compression of the inner margin into a sharp cutting edge, and

disappearance of all except one of the outer radiating ridges.

The upper dental plates differ from the lower in that two, instead

of one, abbreviated coronal ridges are given off from the sharp

angulation of the inner margin. The latter forms a continuous

crest extending nearly to the symphysis anteriorly, and cor-

493

494 THE AMERICAN NATURALIST. [Vor. XXXVII.

responds to both the foremost and hindmost of the coronal ridges

in Ceratodus, plus the intermediate space. Hence it is proper

to speak of an anterior and posterior, and one or two intermediate

coronal crests as the case may be, according as we have to deal

with mandibular or palatine dental plates. All of these coronal

crests are serrated, the anterior one — which is at the same time

the longest — coarsely, and the others finely, with sometimes as

many as six or seven serrations each. The cutting edge in worn

specimens furthermore displays a minutely crenulated appearance,

owing to exposure of the dentine tubules, a condition very

frequently observed in sharks’ teeth. Both sides of the thin

cutting edge exhibit a shin-

ing enameled surface,

which passes gradually

into a narrow base of vaso-

dentine ; and in the case of

the mandibular dental

plates at least, the pair was

suturally united at the sym-

à physis, much in the same

fashion as in Ptyctodus and

Rhynchodus (Fig. 1 a, b).

The six specimens which

the writer has examined

are of comparatively small

size, none exceeding a

total length of 2 cm. anda

height of 0.8 cm. In some

. examples the angulation of

etd Md BR IUE: margiu is copsid

rior (a), inner (6) and outer (c) aspects x 24. The erable, amounting almost

tuc. duds to a right angle, and the

js short intermediate ridges

have a tendency to become slightly curved backward. None of

the dental plates exhibit marks of contact with those of the

opposite jaw, but it is nitural to suppose from the manner in

which the lower pair were united, that essentially the same sort of

contrivance was developed here as we have become familiar with

in Peripristis from the upper Carboniferous, a modification which

No. 439.] PERMIAN DIPNOANS. 495

is truly remarkable. When we recall also the aberrant series of

Edestus-like sharks that flourished contemporaneously, we are

struck with the fact that in two of the most conservative and

persistent groups of fishes, namely the ceratodonts and cestra-

cionts — both of which have had a continuous existence ever

since the Devonian, — the extreme of variation was attained

toward the close of the Paleozoic. ;

Another interesting feature to be brought forward in connec-

tion with the present form is its apparently

wide distribution ; and bearing in mind the P ar

world-wide scattering of the Edestus series A r4

that took place during the late Palzozoic, Se a

we note that the stimulus which quickened

variation and distribution was responded to

simultaneously by the two groups of fishes

exceeding all others in longevity, after

which they relapsed into sluggishness.

The specimens obtained by Dr. Case, and

one or two others belonging to the Münich p, , sagenodus pertenuis

Palzeontological Museum, the latter having nées ettari ipe

been acquired through Mr. Charles Stern-

berg, are all from the Cimarron series (upper Permian) of

Wichita County, Texas. But itis further to be recorded upon

the authority of Dr. Broili of Munich, who recently submitted

the specimens under his charge for identification, that precisely

the same form of tooth occurs in the Permian of Russia. Thus

we have valuable additional evidence from the side of vertebrate

paleontology regarding the homotaxial relationships of the Texas

* Red Beds." The distinguishing features of the above described

species may be briefly summarized as follows : — Dental plates

relatively small, thin, the inner margin strongly angulated and

sharpened into a continuous cutting edge, with a few coarse

serrations in advance of, and finer ones behind the angulation.

Mandibular dental plate with but one, and palatine dental plate

with two short and narrow coronal ridges extending outwardly

from the angulation of the inner margin, their crests finely ser-

rated. Plates entirely without coronal grinding surface. .

CAMBRIDGE, Mass.

NOTES AND LITERATURE.

ZOOLOGY.

Bailey’s Birds of the Western United States.!— It is a great

satisfaction to see a difficult task so well done as in Mrs. Bailey’s

Birds of the Western United States. Owing to the diversified

nature of the area covered by the book, including the Plains, the

Rocky Mts., the Great Basin, and the Pacific Slope, a very large

number of species and subspecies had to be treated. There are

careful descriptions of the different plumages of each species, an

account of its distribution, and of its nest and food. These are

followed by short but graphic biographies. In the case of many of

the larger birds, the accounts of their habits have been supplied by

Mr. Bailey, whose work in the West on the Biological Survey has

enabled him to give great assistance. The introduction is evidence

of the care with which the book has been planned. It contains

directions for collecting birds, accounts of the life-zones and migra-

tion in the West, local lists and much other helpful matter. Atten-

tion is called to the vertical migration due to the height of the

mountains. Certain hummingbirds, for instance, rear a secon

brood at a higher altitude than the first. There are abundant keys

and illustrations. Thirty-three original full page illustrations and

many cuts are by Fuertes. Most of these are well up to this artist’s

high standard. Occasionally as in the case of the Mearns’ Quail, p.

122, and the Pileolated Warbler, p. 428, the effect is marred by

the grotesqueness of some unusual attitude. A great many of the

small cuts and diagrams with which the book abounds are really

illustrative; it is a pity that so many pages are disfigured by the

useless photographs from skins. The students of birds in the West

are to be congratulated on now having a handbook which will prove

as indispensable there as Chapman’s is in the East.

R H.

! Bailey, Florence Merriam. Handbook of Birds of the Western United States,

Boston, Houghton, Mifflin & Co., 1902. 8vo. xc-512 pp., 33 pls., 601 text figs. «

497

498 THE AMERICAN NATURALIST. [Vor. XXXVII.

** The Water Fowl Family''.! — The fourth volume in the Ameri-

can Sportsman’s Library is an attractive book, which will doubtless

prove of much value to those sportsmen who desire to extend their

knowledge of the life habits of the game birds. It deals with the

North American ducks, geese, swans, rails, and shore-birds. There

are general descriptions of the families in each group, and accounts of

each species, including their distribution, nesting and feeding habits.

These have been compiled from standard authorities. There are also

accounts from original observation of the behavior of each well-

known species as it concerns the sportsman, and descriptions of the

various methods employed in hunting it. An encouraging interest

in bird protection is shown throughout the book; the “game hog”

is condemned, and a close season in spring strongly advocated. It

is a pity that a little of the cheap sporting-story element was included.

The chapter on goose-shooting by the man who “hoped to preside

at the obsequies of a goose” might well have been omitted. There

are a number of excellent full page illustrations, three by Bull, and

the rest by Fuertes.

KH.

Ancestral Canide.? — Mr. J. B. Hatcher has published a paper of

unusual interest on the Oligocene Canidz lately discovered in

Nebraska, and now preserved in the Carnegie Museum.? A full

account is given of an almost complete skeleton of Daphenus felinus,

Scott, and two new genera, Proamphicyon and Protemnocyon, are

described. It is held that Daphænus has no known descendant ;

that Proamphicyon is ancestral to Amphicyon; and that Protem-

nocyon is ancestral to Temnocyon. This last animal is of particular

interest, as it seems to be undoubtedly ancestral to Canis ; that is,

to the common dog. The discovery of Protemnocyon carries the

known ancestry of the dog one stage further back; and, in fact, it

was a bed dog-like creature. The sagittal crest is quite as in the

dogs ; two temporal crests of the foxes give their skulls a

ty different appearance. The postorbital processes of the

frontals are essentially as in the dogs, though short. The third

! Sanford, L. C., Bishop, L. B., and Van SIN T.S. The Water Fowl Family.

New York, Macmillan, 1903. 8vo, eras pp. 20 pls

* See also an important article by D: Pai: b. Matthew (Science, June, 5, 1903,

p- 912) published since this notice was w

*Oligocene Canide. Mem. Carnegie sony Vol. 1, pp, 65-108. Pls. XIV-

XX.

No. 439] NOTES AND LITERATURE. 499

molar is wanting, or perhaps occasionally present of very small size,

in the upper jaw; present but very small in the lower. In the dog

the third molar is usually absent in the upper, present in the lower

jaw, but quite numerous cases have been found in which it was

present in the upper jaw. Wortman and Matthew have held that

the modern representative of Temnocyon is Cyon of India, which is

little more than a subgenus of Canis; but as Mr. Hatcher remarks,

this is problematical. Cyon is principally distinguished by the

absence of the last lower molar, which would indicate that it is more

specialized than Canis proper; the same character occurs as an

aberration not rarely in the domestic dog and several wild species of

Canis. It is to be remarked that the genus (Icticyon) which shows

the greatest reduction of the molars (to one above and two below)

is a native of South America. The top of the skull in Protemnocyon

is flat in lateral view; in the coyote it is more elevated, while in the

domestic dog it is strongly convex; these changes no doubt accom-

panying a progressive development of the brain, though partly the

result also of enlarged frontal air-sinuses. In Temnocyon the

absence of the third molar in the lower jaw suggests Cyon rather

than Canis proper, but it does not seem impossible that this molar

might be lost and recovered again (contrary to the doctrine of some),

considering the comparative frequency of an extra (fourth) molar in

the lower jaw of the common dog. However, while Temnocyon

stands nearer to Canis than does Protemnocyon, both in time and by

the auditory bull, it may still be a little out of the direct line.

A. C,

Parker on the Hearing of Fishes.!— Professor G. H. Parker

gives an account of his studies of the sense of hearing in fishes.

Taking the common Killifish, Fundulus heteroclitus as the subject

of experiment, he shows that this fish does actually hear sounds

which may be made by means of tuning forks, and that it becomes

deaf on cutting the auditory nerve. It is possible that fishes of other

species may be actually insensible to sounds as experiments of others

have seemed to show, and in all fishes the ear may be in part an

organ of equilibration.

It is of course not likely that any fish has the ~ to make fine

discriminations in sounds.

! Parker, G. H. Hearing and Allied Senses in Fishes. Bul. U. S. Fish Com.

Jer 1902, pp. 45-64, pl. 9.

500 THE AMERICAN NATURALIST. [Vor. XXXVII.

The lateral line is closely associated with the air, and may also

assist at hearing. The ear is an outgrowth from the tubes of the

lateral line. As Professor Parker aptly observes, in the skin, the

lateral line and the ear, “we are dealing with what may be called

three generations of sense organs: the skin representing the first

generation and giving rise to the lateral line organs, the second,

which in turn produce the ears."

DB. Si pg

Parker on the Optic Nerves of Flounders.'— In the bony fishes,

the optic nerves pass to the optic lobes of the brain, the one passing

to the lobes of the opposite side simply lying over the other, without

intermingling of fibres, such as takes place in the higher vertebrates

and in the more primitive fishes.

According to Parker’s observations, in ordinary bony fishes, the

right nerve may be indifferently above or below the other. In 1000

specimens of ten common species, 486 have the left nerve upper-

most and 514 the right nerve. In most individual species, the num-

bers are practically equal. Thus, in the haddock, 48 have the left

nerve uppermost and 52 the right nerve.

In the unsymmetrical Teleosts or flounders, and soles, this con-

dition no longer obtains. In those species of flounder with the eyes

on the right side, 236 individuals representing sixteen species had

the left nerve uppermost in all cases. :

Of flounders with the eyes on the left side, 131 individuals repre-

senting nine species all have the right nerve uppermost.

There are a few species of flounders in which reversed examples

are so common that the species may be described as having the

eyes on the right or left side indifferently. In all these species,

however, whether dextral or sinistral, the relation of the nerves con-

forms to the type, and is not influenced by the individual deviation.

Thus the stony flounder (Platichthys) belongs to the dextral group.

Fifty normal specimens, the eyes on the right, have the left nerve

dorsal, while the left nerve is also uppermost in 50 reversed exam-

ples with eyes on the left. In rs examples of the California bastard

halibut (Paralichthys californicus) normally sinistral, the right eye

is always uppermost. It is uppermost in 11 reversed examples.

1 Parker, G. H. The Optic Chiasma in Teleosts and its Bearing on the

— of the Heterosomata (Flat Fishes) Bull, Mus. Comp. Zodl., Vol. XL.

©. 5, pp. 221-242, I pl.

No. 439.] NOTES AND LITERATURE. 501

Among the soles this uniformity or monomorphism -no longer

obtains. In forty-nine individuals of four species of dextral soles, the

left nerve is uppermost in 24, the right nerve in 25. Among sinistral

soles, or tongue fishes, in 18 individuals of two species, the left nerve

is uppermost in 13, the right nerve in s.

Professor Parker concludes from this evidence that soles are not

degenerate flounders, but rather descended from primitive flounders

which still retain the demorphic condition as to the position of the

optic nerves, a condition still retained by all bony fishes except the

flounders.

The lack of symmetry among the flounders lies therefore deeper

than the matter of the migration of the eye. The asymmetry of the

mouth is an independent trait, but like the migration of the eye, is

an adaptation to swimming on the side; Each of the various traits of

asymmetry may appear independently of the others.

The development of the monomorphic arrangement in flounders,

Professor Parker thinks, can be accounted for by the principle of nat-

ural selection. In a side-swimming fish, the fixity of this trait

has a mechanical advantage. The unmetamorphosed young of the

flounder are not strictly symmetrical, for they possess the monomor-

phic position of the optic nerve. The reversed examples of various

species of flounders (these, by the way, chiefly confined to the Cali-

fornia fauna) afford “striking examples of discontinuous variation.”

Professor Parker inclines to the opinion that the ancestral floun-

ders were allied to the john dories. This is as plausible a guess as

any. They certainly have no affinity with the cod-fishes.

Dp. S J.

Notes on Recent Fish Literature.— Mr. C. T. Regan (Proc. Zod.

Soc. London) takes up the osteology of the plectognathous fishes

and the classification derived from it. The chief character of the

group as distinguishing it from their ancestors, the Acanthuridæ is

the absence of ribs. He divides the plectognaths into two divisions,

the Sclerodermi and the gymnodontes. To the former group the

Ostracodermi are referred. The supposed families of Chonerhinidæ

and Tropedechebyidæ are regarded as not distinct from Tetraodon-

tidæ and doubt is thrown on the accuracy of the figures of Hollard

which have served as the basis for certain generic distinctions.

The Mexican trigger-fish Balistes naufragium is said to be a spe-

cies of Xanthichthys, a genus rejected by Mr. Regan.

502 THE AMERICAN NATURALIST. (Vor. XXXVII.

In the Annals and Magazine of Natural History (XIL, 459—466)

Mr. Regan discusses the osteology and classification of the anacan-

thine or cod-like fishes. He regards the absence of foramen in the

hypercoracoid, which separates the cod-fishes from the true jugular

fishes (blennies, etc.) as a matter of minor importance, because the

same trait is found in several trachinoid. fishes, which are true jugu-

lar fishes. In the cod-like fishes or Anacanthini, the ventral fins,

sometimes many-rayed, are below or in front of the ventrals, * while

the pelvic bones are posterior to the clavicular arch to which they are

loosely attached by a ligamentous connection." In the true Jugu-

lares the ventrals, with 6 rays or fewer, are jugular, “the pelvic bones

being distinctly and firmly attached to the clavicular symphysis.”

The true Jugulares are, of course, modified Acanthopteri. In Mr.

Regan's opinion *the Gadoids originated from some less specialized

stock," their peculiar features being largely primitive. He suggests

their possible derivation *from some Haplomous stock from which

the Berycidz have also descended, and of which the Stephanoberycidx

are the nearest living representatives."

In Mr. Regan's view the Macrouride are more primitive than the

cod-fishes. In this family, Melanomus and Lyconus should be

placed. Bregmaceros, wrongly placed near the Brotulidz, has the

general structure of the cod-fishes. — Murzenolepis is the type of a

distinct family. Mr. Regan describes a new genus, Gadomus, based

on Bathygodus longifilis. In this genus there is a slit behind the last

gill, and the hypercoracoid unlike that of all other Anacanthini is

perforate. Melanobranchus, another new genus, has the slit behind

the last gill, but the hypercoracoid is as in other Macrouri.

In a recent paper (Ann. Mag. Nat. Hist., XI, 372-374) Mr. Regan

discusses again the skeleton of Zervazes imperialis, deciding finally

that it is a highly aberrant scombroid fish.

Dr. Peter J. Schmidt in Proceedings of the Museum of St. Peters-

burg discusses in Russian, and later in German the fauna of the Seas

of Japan and Okhotsk. In both these seas the species of fishes are

distinctly sub-arctic ; although some shore-fishes enter from the ordi-

nary Japanese fauna, these waters are very rich in agonoid and

Cottoid fishes, far more so than the corresponding latitudes in the

Atlantic. A number of new species are indicated by name, soon to

be described.

In the series of monographic reviews of the fishes of Japan, Méssrs

No 439.] NOTES AND LITERATURE. 503

Jordan and Fowler (Proc. U. S. Nat. Mus., XXV, 939-956) treat of

the fishes known as dragonets, constituting the family of Cad/iony-

mide. Of these fishes 12 species are described, and the new species

and some of the others are well figured by Captain C. B. Hudson.

One species, Draconetta xenica constitutes a new family and a new

genus, Calliurichthys is proposed for the dragonets with spear-like

preopercular spine.

Dis, RA

Hacker’s Autonomy of the Germ Nuclei.’ — This work is in the

main an extension of Hackers earlier papers, (1892, 1896) on the

autonomy of the male and female pronucleiand of their derivatives in

the development of limnetic Copepods. To this central theme he

has added two introductory chapters on the ecological (biologische)

relations and on the general developmental phenomena of copepods,

a chapter on the maturation phenomena of Cyclops and another in

which he seeks to extend the idea of the autonomy of the germ nuclei

to many classes of plants and animals. These nuclear halves he

designates * Gonomeres" while the vesicles formed from individual

chromosomes (chromosomal vesicles) he calls *Idiomeres." These

names are definite, convenient and really necessary to avoid descrip-

tive phrases and it is desirable that they should come into general

use, =

The author thinks it is possible to follow this autonomy of the

gonomeres from the first to the third generation, but his stages are

by no means complete; in fact they consist only of a few cleavage

and gastrulation stages and of the developing gonad. His methods

of distinguishing the cells of the germ track (“ Ae/mbahnszellen’) are

the following: (1) The autonomy of gonomeres is here pre-

served longest. (2) Nuclear divisions are here heterotypic, (3) The

rhythm of division is here slower than elsewhere, (4) Ectosomes

(dark staining granules) are eliminated from the nuclei of the germ

track cells, thus suggesting the chromatic diminution of Ascaris.

The autonomy of the gonomeres is determined chiefly by the pres-

ence of two nucleoli within a nucleus, though in cases where there is

a long resting period this number may be reduced to one. Evidently

the significance of this is that there are as many nucleoli as there

are idiomeres or chromosomal vesicles and when during a long rest-

Ueber das Schicksal der elterlichen und SS

1 Häcker, Valentin.

wate der Vererbungslehre. Jena

Kernanteile, Morphologische Beiträge sum Ausbau

Fischer, 1902. 8vo, pp. 104, 4 plates, 16 text figures.

504 THE AMERICAN NATURALIST. (VoL. XXXVII.

ing period the two gonomeres fuse a fusion of their nucleoli also

occurs.

With regard to the fate of the maternal and paternal halves during

maturation the author says that there are three possibilities ; Either

(1) a complete separation of the halves (Mendel's principle), (2) a

symmetrical mixing of nuclear constituents, or (3) an unsymmetrical

mixing. He concludes that the first maturation is an equation divi-

sion and that the reduction occurs in the second maturation in such

a manner “that the ripe egg cell contains one half of the grand

paternal and one half of the grand maternal chromosomes” thus ful-

filling the second possibility named above.

The conclusion which the author reaches that the reduction is

brought about by a fusion of maternal and paternal chromosomes at

the time of the 2d maturation division is not sufficiently well sup-

ported, especially in view of the fact that recent work, particularly

that of Montgomery and of Sutton, has shown that this fusion

occurs at a period long preceding the first maturation.

BOTANY.

Notes. — The Botanical Gazette for January contains the following

articles: — J. D. Smith, “ Undescribed Plants from Guatemala and

Other Central American Republics, XXIV”; Arthur, “Cultures of

Uredinez in 1902"; Dean, “Experimental Studies on Inulase” ;

Livingston, “The Distribution of the Upland Plant Societies of

Kent County, Michigan”; and Schneider, “Contributions to the

Biology of Rhizobia."

The Bulletin of the Torrey Botanical Club for January contains the

following articles : — Arthur, “ Problems in the Study of Plant Rusts ” ;

Evans, “Hepatice of Puerto Rico — II. Drepano-lejeunea” ;

Underwood, * An Index to the Described Species of Botrychium ” ;

and Kellerman, ** The Effects of Various Chemical Agents upon the

Starch-converting Power of Taka Diastase.

Floral Life is the title of a new journal which begins in January,

its first number being also noted as “Old Series No. 139,” it being a

continuation of Meehan’s Monthly.

No. 439) NOTES AND LITERATURE. 505

The Plant World for December, with a portrait of F. H. Knowlton

as frontispiece, contains the following articles: — Niles, “Origin of

Plant Names — IV; Parish, San Jacinto Mountain "; Wallace, * The

Preservation of our Wild Flowers, Shrubs and Trees"; Williams,

“Where Lichens grow”; Knowlton, “Fossil Mosses”; and Pollard,

" Cocoanuts in Cuba.” As a supplement to this number, — the title

page, etc., of Mr. Pollard's Zhe Families of Flowering Plants.

Rhodora for January contains the following articles: — Collins,

“North American Ulvacez"; Bissell, “A Botanical Trip to Salis-

bury, Ct."; Knowlton, “Flora of Mt. Saddleback, Me.”; Leavitt,

* Outgrowths on the Leaf of Aristolochia”; Pease, “ Erodium mala-

coides at Lawrence, Mass.”; Bissell, * Zycopodium clavatum and its

variety”; and Graves, “ Schwalbea Americana in Ct.”

An article on “The Functional Inertia of Plant Protoplasm,” by

Robertson, is published in Vol. III, No. 3, of the Proceedings of the

Scottish Microscopical Society.

“Plant Physiology for the High School,” by Ganong, and “ High

School botany,” by Syndam, are articles in School Science for February.

A fossil flora of the John Day Basin, Oregon, constitutes Bulletin

No. 204 of the U. S. Geological Survey.

From the structure of their seedlings, Miss Sargant argues, in the

Annals of Botany for January, that the monocotyledons are deriva-

tives of dicotyledons, rather than the reverse.

The anatomy of Macrozamia heteromera is written on by Agnes

Robertson in Vol. XII, part 1 of the Proceedings of the Cambridge

Philosophical Society.

In No. 8 and 9 of the Pharmaceutical Archives for 1902, Per-

rédés and Power respectively discuss the anatomy and the chem-

istry of Derris uliginosa,—an Eastern fish poison ; No. 1 of The

Pharmaceutical Archives for 1903, also, containing the conclusion of

Dr. Power's paper.

Chrysanthemum indicum, one of the original sources of the many

cultivated “Chrysanthemums,” is figured, accompanied by a note by

Sir Joseph Hooker, in Curtiss Botanical Magazine for January.

“Growing Cuban tobacco in the United States” forms the subject

of an illustrated article by Marrion Wilcox, in The World’s Work for

February.

506 THE AMERICAN NATURALIST. [Vor. XXXVII.

Part 4 of. Arthur and Holway's * Descriptions of American Ured-

inex” is published, with line illustrations, in Vol. V, no. 3 of the Bu/-

letin from the Laboratories of Natural History of the State University

of Iowa, dated in October. It may not be known generally that the

exsiccatze of the same authors are further illustrated by excellent

photographic representations of the species distributed.

Torreya for January contains: Gleason, * Notes on Some Southern

Illinois Plants " ; Watterson, * Louise Brisbin Dunn "; Lloyd, “ Vaca-

tion nearer sons — III"; Grout, “ Leaves of the Skunk Cabbage "

Murrill, “A New Family Br the Basidiomycetes " cr opp

based on Xylophagus and allied genera) ; Cockerell, * A New Oak —

Quercus rydbergiana ? ; and Berry, “Insect Visitors of Scrophularia

leporella.”

The 1902 Bericht der Senckenbergischen Naturforschenden Gesell-

schaft contains lectures by Móbius on carnivorous plants, Askenasy

on the phenomena of swelling, and Kinkelin on the development of

the plant world with reference to recent fossil collections.

An article on Droseras, in which several species are figured, is

published in Die Gartenwe/t of January roth.

Additional observations on The Strand Flora of New Jersey, by

Harshberger, have been separately issued from the Proceedings of the

Academy of Natural Sciences of Philadelphia, for October, under date

of December 12th.

Part 16 of J. M. Macoun’s “Contributions to Canadian Botany ”

is published in Zhe Ottawa Naturalist for February.

A flora of the town of Southington, Conn., and its vicinity, by Bis-

sell and Andrews, has been published as Connecticut School Docu-

ment No. 15, issued by the State Board of Education in 1902.

A short article on Santo Domingo, by Harshberger, has been

reprinted from Education of January.

Ginseng culture is the subject of Buletin No. 62 of the Pennsyl-

vania Agricultural Experiment Station.

* How to grow a Forest from Seed” is the title of Bulletin No. Q5

of the New Hampshire Agricultural Experiment Station.

A well illustrated article on “The Mango in Porto Rico,” by G. N.

Collins, is published as Bulletin No. 28 of the Bureau of Plant Indus-

try of the Department of Agriculture.

No 439.] NOTES AND LITERATURE. 507

Arboriculture for January is largely devoted to the hardy Catalpa,

C. speciosa.

Country life in America, for February, contains among other inter-

esting things an article on the orange in Florida, by Webber, and

one on orange growing in California, by Holder.

A well illustrated popular article on the date palm, by Sajo, is con-

tained in Prometheus for January.

A list of American varieties of vegetables for the years 1901 and

1902, by W. W. Tracy, Jr., is published as Bulletin No. 27 of the

Bureau of Plant Industry of the Department of Agriculture, and

forms a closely printed pamphlet of 402 pages.

No. 1 of Vol. VIII of the Anales del Museo Nacional de Buenos

Aires contains Nos. 51 to 190 of Spegazzini’s “ Mycetes Argentinen

es," the signatures of which are dated July 16, 1902.

The Gardeners Chronicle of January 24 contains a portrait and

short obituary of Wendland.

The Botanical Gazette for March contains the following articles:

Thaxter, “New or peculiar North American Hyphomycetes,

III”; Copeland, “Chemical Stimulation and the Evolution of

Carbon Dioxid (concluded) ” » Coulter and Chamberlain, * The Em-

bryology of Zamia”; Fink, “ Some Talus Cladonia Formations ”

Reed, “ Development of the Macrosporangium of Yucca dina”. "

Greenman, “Faxonanthus”; and Hitchcock, “Notes on North

American grasses.”

The Bulletin of the Torrey Botanical Club for March contains the

following articles: Cannon, “Studies in plant hybrids — The sper-

matogenesis of hybrid cotton”; Britton, “Timothy Field Allen

(with portrait) ; Vail, “Studies in the Asclepiadacez, VII. A new

species of Vincetoxicum from Alabama"; and Piper, * A new species

of Waldsteinia from Idaho."

The American Botanist for March contains the following popular

articles: C. F. Saunders, “ Early spring in southern California”; Turn-

bull, “Concerning nomenclature " ; and Goetting, “ A rare perfume."

The first Yearbook of the Carnegie Institution of Washington, re-

cently issued, contains much interesting information concerning the

botanical work being planned, later details of which are noted i in re-

cent issues of Science.

508 THE AMERICAN NATURALIST. [Vor. XXXVII.

The `“ Osservazioni scientifiche eseguite durante la spedizione

polare di S. A. R. Luigi Amadeo di Savoia," Milan, 1903, contains

among other things; chapters on phanerogams by Belli and crypto-

gams by Mattirolo.

The Flora of Tropical Africa, edited by Sir w. T: Thiselton-Dyer,

has reached No. 3 of Vol. IV, comprising Asclepiadeæ, in part, to

Gentianeæ, in part.

A developmental account of African Park-lands, by Professor

Tansley, with illustrations, appears in Zhe New Phylologist of Feb-

ruary 16,

Contributions to Western Botany, No. 11, of. Marcus E. Jones is-

sued April 10, 1903, is largely occupied with Abronia, ed

Eriogonum and Atriplex.

An account of the pine-woods of Florida, by Lepla, appears in recent

numbers of the Bulletin de la Société Centrale Forestière de Belgique.

The question as to what constitutes an “annual” is discussed by

Praeger in the /rish Naturalist for April.

A new Lower-Californian palm, Ezy/Aea brandegeei, is described

and figured by Purpus in Gartenfiora, Vol. LII

A number of new Mexican grasses are described by Hackel in

the opening number of Vol. XVII of the Annalen des K. K. Naturhts-

torischen Hofmuseums of Vienna.

The Ottawa Naturalist for April | contains a paper by Evans on

Yukon Hepatice.

The Bryologist for March contains the following articles: Fink

and Husband, “Notes on Certain Cladonias "; Barbour, “ Hepatics

pésperdi Holzinger, “ Karl Gustave Lisipicht (part 2)"; Holzin-

* Some notes on collecting " ; Grout, “ Pogonatum brevicaule” ;

abie * Psilopilum Dhsehehini ;" and Nicholson, “ Mnium

insigne à l

Separates of Dr. Galloway’s vice-presidential address at the Pitts-

burg meeting of the American Association for the advancement of

Science, on applied botany, retrospective and prospective, have been

distributed recently.

An economic study of Sequoia, published as Buletin No. 38 of the

Bureau of Forestry of the United States Department of Agriculture,

contains the following chapters :— Fisher, “ A study of the redwood” ;

. No. 439.] NOTES AND LITERATURE. 509

von Schrenk, “The brown rot Disease of the Redwood”; and

Hopkins, “Insect Enemies of the Redwood.”

Data on the self-fertility of the grape, comprising studies of the

potency of the pollen of self-sterile grapes, the influence on self-fer-

tility of girdling or bending the canes, and the pollen of the grape,

are published by Beach and Booth in Bulletins No. 223-4 of the

New York Agricultural Experiment Station.

An exhaustive study of the injury of plants by smoke and gases,

by Haselhoff and Lindau, has been issued from the press of. Born-

treger Brothers, of Leipzig.

An account of Polyporus fraxinophilus and its effects on the white

ash, by von Schrenk, constitutes Buletin No. 32 of the Bureau of

Plant Industry of the Department of Agriculture: ,

Professor Arthur's Washington address as President of the Botan-

ical Society of America, on problems in the study of plant rusts,

has been distributed by the secretary of the society.

The relation between frost-injury and parasitic ieie in cereals

is discussed by Sorauer in Landwirtschaftliche Jahrbücher, Vol.

XXXII, Heft 1, issued in March.

Among the complicated series and sub-series of University Bulle-

tins that have appeared in recent years as a means of securing peri-

odical mailing privileges, is to be noted an Ohio Mycological Bulletin,

forming part of the botanical series of the bulletins of the university

of that state.

The Journal of Mycology for February, with portrait of Dr. Far-

low as frontispiece, contains the following articles: Bubak, *Zwei

neue Pilze aus Ohio”; Morgan, “ Lepidoderma geaster” ; Kellerman,

“ A new species of Cephalosporium ”; Kellerman, “ Uredineous infec-

tion rper ene in igos ": : Geren: “Notes on Sclerospora gramini-

cola” ; Atkinson, * A new Mieciss of Calostoma "; Kellerman, “ Ohio

Fungi, fascicle VI, sss and notes] " ; Mica. “ Index to North

American Mycology”; and CERE * Notes from mycological

literature, IV."

The petiolar nectar glands of Viburnum, which form the subject

of a paper by Thouvenin in No. 171 of the Revue Générale de

Botanique, are homologized with leaflets of a compound leaf.

An interesting account of variations in the occurrence of salicin

510 THE AMERICAN NATURALIST. (Vor. XXXVII.

and salinigrin in different willow and poplar barks, by Jowett and

Potter, is issued as No. 28 of the publications of Zhe Wellcome

Chemical Research Laboratories.

The Berichte der Deutschen Botanischen Gesellschaft of March 25,

1903, includes a paper by Rosenberg on the chromosomes of a

Drosera hybrid, and a paper by Correns on the dominating charac-

teristics of hybrids.

From a statement by the Director in Bulletin du Fardin Impériale

Botanique de St. Pétersbourg, Vol. III, Livraison 1, it appears that

35,358 persons visited the extensive plant-houses of that great estab-

lishment in 1902, the yearly average for the past thirty years being

20,655.

The concluding number of Vol. II of the Bulletin of the New

York Botanical Garden, issued in March, shows the incorporation of

‘about 90,000 herbarium specimens, the addition of 1962 bound vol-

umes to the library, and the increase of species of plants cultivated

in the Garden to about 10,600, for 1902.

An account of the Glasgow Botanical Garden is contained in Zhe

Gardeners Chronicle of February 28.

In the recently commenced Buletin du Fardin Botanique de l Etat

à Bruxelles, Professor Massart discusses the problem of gardens for

the class purposes of secondary schools, and gives a list of 72 de-

sirable species, including one Fungus, one Alga, four Bryophytes,

and three Pteridophytes, with instructions for the more difficult

phases of the gardening.

Country Life in America for March is a gardening issue, adequately

. illustrated.

PUBLICATIONS RECEIVED.

(Regular exchanges are not included.)

COHEN, C. Physical t for Physicians and dpi I

from the German by M isc ew York, Henry Holt & vo,

Vili+343 pp., 49 fi qui du uH. Die Seele als deer Nase

Studien iiber die Bewegungen "e t Organismen Leipzig, Engelmann, 1903. 8vo

vi + 97 pp. Marks 1.60. — GARDIN . S. The Fauna and Geography of

R, J

the Maldive and Laccadive elikot Vol. i, pt. iv, pp. 347-471, Pls.

XVIII-XXV, text-figs. 76-119. Cambridge, eiae Press, 1903. 15/. —

HuNTER, S. J. Elementary Studies in Insect Life. Topeka, Crane & Co. 8vo,

xviii + 344 pp., 234 figs. — Nissi, F. Die bus enr und ihre Anhünger.

Jena, Fischer, 1903. 8vo, 478 pp., 2 pls. Mk. 12.— RADL, E. arepe qm

iiber den Phototropismus p Tiere. Lis g ga lmann, 1903 viii

+ 188 pp. Mk. 4.— REDFIELD, C. L. Control of Here edity. A Min p the

Genesis of Evolution by Ditam and Types of Character. Chicago and Phil-

RAR pagpa Book Co., 1903. 8vo, 343 pp. Illustrated. — SHERBORN,

C. Index Animalium sive Index Nominum quae ab A. D. MDCCLVIII

baies et C Soéciebas Animalium imposita sunt. Sectio Prima a kalendis Jan-

uariis MDCCLVIII usque ad finem Decembris, MDCCC. pensi d Univer-

sity Press, 1902. 8vo, lix +1195 pp. 25/.— SNopGRass, R. E. Anat-

omy of the Carolina Locust. (Dissosteira Carolina Linn.. A igit for

Teachers and Students. Pullman, OE , Wash. Agr. Coll. 1903. 8vo., 50

PP. 6 pls. | $0.50. — VERRILL, A. The Be rmuda Islands. An PPN of

their Scenery, Climate, Productions, de Natural History and Geo

with Sketches of their Discovery and Early History, and the changes in =.

Fauna and des due to Man. New Haven, lanig Ae a author, 1902.

8vo, x + 548 , 38 pls. and 250 text-figs.— M Variation in

Animals and | ide New York, Henry Holt & Co., Via o, ix + 415 p

30 figs. — VON ZITTEL, K. A. Text Book of RESTS duis Translated uud

Edited by C. R. Eastman. Vol. II. London, Macmillan, 1902. 8vo, viii + 283

d 3 Bernh $2.7 5.

. W. Notes on Sassafras. au Gazette. Vol. xxxiv, pp. 426-450,

^R " iud 4 i boas T. D. A. A New Cocklebur from New Mex-

ico. Proc. Biol. Soc. Wash. Vol. xvi, pp. 9-10. — CONDRA, G. E. New =

ozoa from the Coal Measures of Nebr ges Amer. Geologist. Vol.

337-359, pls. 18-25. — CuMMINGs, E. R. The nid gun of Platjstrophia.

A Study of the xe ea of a Paleozoic Modi od. Am. Jour. Sci. Vol. xv

63 pp, 27 figs. — DALL, W. H. Review of the S a of dd Crac).

Proc. Biol. Soc Fo . Vol. xvi, pp. 5-8. — DAVENPORT, C. B. The Animal

Ecology of the Cold Springs Sand Spit. Univ. Chicago Decen. Publ. Vol.

X, pp. 157-176. 7 figs. — Davis, B. M. Oogenesis in Saprolegnia. Univ. of

Chicago Decennial Publications. Vol. x, pp. 227-33. pls. 15-16. — pede E C

. Peach Gro wing in West Virginia. Bull. W. Va. Agr. Exp. Sta. o. 82, pp.

339-441. 16 figs.— DEAN, BasHFORD. Japanese Oyster Culture. po US

gtr

512 THE AMERICAN NATURALIST. (VoL. XXXVII.

Fish Com. for r902. Pp. 17-37, pls. 3-7. — DUERDEN, J. E. On the Actinian

Bunodeopsis globulifera Verrill. Zrans. Linn. Soc. London. Vol. viii. Pt. 9,

pp. 296-317. Pls. 25-26.— DUERDEN, J. E. The Morphology of the Madre-

poraria. III. The Significance of Badstine and Fission. Ann. Mag. Nat. Hist.

Ser. 7, vol. x, pp. 382-393. 4 figs. — DUERDEN, J. E. The Morphology of the

Madreporia. IV. Fissiparous Gemmation. Ann. Mag. Nat. Hist. Ser. 7, vol.

xi, pp. 141-155. 7 figs.— DUERDEN, J. E. West Indian | Madreporarin Polyps.

Mem. Natl. Acad. Sci. Vol. viii, pp. 399 + 645, 25 pls.

A Revised List of the Coleoptera observed near te Ohio, with Notes on

Localities, Bibliographical References and Description of .Six New Species.

Four "cin. Soc. Nat. Hist.. Vol. xx, pp. 107-196.. 5 figs. —Fisu, P. A

The Cerebral Fissures of the Atlantic Walrus, Proc. U. S. Natl. Mus. Vol. xxvi,

Pp- 675- 688, pls. 28-29.— GILL, THEO. The Use of the Name Torpedo for the

Electric Catfish. . Proc. U.: S. Natl. Mus. Vol. xxvi, pp. 697-698.— HATCHER,

Be. New Sauropod Dinosaur from the Jurassic of Colorado. Proc. Biol.

Soc. Wash. Vol. xvi, pp. 1-2.— Hay, O. P. On Certain Genera and Species of

North American Cretaceous. Actinopterous Fishes. Bull. Amer. Mus. Nat. Hist.

Vol —95, pls. 1-v-and 72 figs. — HELLER, E. Papers from the Hopkins

Stanford Galapagos Kenedüion, 1898-1899. XIV. Reptiles. Proc. Wash. Acad.

Sez. Vol PP. 39-98.— HOWELL, A. H. Three New Skunks of the Genus

'Bpldguie jy en Biol. Soc, Wash. Nol. xv, pp. 241-242.— JORDAN, D. S. Sup-

Aisi Note on Bleekeria mitsukurii and of Certain Japanese Fishes. Proc.

U. S. Natl.

pp. 16 A 3

FowLER, H. W. r purum o. the Colle or furem of Japan. Proc.

U.S. Natl. Mus. Vol. xxvi, pp.-699-702, 1 fig.— JonDAN, D. S. and FOWLER,

H. W. A Review of the Cobitidz or Loaches of the Rivers of Japan. Proc. U.

S. Natl, Mus. Mee xxvi, pp. 765—774, 2 figs.— JORDAN, D. S. and gre CR

LP of a New Species of Sinne from Japan. Proc. U. S. Natl. Mus.

Vol. «pp. DA HA 1 fig.— KELLOGG, V. p A iy Bares: Midges

(Blepharocéridie) of North America. Proc. Cal. d. Sci. Zool. Vol. iii, pp.

187-232, pls. 18-22. — KRAEMER, H. The Pith pe of Phytolacca fee aig

Torreya. Vol. ii, pp. 141—143.— LYDELI, D. Nie dde and Culture of the

Black Bass. Bul. U. S. Fish. Com. for 1902. pp. 39-44, pl. 8.— iiie, T: H.

On the Identification of a TES of eeni from the Piiisbtuee. Proc

S. Natl. Mus. Vol, x pp. 691-692 de er A.Z. The Atlantic Palolo.

Sci. Bull. Mus. Been inst. Vol. i, No D. 93-103.— MENNEL

The Zimbawe Ruins. Rhodesia Mus. Special d . 16 pp., 6 pls.— MEL

H. o New. Wood Rats (Genus Neotoma) from State of Coahuila,

Proc. Biol. Soc. Wash. Vol. xvi, pp. 47-48. —MiLLER, G. S, Jr. A New Bat

from the Island of Dominica. Proc. Biol. Soc. Wash. o a

MILLER, G. S., Jr. Two New Tropical Old World Bats. Proc. Biol. S

Vol. xv, pp. 245-240.— MILLER, G. S., Jr. arn gie of Eleven New Malayan

Mouse Deer. Zroc. Biol. Soc. Wash. Vol. x P- 31-44.— MILLER, G. S.,

Mammals collected by Dr. W. L. Abbott on rs Coast R Islands of Noui wa

Sumatra. Proc. U. S. Natl. Mis. Vol 437-484, pls.

MILLSPAUGH, C. F. Plante Yucatane (Mágionls Recta Plants of the Insu-

lar, Coastal and Plain Regions of the Peninsula of Yucatan, Mexico. Fasc. I

No. 439.] PUBLICATIONS RECEIVED. 513

Publications PR Colum. Mus. Bot. Ser. Vol. iii, No. 1, 36 pp., many figs.

NEE A Genealogical Study of the Dragon- Fy Wing Venation.

om 5. Natl. Mus. Vol. xxvi, pp. 703-764, pls. 31-54 and 44 text ince

— NELSON A. Two New Plants from Mexico. Proc. Biol. Soc. Wash. Vol. x

pp. 29-30.— NELSON, A. Psilostrophe, a Neglected Genus of Kein

Plants. Proc. Biol. Soc. Wash. Vol. xvi, pp. 19-24.— NELSON, A. an

COCKERELL, T. D. A. Three New Plants fron New Mexico, Proc. Biol. Soc.

Vol Xvi, p 5-46.— OBERHOLSER, H. C. Description of a New

ireo. Proc. Biol. Soc. Was. Vol. xvi, pp. 17-18.— OBE

New Cliff Swallow from Texas. Proc V. Wash. Vol. xvi, pp

Bio. i | 15

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-14.— ScHUCHERT, C. On the Lower Devonic and Ontaric Formations of

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The National Zoo at Washington, a Study of its Animals in Relation to their

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Wash. Vol. xv, pp. 239-240.— STEJNEG A New Min for iba nae

Bird Genus Oreomyza. Proc. Biol. Es are Vol. xvi, pp. 11-12. — STEJ-

NEGER, L. Description of a New Species of Gecko from Cocos Island, Proc.

Biol. Soc. Wash. Vol. xvi, pp. 3-4. — STONE, J. E. Cucumbers under Glass.

Bull. Mass. Agr. Exp. ined No. 87. 43 pp., 12 figs.

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: Vor. XXXVII, No. 440

— The American Naturalist.

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THE

AMERICAN NATURALIST.

VoL. XXXVII. August, 1903. NO. 440.

STUDIES OF GASTROPODA.

II. FuLGUR AND Svcorvrvs.

AMADEUS W. GRABAU.

Tue following brief account of the shell-development in

Fulgur and Sycotypus, and its bearing on our knowledge of the

succession of species in time is a summary of studies carried on

for some years in various museums, especially the National

Museum at Washington, the museums of the Academy of

Natural Sciences and of the Wagner Free Institute, at Phila-

delphia the American Museum of Natural History, the Museum

of Comparative Zoology at Cambridge, and the museum of the

Boston Society of Natural History. To the officers of all these

Institutions I am greatly indebted for many privileges and help-

ful suggestions.!

The fulguroids are a particularly interesting group on account

of their extra-limital distribution. Disregarding a few question-

1I wish € to mention the names of Dr. W. H. Dall, Prof. H.

ap Mr. C. W. Johnson, Prof. R. P. Whitfield, Dr. L. P. Gratacap, Dr. i

Mc. M. Woodworth, Prof. R. T. Jackson, and above all, the late Professor

paces Hyatt, as those who have most materially assisted me in my studies of

the Gastropoda.

545

516 THE AMERICAN NATURALIST. (VoL. XXXVII.

able reports of the alleged finding of these shells in other regions,

we find their geographical range in modern as well as Tertiary

[t otypus canaliculatus jus

before aus from the egg capsule ;

showing the shell, PN heart, and

large velum

times to be along the Atlantic coast

of North America, from the south

shore of Cape Cod, to the Gulf of

Mexico. The peculiarly restricted

distribution of this group.is explained

by the fact, that the animal on leaving

the egg capsule is without a velum.

This latter, though well developed

and large before hatching, (see Figs.

1 and 2) is dropped just before the

: animal emerges from the capsule, and

after the shell is already well devel-

oped. Thus the meroplanktonic

stage, which we may assume existed in the ancestors of Fulgur

and Sycotypus, was apparently suppressed even in the earliest

species of Fulgur, as otherwise the distribution would be more

world-wide. Actual migration is prevented by the differences

in temperature of the water and by the ocean currents. This

condition of affairs has existed in this region in all probability

since Miocene times.

SHELL DEVELOPMENT IN FULGUR AND SYvcoTvPUs.

The earliest shell stage, 7. e. the

protoconch, may be studied easily

in Figur caricum and Sycotypus

canaliculatus. It is generally pre-

served in the adult shell, and may

also be readily obtained from the

egg capsules, particularly in the

autumn, when embryos in all stages

of development are found often in

the same string of capsules. This

protoconch consists of a single

smooth volution, with no apparent

markings ; not even lines of growth

are visible. If however a young

Fic. 2. spe typus canaliculat: Ven

tral view of specimen Fig. 1, showing

foot, teil eyes, and velum

before it is lost. 13.5.

No. 440.] STUDIES OF .GASTROPODA. 517

individual is taken from the egg capsule before the shell has

assumed its distinctive characteristics, very faint lines of growth

may be seen, as well as indistinct radiating lines. These features

have been particularly observed in young Sycotypus canaliculatus,

as noted in a previous paper.! In this early stage the shell is

also umbilicated, and since there is no anterior canaliculation as

yet, this stage has the characteristics of a Natica shell. I have

elsewhere referred to this as the naticoid stage of the proto-

conch, and noted its very general occurrence in gastropods.?

This naticoid stage recalls the characters of early gastropods,

such as Straparollina remota of the Lower Cambric, which I

have tentatively considered as close to the protogastropod.

The protoconch of Fulgur caricum generally merges into the

early conch stage (the nepionic’), without any visible line of

demarkation between the two. Occasionally, however, such a

division does exist, in the form of a strong growth line. In

Sycotypus canaliculatus this line of demarkation appears more

frequently, while in Buccinum undatum it is, so far as noted, a

constant character. In this latter type, the change is often

emphasized by an angulation in the whorl.

The nepionic stage of shell growth begins with the second

whorl. The outermost portion of the lip becomes gradually

extended into the incipient anterior canal so characteristic of the

adults of these shells. In Fulgur and Sycotypus this notch or

incipient anterior canal, occurs at some distance outward from

the umbilical margin of the lip, and hence as growth progresses

the protoconch will have its plane of coiling tilted at an angle

to that of the plane of coiling of the succeeding whorls. This

gives the protoconch the characteristic oblique appearance found

in many canaliculate (siphonate) types. This pattern is less

pronounced in Buccinum and Fasciolaria. In consequence of

1 Studies of Simpie Amer. Nat. vol. 36, p. 925, fig. 8.

s: ae cit. p.

The Ever rno of stages in development [ontogenetic] is: #epronic

or MOM ; meanic or youthful; ephedic or adult, and gerontic or senile. Each

stage is furthermore divisible into 3 substages indicated by the use of the prefixes

ana, meta, or para. The prefix phyl- or phylo- refers to th e corresponding stage

of racial development, 7. e., the phylum; thus: phyloneanic, phylephebic, phyloge-

rontic, etc.

51 8 THE AMERICAN. NATURALIST. [Vor. XXXVII.

this change of plane of coiling the second whorl partly buries

the earliest portion of the first whorl or protoconch.

For about half a volution or less, the shell is smooth, although

lines of growth become more pronounced. At more or less regular

intervals stronger lines of growth appear (ananepionic). In the

later portion of the nepionic stage (metanepionic) longitudinal

wrinkles or ribs appear which characterize the ambital portion of

the whorl, and may be traced upward to the suture between the

two whorls. At about the same time or, in some cases, appar-

ently earlier, faint revolving lines become visible on the shell.

dri dun carica. The young shell taken from the egg cap-

They are nearly uniform, those on the ambitus being slightly

stronger. They represent the earliest part of the primary spirals

so pronounced in later stages. Almost immediately after the

inception of the vertical wrinkles or ribs, an angulation appears a

little above the ambitus of the whorl, which thus becomes divided

into shoulder and body portion. At the same time the ribs

become stronger on the angulation, where they soon assume the

character of tubercles in a single row from the obsolescence

of the ribs above and below the angulation. As the shoulder

above the angulation broadens, two new spiral lines appear on it

between the suture and the shoulder angle. These, with pro-

gressive growth, are augmented by the appearance of new ones

alternately on each side of the first two. At this stage the

operculum is already well developed (Figs. 3 and 4)

No. 440.] STUDIES OF GASTROPODA. 519

In Sycotypus canaliculatus the large velum (Figs. 1 & 2) is

lost at about the time of formation of tubercles on the shoulder

angle, and this may be considered the end of the metanepionic

period, shortly after which the animal passes from the egg cap-

sule. I have not examined Fulgur caricum at this stage, but

there can be little doubt that the conditions are essentially sim-

ilar. The shell-characteristics of these two types and of Buc-

cinum and Fasciolaria at hatching are indicated in figure 18.

The last or paranepionic period, distinguished by a tuberculated

angulation (Fig. 4), is brought to a close in S. canaliculatus by

the formation of the char-

acteristic sutural canal.

This, in some individuals,

appears to have just begun

on the lip at the period of

hatching, but usually it be-

gins only toward the end

of the second volution. At

this stage, the early neanic,

the lines of growth are well

marked and of nearly equal

strength with the revolving lines, the two together giving the

shell surface a reticulated appearance. The sutural canal is

bounded by a strong revolving ridge, between which and the

row of nodes on the angle, lies the sunken and flat or slightly

concave shoulder. On this shoulder three distinct spirals are

defined, at the beginning of the neanic stage, z. e., the third

conch volution. Shortly before the beginning of the fourth

volution of the conch (the fifth counting the protoconch) a

fourth spiral appears on the shoulder, close to the ridge bound-

ing the sutural canal. A little later a fifth appears just within

the line of nodes. Shortly after this, the neanic stage comes to

an end, the shoulder ridge losing its nodes and continuing as a

simple keel. This occurs between the fourth and fifth volutions

of the conch. At the same time the inner ridge becomes less

prominent, and the shoulder consequently level instead of sunken.

At about this time (end of fourth volution) a sixth spiral appears

on the shoulder, next to the inner ridge, and half a volution later

FIG. 4. Sycotypus canaliculatus, tle ycung shell taken

from the egg capsule. X 11.5.

520 THE AMERICAN NATURALIST. [Vor. XXXVII.

a seventh, next to the keel. Finally an eighth spiral appears on

the sutural side before the completion of the fifth conch volu-

tion. All these spirals soon become of nearly uniform size, while

a second cycle of spirals begins to appear between those of the

first cycle. as they diverge with the increase in .width of the

shoulder. A supplementary spiral also appears just within the

sutural canal, next to the bounding ridge. "Where these spirals

are crossed by the lines of growth, short, pointed, horny spines

are found on the periostracum. On the body of the shell inter-

calation begins much earlier than on the shoulder. In accel-

erated individuals intercalated spirals appear earlier on the

shoulder. X

Sycotypus pyrus of the modern Floridian fauna seems to be a

more accelerated type. The smooth and ribbed stages are passed

through quickly, occupying only about half a volution. The

_tubercles continue to the end of the second conch volution, after

which a smooth'keel sucéeeds. When the shell has reached the

size where, in S. canaliculatus, the tubercles change into the

smooth keel (between the fourth and fifth volutions), the keel

in S. pyrus begins to disappear, and the curvature of the outer

lip becomes uniform. In large (adult) individuals the carina has

generally disappeared altogether, and the lip is rounded uniformly.

This produces the “excavatus ” type of lip. The canal begins in

the second volution of the conch. In two old Floridian shells of

S. pyrus, in the collection of the American Museum of Natural

History, the canal becomes narrower and less pronounced in the

last whorl, and in some instances it disappears altogether for a

short space. Ina characteristic specimen five spirals have devel-

oped by the end of the first conch volution. The sixth appears

immediately after the beginning of the second volution, and the

seventh begins in the third volution. Near the end of this volu-

tion intercalated spirals (of the second series) also appear, while

in S. canaliculatus these generally do not appear until the begin-

ning of the sixth volution. In this species the amount of embrac-

ing varies, thus changing the elevation of the spire.

No. 440.] STUDIES OF GASTROPODA. 521

ANCESTRAL SPECIES.

An immediate ancestor of S. pyrus seems to be S. pyriformis

(Conrad) from the late Miocene of the Natural Well, Duplin

Co., North Carolina! In this species the tubercles continue

nearly as long as those of S. canaliculatus; they become obso-

lete on about the fifth volution and are succeeded by a keel.

The last portion of the sixth volution is rounded, as in adult .S.

pyrus. S. pyriformis is therefore a less accelerated type than

S. pyrus, and fulfills all the requirements of the immediate

ancestor of the latter.

The next earlier representative of the series seems to be a

form from the Miocene of Faison, N. C., and probably referable

to S, canaliferus (Conrad) Gill. The type of this species is from

the early Pliocene (Waccamaw beds of South Carolina, Tuomey

and Holmes), and is a more advanced type, not far removed from

the recent S. canaliculatus. (See Busicon canaliculatum, Tuomey

and Holmes, 2/Zoecene Fossil Shells of N. Carolina, pl. 29, fig.

2). This species is considered by Dall as the ancestor of S.

canaliculatus. It fulfills the requirements of such a relation-

ship, in that the keel is tuberculated throughout. While S.

canaliculatus has advanced only one stage beyond S. canait-

Jerus S. pyrus and S. pyriformis have passed two stages beyond

this. Therefore a species of the S. pyrus type only as far

advanced as S. canaliculatus, i. e., one in which the last whorl

is keeled but not rounded, might be looked for in the upper

Miocene. These conditions seem to be satisfied by species

occurring in the late Miocene beds of Faison Mt. Pass, North

Carolina.

Other specimens found in these later Miocene beds of Faison,

N. C., show well the manner of formation of the sutural channel

in retarded as well as primitive types. In the simplest specimen

seen, a flattening appears near the suture in the third whorl of

the conch. This becomes wider, more pronounced, and faintly

depressed, and the shoulder has four simple revolving spirals.

This condition continues to the end of the specimen (a young

! Am, Journ. Conch. Vol. 3, 1867. p. 265, pl. 20, fig. 1.

522 THE AMERICAN NATURALIST. [Vor. XXXVII.

one with five volutions), while the shoulder angle at the same

time retains its tubercles. In more advanced specimens the flat-

tening, which begins earlier, is bordered by a carina arising in

the third or fourth whorl of the conch, while the shoulder

becomes excavated below this. The tubercles of the shoulder

angle cease about a whorl later and a simple carina succeeds.

In a still more accelerated specimen the flattening next to the

suture is depressed on the formation of the bounding carina on

the third whorl, while acceleration is also shown by the presence

of intercalated spirals on the shoulder of that whorl. In the

accelerated types the depressed sutural canal becomes triangular

from the development of the

bounding carina into a strong

posterior notch. This is empha-

sized, as acceleration increases,

by the disappearance of the

shoulder angle; which gives us,

when the spire is low, the type

named by Conrad Busicon exca-

vatus (Fig. 5) but a type very

Fic. 5. AN different in appearance when "55 Sow.

olinensts. After Tuo-

excavatus

Alter Contud- high. (See S. elongatus Gill, mey and Holmes.

Am. Jour. Conchology, Nol. 3, p.

150). Finally in these beds (Magnolia, Duplin Co. Smith-

sonian collection, 114540) occur specimens showing every grada-

tion between S. excavatus and the extremely accelerated type

described by Tuomey and Holmes as Cussidulus carolinensis

(Fig. 6) from the Pliocene (? ) of South Carolina. The “exca-

vatus" type of aperture is, however, not confined to one line of

descent, but crops out in parallel lines, 7. e., is a homoplastic

character. This is shown by old age, or accelerated individ-

uals of modern Sycotypus pyrus and by specimens referred to

S. pyrus from the Pliocene of Shell Creek, Fla. (Natl. Mus.).

These shells can therefore not be classed together as one species.

Apparently S. excavatus led to S. carolinensis, and with that

this branch became extinct in the Miocene. The slight develop-

ment of the sutural channel in some of these specimens is prob-

ably to be explained as a case of retardation in development,

No. 440.] STUDIES OF GASTROPODA. 523

recalling the characters of the late Oligocene ancestors of the

series. (See beyond.) 5

Another branch, which became extinct in the Miocene, is that

of S. zuci/is (Conrad). This is primitive or retarded as far as the

shoulder spirals are concerned, for they never reach the second

cycle, but become obsolete after the third or fourth primary spi-

ral has appeared. The change in this series is toward a progres-

. Sycotypus incilis. An extreme old-age type (a) and an elon-

ate phylogorontic variety (b). From specimens in the palzontologic

collections of Columbia University. X $.

sive widening of the sutural canal, until, in some adults, it is

wider at the lip than the shoulder. The ridge bounding it

becomes very pronounced, so as to form a prominent posterior

notch. This notch separates more and more from the body of

the shell as the sutural canal becomes wider, until finally it occu-

pies nearly the position which is occupied by the shoulder angle in

S. canaliculatus (Fig. 7a). The shoulder angle becomes almost or

524 THE AMERICAN NATURALIST. [Vor. XXXVII.

quite obsolete in the more accelerated individuals, thus giving the

lip a rounded “excavatum’’ form. Here the notch made by the

sutural ridge is the strongest element, having entirely replaced

the original notch of the outer or keel portion. This is com-

monly accompanied by a looser coiling, because the succeeding

whorls clasp below the middle of the preceding one instead of

above it, as in S. canaliculatus. S. incilis developed in the mid-

Miocene Yorktown beds, and terminated in a high-spired and

short-canaled form with prominent sutural channel and obsolete

shoulder angle. (Fig. 7b.) A form of this kind is not unlike in

appearance to that of an old Buccinum in which the last whorl

has been partially separated. The spirals of the shoulder are

always few in number, and generally simple; indicating an off-

shoot from a primitive line with few spirals, or a partial retarda-

tion affecting this feature. This small number of spirals, in S.

zncilis seems to be due to the rapid widening of the sutural chan-

nel and the consequent encroachment of the bounding ridge on

the shoulder space. The latter remains always narrow, thus

preventing a material increase in the number of spirals.

Apparently in the direct line of ancestry of S. incilis is S.

vm alveatus (Conrad) (Fig. 8). This preserves

the keel throughout and has the strong ridge

bordering the sutural channel. With our

present knowledge we may perhaps regard S.

alveatus as the radicle of this line, which

seems to have branched off from the main

line in Miocene time. The changes are

towards a high spire and a profound sutural

canal, terminated by an excavatum-like lip.

ro | S. incilis may therefore be considered as

Fic, 8. Sycotypus alvea- * :

tus. X4. After Cn- Tepresenting a phylogerontic branch in mid-

nd. Miocene time which was not propagated

beyond that period.

In the Duplin beds occurs a high-spired form described by

Tuomey and Holmes as S. conradii, but referred by Conrad to S.

incile. This I believe to bea distinct branch, which seems to

be a derivative from the excavatus line and a parallel to .S.

zncilis. In S. conradii the spirals remain comparatively few and

No. 440] STUDIES OF GASTROPODA. 525

the whorls embrace below the middle. The sutural canal

remains narrow, however, and the shell has more nearly the

aspect of S. canaliculatus. Intermediate types connect this with

S. excavatus and S. canaliferus.

The ancestral type of these branches of Sycotypus is prob-

ably close to S. rugosus (Conrad) in most characters (Fig. 9).

This Mid-Miocene species retains its tubercles

throughout, though these become very promi- `

nent in certain varieties and almost spine like

in the later whorls. The channel already

begins in the late third or early fourth whorl,

hence this is no new character. The number

of spirals is simple throughout, as far as

observed, but there are seven of them fully

formed in the fourth or fifth whorl. In some

specimens the shoulder tubercles begin to E

unite into a strong ridge. UE MERE

In S. coronatus we have a form which com-

pares in the character of the sutural canal with the ancestral

type, but which has the tubercles strongly developed and the

spirals intercalated at an early stage, showing acceleration in

these points. Another terminal type, S. concinnus, Conr. has

simpler spirals and a coronatus-like shoulder angle, which is

however more compressed and projects more than in that

species.

The ancestral type of these species of Sycotypus is probably

to be sought in the Upper Oligocene mutations named by Dall.

tampaensis and perizonatus, and referred as varieties to Conrad's

Fusus spiniger. In the former of these, seven simple spirals

occur on the shoulder and a smooth space between the suture

and the first spiral in front of it. In the latter, intercalation of

secondary spirals begins on the shoulder after the fifth simple

one has appeared, and the smooth space next to the suture

becomes depressed into a canaliculation. The mutation duruszt

Dall of the Upper Oligocene, seems to be still more primitive in

generally having only nodules with coarse and few spirals, and

the suture scarcely channeled. It appears that the specimens

classed together under the name of Fulgur spiniger Conrad are

526 THE AMERICAN NATURALIST. [Vor. XXXVII.

not of monophyletic origin. An examination of Conrad's type

from Vicksburg, Miss., in the collection of the Academy of Nat-

ural Sciences at Philadelphia shows it to have a protoconch of the

Levifusus type. This consists of about two and one-half volu-

tions; the apical whorl is minute, and the succeeding ones

enlarge gradually. Anangulation is formed by two spirals, above

which the shoulder becomes gently concave. The upper spiral

becomes stronger and alone forms the echinations in the later

whorls. Riblets appear early after the shell has become angu-

lated and these are soon reduced to mere tubercles. After two

whorls the latter become strong enough to be called spines.

In the Lower Miocene of the Chipola River, Florida, occurs a

type with true Fulgur protoconch, which has been referred to

F. spiniger. This, together with the species referred by Dall to

F. nodulatum, appear to be lateral branches from the main stem

which led to the modern types. Conrad’s S. nodu/atus has the

aspect of another extreme type of the Levifusus series, though

it may turn out to be a true Fulgur.

At this time, as Dall has well said, the sutural canal was not

a well fixed character, having but just made its appearance. It

was hardly of specific and certainly not of generic value, but

soon after, in the Middle Miocene, it became well established

and fixed in the Sycotypus line of development, which hence-

forth became an independent branch, with only occasional rever-

sions to a faint sutural canal.

On searching for the Eocene ancestor of the fulgurs we

apparently find it in a type from the Lower Clairborne of Texas,

which has been identified with Levifusus pagoda Heilprin. This

type, however, differs from Heilprin's species in having a true

Fulgur protoconch, whereas Z. pagoda has a three-whorled nati-

coid protoconch with gradually enlarging volutions ornamented

in the latter portion by semilunar riblets. This type of proto-

conch is characteristic of many species of Pleurotoma, to which

Levifusus seems to be closely related. But in the Texan type

the obliquely elevated, swollen fulguroid protoconch is smooth

for a little over a whorl and then is furnished with fine vertical

ribs which merge into those of the round-whorled succeeding

portion of the shell. There are at least two round whorls with

No. 440.] STUDIES OF GASTROPODA. 527

simple ribs and simple spirals, after which the shoulder flattens

out and the ribs become faint toward the suture. An angulation

appears on the whorl, formed at first by two strong spirals, but

later, the upper becomes strongest and causes the formation of

rather flattened blunt serrations. Intercalated spirals appear on

the sixth whorl.

This shell, though much smaller than the Fulgurs, has all the

characteristics required for an immediate ancestor. It must, of

course, be separated from Levifusus pagoda, Heilprin, and I pro-

pose to designate it Levzfusus ? harrisi, after Professor Gilbert D.

Harris, whose indefatigable labors in the Tertiaries of the Gulf

region have brought together a wealth of material which may

serve as a basis for further phylogenetic study. Both Dall and

Harris consider Levifusus in the line of ancestry of Fulgur; the

latter, indeed, regards Z. pagoda Heilprin as the prototype of

the Fulgurs. The protoconch of the ordinary form of Z. pagoda

does not satisfy the conditions of such an ancestor, but that of

the Texan form does. Whether or not these two types are to be

considered congeneric remains to be determined ; further investi-

gation may show that the changes from a normal naticoid type of

protoconch to the oblique swollen Fulgur type occurred in this

genus. We may well believe that at first the form remained

unstable, oscillating between the two types, but by the time

Fulgur had developed, that feature no doubt, had become stable.

At the same time fulguroid types were probably developed from

the normal Levifusus pagoda, the result being such types as

* Fuleur" spiniger Conrad, which, as already noted, is not a true

Fulgur. Spiniger-like forms were also developed among the true

Fulgurs, the similarity of forms in both cases being explainable

as an instance of parallelism. “Fusus” guercollis Harris from

the Midway stage, and “Fusus” rugatus Aldrich from the

Lignitic, seem to be related to the ancestors of Fulgur. These

types, for which the generic name Fulgurofusus is proposed

have a fulguroid protoconch, while the early whorls are almost

identical with those of Fulgur. The adult F. guercollis has the

Fulgur characters of whorl grafted upon a Fusus form. This

type is more accelerated in that its whorls become angular as

early as do those of Fulgur, and it is not impossible that this

528 THE AMERICAN NATURALIST. (VoL. XXXVII.

type may be in direct line of ancestry of Fulgur. In that case

Levifusus ? harrisi cannot be considered as in the direct line of

ancestry.

THE SPECIES OF FULGUR.

In turning now to typical species of Fulgur, we find 7. fusti-

formis (Fig. 10) to be the most

primitive Mid-Miocene type.

In this the simple nodules

remain to the end; they are

replaced by a smooth rounded

keel only in accelerated or

old age individuals. This

Oligocene ancestor, which Fic. ıı. Fulgur tu

Fic. 1. Fulgur fusi gave rise to the two lines of nana

SE T MONDE *. Sycotypus and E ae

Fulgur. From Fulgur fusi-

formis on the one hand is derived the large and ponderous /udgur

maximum Conr. of the Yorktown beds

(Fig. 13), in which the tubercles are found

only in the young stage, while the adult is

smooth and round-whorled. Probably a.

phylogerontic derivative of this is the vari-

ety zudiculatum from the upper bed at

Alumn Bluff. In this the final whorl

reaches up so as to rise slightly above the

preceding ones. Fulgur carinatum Conr.

(Fig. 12) on the other hand, is derived

from F. fusiformis probably through F.

tuberculatum (Fig. 11) by a consolidation

of the tubercles into a continuous keel,

which characterizes the last and one or

"e P ME more of the preceding whorls. In some

No. 440.] STUDIES OF GASTROPODA. 529

specimens of F. tuberculatum, from St. Mary's. river, Md., the

nodes are replaced in the last whorl by a continuous strong and

smooth keel, while in others the nodes still persist faintly on

the keel of the last :

whorl. The -more

strongly spinous types

of F. tuberculatum rep-

resent- lateral radia-

tions. In general the

angular character of the

whorl is retained for a

brief period before the

whorl becomes rounded

to the maximum stage,

but no true keel has

been observed between

the tuberculous and the

smooth stages. In Fu-

sus, on the contrary, a

definite keel is generally

formed by the consolida-

tion of the tubercles, as

in Sycotypus canalicula-

tus and Fulgur carina-

tum. In Fusus, a round-

whorled condition follows the keel, but no spines of the /ugur

caricum type are known in any true Fusus. Similarly, a round-

whorled condition follows the keel in advanced species of Sycoty-

pus (S. pyrus, etc.). Such a condition we may expect in .S.

canaliculatus in old age types or in future accelerated descend-

ants of the present types. Examples of Fulgur carinatum may

yet be found with the final whorl rounded.

It appears then, that there are two branches of development in

the Fulgur series, one leading to a keeled condition and probably

never forming spines, the other to a smooth and then -spinous

condition. The latter branch was the successful one among the

fulgurs, while the former condition succeeded in Sycotypus. The

keeled branch occurred in the Miocene fulgurs but did not per-

Fic. 13. Fulgur maximum. X 1. After Conrad.

530 THE AMERICAN NATURALIST. | (Vor. XXXVII.

sist. Some of the early Sycotypus developed echinations by an

accentuation of the tubercles, but these did not prove successful.

Hemifusus and Melongena represent the spinous line among

the Fusidz, which seems to have been equally successful with the

non-spinous line of Fusus proper. In such types as Hemzfusus

colosseus, a strong line or spiral persists for a time after the tuber-

cles have become obsolete. This I have formerly spoken of as

a keel, but it represents merely the natural transition from tuber-

culous to round stage. A similar faintly carinate shoulder angle

“occurs again in most cases, before the spines appear.

In company with Fulgur maximum is a variety (var. A of Con-

rad) in which spines are found on the

last whorl. These spines are a new

feature and are not to be confounded

with the tubercles of the earlier

whorls. The latter are the remains

of the ribs on the shoulder angle,

while the spines are periodic emargi-

nations of the lip on the line of the

shoulder angle. In the early suc-

cessors of F. maximum these spines

occur only on the final whorl, but in

progressively accelerated types they

come in earlier and earlier, having the

appearance of being pushed back on

EY ue TOT the whorls. Fulgur tritonis Conrad

ter Conrad, (Fig. 14) Is a type from the Mid-

Miocene of Yorktown, Va., in which

spines appear after a short period of a round-whorled condition.

The spines gradually increase in size until they are equal to those

of the modern F. caricum. Two branches are met with here:

F. tritonis Conrad with a normal anterior canal, leading to F. cari-

cum; and F. filosum Conrad with a strong oblique fold on the

anterior canal in the later whorls, leading to F. eliceans Montf.

of the modern Fauna.

The modern F. caricum and F. eliceans are produced from

their respective ancestors F. tritonis and F. filosum by an acceler-

ation of the spinous stages, which are crowded back until the

No. 440.] STUDIES OF GASTROPODA. 531

maximum stage is dropped out altogether and the spines follow

immediately upon the tubercles, and even encroach upon them,

thus causing a perfect gradation from tubercles to spines. F.

caricum makes its appearance in the Waccama beds of the Caro-

linas while F. eZzceazs has, so far as I know, been reported only

from the modern Fauna. From A. e/ieaws we finally derive

the variety candelabrum Lam., in which there are only three

or four huge spines in the last whorl.

In the late Miocene (Duplin beds of S. Carolina) and the

early Pliocene (Caloosahatchee

beds of Florida) a parallel series

seems to have developed inde-

pendently from an ancestor of

the F. fusiformis type. In

these formations occur round-

whorled and spiralled shells of

the maximum type (Fig. 13)

except that they are much more

strongly contracted in front of

the body whorl than the York-

town species. This gives the

anterior portion of the shell a

slender fusiform aspect unlike

that of Fulgur caricum. Heil-

prin (Trans. Wag. Free Inst. of

Sci, vol. I, p. 73) describes

typical Fulgur maximum from

the Caloosahatchee beds of Flo-

rida. I have not seen these Fic. rs. Fulgur apum. Smooth variety.

shells, which, if Heilprin's diag- — $527 vani ns Mem

nosis is correct, represent the

last survivors of that Mid-Miocene type. The Pliocene type is a

variety of F. rapum Heilpr. without spines. (Fig. 15.) It is

! Dall states that in al! the members of the genus * the females have a wide

patulous canal, while the canal in male specimens is much narrower and more

cylindrical." (Loc. cit. p. 115.) The difference between the specimens from the

Late Miocene and Early Pliocene, and typical F. maximum from the Mid-Mio-

cene, is greater than can be accounted for by sexual differences alone.

532 THE AMERICAN NATURALIST. [Vor. XXXVII.

the immediate ancestor of typical F. rau», and appears to con-

nect it with F. maximum or F. fusiformis. F. rapum differs

merely in having faint suggestions of spines on the shoulder,

which has again assumed an angular outline; but this is not con-

tinuous. It represents the first stage in the development of the

caricum features. The spines of this species are small and may

be compared to the first formed portion of a spine in adult 7.

caricum, or to abortive spines in the same species. This form is

probably the morphic equivalent of Conrad's F. maximum var. A,

from the Mid-Miocene beds. More accelerated specimens with

a similarly contracted form have the spines fully developed in

the last whorl, and hence parallel 7. £7ztouzs, from which they

differ only in the more slender form. This may be called F.

rapum var. tritonoides. The terminal member of this branch

appears to be the Floridian Fulgur coarctum Sowerby, which has

been considered a dextral F. perversum, but which I believe to

be of independent origin, since its ancestors are found in the

Pliocene strata of the adjoining region. The interior of the

shells of this series is strongly lirate, and the shells are generally

thin.

Turning now to the reversed fulgurs, we find in F. contrarium

(Fig. 16) of the Late Miocene and Early Pliocene

(Duplin and Caloosahatchee beds) a form com-

parable to the smooth variety of F. rapum of the

same horizons. This form is knobbed in the

early whorls, and rounded and smooth (except

for the spirals) in the later ones. It is similar

to F. rapum in having the same slender con-

tracted end. It seems, from our present knowl-

5e KE edge of the shell characters and of the geologic

trarium. x V At. Succession of the mutations, that the separation

mor of the slender series into dextral and sinistral

forms occurred in the Late Miocene, and that the two branches

have remained separate ever since. Leidy and Willcox! have

figured a series of reversed Fulgurs showing the gradual transi-

tion in characters from the smooth type F. contrarium to the

1 Proc. Wag. Free Znst., vol. 3, pl. 9 and to.

No. 440.] STUDIES OF GASTROPODA. $33

modern spinous type, F. perversum. The series is exactly par-

allel to that leading from F. maximum to F. caricum and from

F. rapum to F. coarctum, although the species correspond in form

to the second rather than the first branch, as already noted.

The sinistral type corresponding to F. rapum Heilpr. may be

designated F. oórapum nom. nov.;' while that corresponding to

F. tritonoides is Conrad's Fulgur adversarium (Am. Journ.

Conch. vol. III, p. 185). This species is figured by Tuomey and

Holmes? as Fulgur perversum, and by Leidy and Willcox 3 as F.

contrarium and F. perversum respectively. The final member of

this series is F. perversum (Leidy and Willcox, figs. 6 and 7)

from the Pliocene and modern faunas of the Gulf region. This

corresponds to F. coarctum of the same fauna, which, as already

stated, is considered by Tryon and others a dextral F. perversum.

In Fig. 17 is represented the early nepionic stage of /u/gur

perversum.

In the Miocene marls of Cape Fear river, N. C., associated with

Fulgur filosum, occurs a

reversed type with the

same characters. For

purposes of designation I

will apply to this the name

of Fulgur obfilosum, nom.

nov. This may represent

merely a reversed condition

of F. filosum, or it may be Fig. 17. ^e pem Young shell taken from

a member of a distinct “ort x:

series, developed independently and parallel to the dextral series.

In its earlier stages, it is a Fu/gur adversarium developing the

characteristic fold only in the adult. This suggests that it may

have been derived from that type, though the present unsatis-

factory knowledge of the vertical distributions of these fossils

makes it impossible to state whether or not they are in proper

chronologic relation. A reversed type with the characters of 7.

! Leidy and Willcox, pl. 9 and 10, fig. 3; fig. 2 represents a Perversum type in

which a faint angularity occurs on the shoulder of the final whorl.

2 Pliocene Fossil Shells of N. Carolina, pl. 29, fig. 5-

3 Joc. cit., pl. 9, 10, figs. 4 an

534 THE AMERICAN NATURALIST. [Vor. XXXVII.

eliceans, described by Phillippi as F. £zemer? Phil., occurs in the

modern fauna of the south coast.! This may be merely a reversed

F. eliceans, but it seems more likely that it was independently

derived from the reversed Miocene type just mentioned (F. oġfilo-

sum).

The table at the end of the article indicates the probable

genetic relationships of the species under discussion.

VARIATIONS IN THE SPINES OF ADULT FULGUR CARICUM.

Normally the spines of F. caricum are uniform in the adult, but

individual variation is quite marked here. The final whorls of

this species are generally marked by a series of color bands, or

strong growth lines, which indicate a resting stage following a

growth period. Each growth period begins with a non-spinous

shoulder condition, and the spine begins to form only in the last

half or last third of the period. At the end of the period the

spine is at its maximum, and opens forward. With the begin-

ning of the next growth period, the spine is closed anteriorly, or

more rarely, is abandoned abruptly and floored over at the base,

remaining open in front. Not infrequently the period is short-

ened, when the spine is only partially or not at all developed.

This may perhaps be a pathologic condition. Sometimes

the spine makes its appearance only toward the end of its

normal period, and hence is small. This character, due to

retardation, repeats the ancestral state of the spines in types

like F. maximum var. A Conr. (compare F. rapum), where it

was normal. Again, a period may be lengthened, when the

spine will be larger than the normal. Such is the normal con-

dition in F. candelabrum, where each period constitutes a third

or a fourth of a volution.

In general, when the periods are of normal length, we finda

gradual decrease in the number of periods in a volution as the

shell increases in size. Thus in a number of individuals with

normal periods, the average for the fifth, sixth, and seventh

! Kiener, pl. 9, fig. 2; Tryon, vol. r p 57, fig. 390; F. gibbosum Conrad.

Proc. Acad. Nat. Ph Phila., 1862, p

No. 440.] STUDIES OF GASTROPODA. 535

volutions (including the protoconch) were 14, 13, and 12

respectively. Out of thirteen specimens, the average for the

6th and 7th volutions were 12 and rr: periods respectively,

while 13 for the 5th volution is not uncommon. In old age or

pathologic individuals the number of periods in the last portion

of the final whorl becomes much greater as the periodic additions

to the shell between resting stages are much shorter. Thus it

appears that in normal progressive types the period increases in

length as the shell increases in size, and at a rate more rapid than

the increase of the shell. The number of spines on each suc-

ceeding whorl is therefore fewer. Judged by this standard,

F. eliceans must be considered more accelerated than F. caricum.

In F. eliceans the average periods for the sth, 6th, and 7th

volutions are more nearly 12, 9, and 6 respectively. Finally,

in F. candelabrum we have extreme acceleration in this respect,

as the final whorl is provided with only three periods and spines.

RELATIVES OF FULGUR AND SYCOTYPUS.

Among the near relatives of Fulgur and Sycotypus in the

modern fauna we may mention Fasciolaria and Buccinum. A

ovo

Fic. 18. Outline cf young Fasciolarioid shells. a. Buccinum undatum, b. Fulgur caricum,

= cdd T lS ath d. Fasciolaria gigantea, X 10.

comparison of the early (ananepionic) shell stage of these two

forms with Fulgur and Sycotypus in the same stage of develop-

ment shows a very close similarity. (Compare Fig. 18.) Leav-

536 THE AMERICAN NATURALIST. | [Vor. XXXVII.

ing aside differences in size, we find that Fulgur has the longest

anterior canal, Buccinum the shortest, while that of Fulgur is

the narrowest, that of Sycotypus is the widest and most patulous.

Fasciolaria is the most ventricose of the series. In spite of

these differences, there is an unmistakable family resemblance

between the four young individuals, a resemblance, which in the

case of Fulgur and Sycotypus is retained even in the adults.

In Buccinum undatum spirals appear in the metanepionic stage,

and ribs at a later period. In Fasciolaria gigantea ribs appear

first (Fig. 19), the spirals afterwards. This therefore seems to

be nearer to Fulgur in which the same order of succession of

features obtains. It is therefore evident, that we must look to

the Mesozoic fasciolarioid shells for the ancestors of these

types.

The development of plications on the columella has been

shown by Dall to be due to a relative retreat of the muscle of

Fic. 19. gig toc spend ? Young shell taken from the

egg capsule. T"

fixation from the aperture, and the consequent crowding and

folding of the mantle. This feature appears in a large number

of phyletic series, and must be regarded as a secondary acquisi-

tion. In the fasciolarioid ancestors of the present types, such

plications could not have existed, since they do not occur in

Fulgur and Sycotypus, nor in the young Fasciolaria. We know

too little about the Cretacic forms at present to determine what

species are in the line of ancestry of the types under consid-

eration. Pyropsis or Pyrifusus, both of which are well repre-

No. 440.] STUDIES OF GASTROPODA. 537

sented in the Atlantic coast Cretacic, may represent the Miocene

progenitor, both having so far as I have been able to ascertain,

the characters requisite for such ancestry. These may lead on

the one hand to the Eocene levifusoid ancestor of the Fulgurs,

and on the other hand to Fasciolaria through such forms as

Odontofusus, with but a single columellar plication.!

Fusus, Hemifusus and Melongena are not directly related to

Fulgur. Fusus at least existed before Fulgur, having already

acquired its highly accelerated protoconch in the Eocene.

Fusus seems to be an Old World type, which did not reach the

American coast until Post Eocene time, so far as indicated by

evidence now available? Hemifusus is an accelerated descend-

ant of Fusus, with fulguroid form,? while Melongena is a phylo-

gerontic terminal member of that series. Thus it appears that

the Fusidze and the Fasciolaridze (the latter with branches lead-

ing to Fulgur on the one hand and to Buccinum on the other)

have been distinct since pre-Eocene time, and that their common

ancestor must be looked for in the Cretacic if not earlier.

I am well aware that to class Buccinum and Fulgur with

Fasciolaria, and Melongena and Hemifusus with Fusus, subordi-

nates lingual dentition to shell characters. The odontophores of

Fusus and Fasciolaria are similar, and different from those of the

other genera mentioned. In these again the type of dentition is

similar, though minor significant differences occur. It seems to

me that where the shell characters point the other way, mere sim-

ilarity of dentition is insufficient to establish relationship, but

must be explained rather as parallelism.

We know nothing of the lingual dentition of the Tertiary and

earlier species, and but little of that of modern types. Further-

more we know nothing of the changes, if any, which the denti-

tion undergoes in the development of the animal and so cannot

use lingual ontogeny as index of genetic relationship. The den-

!See Whitfield: Gastropoda amd Cephalopoda of the Raritan Clays and Green-

sand Marls of New Jersey. Monograph 18 U. S. Geol. Survey.

2 The American Eocene species of Fusus so far described do not belong to

that genus. See Grabau: American Naturalist, vol. 36, p. 922.

3 For illustration of protoconch and early conch of Hemifusus see Grabau /o-.

tit. p. 921; fig. 5.

538 THE AMERICAN NATURALIST. (Vor. XXXVII.

tal apparatus of Buccinum and Fulgur is very similar, and that

of Hemifusus and Melongena is nearly identical. In both groups,

the outer members or marginals of the adult odontophore bear

each one large and one or more small denticles, while the median

is supplied with small denticles only. In Fasciolaria and in

Fusus so far as known, the margina!s are furnished with numer-

ous nearly equal denticles. If we- assume that in the ancestral

type both marginals and median had one or at the most only a

few denticles, it is easy to see how the multidenticulate type

of modern Fasciolaria may be developed along one line, and the

similar type of modern Fusus along a parallel line. It is also

easy to see, that the heterodenticulate types of modern Buccinum

and Fulgur could branch off from the primitive fasciolarioid

ancestor, by the accentuation of the outer denticle of each mar-

ginal. Again the simple melongenoid type could branch off

from the primitive fusoid type, and develop by a similar accen-

tuation of the outer denticle. I see therefore no sufficient rea-

son in the similarity of odontophores for the present classifi-

cation of these genera, and instead of uniting Fulgur and

Hemifusus-Melongena in one family, and Fusus and Fasciolaria

in another, I feel that the development of the shell characters

show very clearly the close relationship of Fusus and H emifusus-

Melongena, and of Fasciolaria and Fulgur, with Buccinum not

far removed.

PALAONTOLOGICAL LABORATORY,

COLUMBIA UNIVERSITY.

No. 440.]

STUDIES OF GASTROPODA.

539

TABLE OF THE GENETIC RELATIONSHIPS OF THE SPECIES

oF FuLGUR AND SYCOTYPUS.

S. pyrus F. candelabrum

: (excavatus var.) Sere o

> FEET AA

B S. canaliculatus F. caricum

E - M ee

S. pyrus F. kieneri F. eliceans F. coarctum F. perversum

f F. adversarium

u F. tritonoides

T TA N cimi

E S. canaliferus F. caricum F. obrapum

F. contrarium

k F. rapum

2 S. carolinensis

F. rapum (var.)

S. elongatus F. obfilosum

& |S. «conradii S. excavatus F. filosum

Z< | |

= S. iformi

Ss 2 iin

S. canahiferus ,

" Faison var.

S. incilis var. F. tritonis F. carinatum

S. incili

mus F. maximum var. A F. tube rculatum

S. alveatus d i (carinate var.)

PROC 3 = ust F. tubercula-

~ e erein i tum

n | (echinate var.)

3 S. rugosus S. rugosus : «uu RUE

z (carinate var.) (echinate var.) |

4 painaen ina ER 4 haie teo STS

i F. maximum F. tuberculatum

a pU

z S. coronatus S. rugosus L

S. concinnus F. fusiformis

S. rugosus, primitive

z S. tampaensis F. spiniger ?

$= F. nodulatum ?

.É S. perizonatus S. burnsii |

ka |

Pz UL eueisisquiecii SPU RET crac ean cce

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF THE

MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L. MARK, Director.— No. 144.

ON THE STRUCTURE OF THE OUTER SEGMENTS

OF THE RODS IN THE RETINA OF

VERTEBRATES.

ARTHUR D. HOWARD.

SOME forty years ago the structure of the rods in the retina

of vertebrates excited a lively interest and received the attention

of many skilled observers, whose researches were directed prin-

cipally to the rods in the amphibians because of the advantage

offered by the large size of these organs. Good summaries of

these early investigations have been given by Hoffmann ('73-

78), and by Krause ('92).

From the first of these sources the following description of

the outer segment of the rod is taken. The outer segment is

composed of a highly refractive substance staining black with

osmic acid. In form it is cylindrical with a hemispherical,

slightly bulging, distal end. Under high magnification its outer

surface is seen to be marked by parallel striations which deviate

from a strictly longitudinal course only in that they are very

slightly spiral. This appearance is due to superficial furrows

alternating with ridges. The form of the outer segment may

thus be well compared to that of a column with a slightly spiral

fluting. In addition to the longitudinal striations, transverse

bands are present at regular intervals. These are surface indi-

cations of a plate structure. The plates of which the outer seg-

ment is composed are constant in thickness for a given species

and show little variation even in the whole vertebrate series.

. They are held together by a cement lying between their approxi-

mated faces. This cement is affected rapidly by certain reagents

and by its swelling causes a characteristic disintegration of the

outer segment into disks. This disintegration occurs earlier at

541

542 THE AMERICAN NATURALIST. [Vor. XXXVII.

the distal end of the segment than at the proximal one, the dif-

ference having been attributed to a protecting sheath over the

latter. The presence of an axial fiber, as maintained by Ritter

and others, was discredited because of much negative evidence.

In contrast to this general view Krause ('92), whose compara-

tive studies included particularly Rana and Salamandra, main-

tained that the outer segment consisted of a bright ** Grundsub-

stanz," in the periphery of which fibrils were imbedded and

twisted in so close a spiral that they gave the appearance of

transyerse striations, thus producing a condition favorable to

apparent transverse fracture. Greeff (:00, p. 103), however,

has recently expressed himself in favor of the older view that

the outer segment consists of a series of plates with an envelop-

ing sheath. “Jedes Aussenglied besteht 1. Aus einer Hülle

(Mantelschicht, Rinde, Haut) und 2. einem aus Plattchen

und Zwischensubstanz gebildeten ZzZa//£.' This view is also

accepted by Levi (: o1).

Bernard (:00, :01), who has published an account of the

embryonic as well as of the adult rods in amphibians, has main-

tained that these structures are protrusions from a syncytial

retina and that each rod is a delicate protoplasmic vesicle

traversed by a reticulum which eventually becomes condensed

into the axis of the rod by the absorption of a colorless

refractive and amorphous matter from the pigment cells. Thus

even among recent investigators much difference of opinion

exists as to the structure of the vertebrate rods.

The progress made in the study of the rods in vertebrates

during the last twenty-five years stands in considerable contrast

with that made in the investigation of the terminal optic organs

of invertebrates. In the arthropods, for instance, the rhabdomes,

the analogues of the rods and cones of vertebrates, were sup-

posed by most of the earlier writers to have been formed by

secretion, and in fact Watase ('90) went so far as to compare

them with surface cuticula. Their fibrous character, however,

. was observed by Patten ('86) and others and it was demonstrated

by Parker (95) that in the crayfish the fibrils composing them

are neurofibrils, and that the substance of the rhabdome is

more correctly described as differentiated living material like the

No. 440.] STRUCTURE OF OUTER SEGMENTS. 543

contractile substance of a muscle fibre, than as a secretion. This

view that the rhabdome is composed of neurofibrils has been

greatly extended among the invertebrates by the recent work of

Hesse (: 00, : 01).

Since in some crustaceans the rhabdomes are not only fibrous

but are composed of plates not unlike the so-called plates in the

rods of vertebrates, it is natural to ask whether the rods in ver-

tebrates may not also be fibrous, and with this question the pres-

ent paper is chiefly concerned. The ease with which frogs could

be obtained at all seasons and the comparatively large size of

their rods led me to investigate the retinas of these animals and

I chose in particular the common leopard frog, Rana pipiens

Schreber, as a favorable species.

For a satisfactory study of the rods it was necessary to free

them from their surrounding pigment. This was done in the

usual way by keeping the animal two or three hours in the dark.

In such “dark frogs” the retinal pigment completely withdraws

from the region between the rods into the bodies of the retinal

pigment cells.

Of the various methods for obtaining unwrinkled retinze the

following was used with success, and I know of no other which

preserves the eye in as natural a condition as this does. Frogs

kept the usual length of time in the dark were etherized, their

hearts were exposed, and fixing fluid was injected into their

arteries as in ordinary injections to demonstrate the arterial

system. The fluids used were Vom Rath’s picro-platino-osmo-

acetic mixture, j^; osmic acid, corrosive-acetic mixture, and

Perényi's fluid.

The last two penetrated most successfully. The osmic

preparations were only partially successful, for, owing apparently

to the rapid constriction of the blood vessels, a smaller amount

of the fluid reached the interior of the eye than by the other

methods. After injection, the whole head was immersed in the

fixing fluid and the eyes were not opened until they were more

orless fixed. Eyes thus prepared were embedded in paraffine

and cut for longitudinal or transverse sections of the rods. The

sections were stained in Heidenhain's iron-haematoxylin, Bóhmer's

haematoxylin, Mayers haemacaleium, and by Kupffers and

544 THE AMERICAN NATURALIST. [Vor. XXXVII.

Bethe’s methods for neurofibrils. Preparations were also made

by the cover-glass method for blood technique. This is well

adapted for experimenting with a large number of reagents and

stains, and has the advantage of insuring immediate fixation.

The examination of material prepared in the ways enumerated

gave evidence of a well marked axial core in the outer segment

of each rod. This core was seen in both longitudinal and

transverse sections of rods fixed in the various fluids already

mentioned and measured about one fourth the diameter of the

rod. It took none of the stains which I have tried with the

possible exception of picric acid. It is probably the structure

long ago seen by Dreser ('86) and recently identified by Bernard

(: Ot, p. 465) as the condensed reticular portion in the axis of

the rod. Its relative thickness precludes the possibility of its

being the so-called fibre of Ritter, if in fact this fibre exists.

At present I do not wish to express any opinion as to the exact

nature of this core.

As previously stated the substance of the rods has been

variously described as lamellar, spirally fibrous, etc. Since rods

prepared by different methods showed much difference in

structure, it was necessary to study fresh ones as a means of

interpreting what was seen in the preserved preparations. But

under the ordinary microscope the substance of fresh rods

appeared to be almost homogeneous and I was, therefore, obliged

to seek other means of studying these bodies. The problem

thus resolved itself into a search for conditions which would

bring out optical differentiation in an object which under

ordinary circumstances seemed optically homogeneous. Polar-

ized light seemed the most likely means, for, if the rods are

fibrous not only ought this to be open to determination by a

polarizing microscope, but it ought also to be possible by the same

means to ascertain the direction of the fibrils.

A polarizing microscope was used with a powerful artificial

white light and a gypsum interference plate inserted between

the Nicol prisms. The prisms were placed at such an angle to

each other as to give an interference color of a sensitive violet

of the first order. With the apparatus thus set up fresh prep-

arations of the retina more or less teased out were examined. -

No. 440.] STRUCTURE OF OUTER SEGMENTS. 545

In such preparations fields may easily be found containing

detached rods with their longitudinal axes lying in various

directions.

Outer segments lying parallel to the a axis of the gypsum

plate, 4- 45^ to the cross-hairs, showed a bright yellow color,

while those at right angles to this were bright blue. The colors

of an individual rod could be reversed by turning the preparation

so as to bring the rod into a line at right angles to its former

position. The inner segments of the rods are not highly refrac-

tive. These observations were made on the rods of Rana pipiens

but I have also tested the outer segments of the rods or cones,

as the case may be, in the mudpuppy (Necturus), turtle, snake,

lizard (Anolis), guinea pig, mouse, and ox, and with wholly con-

firmatory results.

This definite reaction demonstrates that the substance of the

outer segments is positively doubly refractive or anisotropic, 7. ¢.,

as regards their optical properties the outer segments have axes

of maximum elasticity at right angles to their lengths. To

obtain an immediate basis for comparison I made similar tests

of other tissues. Thus bundles of naked axis cylinders from the

inner surface of the vertebrate retina gave light reactions exactly

like those given by the outer segments of the rods and the same

was true of striped muscle fibres from the crayfish, frog, and ox

as well as of connective tissue fibres from the ligamentum nuche.

The rhabdomes from the compound eye of the crayfish were,

however, negatively anisotropic, but when it is remembered that

the fibrous structure of these bodies is at right angles to their

length instead of being parallel to it as in all the other bodies

tested, this apparent exception disappears. Since the neuro-

fibrils are known to run lengthwise the axis-cylinders of nerves

and since naked axis-cylinders and the outer segments of the

rods give the same color reactions in the polarizing microscope, .

I believe I am justified in concluding that fresh outer segments

of the rods of vertebrates like axis cylinders of nerve fibres pos-

sess a longitudinal fibrillation.

The color reactions just recorded are directly opposed to

Krause's conception of the rods as made up of spirally twisted

fibrils. Such a structure would give color reaction the opposite

546 THE AMERICAN NATURALIST. (VoL. XXXVII.

to those actually seen, for the fibrils would be nearly at right

angles to the longitudinal axis of the rod. Nor do these reactions

favor the view held by Bernard (:01) that the rods are proto-

plasmic vesicles filled with an amorphous refractive substance,

for the material is not amorphous but gives evidence of longi-

tudinal fibrillation. Patten’s ('98) hypothesis that the outer

segments are made up of minute fibrils at right angles to their

longitudinal axes is also inconsistent with these observations.

Although the evidence I have advanced cannot be said to be

opposed to the generally accepted view that the rod is made up

of many disk-shaped plates, I ‘am not inclined to place so much

emphasis on this as some have done. I have obtained abundant

evidence for the presence in fresh rods of transverse bands about

equal to each other in thickness and held together by an inter-

mediate substance of different optical behavior. But I have not

found the evidence for the disintegration of a rod into disks at

all convincing. There were certainly frequent instances of trans-

verse breaking, but it was seldom clear cut and there were often

signs of longitudinal splitting and of spreading at broken ends.

I believe we have in the rod certain conditions analogous to

those of striped muscle fibres. Both bodies are positively refrac-

tive, both possess a transverse lamellar arrangement of optically

differing substances, and under the action of certain reagents

both are said to break into transverse segments.

The structure of the muscle fibre is essentially fibrillar not-

withstanding its transverse fracture and I believe the structure

of the outer segment of the retinal rod to be in this respect like

that of the muscle fibre.

Having given the evidence for the longitudinal fibrous struc-

ture of the rods as I have found it by the use of polarized light,

I wish to discuss some contradictory results already recorded as

having been obtained by this method. Valentin (62) investi-

gated with polarized light a large number of animal tissues

including the rods of the retina and the axis cylinders of nerves,

and, as the following quotations show, he found that the reactions

of these two bodies were not similar but opposite. “ Die

nühere Verfolgung des Gegenstandes zeigt, das die optische

Axe der Langsaxe der Nerven parallel geht, man also hier

No. 440.] STRUCTURE OF OUTER SEGMENTS. 547

einen wahrhaft negativen Körper vor sich hat und die ganze

Erscheinung nur von dem Marke herrührt" (Valentin, '62, p.

123). “Man könnte theoretisch annehmen, das die Stäbchen

an und für sich nicht anders, als die markigen Nervenfasern

wirken" (p. 136). “Jene (Stäbchen) wären aber wahrhaft

positiv und das Ceres Mark von diesen wahrhaft negativ "

(p. 136).

It is thus evident that Valentin believed that the optical axes

of the rods and of the nerve fibres were not in agreement but

were at right angles to each other, and this opinion was accepted

by Max Schultze ('67), Krause ('92),! and Greeff (: oo).

It is not easy to account for Valentin’s statement that the

axis cylinders of nerves are negatively anisotropic unless we

assume that in consequence of the imperfect knowledge of nerve

Structure at his time he has recorded the reaction of the med-

ullary sheath, which is negative, instead of that of the axis cylin-

der. Valentin's work was done on Torpedo marmorata and

shows that his observations were made almost entirely upon

medullated nerves. It is quite evident that what he refers to as

sheaths of the nerve must have been the positively reacting con-

nective tissue of the peripheral nerves, for he makes no mention

whatever of the brilliantly conspicuous medullary sheath as such.

He does, however, speak of pressing out the retina of a frog

with a cover-glass and finding fibres which he considers to be

parts of the optic nerve. These, he states, also showed negative

reactions, but there is no certainty that what he described were

really optic nerve fibres.

In my tests of nerves I found medullated fibres unsatisfactory

objects for clear demonstration of optical properties in the axis

Cylinder, because of the strong predominance of the reaction

color of the medullary sheath. The non-medullated fibres from

invertebrates (crayfish) were more satisfactory, but even here

the presence of the positive Schwann's sheath, though compara-

tively thin, made conclusive observation out of the question for

the color of the sheath was projected on the less strongly react-

ing axis.

! * Die Aussenglieder sind femer positiv doppelbrechend, die optische Axe liegt

in ihrer Längsrichtung und es ist bemerkenswert das sie sich entgegengesetzt wie

-das bekanntlich negativ Nervenmark verhalten.” (Krause, '92, p. 159.)

548 THE AMERICAN NATURALIST. [Vor. XXXVII.

It was, therefore, necessary to üse nerve fibres without pro-

tective coverings. The naked axis cylinders radiating from the

entering optic nerve in the fibre layer of the retina, met this

requirement. In order to get a clear demonstration of these, I

made tests upon the retina from a perfectly fresh ox eye where

the large size of the eye made manipulation comparatively sim-

ple. In this case there was little difficulty in identifying the

radiating bundles of nerve fibres which were readily distinguish-

able from small blood vessels and other structures of a fibrous

nature. The bundles of naked axis cylinders proved to be dis-

tinctly positive, thus agreeing with the rods and I am conse-

quently forced to conclude that in some way Valentin's obser-

vations were in this respect erroneous.

Summary. The outer segments of the rods in the retina

of the frog contain each an axial core that differs from the

peripheral substance, but the exact nature of this core has not

yet been made out. The outer segments, as demonstrated by

the use of polarized light, are positively anisotropic and agree in

this respect with the axis cylinders of nerves. These outer

segments therefore, give evidence of containing longitudinal

fibrille. Since they also show in the fresh state a transverse

banding, their structure is in some respects not unlike that of

a cross-striped muscle fibre in that in addition to a cross band-

ing they also possess a longitudinal fibrillation.

No. 440.] STRUCTURE OF OUTER SEGMENTS. 549

BIBLIOGRAPHY.

BERNARD, H. M.

:00. Studies in the Retina: Rods and Cones in the F rog and in some

other Amphibia. Part I. Quart. Jour. Micr. Sci., new ser., vol.

43, no. s pP- 23-47, pl. 3.

BERNARD, H.

O01. Studies in the Retina: Rods and Cones in the Frog and in some

other Amphibia. Part II. Quart. Jour. Micr. Sci, new ser.,

vol. 44, no. 175, pp. 443-468, pls. 30-31.

DREsER, H.

'86. Zur Chemie der Netzhautstábchen. Zeit. f. Biol., Bd. 22, pp. 23-39.

GREEFF, R.

:00. Mikroskopische Anatomie des Sehnerven und der Netzhaut. Graefe-

Saemisch Handbuch der gesamten Augenheilkunde, Auflage 2,

Bd. 1, Kap. 5, 212 pp., 2 à

HESSE, R.

:00. Untersuchungen über die Organe der Lichtempfindung bei niederen

Thieren. VI. Die Augen einiger Mollusken. Zeit. wiss. Zool.,

Bd. 68, Heft. 3, pp. 379-477, Taf. 25-32.

HESSE, R.

:01. Untersuchungen über die Organe der Lichtempfindung bei neideren

Thieren. VII. Von den deo Rp Zeit. wiss. Zool.,

d. 70, Heft. 3; PP- 347-473, Taf. 1

iren CK.

78. Klassen und Ordnungen der Amphibien. Bronn Klassen und

Ordnungen des Thier-reichs, Bd. 6, Abt. 2, 726 pp., 52 Taf.

KRAUSE, W.

'92. Die Retina. Monatschr. f. Anat. u. Physiol, Bd. 9, pp. 150-155,

157-236.

LEvi, G.

:01. Osservazioni sullo svilleppo dei coni e bastoncini della retina degli

urodeli. Lo Sperimentale, anno 54, pp. 521—539, I tav.

PARKER, G. H.

'95. The Retina and Optic Ganglia in Decapods, especially in Astacus.

Mitth. Zool. Stat. Neapel, Bd. 12, Heft. 1, pp. 1-73, pl. 1-3.

PATTEN, W.

'86. Eyes of Molluscs and Arthropods. Mitth. Zool. Stat. Neapel, Bd.

6, Heft. 4, pp. 542-756, pls. 28-32.

PATTEN, W.

'98. A Basis for a Theory of Color Vision. Amer. Nat., vol. 32, no. 383,

pp. 833-857.

550 THE AMERICAN NATURALIST. [Vor. XXXVII.

SCHULTZE, M.

'67. Ueber Stábchen und Zapfen der Retina. Arch. mikr. Anat., Bd. 3,

pp. 215-247, Taf. 13.

VALENTIN, G.

'62. Histologische und physiologische Studien. Zeit. f. rationelle Medi-

cin, Reihe 3, Bd. 14, pp. 122-181.

WATASE, S.

:90. On the Morphology of the Compound Eyes of Arthropods. Studies

Biol. Lab. Hopkins Univ., vol. 4, no. 6, pp. 287-334, pls. 29-35.

VERNACULAR NAMES OF ANIMALS.

EDWIN W. DORAN.

THE interests of popular science demand the introduction of

some system in writing the common, or vernacular, names of

animals and plants. There is not only a great lack of uniformity

among authors as to the correct form of these names; but in

case of some approach to an agreement with regard to certain

forms, often the agreement is without reference to the principles

that underlie the formation of such words.

From an examination of many thousand names of animals it

appears that only about ten per cent. of the names consist of a

single word ; about thirty per cent. consist of two or more words

which should be written in separate form ; about sixty per cent.

consist of two or more words which should be written in com-

pound form. (Not all of these should have a hyphen, however,

as will be shown later). What names should be included in this

sixty per cent. is a problem upon the solution of which this paper

is designed to shed some light.

There are those who say the solution of the problem lies in

abolishing the use of vernacular names entirely, and using in

their stead the more exact technical names. There is little hope

of getting any except the most rigidly scientific, to adopt for

everyday use the unfamiliar forms derived from foreign lan-

guages, no matter how exact they may be. Vernacular names

always have been in use and no doubt always will be. No one

expects to live to see the day when we shall discard the familiar

names of dog, cat, rabbit, squirrel, cow, and sheep, and use in

their places the high-sounding though exact names derived from

the Latin or the Greek. If we retain these common names, —

household words, — we may expect also to retain such other

names derived from them, as prairie-dog, wildcat, jack-rabbit,

ground-squirrel, sea-cow, and mountain-sheep. Then we are

likely to distinguish still farther between the many closely-related

forms by using Missouri prairiedog, American wildcat, white-

55!

552 THE AMERICAN NATURALIST. [Vor. XXXVII.

tailed jack-rabbit, striped ground-squirrel, Dall mountain-sheep,

etc.

These terms are less exact for English-speaking people than

the corresponding technical ones simply because of the confusion

that has always existed in writing them; because of the many

names that have been applied to the same animal. The same

confusion would exist with reference to the technical names but

for the carefully prepared synonymies. I believe that I am the

first’ who has thought it necessary to prepare a synonymy of

the vernacular names of animals.

Some writers appear to have an antipathy for the much-abused

little character, known as a hyphen. This seems to be true also

of many printers and publishers. As one writer puts it, “A

hyphen affects some printers very much as a red rag does an

angry bull" ; hence they omit the greater number of hyphens in

a manuscript. Many writers on science relegate to the printer

or to their amanuensis such details of manuscript-revision and

proof-reading, owing to a lack of authorities on the subject.

Dr. J. A. Allen says?: “If the use of the hyphen could be per-

manently regulated by the formation of a set of rules, how great

a boon would be conferred upon writers, and particularly upon

editors! As, however, the use of the hyphen varies within wide

limits, in accordance with the radically different rules enforced

by editors and publishing houses, from its practical non-use to

its employment to connect remote elements into a compound

word, there is little hope of securing a uniform system of hyphen-

ization in the construction of bird names..... In publications

which allow the hyphen its time-honored function, great diversity

is met with in just the class of cases to which Dr. Doran has

called attention.”

Here is the difficulty. Every writer has some system of rules

which he follows in compounding words (provided he gives any

attention to the subject at all) ; but too often these rules are

formulated without regard to language-principles or reason.

1 The author has in preparation a synonymy of all the vernacular names of

vertebrates. Mr. Robert Ridgway many years ago suggested the need of some-

thing of this kind for the names of birds.

? The Auk, Jan. 1905, discussing my article on Vernacular Names of Birds.

No. 440.] VERNACULAR NAMES. 553

Some have not taken the trouble to reduce their rules to tangible

form or to a system, and thereby get rid of their inconsistencies.

Now, I believe that a system of rules may be formulated and

in accordance with the established principles of the English lan-

guage, by means of which we may be rid of the present chaos in

compounding the vernacular names of animals.

The writer has previously attempted this for certain groups of

animals! and in this paper the discussion is extended so as to

include all classes of vertebrates and the insects.

Before formulating a set of rules for compounding the vernac-

ular names of animals, it is necessary to give attention to a few

well-established general principles — principles which are recog-

nized by all the great masters of English, though expressed

concisely by few writers. In fact, the literature of the subject

is very meager, and only within the last twenty years has there

been any serious attempt to evolve a system of writing such

words,

The Standard Dictionary lays down three general principles

for compounding English words,? the second of which is as

follows: * Abnormal association of words generally indicates

unification in sense, and hence compounding in form.” In

accordance with this principle I submit the following rules for

compounding the vernacular names of animals: Following each

rule are given numerous examples taken from the different

groups of animals to show the application of the rules. The

first rule is more general than the rest, and to some extent

includes the others.

Write in compound form, —

I. Any pair of names or words in joint arbitrary use ; as, leaf-

roller, black-nosed dace, four-toed salamander, red-bellied

snake, blue-throated lizard, soft-shelled turtle, whippoorwill,

polecat.

1See ‘ Entomological News,’ Nov. 1902, for a discussion of the vernacular

names of insects, and ‘ Zhe Auk, January, 1903, previously cited, for a similar

treatment of the names of birds.

? Consult also the works of F. Horace Teall on compounding English words.

? Thes es are intended to apply to insects and vertebrates only, but will be

found serviceable for all forms.

554 THE AMERICAN NATURALIST. (Vor. XXXVII.

2. A general name used with any other name prefixed for

specification denoting,

a. Food or prey; as, potato-beetle, spawn-eater, chicken-

snake, duck-hawk, rice-rat.

b. Host; as, horse-fly, dog-flea (chiefly parasites).

3. A general name used with any other name prefixed for

specification denoting,

Similarity ; as, mole-cricket, alligator-gar, cricket-frog,

garter-snake, box-turtle, turkey-vulture, fox-squirrel.

b. Habit; as, army-worm, pilot-fish, rattlesnake, snapping-

turtle, butcher-bird, flying-squirrel.

c. Habitat ; as, house-fly, brook-trout, tree-frog, water-moc-

casin, fence-lizard, land-tortoise, wood-duck, prairie-dog.

Characteristic ; as, scale-insect, sword-fish, spade-foot

frog, horn-snake, spine-tailed lizard, map-turtle, song-

sparrow, musk-ox.

4. A phrase consisting of an adjective and a noun together

used as a mere name, formed by writing (generally in solid

form),

a. An adjective with the name of an animal ; as, whitefish,

blacksnake, redbird, wildcat.

b. An adjective with the name of some characteristic feature

of the animal; as, longsting, blackfin, bluetail (lizard),

yellowlegs, bighorn.

I feel safe in affirming that the foregoing rules are so simple

that anybody can apply them; that they are sufficiently compre-

hensive to include all names of the groups indicated; and that

they are in accord with reason, language-principles, and the

usages of the highest authorities! I admit that the rank and

file of investigators in any particular department of science may

vary widely in usage from these rules, or any other set of rules

that might be formulated.

All the vernacular names can not be found in any one diction-

! As representative of the ‘highest authorities’ on this subject I would include

the Standard Dictionary, the Century Dictionary, Murray’s New English Diction-

ary, and such special works as Newton’s Dictionary of Birds, Webster's ‘ Inter-

national" and other older dictionaries will not always be found in accord with the

authorities just named in compounding words, nor with these rules.

No. 440.] VERNACULAR NAMES. 555

ary, nor in all combined, perhaps ; neither do these authorities

cited always agree in compounding words, just as they do not

always agree in spelling and pronunciation ; but they are the

best reflectors of good usage we have. In fact, the chief func-

tion of the dictionary is to exhibit the usages of the best speakers

and writers.

There is one phase of the question on which it is somewhat

difficult to secure uniformity. (Compound words are divided

into two classes: (1) separable compounds, written with the

hyphen, as ‘tent-caterpillar’; and (2) solidified compounds,

written without the hyphen, as ‘ladybird.’ It is not easy to

formulate rules for determining just when the hyphen may be

omitted from a compound, and the word written solid. Gen-

erally the old and familiar forms are solidified, while the newer

and unfamiliar names retain the hyphen; but age and frequency

of use alone can not determine. However, this is not so impor-

tant as some other phases of the subject. "Words are compound

whether hyphened or solid, and the choice of forms depends more

upon taste, since no fundamental language-principle is at stake.

But we should strive for uniformity in this respect also. The

limits of this paper will not admit a fuller discussion of this

feature.

There should be some concerted attempt to secure uniformity

in writing the vernacular names of animals. The principles and

rules which govern in these matters are just as binding with

regard to the hyphen as with regard to capital letters, punctua-

tion, or the construction of sentences. There is no more reason

for a writer's ignoring the correct use of the hyphen than for his

ignoring any other essential to good orthography or correct

syntax.

CHAMPAIGN, ILLINOIS.

NOTES AND LITERATURE

ZOOLOGY.

" American Food and Game Fishes”? is the title of an admirable

volume of about 600 pages, written by Dr. D. S. Jordan, President of

Leland Stanford University and Dr. B. W. Evermann, Ichthyologist

of the U. S. Fish Commission.

This book is the only attempt yet made to give an account of all

the species of American fishes north of Panama used by man as

food or sought by anglers for the sport which their capture affords.

About rooo species of fish are described, nearly half of which are

illustrated by colored plates, half-tones and text figures. The text

figures are excellent reproductions from drawings which were origin-

ally used to illustrate the species in more technical papers, and are

as exact as it is possible to make them. The same is true of the

colored plates. The half-tones, of which there are more than 100,

are made from photographs of live fish in the water. They were

made expressly for this work at Key West, Lake Maxinkuckee, and

the Pan American Exposition at Buffalo. The artistic skill displayed

in making these photographs has certainly not been excelled in the

line of animal photography. The half-tone reproduction of these

photographs is seldom equalled.

The fact that the book is written by two of our most active stu-

dents of American fishes is a guarantee of its excellence. The

important feature of the entire book is its accuracy. The descrip-

tions of the different species, their life histories and geographic

range are correctly as well as interestingly give

It is the only book that gives the sited fisherman accurate

and detailed information about all American fishes which find their

way into our markets. As a literary and scientific production one

cannot praise this book too highly. It is certainly the work of

masters. Zodlogists will no doubt regret that room was not found

! Jordan, D. S. and Evermann, B. W. American Food and Game Fishes. A

Popular Account of all the Species found in America North of the Equator,

with Keys for ready Identification, Life Histories, a Methods of Capture.

New York, Doubleday, Page & Co., 1902. 8vo, 1+ 573 pp., plates and text

figures.

“I

558 THE AMERICAN NATURALIST. [Vor. XXXVII.

in the book for an account of the gar pike and the dogfish or Amia

because of their special interest in these fishes. They are, however,

neither food nor game fishes, and are so well known in special works

in zodlogy that ready reference to them can easily be had elsewhere.

While this work will not fail to interest the general reader, it will

be most highly appreciated by those who really wish to know fishes.

To the students of natural history in our high schools and colleges

the book will be especially useful, as it is a strictly up-to-date syste-

matic treatise and contains a vast amount of accurate natural history

information. By means of keys, descriptions and the illustrations

any student of ordinary intelligence can easily identify all of our

food and game fishes which he may have.

To the angler it is indispensable, for it tells him where the different

game fishes are to be found, and will enable him to readily identify

any fish he may catch, and to know it by its correct scientific as well

as its common name.

S. E. MEEK.

Notes on Recent Fish Literature.— In connection with the

elaborate investigation of the fishes and fisheries of the Hawaiian

Islands, undertaken by the United States Fish Commission in 1901

and r9o2, Messrs. Jordan and Evermann, who have the work in

charge, have published accounts of part of the new species

obtained. In this preliminary paper (Bull. U. S. Fish. Comm. for

1902) 56 new species are described. "These will be figured with all

the known species in a final report. The Hawaiian Fauna is much

less rich in the species than that of Samoa, and while the Genera are

all distinctly tropical, the species are very largely different from those

found in Polynesia and Micronesia. This isolation of the Hawaiian

Fauna is in part explained by the direction of the ocean currents,

which set to the eastward in Samoa and Tahiti, but to the westward

about Hawaii. The new Genera in this paper are the following:

Fowleria (Apogonidz), Bowersia (Lutianide ), Verriculus (Labridz)

Quisquilius and Vitraria (Gobiida), and Osurus (Pteroprarida).

In the Annals and Magazine of Natural History (Vol. II, No. 63),

Mr. C. Tate Ragan of the British Museum gives a review of the

angler fishes of the family of Laphiide. Three genera are recog-

nized: Chirolophius (ares), Lopheonius and Lophius. ZopAeonius

spelurus Garman, from off Panama, is referred to Chirolophius. The

species described by the present writer from Japan as Lopheonius

litulon, is as Ragan suggests, a true Lophius, having 27 vertebra,

No. 440] NOTES AND LITERATURE. 559

while the other Japanese species, externally extremely similar,

Lopheonius setigerus has but 17 or 18. The former is a species of

northern distribution, It is remarkable that so great a difference in

the vertebral column should be developed without any considerable

external distinction. About 15 species of anglers are now known.

Dr. Tarleton H. Bean has published a useful account of the habits

and distribution of the fishes found on Long Island. It occurs in

the Annual Report of the Forest, Fish and Game Commission of New

York, 1901.

A new writer on Ichthyology, Senhor Alipio de Miranda Ribeiro,

gives an account of eight fishes from Rio Pomba in a report of the

Sociedade Nacional de Agricultura for 1902. For the genus usually

called Centropomus, Ribeiro uses the name Platycephalus, “in obser-

vance of the laws of priority."

Mr. Cloudsley Rutter, naturalist of the “ Albatross,” describes the

fishes obtained by him in the lakes of North Eastern California

(Bull. U. S. Fish Commission for 1992). Most of these lakes are

without outlet, although formerly tributary to the post-glacial Lake

Lahontau. The new species are Pautorteus lahontau from Susan

River, Chasmistes chamberlaini from Eagle Lake, and Agosia robusta

from Susan River and other streams.

Mr. Rutter also gives a report (Bull. U. S. Fish Commission for

1902), on five years’ work (1896-1901) of observation on the habits

of the Sacramento salmon. It is a very complete and valuable piece

of work. Among the important conclusions are these :—

There is no evidence that salmon returning to spawn seek their

native stream. Probably most salmon do not wander far into the

sea, and the native river is the nearest one when the spawning season

comes. There is no evidence that they remember any particular

branch of the river basin in which they were spawned. Most salmon

return to spawn aftertwo years in the sea, some remaining three and

others four years.

All die after spawning, none making any effort to return to the

sea. Most of them die within 5 to 16 days after reaching the

spawning grounds. They will not survive if placed in salt water.

The Pacific salmon are like cast off leaves of a tree, when their

period of usefulness to the species is past.

In the Proceedings of the United States National Museum (Vol.

XXVI) Messrs. Jordan and Fowler continue their discussions of

560 THE AMERICAN NATURALIST. (Vor. XXXVII.

Japanese fishes. The Elasmobranchs, or sharks, rays and chimeras

are represented by 56 species. Other papers are on the Cobitidz or

Loaches, six species in Japan, and on the Cepolidz or Band-fishes,

of which Japan has three. The genera, Embolichthys and Zen are

subjects of a special paper.

In the Annotationes Zoologice Faponenses (Vol. IV) of the Imperial

University of Tokyo, Dr. Bashford Dean gives an account of the

cleavage of the egy in the cestraciont shark Heterodontus japoni-

cus. He finds on the egg certain marks or lines reminiscent of

holoblastic cleavage.

Gardiner's *« Maldive and Laccadive Archipelagoes,’’ Part IV.'—

The fourth part of Gardiner's Fauna and Geography contains

seven papers including a detailed description, with charts, of the

Atolls and Banks— a valuable contribution to geography but not

abstractable. In his concluding notes Gardiner touches on the

causes of deaths of parts of the coral reefs. Silting up is destructive

and senile decay, after the colony has reached a great size, causes

great mortality.

The Cephalochorda are described systematically and anatomically

by C. F. Cooper while R. C. Punnett considers their variation. A

new species, ZZe/eropleuron maldivense is described. In the conclu-

sions as to the great variability of Cephalochorda based on the num-

ber of myotomes the possibility of an increase in the number of

myotomes throughout life is not sufficiently considered.

The Avifauna is analyzed by Gadow. Twenty-six species were

examined; none peculiar to the islands. The permanent residents

are, excepting the Indian crow, Corvus splendens, all water birds,

mostly of wide distribution in the Old World. Eight genera of birds

are winter visitors from the Asiatic continent and a few species are

wanderers from India and Ceylon. Finches, starlings and pigeons

are wholly absent. At one point in the Archipelago it was observed

that all birds retired daily from 11 a. M. to 3 P. M.

The earthworms are reported upon by Beddard. He comments on

the favorable material afforded by this group for studies in geographic

variation owing to impracticability of their unassisted migration over

a tract of sea. Three species are recorded, two are very common

! The Fauna and Geography of the Maldive and Laccadive Menon ing etc.

Edited by J. Stanley Gardiner. Cambridge Bree Press, 1903. i, pt.

iv, pp. xix 4- 348-47 1, pls. 18-25, text figs. 76-11

No. 440.] NOTES AND LITERATURE. 561

and widespread oriental species; the third is a new Pontodilus — a

seashore inhabitant.

The classification of crabs is undertaken by Borradaile, who

thereupon describes the crabs from the Archipelagoes belonging to

the Catometope and Oxystomata. As before, especial stress is laid on

bionomics and adaptations. A collection of 16 species of Barnacles

is also described. Most of them are Indo-Pacific species, but two

occur in the West Indes. Lanchester’s study of the Stomatopods,

based as it is on the study of individual variation will be of interest

and, it is hoped, of instruction to the species splitter. Excepting two

specimens of Pseudosquilla ciliata all the adults belong to the world-

wide Gonodactylus chiragra, including seven synonyms. Gonodac-

tylus lives on the surface of reefs and is abundant here. Squilla,

which demands mud to burrow in, is absent; because the mud is.

This is another illustration of the law that the habitat of a species is

determined by its instincts. The author introduces “term” to

express the extremes of structural type in a continuous variation.

Finally the Lithothamnia are described by M. Foslie, who combats

the erroneous notion that these Algz are more abundant in tropical

than in northern seas.

C.N I

Webster's ** Diffusion of Insects in North America."— In the

April number of Psyche we have a very interesting article on the

above subject, from the pen of Professor F. M. Webster, who has

already contributed various papers of the same general nature to our

entomological journals.

The problem of the nature and extent of destruction of life during

the Glacial epoch is but barely touched upon, the intention of the

author being to show how post-glacial distribution has been accom-

plished. He points out that with the retreat of the ice three gate-

ways for the introduction of species into this continent were open —

(1) the Alaskan chain for Asiatic insects, (2) Central America for

forms from South America, and (3) Florida, by way of the West

Indies as an alternative to the Central American route. All new

life depending to any large extent upon land for its introduction

must come by some one of these three roads, the agency of man in

the matter being of so recent an origin as to need separate consider-

ation.

The northwestern sstetingyi leading from Asia, seems to have been

taken advantage of by numerous Coccinellidz and certain Chrysome-

562 THE AMERICAN NATURALIST. [Vor. XXXVII.

lide. Some of these have a much more extended range in this con-

tinent than in the Old World, Zina lapponica for example being found

as an European species only in the extreme north, while with us it

reaches south as far as Texas. ‘This adaptability is a potent factor

in distribution and the readiness with which a given species assimi-

lates with new surroundings has a powerful bearing upon its final

geographical range. Insects coming to us from Asia by the path

mentioned may spread to the south through the great valleys lying

between the Rocky Mountains and the Cascades or by use of the

passes in the former range gain the great plains of Canada and

eventually appear on the Atlantic coast. If their nature is such as

to enable them to bear an arid or warm climate they may reach

points far to the south since there are no great natural barriers in

the way.

The southwestern gateway is, in Professor Webster's opinion, by

far the most important since through it we receive accessions from

the rich fauna of Central and South America. Asa striking example

of a faüna received through this medium he cites Halisidota, a genus

of moths apparently South American in type. Some of our forms

are supposed to have had their origin in the South and to have come

to us by way of Central America. A certain species with a present

range from Argentina to Costa Rica is thought to be the stem of our

HË. tesselaris and H. citripes, which seem to have reached us by two

different courses — the one coming north by way of Texas, the

other east by way of Yucatan and Cuba. The distribution of Pacific

and northern Halisidotae indicate to the author the probability of

certain species having reached (through stem forms) the New Eng-

land coast by following the Pacific Maritime trend from Central

America through California, Oregon, Washington and British

Columbia, thence traveling eastward along the lines indicated for

the immigrants of Asiatic origin. ‘The genus Diabrotica, among the

beetles, warrants the belief that offshoots of a common Central

American stem may become separated far to the south, one going to

the Pacific district while its fellow inhabits the Atlantic slope.

But little matter concerning the antillean trend is presented, this

being thought of less importance than the others, though many

insects appear to have come in that way.

Concerning the route taken by insects introduced through the

eastern ports by agency of man, Professor Webster has ascertained

that they follow a path which he calls the transappalachian trend —

reaching the fertile prairies of the interior through the great gap in

New York which forms the principal thoroughfare for insect invasion.

No. 440] NOTES AND LITERATURE 563

A glance at the map with which the paper is illustrated suggests

the thought that we have in Professor Webster's article a strong

defense of the Wallacean theory of distribution and of the important

part played by barriers in determining the spread of animallife; it is

to be regretted that the recent propaganda of the theory of isothermal

distribution of organisms, while undoubtedly of great value in indicat-

ing the possibility of artificial cultivation, has tended to obscure the

importance of geographical features under really natural conditions. |

H. F. WICKHAM.

BOTANY.

Livingston’s Osmotic Pressure and Diffusion in Plants.! — The

author begins with a treatment of the purely physical phenomena con-

nected with diffusion and osmosis, rightly holding that it is difficult

for the student of physiology to easily obtain the information he needs

in compact form. Such a treatment covering, as it does but forty-

four pages could not possibly be detailed, nor does the author main-

tain that itis. Nevertheless a summary of this kind can be exceed-

ingly useful, if in no other way than to stimulate the student to

further reading. This résumé is clear and sufficiently full to give an

adequate conception of the theories concerned. There are six chap-

ters in the first part, which treat of the fundamental theories of the

nature of matter, of diffusion and diffusion tension, of solutions and

ionization, of osmotic phenomena and the measurement thereof.

In the second part on the physiological aspect of the matter, the

author continues his summarization of the work which has been

done, but of course in far greater detail than in the previous half,

since the field is smaller and since this is the real object of the book.

In the first chapter the question of turgidity is taken up. The impor-

tance of this subject demands full treatment, and forty-two pages

are devoted to it; the author does not develop anything new, how-

ever. In the even more complicated, and certainly more dubious,

matter of the absorption and transmission of water, which is consid-

! Livingston, Burton E. Zhe Role of Diffusion and Osmotic Pressure in Plants.

The Decennial Publications of the University of Chicago. Second Series, vol.

7. The Univ. of Chicago Press, 1903. 8vo, pp. i-xiii, 1-149.

564 THE AMERICAN NATURALIS1. [Vor. XXXVII.

ered in the second chapter, there is hardly space enough allowed for

a very complete discussion of the question. The optimistic point of

view is taken that when the nature of the plasmic membranes is

known, *it is not improbable that the solution of the problem of

water transport will follow as the simplest corollary.” In the next

chapter the absorption and transmission of solutes is treated of, the

important matter of quantitative selection is somewhat lightly referred

to, and one is lead to the belief that it is either taken as a matter of

course, or that the author does not consider it of the same importance

that many physiologists do. Simple diffusion is said to be the great-

est factor in the distribution of solutes in the plant body. As to the

influence of the osmotic pressure of the surrounding medium upon

organisms, which is the subject of the final chapter, the author gives

a summary of his own work in that line and of that of other investi-

gators. Investigations have shown that growth is accelerated in weak

solutions of various substances and retarded in concentrated ones.

Cell division may also be influenced by the osmotic pressure of the

surrounding medium, and reproduction being a peculiar form of cell

division is apparently dependent, in some cases, upon the pressure.

Whether the effect is due merely to the extraction of water, or to a

strictly chemical influence the author does not decide.

While there is not a great deal that is new in the volume it is an

excellent brief review of the various questions as they stand to-day.

As a reading book for more advanced students in physiological

botany the book will have considerable value, and it is one which

may well be placed upon the shelves of the laboratory reference

library. As a final source of information it cannot of course be sat-

isfying, but the copious references to the literature make it valuable

in this respect also. "There is a distinct place for a book of this

character.

Pierce's Plant Physiology.!

his desire to fill the gap in text books which exists between the mon-

umental work of Pfeffer on the one hand, and the clever but short

account of plant physiology which is found in Stasburger's text-book.

The intention to furnish a good reading book of this character is cer-

tainly a laudable one, and in so far as the author has succeeded, he

is to be congratulated. It is further stated that only “safe views”

! Pierce, G. J. A TZeat-Book lii Plant Physiology.

New York, Henry Holt &

o. 1903. 8vo., vi-291 pp., 22 figs

No. 440.] NOTES AND LITERATURE. 565

of the various physiological processes will be given, and such a posi-

tion may also be regarded as an excellent one. ‘Too great conserva-

tism, however, may easily lead to two serious difficulties, one is that

the style may readily become unsuggestive and the other is that these

very same “safe views” may become almost dogmatic. After an

introductory chapter on the general problem of physiology the ques-

tion of respiration is at once entered into and with it the correlated

phenomena of fermentation. There may be some who would be

inclined to doubt the entire wisdom of at once plunging into these

complicated matters, before the student has been informed of what is

known as to how or where the materials concerned in respiration are

produced. This is especially true of the subject of enzyme action,

and as a consequence the handling of this important topic is hardly

satisfactory.

The space devoted to it is chiefly taken up with a consideration of

yeast fermentation, and the great classes of intra-cellular enzymes

are scarcely mentioned. The third chapter is on Nutrition and in

the next, absorption of water and food is treated of. Here for the

first time the fundamental question of osmosis is explained. The

different, necessary chemical elements, are taken up in turn, In

passing it may be mentioned that the literature quoted does not

always include the latest contributions to the subject, by the authors

named. The fifth chapter is devoted to the consideration of the

primal phenomena of growth, and following it is a long one, entitled

Irritability, under which head all growth responses, as well as the

movements of nature organs, are taken up. As an example of

unfortunately dogmatic statements may be mentioned the following,

which is given as an explanation of phototropism: “The cells on the

side of the stem away from the window receive less light and are

less checked than those on the opposite side, and hence push the

tip of the stem over towards the window.” Such a definite explana-

tion, on the basis of etiolation, as the cause of phototropic curvature

would not be accepted by many physiologists, and is perhaps too

“safe” a view to take of this perplexing response. Attention may

' also be called to the fact that the familiar term, etiolation is not used

at all by the author, and that the term heliotropism is preferred to

the generally admitted better one, phototropism. The last chapter

deals in the compass of thirty pages with the subject of reproduction,

including a three and one half page consideration of the problems

connected with heredity. The index leaves something to be desired,

not infrequently one must look in vain for references to familiar

terms, such as hyponasty, epinasty, etiolation, etc.

566 - THE AMERICAN NATURALIST. [Vor. XXXVII.

It is perhaps unfair to the book to have picked out such passages,

where the handling of the subject is not in accord with the ideas of

the reviewer although many more might be cited. In the main it is

a careful and conservative — almost too conservative — treatment of

the subject of plant physiology. According to one's point of view, its

faults or its virtues lie in the very definite, perhaps non-stimulating

style, in which it is written. It will no doubt prove a useful addi-

tion to the somewhat slim stock of reading books in plant physiology

which are at the disposal of the English reading student.

M. R.

Notes.— Zhe American Botanist for April contains the following

articles: — Bailey, “ Violets”; Bradshaw, “The Chilicothe Vine ”

Blight, “What is American Weed?”; Fetherolf, “Among Texas

Ferns”; and Steele, “Species or Varieties ? "

Part IX of Hough’s American Woods, published at Lowville, N. Y.,

comprising nos. 201 to 225 inclusive of his admirable sets of radial,

tangential, and cross sections of each species, is devoted to Pacific

Coast species, and is accompanied by a text brochure including, in

addition to an account of each of the species represented in this part,

leaf and fruit keys to the entire series thus far issued.

The Botanical Gazette for April contains the following articles : —

Davis, * Oogenesis in Saprolegnia"; Mottier, “ The behavior of the

chromosomes in the spore-mother-cells of higher plants and the

homology of the pollen and embryo-sac mother cells"; Hitchcock,

* Notes on North American grasses — III, New species of Will-

kommia," and Bower, “ The morphology of spore producing members.”

The Bryologist, for May, contains the following articles : — Grout,

“Some interesting forms of Polytrichum”; E. G. Britton, * Notes on

nomenclature — II”: Bailey, “An interesting tree”; Grout, “Sun

prints in bryology — additional notes”; Holzinger, “Obituary, M

Emile Bescherelle,” and “ Se/igeria tristichoides in southern France”

and Williams, “ Oedipodium Griffithianum.”

The Bulletin of the Torrey Botanical Club for April contains the

following articles: — Goebel, “ Regeneration in plants”; Morgan,

“ The hypothesis of formative stuffs”; Howe and Underwood, * The

genus Riella, with descriptions of new species from North America

and the Canary Islands”; Murrill, “The Polyporaceae of North

America — III, the genus Fomes”; Piper, “Four new species of

grasses from Washington”; Osterhout, “New plants from Colo-

No. 440] NOTES AND LITERATURE. 567

rado”; Eastwood, “New species of Oreocarya”; and Rydberg,

“ Studies on the Rocky Mountain flora — X."

The Bulletin for the Torrey Botanical Club, for May, contains the

following articles: — Rydberg, “ Some generic segregations”; Harper,

“ Botanical explorations in Georgia during the summer of 1901 — 1”;

Murrill, “The Polyporaceae of North America — IV, the genus

Elfvingia”; Kunzé, *'The desert flora of Phoenix, Arizona”; and

Sheldon, ‘‘ New species from the Pacific Coast — I.”

The Fern Bulletin for January, with portrait of J. A. Graves as

frontispiece, contains the following articles: — Clute and Cocks,

“The fern flora of Louisiana"; Waters, “My indoor fernery "

Eaton, * The genus Equisetum in North America, XII, the sub-genus

Hippochaetae”; Clute, * Fernwort notes — I"; Buchheister, “ Notes

from the Catskills”; Cocks, “ Eguisetum robustum" ; Flett, “ Varia-

tions in the habitat ut two ferns ”; Clute, Cultivation of our hardy

ferns”; and Anthony, “ Notes on the ferns of Florida, East Coast.”

The Fern Bulletin for April, with portrait of R. R. Scott as frontis-

piece, contains the following articles: — Reverchon, * The Fern flora

of Texas"; Maxon, “Notes on American Ferns — VI”; Eaton,

“The genus Equisetum in North America — XIII: Æ. Zevigatum;

Osmun, L£guisetum scirpoides in Connecticut "; Clute, * Fernwort

notes — II"; Eaton, “ SE Nephrolepis from spores"; Druery,

* New forms a Ferns”; and Clute, “List of Fernworts collected in

Jamaica — conclusion.”

Under the title Aora and Sylva, Mr. Robinson, an enthusiastic

student and cultivator of plants, has begun the publication of a new

monthly journal, beautifully illustrated. The editorial offices are at

63 Lincoln’s Inn Fields, London. ‘The first number contains exqui-

sitely colored plates illustrating species of Magnolia and Calochortus.

The Journal of the New York Botanical Garden for May contains,

among other things, reports on a trip to eastern Cuba; the Jenman

collection of ferns; and a large collection of conifers for the pinetum

of the institution.

Like earlier volumes, the fiftieth volume of Proceedings of the

American Pharmaceutical Association contains a considerable number

of scientific papers dealing with the structure and particularly active

principles of plants which furnish medicinal products.

568 THE AMERICAN NATURALIST. [Vor. XXXVII.

Rhodora, for May, contains the following articles : — Collins,

“ Lorin Lowe Dame”; Blankinship, “ Plant formations of eastern

Massachusetts ” ; Sargent, “ Recently recognized species of Crataegus

— III”; Bissell, “ Ajuga Genevensis in New England”; Fernald,

* Red-tlowered Anemone riparia”; and Robinson, “ Viola arvensis

in New England.”

Torreya, for April, contains the following articles: — House,

* Notes upon the orchids of central New York”; Ross, “ Vagaries

of Hepatica”; Earle, “ Key to the North American species of Len-

tinus —II”; and King, “Explosive discharge of antherozoids in

Conocephalum,”

Torreya, for May, contains the following : — Harshberger, “ Notes

on the strand flora of Great Inagua, Haiti and Jamaica”; Curtis,

* Observations on etiolation ?; and Berry, “A triple samara in Acer

rubrum.”

Part 2 of Trees and Shrubs, issued from the Riverside Press of

Cambridge, like its predecessor contains illustrations and descriptions

of a number of species of the now popular genus Crataegus, as well

as of other genera of interest.

In the Bulletin de / Herbier Boissier, Dr. Hallier has recently pub-

lished a preliminary scheme for a new phylogenetic classification of

the flowering plants. Dr. Engler, in the new edition of his Sy//adbus,

also considerably modifies his previous treatment of the larger groups.

A practical application of the various ways of treating botanical

nomenclature is given by Hitchcock, apropos of Festuca spicata,

Pursh, in Science, for May 22, 1903.

An important, though not lengthy, paper by Van Tieghem, on the

structure of the stamen in Scrophulariaceae, is contained in No. 8 of

the Bulletin du Muséum d'Histoire Naturelle, for 1902.

The Morphological propriety of designating stamens and pistils

sexual organs, is discussed by Ganong in Science of April 24, and

MacMillan in the same journal for May 15.

Cotyledonary studies of Pinus maritima, by Chauveaud, are pub-

lished in No. 7 of the Bulletin du Muséum a’ Histoire Naturelle, for

1902.

Ramaley has a note on the cotyledons and leaves of certain

Papilionacez, in No. 3 of Zhe University of Colorado Studies.

No 440.] NOTES AND LITERATURE. 569

The Botanische Zeitung of April 16 is occupied by an analysis of

recent hybridization literature, by Correns, who also contributes two

papers on the same subject to the Berichte der Deutschen Botanischen

Geselischaft of April 23.

Professor Fernow has an article on “applied ecology” in Science’ of

April 17.

“New Hampshire Wildflowers” is the title of Mature Study

Leaflet No. 4 of the New Hampshire College Experiment Station, by

Professor Weed.

An account of a biological reconnoissance in the vicinity of Flat-

head Lake, by M. J. Elrod, is published as No. io of the Bulletin of

the University of Montana,— No. 3 of the biological series.

Professor Ganong contributes a preliminary synopsis of the group-

ing of the vegetation (phytogeography) of the Province of New

Brunswick to No. XXI of the Bulletin of the Natural History Society

of New Brunswick.

A list of plants from Labrador, by Mackay, is contained in the

Proceedings and Transactions of the Nova Scotian Institute of Science,

Vol. X, part 4, issued in March.

An account of the vegetation of one of the Amazon districts, by

Huber, is published in the December number of the Boletim do

Museu Paraense, and includes 189 entries. The article is followed

by an account by the same author of the physical geography of the

region, the “Furos de Breves" or communications between the

Amazon and the Para.

With the third fascicle, issued under date of January, 19o3, De

Wildeman’s Études sur la flore du Katanga, constituting “Series IV

— Botanique " of the Annales du Musée du Congo, is completed. The

work forms a folio volume of xii + 241 pages, and 46 plates.

A monograph of Cardamine, by Schulz, has recently been published

in Engler's Botanische Jahrbücher.

Under the title “Leguminose Langlasseane” the Société de

Physique et d' Histoire Naturelle de Genéve has published, as Volume

XXXIV. Fascicle 3, of its Mémoires, a beautifully illustrated account

of certain Leguminosz collected in 1898 and 1899 by Langlassé

through the Mexican states of Michoacan and Guerrero. The man-

570 THE AMERICAN NATURALIST. [Voi XXXVII.

uscript was prepared, shortly before his death, by Micheli, whose

portrait forms a frontispiece to the paper.

A short note on New Brunswick violets, by Vroom, is printed in no.

XXI of the Bulletin of the Natural History Society of New Brunswick.

A critical revision of the genus Eucalyptus, by J. H. Maiden,

Government Botanist of New South Wales, is in course of publica-

tion at Sydney. The first part, comprising 47 pages of text and 4

plates, bears date of January, 1903, at end of the preface.

* A research on the Eucalypts, especially in regard to their essen-

tial oils,” by Baker and Smith, is published from the Zechnological

Museum of New South Wales, as No. 13 of the Technical Education

Series of its publications.

A revision of Sebaea, § Eusebaea, by Schinz, is distributed from

the Mittheilungen der geographischen Gesellschaft in Lübeck, Heft 17,

1903.

An analysis of the relationships of the vernal group of Primula,

comprising the oxlip, cowslip and primrose of England, by Bailey,

is printed in ZZe Journal of Botany for May.

Betula papyrifera is figured in Vol. IV, no. 1, of the cones Selecte

Horti Thenensis, published by M. Leon van den Bossche of Tirlemont,

Belgium ; — one of the most accurate and attractive of current works.

devoted to the illustration of the higher plants.

A catalogue of the Mosses of Australia and Tasmania, by White-

legge has been started, the first part appearing as a supplement to

no. 107 of the Proceedings of the Linnean Society of New South Wales.

Some points in the structure and life history of diatoms are dis-

cussed by F. R. Rowley in an illustrated paper in Zhe Journal of the

Quekett Microscopical Club for April.

An exhaustive and beautifully illustrated account of “blueing ” and

“red rot” of the western yellow pine,— caused respectively by

Ceratostomella pilifera and Polyporus ponderosus, is published by

von Schrenk as Bulletin no. 36 of the Bureau of Plant Industry of

the U. S. Department of Agriculture.

The synonymy of Gloecsporium fructigenum is discussed by von

Schrenk and Spaulding in Science for May 8.

No. 440.] NOTES AND LITERATURE. 571

An illustrated account of the coffee disease caused by Stilbella

Jfavida, is published by Kohl in the Beihefte zum Tropenpflanzer for

May.

A destructive apple rot, caused by Cephalothecium roseum following

the attacks of Fusicladium, is described by Eustace in Buletin of the

New York Agricultural Experiment Station No. 227, and the preced-

ing Bulletin of the same station deals with the obscure cane blight

and yellows of the raspberry.

What appears to be an important study of the cancer of trees,

referred to a bacterial cause, by Brzezinski, is contained in the

March number of the Bulletin International de P Académie des sciences

de Cracovie, classe des sciences mathématiques et naturelles.

Dr. G. U. Hay, in No. XXI of the Bulletin of the Natural History

Society of New Brunswick, records over 180 species of higher fungi

for that Province, in addition to an earlier list which is reprinted.

An important paper on the Algz of northwestern America, illus-

trated by eleven plates, is published by Setchell and Gardner in Vol.

I. of the University of California Publications — Botany, issued on

March 31.

The Hokkaido governmental fishery bureau is publishing in

Japanese a well illustrated series of reports on the marine resources

of Hokkaido, the third of which deals with the Laminariacez and

Laminaria industries.

Under the title “ The Fower Beautiful,” Professor Weed has writ-

ten an attractive and daintily illustrated little book on the decorative

use of flowers, published by Houghton, Mifflin & Co., Boston.

An article on the forest policy of Pennsylvania, by G. W. Wirt, is

published in the Journal of the Franklin Institute for May.

Some useful shade-tree suggestions, for Wyoming, are published

by Aven Nelson as Bulletin of the Agricultural Experiment Station,

no. 57, of that state.

An economic account of Manihot, by Tracy, forms Farmers’ Bulle-

tin of the U. S. Department of Agriculture, no. 167.

An account of olives and olive oil in France, by R. P. Skinner, is

published as no. 1639 of Advance Sheets of Consular Reports, dated

May 6

572 THE AMERICAN NATURALIST. ([Vor. XXXVII.

The Kumquat and Mandarin groups of Citrus are discussed by

Hume in Bulletin of the Florida Agricultural Experiment Station,

Nos. 65 and 66.

Some conditions of stock poisoning in Idaho are discussed by

Professor Slade in Bulletin of the Experiment Station, No. 37, of that

state.

The poisoning of cattle by Sorghum vulgare is considered by Peters,

Slade and Avery in Bulletin of the Agricultural Experiment Station of

Nebraska, No. 77.

The first volume of “ Datos para la materia médica argentina,” by

J. A. Dominguez, has been issued by the Museo de Farmalogia of

Buenos Aires.

The value of making an herbarium is the subject of an article by

Cameron in School Science for May.

“More Letters of Charles Darwin,” published by the Appletons

(New York) in two volumes, contain a number of letters of interest

to botanists.

CORRESPONDENCE.

To the Editor of the American Naturalist:

SIR:—A little over a year ago Dr. William Patten’ claimed to

have discovered evidence of * numerous pairs of jointed oarlike

appendages " in Tremataspis and related forms, in none of which

had similar indications been previously detected. Nor, for that

matter, had anyone previously had the temerity to attribute more

than two pairs of limbs to any vertebrated creature, living or

fossil. Beyond Dr. Patten's assertion that Cephalaspis is pro-

vided with “a fringe of jointed and movable appendages (25-

30 pairs) along the ventral margin of the trunk," we have had

from him no further enlightenment as to the ambulatory equip-

ment of this Ostracophore, but there has recently been heard

from Dr. Gaskell? an absolute denial of the existence of seg-

mental appendages in that genus.

Now, since it appears that segmented appendages are wanting

not only in Cephalaspis, but universally amongst the Heteros-

traci and Osteostraci so far as known, the presumption is certainly

very strong that they do not occur in Tremataspis, whose close

relationship with Cephalaspis none will question. Professor

Patten, however, in returning to the matter de novo,3 combats

against this presumption, and affirms his belief in the existence

of not only one, but possibly several pairs of jointed appendages

in Tremataspis. Naturally he must have strong reasons for

inclining him to so heterodox a notion, and what are they,

palzeontologists are curious to know? Has he discovered speci-

mens showing one or more pairs of these chimzrical organs

10n the Structure and Classification of the Tremataspide. Amer Nat., vol.

36. 1902, pp- 379-393 :

? On the Origin of Vertebrates, Journ. Anat. and Phys., vol. 37. 1903, p. 198:

text fig. 5.

3 On the Appendages of Tremataspis. Amer. Nat., vol. 37. 1903, pp. 223-

42.

2731

3423

574 THE AMERICAN NATURALIST. [Vor. XXXVII.

in situ? Or has he found detached appendages positively deter-

minable as belonging to Tremataspis, to the exclusion of all

other accompanying fossil remains? Professor Patten answers

the first of these interrogatories in the negative, the second

affirmatively. He has obtained in all four detached plates of

small size and mediocre preservation, which he regards as

portions of as many “paired cephalic appendages” ; these he

figures of ten times the natural size and describes in praiseworthy

detail. But by what process of reasoning he is able to identify

them as belonging indubitably to the genus Tremataspis he does

not take the trouble to state, leaving the reader to take it for

granted that his determination is correct.

Not all readers, however, will be prepared to accept a deter-

mination so utterly at variance with analogy. On the contrary,

rational students will maintain that inasmuch as certain fish

fragments are identifiable as parts of jointed oarlike appendages,

that fact is príma facte evidence of their pertaining not to Trem-

ataspis, but to an entirely different order of Ostracophores —

the same to which Pterichthys, Bothriolepis and Asterolepis

belong. This indeed was the view taken by Pander as early as

1856, who, so far from associating certain fragmentary append-

ages from the Baltic Silurian with Tremataspis, referred them to

the Pterichthyid order, whose presence in those beds is not

otherwise indicated. This procedure is entirely justified by the

fact that structures of this nature are known to be present in the

group typified by Pterichthys, but not in that to which Trema-

taspis and Cephalaspis belong. The danger of a reliance on

merely superficial characters for the determination or theoretical

association of detached fragments is well illustrated by Cope's

confusion of an appendage of Bothriolepis with Holonema

remains,' certainly no trifling error, and other instances are but

all too common. To cite an analogous case borrowed from

invertebrate paleontology, it is the same as if Professor Patten

had found parts of a Eurypterus- or Pterygotus-like swimming

appendage, and was misled by superficial resemblances into

supposing it to have belonged to some genus of Trilobites

! On the Characters of some Paleozoic Fishes. Proc. U. S. Nat. Mus., vol.

14. 1891, p. 456, pl. XXX. fig. 7.

No. 440.] CORRESPONDENCE. 575

occurring in the identical formation, and exhibiting similar tex-

ture or surface markings. We wonder how many would be con-

verted to this belief until a Trilobite had first been discovered

with the actual members preserved zz situ?

In precisely the same way we must reject Professor Patten's

theoretical association of parts as erroneous, and deny that forms

like Tremataspis, Cyathaspis, Cephalaspis, Tolypelepis, etc.,

possessed paired segmented appendages until at least one single

individual shall have come to light having the postulated organs

Fic. 1. Ventral aspect of the head-shield of Tremataspis (from Patten).

preserved zz situ. We can also afford to await the discovery of

a six- or eight-limbed vertebrate before abandoning a rather deep-

seated prejudice in favor of a maximum number of two pairs of

limbs

Professor Patten calls attention to the marginal openings on

the visceral side of the head-shield in Tremataspis, arguing that

their ventral position furnishes strong reason for believing that

they served for the attachment of appendages. But this argu-

ment, such as it is, is negatived by the fact that a row of

576 THE AMERICAN NATURALIST. (VOL. XXXVII.

precisely similar openings occurs in Birkenia on either side of

the body at its junction with the head, — that is to say, in a

position where they would zo? be serviceable for the attachment

of such organs. These openings are commonly regarded as

branchial in function, in which case it is easy to conceive of them

as having been diaposed in the two genera relatively as in the

skate and shark.! It is the veri-

est flight of fancy to suppose

that the series of six lateral open-

ings in Birkenia, and nine ventral

ones in Tremataspis, imply the

existence of as many pairs of seg-

mental appendages in the respec-

tive. forms. And were we to

temper our imagination so far as

to conceive that only two pairs of

incisions served for this purpose,

or indeed even one pair, what

function should we assign for the

rest, since they are all alike?

Given an integral series, why com-

plicate matters by postulating a

: multiple function? Either let us

gorel Pie sig aspect of the bodyarmor regard them provisionally as gill-

openings, or else declare that their

nature is entirely problematical; Patten’s guess we are not pre-

pared to take seriously, even as a guess.

One point further must not escape attention. Our denial of

the existence of appendages in Tremataspis is supported not only

by analogy and negative evidence, but by the general configura-

tion of the head-shield itself. The latter agrees with that of

Cephalaspis, Auchenaspis, Drepanaspis and the like, in its convex

or regularly elliptical outline, especially as seen from the ventral

aspect, where according to Patten the appendages had their

attachment (Fig. 1). But if we examine the visceral surface

tree

rte Pa

--4-

pou b

1 Traquair, R. H. Report on Fossil Fishes. Zyans. Roy. Soc. Edinburgh, vol.

39, 1899, p. 859.

No. 440.] CORRESPONDENCE. 577

of Pterichthys, Bothriolepis or Asterolepis, we note at once that

the contour of the body-wall is inflected, or even indented, on

either side at the point where the pectoral limbs peculiar to

these forms were attached (Fig. 2). This latter condition being

governed by mechanical principles, as anyone can perceive, it is

common only amongst those forms which possess swimming

appendages. And its absence in Tremataspis and generally

throughout the order to which it belongs fortifies us in our con-

clusion that in this group paired segmental appendages were

wanting.

C. R. EASTMAN.

CAMBRIDGE, MAss.

(No. 439 mailed September 5, 1903.)

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THE

AMERICAN NATURALIST.

VoL. XXXVII. September, 1903. NO. 441.

A CONTRIBUTION TO THE MORPHOLOGY

AND DEVELOPMENT OF CORYMORPHA

PENDULA AG.

ALBERT J. MAY.

Tue following paper on the morphology and development of

Corymorpha pendula was undertaken at the suggestion of Dr. C.

W. Hargitt, for whose kindly criticism and suggestions throughout

its progress it is a pleasure to express my obligation.

The purpose of these observations is to investigate some of

the more fundamental morphological features of this hydroid ;

to trace the origin and development of the Medusa and to inquire

briefly into the origin of the sex cells and the phenomena associ-

ated with oógenesis.

Corymorpha, in all the synopses of the Hydroidea which I have

examined, is described as a solitary form, but in the material

which was placed at my disposal, my attention was early called

to the presence of what appeared to be definite colonial buds

arising from the hydrorhiza. Acting upon the suggestion of Dr.

Hargitt that it might be an interesting problem to determine

whether these associated forms were true buds or mere parasites,

I have begun my study with an investigation of the nature and

relation of these forms.

579

580 THE AMERICAN NATURALIST. [Vor. XXXVII.

Methods. — The specimens at my disposal were collected by

Dr. C. W. Hargitt at Wood's Holl, Mass, during the summer

of 1900. Several methods of killing and preserving were used.

My best results were obtained from material killed in corrosive

acetic acid. Formalin and Flemming's solution gave less satis-

factory results.

In the study of general histological features I found that zz

toto staining in borax carmine was both a satisfactory and conven-

ient method. Specimens might be left in the stain from 10 to

48 hours and then the stain extracted to any desired degree in

1% acid-alcohol. Dehydration required from one to two hours.

For clearing, cedar oil, clove oil, xylol or turpentine were used;

xylol or turpentine proving most satisfactory.

In the study of the developmental features a number of sec-

tions were stained on the slide with iron-hematoxylin. Combi-

nations of iron-haematoxylin and Bordeaux red, and of eosin and

haematoxylin were used with good success.

In the former combination sections were placed in a 2% solu-

tion of ammonio-ferric-alum for from thirty minutes to four hours.

After this they were washed for several minutes in running

water and then stained in a 5% aqueous solution of haematoxylin

for from one to three hours. They were then again washed in

running water and again treated with the iron solution, which

slowly washed out the stain, until a satisfactory differentiation

was obtained. After rinsing the sections in water they were

stained from fifteen minutes to one half hour in Bordeaux red,

carried up through the alcohols and mounted in balsam.

In the eosin-hzematoxylin method sections were first stained

from one to two hours in a 2% solution of alcoholic eosin and

then from five to fifteen minutes in a weak solution of Delafield's

haematoxylin.

Associated forms.— As stated in the introduction, one of the

first things that was called to my attention in taking up the

study of the morphology of Corymorpha pendula, was the presence

of a smaller hydroid associated with it. These smaller forms have

all the appearance of true buds. In some cases they are found

attached to the hydrocaulis, and in others they are observed

springing up from within the tangled network of rhizoidal fila-

No. 441] CORYMORPHA PENDULA AG. 581

ments. Wishing to discover something as to the true relations of

these forms, I made a series of transections of the stem of Cory-

morpha in the region of these supposed buds in order to deter-

mine whether, in the first place, there was any organic connection

between them. A study of these sections revealed no such

organic connection whatsoever. The base of the smaller hydroid

was in all cases buried only in the perisarc of Corymorpha and

generally separated from the ectodermal layer by a considerable

thickness of perisarc.

Upon taking off some small portions of the perisarc of Corymor-

pha bearing these supposed buds, staining with borax-carmine

and mounting whole, a peculiar modification of the base of these

smaller hydroids was observed. The stem after it has pene-

trated the perisarc for a short distance, expands and gives off a

number of lateral finger like projections extending through the

perisarc in various directions.

Further examination also proved that the stem is annulated

for a considerable portion of its length and that the coenosarcal

canals so characteristic of Corymorpha are entirely lacking.

From these observations it is quite clearly evident that we have

here distinct forms instead of true colonial buds, thus confirming

the usual descriptions of Corymorpha as a solitary form.

This species, however, I have not been able to find described

in any of the synopses of the Hydroidea. From its morphology

it is evidently a tubularian. Dr. Hargitt has suggested for the

species the name parasitica, and as such it has recently been

described by him. Torrey (:02) has expressed the opinion that

these forms are undoubtedly clusters of young individuals of

Corymorpha. This view would seem to be precluded not only

from a consideration of their morphological characteristics, but

from the fact that all the specimens examined were of about the

same size, and some of them even began to show signs of sexual

maturity.

Corymorpha pendula.

External Morphology. — L. Agassiz (62, p. 276) says, “ This

hydroid is not found along our shores as are the other tubula-

rians, but may be obtained by dredging in deeper water on a

582 THE AMERICAN NATURALIST. [Vor. XXXVII.

sandy or muddy bottom. In some localities it is quite plentiful.

It has been collected in three different places, all within Massa-

chusetts bay." Specimens have been taken off “Crab Ledge,”

Chatham, in the deep waters of Muskeget Channel and in Vine-

yard Sound. The natural position of this hydroid is an upright

one. Hargitt (:01, p. 313) describes this hydroid as “bright

pink in color, medusze light yellowish, manubrium, tentacles and

bulbs pinkish.”

Corymorpha is always found as a solitary form, never budding

nor giving rise to colonies. The stems grow to a height of from

six to ten centimeters and reach a maximum diameter just a

little above the proximal extremity. Toward both extremities,

the diameter decreases ; the stem tapering gradually toward the

distal end, where it terminates ina short blunt point. The stem

is invested in a very delicate, filmy, colorless perisarc, which in

the upper region of the stem is very closely applied, but below

becomes separated from it by a very considerable space, forming

here a sort of loose fitting corrugated sac over the blunt point of

the basal extremity. The stem is seen to be traversed by a series

of longitudinal bands which anastomose with one another here

and there, and which represent, as we shall see later, a series of

canals hollowed out in the entoderm. From the lower region

of the stem there grows out a large number of short papilliform

processes. The hydrorhiza consists of a tangled network of

numerous thread-like filaments.

The hydranth is somewhat flask shaped, the neck of the flask

being represented by the flexible proboscis at the distal extrem-

ity of the hydranth. In the centre of this proboscis is found the

mouth surrounded by from seventy-five to eighty-five closely set

tentacles arranged in a number of irregular whorls, and having

somewhat the appearance of a brush. About the base of the

hydranth is a single circle of very much larger tentacles varying in

number from twenty-five to thirty.

The medusoids are arranged in closely crowded clusters borne

upon slender branched pedicels which arise from the body of the

hydranth immediately above the proximal set of tentacles.

There are from fifteen to twenty-five of these medusoid-bearing

branches.

No. 441.] CORYMORPHA PENDULA AG. 583

Hisrorocv or Parts.

(a.) Hydrocaulis.— The perisarc: — The perisarc presents

the appearance of a thin, semi-transparent film. This film is

considerably thicker about the basal portion of the stem than

it is in the distal region where it becomes so thin and delicate

as to be almost lost to sight.

Ectoderm :— The ectoderm, although its cellular structure

was not especially well defined in some sections, is composed of

a single layer of somewhat columnar cells with large and dis-

tinct nuclei, and filled with numerous granules. Among the

cells of the ectoderm are to be found great numbers of small

transparent cells, the developing nematocysts. These nema-

tocysts seem to be most numerous and best developed in

the tentacles.

Entoderm:— The entoderm is composed of two distinctly

different kinds of cells whose disposition through the stem may

be best understood by calling attention here to the peculiar modi-

fication which Corymorpha presents as regards the enteric cav-

ity. In most hydroids this is represented by a hollow coenosarc,

its tubular cavity being central in position and communicating

directly with the cavities of the polypes. In Corymorpha, how-

ever, this cavity is represented by a number of intercommuni-

cating canals running longitudinally through the entire length of

the stem. (Fig. 1, m.) The whole central axis of the stem is

occupied by a column of parenchymaike tissue composed of

large, loose, vacuolated cells of an irregular polygonal shape.

(Fig. r, 2.) These cells have very little contents with the excep-

tion of here and there a number of small nuclear corpuscles.

Torrey (:02) refers to these entodermal cells as “skeletal cells ” ;

and, as he suggests, this condition of peripheral canals and a solid

entodermal axis, would seem to be, in view of the large diameter

of Corymorpha, a “direct adaptation to size." The very deli-

cate character of these cells might make the propriety of the

term *skeletal" somewhat questionable. It seems quite proba-

ble, nevertheless, that these parenchyma-like cells do afford some

support to the stem, but rather by imparting to it a sort of

584 THE AMERICAN NATURALIST. (VOL. XXXVII.

turgescence, than because they serve as a true, supporting

skeletal framework.

The peripheral layer of entoderm, on the other hand, is com-

posed of small sub-spherical cells filled with granular contents and

possessing distinct nuclei. (Fig. 1, 2.)

Fic. 1. ewig tion of stem of Corymorpha showing one of the ccenosarcal canals at

nner layer of entoderm ; 4, peripheral layer of entoderm; c, mesoglea; 4,

fing é, perisarc.

It is in a zone between these two kinds of entodermal cells

that the longitudinal canals are excavated. These canals are

simple tubular spaces lying between some of the entoderm cells

and entirely devoid of. anything like a specially differentiated

wall. Above, these canals pursue their course through a cone

like projection of the loose entoderm cells which extends for a

considerable distance into the cavity of the hydranth proper.

(Fig. 2, 4.) Because of the lacunar nature of these canals, their

course through the entoderm cone is somewhat indefinite. At

the base of the cone, the canals converge and finally empty into a

single median canal which is continued upwards to the apex of

the cone. (Fig. 2, 6.) I have not been able to get sections

No. 441.] CORYMORPHA PENDULA AG. 585

which show satisfactorily the course of this median canal, but I

am inclined to think that it is continued up through the apex of

the cone, thus finally communicating directly with the hydranth

cavity.

Towards the base of the stem these canals become fewer and

larger by union with each other, but finally disappear in a region

just a little below the belt of papillae.

VES

E gerer

Fic. 2. Longitudinal section of hydranth showing core of entodermal cells at a,

through which pass the longitudinal canals, 4; c, gland cells of entoderm; d, ecto-

derm; e, tentacle.

For the purpose of comparison a series of transverse and of

longitudinal sections of Tubularia couthouyt were made, and

here was found a structure which in all essential features resem-

bles what we have described in Corymorpha. The canals of

Tubularia, however, are distinctly wedge shaped in transection

with the apex of the wedge directed toward the center of the

stem. In Corymorpha, the canals are oval or elliptical in tran-

section. Moreover in Tubularia, the pith like core occupying

the center of the stem is composed of small nearly spherical

586 THE AMERICAN NATURALIST. | (Vor. XXXVII.

cells well filled with granular contents. There is no projection

of the entodermal axis into the hydranth cavity as we found in

Corymorpha.

A number of sections of Tubularia larynx were examined for

further comparison. The structure of this species agrees in con-

siderable detail with that of T. couthouyt.

Allman (71, p. 205) describes a similar condition in Tubularia

indivisa. Me says: “The stem of Tubularia indivisa presents

immediately within the perisarcal tube a continuous layer of

ectoderm enclosing the endoderm, which extends to the very

center of the stem and thus obliterates all trace of a central

somatic cavity. The place of the cavity however is supplied by

numerous canals which are excavated in the endoderm and take

a longitudinal course through the stem, occasionally communicat-

ing by lateral offsets with one another and finally all merging in

a common central cavity at the base of the hydranth." He

further states that the canals are of unequal size, **one of them

especially being in almost every instance considerably larger

than any of the others." L. Agassiz (662) says that a similar

condition as to the size of the canals may sometimes be detected

in 7. couthouyi, although I was. unable in my observations to

perceive any appreciable difference in size.

(b) Papilliform processes. — The lower part of the stem of

Corymorpha is covered with numerous short conical papilla

arranged in an irregular longitudinal series and apparently fol-

lowing the course of the canals. Lower down toward the base

of the stem these papillz increase in length and in many speci-

mens these small processes were seen in all stages varying from

very short blunt papillae above, to extremely elongate filaments

below. Immediately below these processes are found the numer-

ous filamentary rhizoids which cover the saccular portion of the

perisarc. The presence of these two structures at the base of

the stem and the apparent merging of the one into the other

naturally suggests the problem as to their relation and derivation.

Upon examination of transections of the stem in this region we

find that these papillae begin as simple outgrowths of the ecto-

derm. (Fig. 3.) At the same time the mesogloea directly under-

neath these outgrowths is seen to grow very much thinner, and

No. 441] CORYMORPHA PENDULA AG. 587

in the course of development, to extend up into the papilla for a

short distance as seen in Fig. 4 v. As the layer of mesogloea

is thus interrupted, the peripheral entodermal cells extend up

into the papilla, forming a central axis of entoderm. L. Agassiz

(62) states that these papillae are hollow and are permeated by

prolongations of the chymiferous tubes of the stem. Allman

(71, p. 209) in a description of C. nutans, modifies this state-

ment somewhat and says, * They apparently communicate with

FiG 4.

Fics. 3, 4, 5 : — Successive stages in the development of the papilliform processes: 4, ento-

derm ; c, mesoglea; d, ectoderm.

the canal over which they lie." From a study of a number of

sections I have been unable to verify this statement of Agassiz.

Not only do the papillze very often lie in regions other than over

the chymiferous canals — I have frequently found them growing

out of an area of the stem between two canals — but all of the

sections show the papillae to be solid, composed of an ectoderm

and a central core of entoderm. They are not hollow, but main-

tain this solid character even after they have grown into com-

paratively long filaments. (Fig. 5.)

As these papilla continue to grow and elongate, they finally

588 THE AMERICAN NATURALIST. (VOL. XXXVII.

break through the perisarc; a perisarcal sheath begins to be

secreted about them, and their cellular structure becomes less

distinct. As the hydroid now continues to grow, the perisarc,

which in the younger stage was somewhat closely applied to the

base, now becomes separated from it by a considerable distance,

forming a loose corrugated sac over the proximal end. With

the withdrawal of the coenosarc, the papillae also sever their

connection with the coenosarc. The living cellular portion now

being separated from that of the main stem and hence devoid of

any means of nourishment, disintegrates, leaving merely a hollow

perisarcal tube in its place. Upon examining sections of the

filamentary rhizoids, we find that they are mere hollow tubes of

perisarc, terminating in an imperforate and somewhat clavate

extremity. There thus seems to be no doubt that these rhizoids

» are but the products of secretion of the papilliform processes.

Allman. (71, p. 209) referring to them says that these proc-

esses *have never been seen to act as organs of adhesion nor

have we yet any evidence of the office they may serve in the

economy of the animal, but it is impossible not to recognize

in them structures having a close relation to the filaments of

attachment which are given off from the stem a little lower

down."

(c.) Hydranth.— The most striking histological feature of the

hydranth is the presence of numerous and highly developed

gland cells (Figs. 6, 7). The development of this remarkably

specialized structure leaves little doubt that the function of diges-

tion devolves especially upon this part of the hydroid cavity and

that there are here structures set apart for the distinct purpose

of secreting the digestive fluid. In the distal part of the

hydranth cavity, the entoderm is thrown into a series of large

and complicated folds or ridges whose surfaces are covered with

these digestive gland cells. In the intervals between the ridges

are often found smaller and less numerous cells whose protoplasm

takes a denser stain than that of the surrounding cells. The

free ends of the gland cells are seen projecting out into the

hydranth cavity where their walls may often be seen ruptured

and their contents sloughing off into the cavity of the hydranth.

(Fig.6, a.) Often the gland cells may be seen entirely separated

No. 441.] CORYMORPHA PENDULA AG. 589

from the parent tissue and projected bodily into the hydranth

cavity. (Fig.6, 5). Toward the center and base of the hydranth

cavity, the folds of the entoderm disappear, but the gland cells

become decidedly larger and are found covering a considerable

portion of the pith-like core which projects into the cavity as

described above. The cells in this region are more elongated,

with their broad ends projecting into the digestive cavity ; the

outer ends are rather narrow and tapering, and become merged

in the converging ends of the neighboring cells. (Fig.6.) These

Fic. 6.— Portion of a transection through the basal region of hydranth showing a, gland

lls sl hi ffi hvd th ity d b, others projected bodily int ity ; M,

mesoglcea.

gland cells possess a distinct nucleus and nucleolus. The pro-

toplasm often presents a finely alveolar structure. Lying in the

protoplasm are numerous minute granules and very often gran-

ules of a larger size. The whole structure presents a striking

similarity to the digestive epithelium of the larva of some of

the insects. Needham gives a description of the digestive

glands of dragon fly nymphs which corresponds in considerable

detail with what we find in Corymorpha.

In Hydra and in most of the simpler hydroid polypes, the

590 THE AMERICAN NATURALIST. [Vor. XXXVII.

digestive cavity consists of a simple tube traversing the long

axis of the body, but dilated in the region of the hydranth. The

function of digestion is not limited to any particular region of

this enteric cavity, but is carried on to a greater or less extent

in all its parts. In Corymorpha, however, the enteric cavity

Fic. 7. — Portion of a transection through the distal a of inen showing

character of gland cells. v, vacuoles; 7z, mesogloea

does not remain a single tube, but becomes differentiated into

secondary cavities having the character of pouches and canals.

From the histological character of these extensions of the primi

tive digestive cavity, we see that they are not only morphological

differentiations of the latter, but that they also correspond to

distinct physiological differences. As stated above, the gland

cells are highly developed in the hydranth, but are entirely absent

from the ccenosarcal canals ; that is, the physiological activity of

the digestive cavity is not shared by the canals which arise from

it, and the function of digestion has become localized. It would

seem to be strongly probable that we have here a specific case

of localized digestion and the subsequent distribution of its prod-

ucts by means of the coenosarcal canals. The central primary

space together with the accessory spaces constitutes what might

be termed a gastro-vascular system, the coenosarcal canals under-

taking the function of a circulatory system. The gastric sys-

tem in Corymorpha then may be distinguished from that of

Hydra and of the simpler hydroid polypes by this exhibition of

a higher differentiation.

(d.) Tentacles —The ectoderm of the tentacles consists of short

columnar cells with large distinct nuclei. Scattered among

No. 441.] CORYMORPHA PENDULA AG. 591

these cells are numerous large thread cells. The entoderm con-

sists of large irregular polygonal cells which entirely fill up the

axis of the tentacle. Toward the distal end, the entoderm cells

become fewer and larger, their boundaries stretched transversely

across the tentacle in longitudinal section, thus giving it a

peculiar’ septate appearance.

The ectoderm and entoderm are separated from each other by

a thin supporting lamella, the mesogloea, which also separates

the two layers of cells in other portions of the hydroid.

ORIGIN AND DEVELOPMENT OF THE MEDUSA.

The medusoids in this species are found at the extremities of

a number of hollow branched peduncles. These peduncles are

from fifteen to twenty-five in number and are arranged in two

circles about the hydranth just above the proximal set of ten-

tacles. On the same specimen will usually be found medusoids

in various stages of devel-

opment, from buds just

forming, to rather mature

Medusze, together with in-

termediate stages. The

general order of develop-

ment of the medusa buds i.

is centripetal, that is, the — 2 .....M

younger buds are usually

found at the bases of the

branches, while the older

ones appear at the distal

extremities of the branches. 1G. 8.— An early stage in the development of the

The ~ pedicles upon 7 Béenle Me ur ird M ente

which the Medusse are

borne are simple hollow outpushings of the hydranth and are

composed of the same layers — ectoderm, entoderm and meso-

gola. In their development the medusa buds present essentially

the same succession of phenomena which we find in other

Tubularie. They begin as simple evaginations of the wall of

the penduncle. (Fig. 8. By a proliferation of the ectoderm

592 THE AMERICAN NATURALIST. [Vor. XXXVII.

cells in the distal end of the bud, a plug of ectodermal cells is

formed which grows down into the medusoid cavity, forcing back

the entoderm as it advances. This retreating fold of entoderm,

as it doubles upon itself by the increased growth of the ecto-

dermal plug, presses closely upon only four. meridional areas of

the stationary layer of entoderm. By this process, four equi-

distant spaces or chymiferous channels are left in direct com-

munication with the medusoid cavity. These spaces constitute

the beginnings of the radial canals. (Fig.9, c.) As these chan-

nels continue to elongate, they become dilated at their extremities

into bulb like ex-

pansions which

evidently give

rise to the cir-

cuar canals.

Earl in the

process of de-

velopment, the

entoderm, forces

its way back

through the

center of the

ectodermal plug

to form the ma-

nubrium. (Fig.

G m) Ihe

mass of cells

lying between the bell and the manubrium, and which are of

ectodermal origin, give rise to the future reproductive elements.

(Fig. 9, g.) The inner ectodermal layer of the bell, and the

ectodermal layer of the manubrium arise as successive differ-

entiations of the germinal layer. (Fig. 10, t. c.) ;

L. Agassiz (/62, p. 278) says, “the medusa buds of this

Hydroid do not become free individuals, but remain attached,

develop their generative material and then wither and die.”

A. Agassiz (65, p. 193) on the other hand, in speaking of

the Medusa of Corymorpha pendula says, ** Although the separa-

tion of this Medusa from its Hydrarium has not been observed,

Fic. 9. — A later stage in the development of the medusa bud, show-

1 +h A £ £, M "* kat A *h Aint 1

€; g, germinal layer; 2, entoderm; 2, ectoderm.

No. 441. CORVMORPHA PENDULA AG.

593

yet their similarity to the most advanced Medusa buds observed

on our Corymorpha, leaves but little doubt on this point."

Torrey (:02, p. 38) in his observations on C. palma states that

the gonophores do not become free. The eggs “ drop from the

Manumbrium of the attached Medusa " and *there is no free

swimming larva." Dr. Hargitt tells me that in his mind there

is no doubt that the gonophore sometimes becomes free. Ona

number of occasions hydroids have been taken by him, which

DNIH

Fic. 10. — An advanced stage in the development of the medusa bud; 7, tentacles; », circular

canal; 7, radial canal; e, germinal layer; æ, ectoderm; 4, entoderm; 7, inner ectodermal

layer of bell; c, ectodermal layer of manubrium.

bore Medusze in all stages of development, and at the same time,

from waters close about where the hydroids were dredged, free

medusz. These Medusz unmistakably belonged to Corymorpha

and there seems to be but little doubt as to the indentity of their

relations. We have here a condition just intermediate between

the fixed and the free gonophore phases, a phenomenon not very

uncommon among the Tubularidz.

594 THE AMERICAN NATURALIST. (Vor. XXXVII.

Oögenesis. — Between the manubrium and the bell is a large

mass of cells, which as we have seen above, is derived from the

ectoderm and is destined to give rise to the future reproductive

elements. Brauer ('91t, p. 575) in speaking of the origin of the

genital products in Tubularia mesembryanthemum says, “ Die

Geschlechtsprodukte von Tubularia entstehen aus interstitiellen

zellen des Ektoderms des Gonophorentragers, sie treten nahe der

Ursprungstátte eines Gonophors ins Entoderm über, wandern

hier ihrer Reifungsstátte, dem ektodermalen Glockenkern, zu."

Weismann ('83) states that in some hydroids, the reproductive

elements may originate in the coenosarc of the trophosome.

While this is very rarely the case in hydroids where a definite

medusoid is developed, I have found one case where there

appeared to be a small egg cell in the ectoderm of the stem.

The cells of the germinal layer, especially in female gonophores,

are a great deal

larger than those

of the surrounding

1 š layers. They are

uS — 1 "48 - closely packed to-

gether, nearly

spherical in form,

and possess large

p.09 distinct nuclei with

prominent nucleoli.

(Fig. 11, 4) For

a time these cells

all increase slightly

in size, the proto-

Fic. r1.— Portion of a transection through a late medusa bud plasm in the mean-

showing the growth of the ovum by absorption of primitive ‘ :

egg cells; o, ovum; », nucleus of ovum; e, entodem; 2, time becoming

ectodermal layer of manubrium. somewhat denser

from the periphery toward the center of the cell. At this stage

may generally be found four or five cells which are outstripping

their neighbors in growth and are attaining a considerably greater

size. These cells are destined to develop into mature ova and are

found for the most part occupying a position next to the manu-

brium. They now assume a distinct amceboid form sending out a

No. 441.] CORYMORPHA PENDULA AG. 595

number of pseudopodia-like processes among the adjacent cells

and finally absorbing them into their own substance. (Figs. 11,

12.) The phenomenon as I have observed it is essentially as

described by Doflein ('96) for Tubularia larynx.

The boundaries between one of the large cells and those adja-

cent toit begin to break down. This large cell which, according

to Doflein has had the advantage of position and nourishment,

at once appropriates to itself the protoplasm of these surrounding

cells. A syncytium with irregular outline is thus formed from

the fusion of these cells, and in it may be detected for some time

the disintegrating nuclei of the absorbed cells. (Fig. 12, 2.)

Fic. 12. — Absorption of primitive egg cells by growing ovum; 4, Poi gs of qe sit

tate of Pee ian treads d, entire cells lying within the

7, ies

In regard to the process of absorption, there are two distinct

views. In Doflein’s words ('96, 2. 1):

“Die einen Forscher nehmen an dass die Eizelle die umgeb-

enden Nihrzellen aktiv auffrisst, sich von denselben, wie eine

Amóbe von anderen Organismen ernahrt. Dagegen behaupten

Andere eine Auflósung der Nahrzellen und eine Aufnahme der-

selben in flüssigem Zustand."

Doflein (96) contends that the process of absorption described

by Balfour and Tichomiroff as “amæboides Fressen” is decid-

edly incorrect and that the protoplasmic processes of the devel-

oping ova do not function as true mouths which bodily engulf

the adjacent primitive ova. While my results in general con-

596 THE AMERICAN NATURALIST. (VoL. XXXVII.

firm this view of Doflein’s, nevertheless in one or two cases there

seemed to be a distinct engulfment of the primitive ova. It

appears to me that neither theory alone explains all the phenom-

ena involved in the growth of the ovum, but that a combination

of both theories would better explain the facts. My observa-

tions, however, lead me to agree with Doflein in his objection to

the term “amoeboides Fressen." If we watch an Amoeba in its

movements, we will observe that when a process of the proto-

plasm presses against certain foreign particles of organic nature,

they become sunk in the substance and pass gradually into the

interior. Here they become surrounded by a little globule of

watery fluid, a vacuole; and by degrees these particles partially

or wholly disappear. All the matter ‘which is capable of it

becomes digested and assimilated by the protoplasm. It is very

probable that the vacuole contains some ingredient of the nature

of a ferment which is capable of acting upon these foreign sub-

stances and rendering them more soluble. These are the phe-

nomena involved in the process of amoeboid eating. Yet, while

they agree in a few respects with those phenomena which are

exhibited in the growth of the ovum, on the other hand, it seems

to me that they present such fundamental differences as not

entirely to warrant the statements of Balfour and Tichomi-

roff. In the first place, there is no formation of a vacuole

about the absorbed cells, with the exception of, later on, a small

one about their nuclei. I did not find these vacuoles in any sec-

tions of Corymorpha, but in examining some slides prepared by

Miss Allen (:00) in her study of the development of Tubularia

crocea, 1 found numerous cases of these vacuoles containing,

from one to as many as seven or eight nuclei. These vacuoles

however, were all found in ova which had begun to segment, and

undoubtedly the same phenomenon would have presented itself

in Corymorpha, if the material used had been of a later stage

of development. Doflein ('96) describes the same thing in

Tubularia larynx.

Furthermore, the cytoplasm of the Mbecchad cells simply min-

gles with that of the growing ovum, and undergoes no apparent

change whatsoever. There is no process of absorption, the cell

walls disappear gradually and the nuclei appear to have been

No. 441.] CORYMORPHA PENDULA AG. 597

carried along in all directions by a “streaming motion of the

plasma of the ovum." While I could not observe such a process

in the preserved material, the “ general agreement of authors on

this point" (as Doflein states) makes it quite certain that this

phenomenon of amoeboid movement does exist.

Miss Allen (:00) in her observation on Tubularia crocea, says

that in numerous cases the outlines of the absorbed eggs could

be distinguished in the protoplasm of the absorbing egg. Ina

very few favorable sections, I have been able to detect the out-

lines of these absorbed cells (Fig. 12, 7.), but in most cases only

the nuclei were distinguishable.

A great many of the primitive ova do not thus become

absorbed, but remain scattered among the mature eggs in

their original undeveloped state. Doflein suggests that possibly,

after the larvae have left the gonophore, these remaining germ

tissue cells unite to form new eggs.

SUMMARY.

In summarizing the results obtained in this study, the fol-

lowing points should be noted : :

I. Corymorpha, in keeping with its usual description, is a

solitary form.

2. 'The rhizoidal filaments of attachment are formed as

secretions from the papilliform processes.

Both filaments and papilla are modifications of the same

structure.

3. The central axis of the stem is filled with a mass of

parenchyma-ike cells in which is excavated a number of longitu-

dinal canals. The longitudinal canals are extensions of the

hydranth cavity.

4. Gland cells are highly developed in the hydranth cavity,

but are entirely absent from the ccenosarcal canals. The

function of digestion thus becomes localized in one portion of

the enteric cavity while the caenosarcal canals function as a

circulatory system. A fairly well differentiated gastro-vascular

system thus becomes developed.

598 THE AMERICAN NATURALIST. [Vor. XXXVII.

5. The medusoid is developed from a bud which is formed

by a simple outgrowth of the wall of the peduncle.

6. The chymiferous canals of the medusoid are formed by a

fusion of the two layers of entoderm throughout the inter-radial

areas. i

7. The sex cells are derived from a plug of ectodermal cells

formed at the apex of the bud.

8. The eggs develop by a process of absorption of the cells

of the germinal tissue, thus giving rise to an amoeboid syncytium.

The nuclei of the primitive germ cells persist for some

time in this syncytium, but gradually disintegrate.

THE TONEDOTENE LABORATORY,

USE UNIVERSITY,

e 25, 1903.

BIBLIOGRAPH Y.

AGASSIZ, A.

'65. Illustrated Catalogue of the Mus. Comp. Zoól No. u, North

American Acalephe. Cambridge.

AGASSIZ,

ALLEN, CARRI

Development of Parypha crocea. Biol. Bull., Vol. 1, pp. 291-315.

PROC

'63. Reproductive System in the Hydroids. Rept. British Assoc. Adv.

Sci.

ALLMAN, G. J.

"71. Monograph of Gymnoblastic Hydroids. London.

BRAUER, A.

'91. Ueber die Entstehung der Geschlechts producten und die

Entwicklung von Tubularia mesembryanthemum Allm. Zeitschr. wiss.

Zool., Bd. 52, pp. 551-579.

BOURNE, A. G.

'83. Recent Researches upon the Origin of the Sex Cells in Hydroids.

Quart. Jour. Micr. Sci., Vol. 23, pp. 617—622.

BUNTING, MARTH

‘94. The Origin: of Sex Cells in Hydractinia and Podocoryne. Jour.

Morph., Vol. 9, pp. 203-2536.

DOLFLEIN, F.

'96. Die Eibildung bei Tubularia. Zeitschr. wiss. Zool, Bd. 62, pp.

61—73. S

Cobitelbadans to the Natural tiay of the United States. Boston.

E M.

No. 441.] CORYMORPHA PENDULA AG. 599

HARGITT, C. W.

: 00. Contribution to the Natural History and Development of

Pennaria tiarella. Amer. Nat., Vol. 34, pp. 387-414.

HARGITT, C.

: O1. Srani of North American Invertebrates, XIV. Amer. Nat.,

Vol. 35.

Hincks, I.

'68. A History of the British Hydroid Zoophytes. London.

KoRSCHELT & HEIDER.

'75. Text-Book s the Embryology of Invertebrates. Translation by

Mark & Woodworth. London.

LANG, A

'5. Comparative Anatomy. Translation by Barnard. London.

McMuvunnich, J. P.

A Text-Book of Invertebrate Morphology. New York.

MuMMERY, S. R.

'58. On the Development of Tubularia indivisa. Quart. Jour. Micr.

Sci., Vol. t, pp. 28-30.

Genes. H. L. & HanairT, C. W

'94. Perigoniums jonesii; a Hydroid supposed to be new from Cold

Spring Harbor, Long Island. Amer. Nat., Vol. 28, pp. 27-34.

Sans, M.

'58. On the Nurse Genus Corymorpha and its Species together with the

Medusae produced from them. Ann. Mag. Nat. Hist., Vol. 8, pp. 353-

360.

SCHNEIDER, C. C.

:02 Moe Histologie der Tiere. Wien.

SMALLWOOD

'99 quisa to the Morphology of Pennaria tiarella McCrady.

Amer. Nat., Vol. 38 pp. o

TORREY, H. D.

02. Hydroidea of the Pacific Coast. Univ. Cal. Publ, Zool. Vol. 1,

pp. 1-194.

WEISMANN, A.

83. Die Entstchung der Sexualyellen bei den Hydromedusen. Jena.

THE HABITS OF CAMBARUS.

J. ARTHUR HARRIS.

Ir 1s the purpose of the present paper * to present in a brief

way some of the main points which have been collected on the

habits and distribution of North American crayfishes.

Our crayfishes offer a particularly inviting and important field

for ecological work. A fuller knowledge of the habits of the

different species may contribute to a more complete under-

standing of the very remarkable sexual dimorphism occurring in

the adult males. Individual variability in the species of Cam-

barus is very great and puzzling, and the differentiation into

species as compared with the other genus Astacus, of the sub-

family to which it belongs is extensive. The great differences

in environment to which the species are subjected is apparent to

anyone who is acquainted with the physiographic conditions

prevailing in the vast stretch of territory over which this genus

is distributed and an examination of these conditions and the

adaptation of the animals to them would doubtless yield interest-

ing results.

The habits of any species necessarily depend largely upon the

character of its environment, and from an ecological or biological

standpoint it is impossible to consider the two separately. In

this place space cannot be given to a discussion of the importance

of a correlation of physiographic features with floral and faunal

distribution as it has been emphasized by Woodworth, Hays and

Campbell, Simpson, Cowles, Adams and others.

It has long been known that the fishes occurring in the upper

course of a stream of considerable size are different from those

found in its lower course, and more recently the ecological factors

concerned have been moré fully discussed. In the crayfishes as

in fishes the Fauna of different parts of a stream is not the same,

: Wn is an abstract of a part of an ecological catalogue of the crayfishes

ing to the genus Cambarus which is being published in the Xansas Univer-

i grad Bulletin

601

602 THE AMERICAN NATURALIST. ([Vor. XXXVII.

but in the crayfishes the problem is not a simple one since the

animals may remain out of water for a considerable length of

time and it is to be expected that the conditions under which

they may be found are not always the same. C. virilis usually

occurs in running streams but may also be found in stagnant

ponds with C. immunis and C. gracilis and has been known to

resort to burrowing. C. bartonii seems to be characteristic of

the cooler mountain springs and streams, but it is also found in

limestone caves with C. pellucidus and associated with C. dioge-

nes, a burrowing species, with the same habits. C. affinis and

C. blandingii may also be used as illustrations of the same point.

While an examination of a table of distribution of species

offers some interesting suggestions, it is as yet impossible to

prepare a list of species characteristic of certain types of local-

ities, though many species may be accurately assigned.

Several species appear to be confined to mountain streams.

The crayfishes of the Ozark Mountains are little known, but the

point is illustrated in the Appalachian tributaries of streams

emptying into the Atlantic Ocean and the Gulf of Mexico. The

lower courses of these streams are very different from their

sources and the general Fauna is very different. C. extraneus,

C. spinosus, C. bartonit, C. acuminatus, C. forceps, C. longulus

and other species seem to be confined for the most part to

mountain streams. As is the case in fishes, the same species

may occur in the head waters of streams originating on opposite

sides of a divide and debouching at widely separated points.

Some species found in the lowland portions of the same streams

are characteristic so far as yet known, and some idea of the

species characteristic of the lower portions of the course of a

stream — the lowland forms — may be gained from an examina-

tion of a list of the forms reported from the Atlantic coast plain

and the lower portions of the Gulf States. It must be remem-

bered, however, that elevation above the sea is not the only

factor producing lowland conditions so far as faunal and floral

distribution is concerned.

Faxon pointed out that the greater the fall of a stream the

greater the difference between the species of the upper and

lower portion of its course and while the question needs careful

No. 441.] HABITS OF CAMBARUS. 603

investigation the same seems to be true of the number of species

found in a given stream ; a river with a heavy fall having a larger

number of species than one with the same or greater length, but

having a more limited vertical range. It is apparent that the

former would present much more diverse life conditions than

the latter.

The habits of certain of the North American species of cray-

fishes have attracted considerable attention. These are the

burrowing species. As burrowing species are to be designated

only those forms which show an especial dependence upon this

method of life, since it is reasonable to suppose that most, if not

all the stream-inhabiting species dig short burrows in the banks

at least in certain localities. The species which seem to be most

dependent on this habit are C. diogenes, C. gracilis, C. carolinus,

C. argillicola, C. simulans and C. immunis.

C. diogenes has well been characterized as preéminently a

burrowing species. Its presence is usually indicated in the low

places where it is most frequently found by the large number of

mud * chimneys," about a foot in height, sometimes scattered

over several acres, radiating from some sluggish stream, ditch,

brook or lower, moister portion of the area, the animals being

frequently found at a considerable distance from any permanent

body of water. Of the other species C. gracilis seems to be as

typically a burrowing species as C. diogenes, and is generally

reported as an inhabitant of prairie regions. Adults are to be

found in open ponds only in the early spring, and the burrows

are often found at long distance from any permanent body of

water. C. simulans has been reported from streams and ponds

and from burrows in a slough. C. immunis is known principally

from stagnant ponds, resorting to burrowing upon the drying up

of the ponds and upon the approach of winter. Of C. azgz//icola

and C. carolinus the nature of the habitat has not been described.

All of the above are known to be ** chimney-building " species.

The *chimney," very aptly so called, is a mound of mud of a

quite regular, conical or pyramidal form constructed at the

mouth of the burrow, with a smooth, internal opening which is

merely a continuation of the shaft of the burrow, and is some-

times sealed at the top. The chimneys of all the species have

604 THE AMERICAN NATURALIST. . (VoL. XXXVII.

not been carefully described, but so far as known they are

essentially the same. That of C. diogenes has a maximum

height of twelve inches, but is usually lower, is in shape, like a

truncated cone often somewhat higher than broad. The most

remarkable difference being that of chimneys two inches in diam-

eter and eight to eleven inches in height, described by Abbott,

who states that those found in meadows at a distance from run-

ning water were invariably broader and not so high as those

erected near running water. The chimney is composed of

pellets of clay firmly cemented together, owing to the moist

condition in which they are laid on, giving an irregular, nodular

appearance to the outside of the structure. This brief descrip-

tion applies to the more perfectly formed or typical chimney.

We will recur to this subject later, after the form of the burrow

has been mentioned. The actual process of construction has

been observed only once.

As has been suggested above, where the habits of the differ-

ent forms was mentioned, the burrows are made at the time of

the drying up or the lowering of the body of water in which the

animals are found. At the edge the burrow may be a simple

shaft, a foot, more or less, in depth, ending below in a cistern-

shaped enlargement, in which the animal, usually only one but

sometimes two, is found. Farther back from the stream in

moist meadows where the burrows may have been begun at the

time of high water, the depth must necessarily be much greater

to reach soil water during the prolonged heat of summer. These

burrows are not uniform in structure but as yet data are lacking

for the determination of any plan, other than the one of reaching

water in the most direct manner. The presence of more than

one opening to the same burrow has often been noticed. They

are usually quite simple but occasionally are branched in various

ways. When many burrows occur in a limited area they may

easily become connected accidentally. Enlargements in the

shaft of the burrow have been noticed and attributed to the

original enlargement at the bottom of the burrow which has

been repeatedly carried deeper as the water in the soil became

lower. In recent observations on C. carolinus this explanation

does not seem to hold.

No. 441.] HABITS OF CAMBARUS. 605

Too much importance, it seems to me, has been attributed to

the chimney in discussions of the habits of Cambarus. While

the chimney is usually a very regular and well-built structure, it

is often found, in some species at least, as a more or less irreg-

ular heap of clay pellets, and so far as our present knowledge

extends, can hardly be regarded as anything more than the result

of the easiest method of disposing of the material removed in

excavating the burrow. On this point, however, further obser-

vations are desirable. The purpose of the sealing of the burrow

is not so clear. The prevention of material falling into the bur-

row from the surface and possibly (?) protection against enemies

may account for it. That the sealing is not a matter of the acci-

dental falling together of the upper edges of the chimney while

in a moist condition is evident from the fact that the opening is

sometimes filled to below the surface of the ground and, as some-

times happens, with clay of a different nature from that compos-

ing the rest of the chimney. "

Concerning the purpose of burrowing there can now be no

question. Some species of Cambarus seem never to reĝðrt to

the habit, in the restricted interpretation of the term, while

others, C. immunis, and, to a less extent, C. virilis, are inhab-

itants of ponds or streams and resort to burrowing only upon the

drying up.of the ponds or the approach of winter, while C. diog-

enes and C. gracilis have adopted this mode of life almost entirely,

being found in the open water during but a very small part of

the year. That the burrows are not for retreats while the eggs

are being hatched has been conclusively shown. That they

serve as a place of protection against enemies has been sug-

gested and while it cannot be stated that the burrowing species

are not better protected against animal enemies than are the

forms inhabiting open. water this cannot be the primary purpose

of the burrow. The burrows are almost invariably described as

extending to the water in the soil, and while the water in the

enlargement at the bottom of the burrow is usually very muddy,

it enables the animal to keep its gills moist. A point of inter-

est in connection with the burrowing species is the range of the

species. C. diogenes seems to be the most widely distributed

species. C. argillicola has a wide distribution, and C. carolinus

606 THE AMERICAN NATURALIST. [Vor. XXXVII.

has been reported from widely separated localities. To what

extent this is dependent on the habits of the animals is difficult

to say, but the burrowing species obviously have a great advan-

tage over the others in their ability to occupy territory which

would be habitable to many of the species for but a very small

portion of the year.

Only one species is found in salt water, C. wh/eri, a species of

limited range, is found in salt marshes covered twice daily by the

tide, and also in brackish and fresh water where C. blandingii is

sometimes found associated with it. C. montezume is said to

occur in salt water.

Many observations have been made upon the blind species,

inhabiting caves and underground streams in Kentucky, Ten-

nessee, Indiana, Missouri and Florida, but they are not of a

nature to be easily summarized. The blind forms are not

confined to one group or section of the genus. The species are:

C. acherontis, C. setosus, C. hamulatus, C. pellucidus and C.

pellucidus testi.

Parasites, various species of Branchiobdella, have been noticed

on C. affinis, C. bartonii, C. digueti and Cambarus sp. and will

doubtless be found on many other forms, and C. digueti is

recorded as being attacked by Temnocephala.

Little has been recorded of the habits of the crayfishes dude

the winter. It seems most probable that the stream inhabiting

species pass the winter in burrows in the bank or under stones,

etc., in the bed of the stream. The latter is sometimes the case

with C. virilis. The burrowing species seem quite generally to

spend the winter in the burrows, coming out early in the spring

and returning again when the water begins to become low as the

summer progresses.

Observations have been made on the colors of the crayfish in

. relation to its environment. One observer, working, for the

most part, on C. immunis, with fewer observations on C. pro-

pinquus, C. bartonit and C. diogenes, concludes that the coloring

closely resembles the environment and has a protective function.

According to him, the colors in all cases were similar to the

environment except in those with a red coloration. The red

color, he concludes, is due to the immediate effect of the sunlight.

No. 441.] HABITS OF CAMBARUS. 607

He found that young crayfishes which are red, due to the pre-

sence of large chromatophores, changed to blue or black or

suffered no change as the adults of the locality were blue, black

or red. He finds that the burrowing crayfish, C. diogenes, comes

out in the spring much the color of the soil, but this color is

gradually changed to red in the open sunlight. Other obser-

vations indicate that in the case of C. gracilis, as typically a

burrowing species as C. diogenes, the females are always olive-

green no change taking place during the time they are to be

found in the ponds in the spring, while the few males which

have been taken are a marked salmon red, although they had just

left the burrow. In C. carolinus, another burrowing species,

“red” and “blue” individuals seem to occur. While it is

undoubtedly true that individuals of a species taken from dif-

ferent localities may show marked differences in coloration, cau-

tion.must be exercised in designating all the differences as pro-

tective adaptations.

Observations on the breeding habits are very limited. As to

the time of copulation and oviposition a few data have been

recorded. In C. diogenes, upon the habits of which more has

been written than any other species, copulation and oviposition

seem to occur in the spring. One observer found females in

burrows carrying eggs in March and April, while another gives

the middle of May as the approximate time of hatching of the

eggs. Another observed C. diogenes and C. gracilis, kept in

aquaria, copulating in the spring and never found crayfishes

(sp?) mating except in March, April and sometimes May, and

was able to get reports of females **in berry " later than june in

only two instances. Another observer reports the species as

copulating in the open water April 2nd and laying eggs April

18th to 30th. A female with eggs in an early stage of develop-

ment has been reported May 3rd. These observations were

made over a wide range of territory. An interesting exception

is the taking of a female with eggs nearly ready to hatch, on

January rst and might suggest autumn oviposition, as has been

observed in some other species. Females of C. gracilis have

sometimes been found in open ponds in early spring with a few

young and it may be that the young leave the parent immedi-

608 THE AMERICAN NATURALIST. [Vor. XXXVII.

ately after they quit the burrows in the spring. In southern

Kansas, C. simulans has been taken from burrows, with eggs

apparently recently laid, late in August, and in New Mexico,

with the swimmerets loaded with eggs, in May. In C. immunis

the females are found with eggs in stagnant ponds in the fall ;

they pass the winter in burrows and appear again in ponds,

where the process of hatching is completed, in the early spring

— about March 21st. C. argillicola has been reported with

young as early as April 2nd. C. neglectus was found with eggs

and young in the cold water at the mouth of a large limestone

cave in the Ozarks early in June, and since those taken at the

same time in various other places in the neighborhood had

neither eggs nor young, the lateness of the date may be attrib-

uted to the retarding effect of the low temperature upon the

hatching of the eggs. In another locality C. neglectus (?) was

taken with eggs April 13th. In C. virilis the females are found

with eggs in the spring, but not during the winter. ;

Of the above species C. diogenes and C. gracilis are preëmi-

nently burrowing forms, C. argillicolais a burrower, C. simulans

burrows extensively as does also C. immunis, and C. virilis some-

times resorts to the habit. C. neglectus seems to be found prin-

cipally, in clear rocky streams. In regard to the burrowing spe-

cies it has been suggested that the burrow is designed as a retreat

while the eggs are being hatched, but this is not very likely,

although the eggs may undergo a very large part of their develop-

ment in the burrow.

Observations on the habits of the young crayfish are few.

Two suggestive ones are that in C. diogenes the neatest chimneys

are those constructed by the smallest individuals and that the

young of C. gracilis are the first to appear in the stagnant ponds .

which are frequented by this species in early spring and are also

found there in the late summer after other forms, C. immunis,

C. virilis, and the adults of C. gracilis, have gone into their bur-

rows.

St. Lours, Mo., Mar. 22, 1903.

SYNOPSES OF NORTH AMERICAN

INVERTEBRATES.

XIX. Tue TrICHODECTID#,!

MAX MORSE.

Tur genus Trichodectes, the only genus of the family Tricho-

dectidæ, is one of the two genera of the order Mallophaga

which are found on mammals. From the other genus, Gyro-

pus, Trichodectes is distinguished by the fact that its members

have a three-jointed antenna. The order Mallophaga has been

divided by Piaget (80) into two families, vzz., the family Liothe-

idz and the family Philopterida — the two being distinguished

by the character of the legs which are modified either for run-

ning (the former) or for clasping (the latter). Kellogg (99) has

separated the genus Trichodectes from the Philopteridz by the

erection of the family Trichodectidae which he attributes to

Burmeister. His authority for this attribution is not evident.

Of the 48 species in the genus, 18 are considered here as hav-

ing been taken on North American Mammals.

The species of Trichodectes are distinguished from one another

by such characters as the size and shape of the antenne, the

character of the setze, or hairs that clothe the body, the shape of

the thorax, the genitalia of both male and female, etc. In the

present paper, at the suggestion of Prof. Herbert Osborn, much

attention has been paid to the so-called “abdominal appendages "

of Piaget. This organ, for it is all one organ, is a growth of the

posterior ventral edge of the antepenultimate segment of the

abdomen, in the female. The extension grows backward, under-

lying the major portion of the last abdominal segment and grow-

ing upward at the sides to reach the level of the tergum of that

segment. The median portion of the extension, however, does

not grow very far backward and the result is the formation of

' Contributions from the Department of Zoólogy and Entomology of the Ohio

State University, Number 12.

609

610 THE AMERICAN NATURALIST. [Vor. XXXVII.

two lateral flaps as can be seen in the figures. Dorsally it has

the appearance of forming a prominent curved hook at either side.

Much variation in this organ in the several species was found and

its general shape, together with the presence or absence of setae

on it afford excellent criteria for the separation of species. The

function of the abdominal appendage is partly in clinging to the

hairs of the host but more especially in the adjustment of the

eggs to the hairs.

A word may be said concerning the general habits of these

insects. Their food consists of scales and epidermal excretions

from the host. The mouth-parts are fitted for biting and the

mandibles are well developed. They cling to the hairs of the

host by means of the mandibles, which are set at the posterior

end of a clypeal groove running longitudinally along the ventral

side of the head, into which groove may be fitted a hair and this

then grasped by the mandibles. It is probable that the sides of

the groove are capable of being closed down over the hair and

thus anchor the Mallophagan to the host, without the assistance

of the mandibles. The legs, also, assist in holding the insect to

the hair. The office of the abdominal appendage has been men-

tioned. The eggs are often seen in the body of the specimen.

There is a well-developed lid fitting over a chitinous capsule.

The capsule is glued to a hair and development occurs there,

the lid being shoved off at the emergence of the larva. Most

of the species are confined to one species of host, although

exceptions are met with.

Only those species known to occur in North America are con-

sidered here. It is hoped that by the aid of the keys, the

figures and the descriptions, any member of the group in the

region defined may be identified, even if the host is not known.

The characters in the key are given in the main for either sex and

none of these characters are difficult of examination. A ready

method of preparation is to boil a few minutes in a solution of

potassic hydrate in water, clear in carbolic acid and mount in

balsam.

No claim to completeness is made for this paper, for it is

improbable that it includes all the species to be found on North

American mammals. Doubtless there are species imported from

Europe that have thus far escaped our notice.

No. 441.) WORTH AMERICAN INVERTEBRATES. 611

Acknowledgments are due Professor Vernon L. Kellogg,

Professor Lawrence Bruner, Dr. L. O. Howard, and Dr. D. E.

Salmon, for material. My thanks are especially due Professor

Herbert Osborn of the Ohio State University, under whom the

work was done and who has offered valuable suggestions in the

course of the work. The private collection of Professor Osborn,

which embraces nearly all the species considered in this paper

was put at the disposal of the writer.

The literature of the group is not large, but I append only the

more important papers from the point of view of the worker in

North America.

CHAPMAN, BERTHA.

'97. Two new Species of Trichodetes. Ent. News, Vol. 8, pp. 185.

DENNY, H.

'42. Monographia Anopleurorum Britanniz. London.

GIEBEL, C. G

'74. Insecta epizoa. Die auf Sàáugethiere und Vögeln schmarotzenden

Insecten. Leipzig.

KELLOGG, V.L

'99. A List of the Biting-Lice (Mallophaga) taken from Birds and Mam-

mals of North America. Proc. U. S. Natl. Mus., Vol. 22, pp. 39-

100.

OSBORN, H.

'91. The Pediculi and Mallophaga Infesting Man and the Lower Ani-

mals. U.S. Dept. Agr., Div. Ent., Bull. No. 7.

OSBORN, H.

'96. Insects Affecting Domestic Animals. U. S. Dept. Agr., Div. Ent.,

Bull. No

OSBORN, H.

:02. Mallophagan Records and Descriptions. Ohio Nat., Vol.2. p.

PIAGET, E. )

‘80. Les Pediculines. Essai Monographique. Leyden.

PIAGET, E.

'85. Zdem. Supplement.

For further treatment the reader is referred to the works of

Osborn and Kellogg as well as those of Piaget that have been

cited above.

612 THE AMERICAN NATURALIST. (VoL: XXXVII.

Key.

a. Setæ short and delicate, not reaching middle of the succeeding segment.

emale abdominal appendage with sete . . (RUMINANT TYPE.)

6. Head convex in front.

c. Head pointed, spines on the anterior border of the female abdom-

inal appendage small and inconspicuous . . scalaris.

cc. Head rounded, spines on anterior border of the Man abdom-

inal appendage equalling in size those on the posterior.

d. Form elongated, setze of antenne strong and conspicuous.

e. First tarsal joint of third leg extended ; portion of head

anterior to antennz wider than posterior ; inner border

of female abdominal appendages lobed.

spherocephalus.

ee. Notas above, inner border of female abdominal append-

age straight . . . parumpilosus.

dd. Form short and thick, inteni setae ‘dalitate and incon-

spicuous.

f Abdomen wider than head, thoracic suture evident,

ventral border of penultimate segment of the abdo-

men with deep emargination in the median line.

limbatus.

ff. Abdomen width of or less than head, thoracic suture

not evident, border "mr slightly emarginate.

climax.

bb. Head emarginate in front.

£. Distal portion of inner border of tibia of third legs translucent,

with setze short and delicate.

A. Prothoracic spiracle situated internally, not on lateral edge

of the prothorax . . tibialis.

hh. Prothoracic spiracle ada Literally beyond edge of

segment . . parallelus.

Same chitinized and fined with eioi lods sete. vi SUS.

aa. Setæ long, linear, reaching or extending beyond the middle of the next

posterior segment; female abdominal appendage without setæ.

(CARNIVORE AND RODENT TYPE.)

t Head pointed > < roo 9 33 e. e Ssubrostratus.

ij. Head not pointed

7. Antennz with proximal joint in the male swollen and having the

` whole antenna long.

&. Second antennal joint in the female with posterior process.

geomydis.

kk, Second antennal joint in female without process.

/. Prothorax long, twice the length of the meso-metathorax.

No.441] NORTH AMERICAN INVERTEBRATES. 613

m. Anterior border of the head converging to a point.

californicus.

mm. Anterior border of the head convex-rounding.

castoris.

Jl. Prothorax short, equalling in "a the posterior part.

4. Length under one mm. . . mephitidis.

nn. Length over one mm. . thoracicus.

Jj. Antenne linear, the proximal joint not ana

o. Aserjes of gan strong spines on the posterior border of the

metathora nasuatts.

00. Spines linseed to the literal PU 5i the postei ii bor-

der of the metathorax.

f. Præocular sinus in the male wide, the præantennal spur

being widely separated from the eye.

gp. Bength over t.s6mm. =: o s o alis.

gg. Length under 1.50 mm . retusus.

pp. Sinus narrow, the eye being dai to the tien nal

EDEK s. Sov 9o guadraticeps.

DESCRIPTIONS.

Family Trichodectide. Kellogg.

Genus TRICHODECTES. Nitzsch.

T. scalaris Nitzsch: Female: Length,

1.368 mm.; width of abdomen, .666 mm.;

head, .342 mm.; thorax, .180 mm.; anten-

ne,.154mm. Outline of body elliptical.

Head not as broad as abdomen, roughly

triangular in outline, front converging to

a bluntly-pointed anterior margin. Anten-

næ linear and small. Thorax trapezoidal.

Pro-mesothoracic suture distinct. Meso-

metathorax with lateral borders salient

and prominent, with strong sete; poste-

rior border of thorax sinuous. Abdomen

widest on segment 3. Lateral borders of

segments dark, heavily covered with setze.

Segments 1-8 with transverse, fuscous

bands. Setz strong, but short and blunt.

Head covered with sete as is also the

case with the abdomen. No male speci- * Nd a Rss: f. adus ppon-

men available. On domestic cattle. dage of female.

614 THE AMERICAN NATURALIST. [Vor. XXXVII.

T. spherocephalus Nitzsch. Female:

thorax.

concave.

Length,

1.764 mm.; width, .63.;

.396; thorax, .27; Antenna, 27.

Body elongated, narrow. Head not as

broad as abdomen, squarish, semicircular

in outline in front of the antenna. Pro-

thorax not well marked off from rest of

thorax and narrower than meso-meta-

Posterior border of metathorax

Abdomen elliptical, widest on

segments 3 and 4. Middle of segments

with smoky brown bands running trans-

versely. First tarsal joints of second and

third legs with inner border extended

T. spherocephalus. a, adult female; 6,

abdominal appendage of female.

sete. Basal joint of antenna larger

than other two joints, distal segment

clavate. Tarsal claws long and strong.

2.127 mm.; width, .81; head, .504;

thorax, .306; antenna, .27. Form

large. Head convex in front, slightly

longer than wide, about two thirds as

wide as the abdomen. Antenne

stout, linear, proximal joint but little

larger than others. Mandibles small.

Thorax with the suture indistinct, the

two segments being of about equal

width. Posterior border convex. Lat-

stigmata surrounded by dark blotches.

mpilosus. a, adult female; 4, abdom-

female.

T. paru

taal appendage of

No.441] WORTH AMERICAN INVERTEBRATES. 615

Interior of segments transversely banded. Setz delicate, short and incon-

spicuous. Tarsal claws slender. Male not in hand. Found on the horse.

This is a synonym of egui Denny.

T. limbatus Gervais. Female: Length, 1.71 mm.; width, .828; head,

.45; thorax, .22; antenna, 27. Body large, elliptical in outline. Head

square, with anterior portion trapezoidal, the anterior border being truncate.

Preantennal spur well-developed. Eyes prominent. Antennz linear.

Thorax short, narrow, with antero-lateral angles produced. Abdomen

widest on segments 3 and 4, the outline ovate. Lateral borders of the seg-

ments dark and heavily chitinized. Tarsus normal, claws strong, slightly

T. limbatus. a, adult female; 4, adult male; c, abdominal appendage of female.

curved. Seta scant and delicate. Male; Length, 1.206 mm.; width, .54 ;

head, .324; thorax, 198; antenna, .234. Smaller than the female, but gen-

eral shape the same, except that the abdomen converges more abruptly pos-

teriorly from the 3 and 4 segments. Sete as in the female. From the

Common Goat and the Angora Goat.

T. climax Nitzsch. Female: Length, 1.693 mm.; width, 792; head, 451

circle. Antennal pit deep, the angle well defined. Antenne linear, distal

segment slightly swollen in the middle. Thorax trapezoidal, the suture not .

THE AMERICAN NATURALIST.

[VoL. XXXVII.

T. tibialis. a, female; 4, male.

No. 441.] WORTH AMERICAN INVERTEBRATES. 617

very distinct. Posterior border convex. Abdomen widest on segments 3

and 4. Lateral borders of the segments dark. Middle of the segments

transversely banded. Seta delicate but profuse. Tarsal claws long and

strong. Male: Length, 1.35 mm.; width, .702; head, .378; thorax, .18;

antenne, .27. Head as in the female. Thorax longer in proportion than

in the female, but similar in outline. Posterior border slightly concave.

Abdomen widest on segments 2 and 3, whence the body outline curves

rapidly to the end of the abdomen. Lateral borders of the segments dark

and middle of the segments banded transversely. . Tarsal claws longer and

less curved than in the female. On the Domestic Goat.

T. tibialis Piaget. Female: Length, 2.016 mm.; width, .756; head, .595 ;

thorax, .288; antenna, 360. Head about as wide as the abdomen, elon-

gated, eyes prominent, preantennal spur obtuse. Mandibles long and

strong. Antennz with distal joint swollen. Thorax narrower than head,

margin continuous with the contour of the abdomen. Prothoracic spiracle

distinct. Suture fairly evident. Posterior border convex. Abdomen ellip-

tical, with dark blotches in front of the spiracles. Segments with transverse

bands. Seta short and thick, in single transverse rows on the abdomen.

Shape as in the female. Proximalsegment of the antenna enlarged. Length

of the antenna in the male greater than in the female. Trabeculz in the head

conspicuous. Thorax as in the female, but the lateral angle more extended.

bdomen elongate, with segments trans-

versely banded. Spiracles with blotches as

in the female. On the Black-tailed Deer.

Trichodectes parallelus Osborn. Female:

Length, 1.56 mm.; width, .45; head, .45;

thorax, .21; antenna, .28. Body elongated,

abdomen narrow Leipurus-like, the ratio of

the length to the width being about 3.1.

antennz converging forward. Anterior bor-

der of head emarginate. Head heavily chiti-

sparse and thinly distributed. Spiracles of

the abdomen capped anteriorly with black

blotches. Tarsal claws long. Male not

known. Ona species of deer, the species not being known to the describer.

T. parallelus. Female.

618 THE AMERICAN NATURALIST. [Vor. XXXVII.

Trichodectes setosus Giebel. Female: Length, 2.304 mm. ; width, .846 ;

head, .594 ; thorax, .396; antenna, .324. Large, elongated. Width of the

head and that of the abdomen nearly equal, the latter slightly the greater.

Head large, sides in front of the antennze converging to the slightly concave

front border. Antenne filiform. Thorax large, prothoracic spiracle very

prominent. Sutureevident. Posterior border slightly concave. Abdomen

with the lateral border of the segments chitinized and dark. Middle of the

UW.

T. setosus. a, female; 4, male; c, abdominal appendage of the female.

segments pigmented. Abdomen widest on segments 4 and 5. Seta short

large. Abdomen terminated in a point. Pro-mesothoracic suture more

distinct than in female. On the porcupine, Erethizon ermineus, Neb.

Trichodectes subrostratus Nitzsch. Female: Length, 1.116 mm. ; width,

No. 441.] NORTH AMERICAN INVERTEBRATES. 619

.54; head, .36; thorax, .og; antenna, .114. Head acutely pointed ante-

riorly. Antenna slender. Thorax short. Suture between prothorax and

mesothorax distinct. Posterior border of thorax

concave. Abdomen elliptical, obtuse and broad,

broadly rounded posteriorly. Segments banded

M not i

Length, 1.134; width, .66; head, .306;. thorax,

Antenna thick and short, heavily chitinized, with

second joint provided with a posterior extension.

Prothorax narrower than meso-metathorax, the lat-

ter with lateral borders extended. Posterior bor-

der of the thorax concave. Abdomen broad,

widest on segments 3 and 4, whence the sides of

the abdomen converge gently posteriorly. Seta

T. subrostratus. Female.

width, .63; head, .23; thorax, .114; antenna, .396. Similar to female, but

Sree urahaits $

PEETA ee

T. geomydis. a,female; 4, male.

Trichodectes californicus Chapman. Female: Length, 1.37 mm. ; width,

.84; head, .33 ; thorax, etc., not given. Form “ short, broad, pale yellowish

white without definite markings, except on the front of the head." “ Head”

with “anterior margin with a deep incision; sides of the front receding

rapidly to the sharply angulated trabeculz;....antennz tud and large,

— beyond the temporal margin when extended back ;... eye prom-

inent ;....occipital margin nearly straight and without hairs or spines ;... .

620 THE AMERICAN NATURALIST. [Vor. XXXVII.

prothorax long and wide; sides slightly rounding ; posterior margin medially

convex ;....abdomen broadly oval, reaching

its cuentas width at the third segment ; .... "

Chapman (97). Male not known. From a

pocket-mouse, Perognathus, sp.

Trichodectes castoris Osborn. Female:

Length 1.20; width, .52; head, .32; thorax,

.16; antenna, .19. Head broader than long.

Antenne slender, basal joint not swollen.

Thorax not showing the suture distinctly.

Thorax short, posterior border concave.

Abdomen elliptical, widest on segments 4

and 5. Sides straight. Setz long and deli-

cate on the posterior lateral borders of the

segments, shorter on the middle of posterior

er. Tarsal claws short and delicate.

foe» Vee Mau RN Male: Length, .99; width .48; head, .21;

man, Ext, News, (97). thorax,.126; abdomen,.252. General shape

as in the aa, Antenne longer, with

proximal segments swollen. Suture in the thorax more distinct. Abdomen

broadest on the third segment, whence the sides converge posteriorly. Seta

on the abdomen long. Genital hooks well developed, lying parallel with

one another, but slightly curved. From the Beaver

T. castoris. a, female; 4, male; c, abdominal appendage of female.

Trichodectes mephitidis Osborn. Female: Length, .9o mm. ; width, .50;

head, .30; thorax,.12; antenna, 18. Form short and thick. Head broader

than long and as wide as abdomen. Antenne thick, segments equal in

diameter. Thorax short and wide, prothorax narrower than remainder of

orax. Metathorax with antero-lateral angles extended. Abdomen thick

No. 441) NORTH AMERICAN INVERTEBRATES. 621

and broad, widest on 4. Seta long and delicate. Tarsal claws of medium

.27; thorax, .18; ; antenna,

-19. Shape as in the

female except that the

front of the head is more

acute. Antenne longer

than in female, with e f i Y

P ins E» ETT TTE EE TRO

proximal joint swollen. ipte

Thorax longer, narrower Abi E Ed sy

delicate as in the female.

On the Skunk, Mephitis

mephitica, la. an e

Trichodectes Menace: mum 6$

Osborn. Female: ipd T. méphitidis. a, female; 4, male; c, abdominal appendage

UM 183 mm.; width, of female

head, .324; thorax, Jas a ntenna, .180. Head broad. Antenne filiform.

Thorax with two parts well distinguished by suture. Lateral border of

meso-metathorax widely extending. Posterior

border of thorax concave. Abdomen ovate,

widest at segments 2 and 3, converging very

gently towards the posterior end of the abdo-

men. Sete slender and long. Abdominal

appendage with flaps provided with extensions

toward median line of the abdomen. Male:

Length, 1.15 mm.; width, .58; head, .32;

thorax, .18; antenna, .21. Head more elon-

gated than in the female. Thorax and abdo-

men as in the female, save that the abdomen

converges more abruptly posteriorly. Prox-

imal joints of the antennae enlarged. On

Bassaris astuta, Palo Alto, California.

rower than the a nte slender,

proximal segment slightly enlarged

acicus. a, female; 4, ab- short, posterior bo: ve. bdomen

oni np THING ad anteriorly, converging from segment 3

to the posterior end. Sete long and delicate. Tarsal claws long and

slender. From Nasua, Costa Rica.

622 THE AMERICAN NATURALIST. [Vor. XXXVII.

Trichodectes latus Nitzsch. Female: Length, 1.890 mm.; width, 1.116;

head, .414; Thorax, .27 ; antenna, .2888.

Head wider than long, front margin

flattened-convex, arching forwards but

slightly from the antenne. Eyes promi-

nent. Posterior border of the head as

well as the postero-lateral border pro-

vided with six long setz on either side.

Antenne of moderate thickness and

length, terminated by a tuft of delicate

sete. Antennal sulcus moderately wide.

Prothorax well differentiated from the

remainder of the thorax by the suture.

Prothorax the length of the meso-meta-

thorax, sides diverging from the attach-

ment with the head, posteriorly, to meet

the expanded antero-lateral knob-like ex-

b than the head, without transverse bands

on the segments. Sete in general long

and delicate. Genetalia the type of

Retusus, but the delicate inner border

T. nasuatis. a, temale; 4, abdominal

A £K f, 1

er >

T. latus. a,female; 4, male; c, female abdominal appendage.

No. 441.] NORTH AMERICAN INVERTEBRATES. 623

wider and sinuous. The strong spines of the forward portion of the geni-

talia point in a postero-median direction as well as outwardly as in Refusus.

Male: Length, 1.638

mm.; width, .810; head,

404s thorax, 27;

tenna, 36. Antennal sul-

cus very wide. Eyes less

prominent than in the fe-

male. Antennz with the

basal joints swollen. Ab-

domen pointed posteri-

line, but slightly curved.

On the Domestic Dog.

Trichodectes retusus

Nitzsch. Female: length,

1.206 mm. ; width, .576;

head, .324 ; thorax, .162 ;

antenna, .162. Small,

head narrower than the

T. retusus. a, female; 4, male; c, abdominal appendage of

abdomen. Sides of head, in front of antennz, converging anteriorly, the

contour of the clypeus being a semicircle.

inconspicuous. Antenne chitinized, strong

T. quadraticeps. a, female; , male; c, abdominal appendage of

emale.

Trabecule prominent. Eyes

Distal segment of the antenna

clavate and provided

with a wart-like inner

extension, Thorax

with suture definite.

Mesothorax with lat-

eral angles widely ex-

panded provided with

t Hind

short and delicate.

Tibia with distal angles expanded. Male: Length, .828 mm.; width, .450 ;

head, .288; thorax, 144; antenna, 180.

General shape as in the female.

Abdomen shorter, rounded posteriorly; generative organs inconspicuous.

624 THE AMERICAN NATURALIST. [Vor. XXXVII.

Sete delicate. Tibia with distal edges expanded as in the female. Tarsal

joints small. On the Weasel Putorius ermineus.

Trichodectes quadraticeps Chapman. Female: Length, 1.08 mm.

width, .63; head, .324; thorax, .114; antenne, .18. Head quadrangular

widest along the posterior border. Antennz slender and long. Thorax

short, with metathoracic segment extending laterally. Spiracles prominent.

Abdomen wide, much wider than head. Widest portion of the abdomen

on the fourth segment. Posterior part of abdomen squarish, the sides but

little converging. Generative appendages long and prominent. Male:

Length, .918 mm.; width, .54; head, .27; thorax, .126; antenna, .114.

General shape as in the female. Abdomen wider in proportion than in the

female. Setz short, delicate and sparse. Tarsal claws long, straight and

narrow. On the fox, Urocyon virginianus.

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Vernon's Variation.'— Mr. H. M. Vernon of Oxford, England, has

summarized in a handy octavo volume the most important observa-

tions on variation made since the publication of Darwin's great work

on Lhe Variation of Animals and Plants under Domestication. The

book will form a valuable student’s manual in the field of general

biology. It is clear and concise in style and is remarkably free from

technical terms and mathematical formulz, considering the fact that

it deals largely with statistical methods.

It is divided into three parts which treat respectively of “The

Facts of Variation,” “The Causes of Variation” and “ Variation in

its Relation to Evolution.”

Part I includes a brief explanation of the statistical methods

employed in the study of variation, a remarkably clear presentation

of the difficult subject of correlated variations, and a discussion of

dimorphism and discontinuous variation, in which the ideas of Bate-

son and de Vries receive special attention.

Part II treats of the effects on organisms of external conditions,

such as temperature, light and moisture, a subject discussed more

exhaustively by Davenport in his Experimental Morphology and by

Verworn in his Ad/gemeine Physiologie. Two chapters devoted to

blastogenic variations contain, along with much other material, an

account of important experiments made by the author in the hybridi-

zation of various species of echinoderms. Accepting as probably

correct the idea of Weismann that variations which are hereditary are

of germinal origin, Vernon believes that the heritage borne by the

germ-cell is not at all periods of its existence the same, but that it

changes as the germ-cell changes in maturity. Thus when two

species of echinoderm are crossed, which ordinarily breed at different

seasons of the year, that species impresses its characters most

strongly on the offspring which is (at the time the cross is made)

nearest the height of its breeding season. ‘This conclusion is based

1Vernon, H. M. Variation in Animals and Plants. 8vo., ix+ 415 pp., 30

figs. New York, Henry Holt and Co., 1903.

625

626 THE AMERICAN NATURALIST. (Vor. XXXVII.

on the averages of large series of measurements of hybrid offspring,

but is very probably vitiated by the occurrence of artificial partheno-

genesis so easily produced in the case of echinoderm eggs. Another

series of experiments cited by Vernon in support of his view is hardly

more convincing. It consists in experiments made by Ewart with

mating rabbits early or late in rut.

* Mendel's Law” is treated as a law of “hybridization " only, its

profound significance as a general law of heredity being unnoticed,

while the Galton-Pearson “ Law of Ancestral Heredity " is treated as

the law of heredity. To many biologists the evidence for the Men-

delian principles is too strong and too clearly counter to the Galton-

Pearson law to be thus lightly brushed aside. It also raises a strong

presumption against Vernon’s idea of a heritage gradua//y changing

during the ripening of the sexual products.

Part III, contains a brief survey of a familiar field. Natural

selection is recognized as the efficient agency in evolution. Adaptive

variations are discussed at some length and the evidence for and

against their inheritance are considered. Environment is regarded

as directly inducing germinal variation.

PHYSIOLOGY.

Von Fürth's Comparative Chemical Physiology of the Lower

Is. — Perhaps the most important general advance made by

physiology in the last ten years has been the inclusion of the lower

animals within its field of research. Just as anatomy was immensely

illuminated by a thorough investigation of the structure of the lower

forms and thus became truly comparative, so physiology will gain a

clearer and more certain insight into life processes by a study of

these where they occur in greatest simplicity, What has already

been done in this direction especially from the chemical standpoint

is scarcely accessible to the student except through the original

sources of publication for ever so excellent a book as Verworn's

General Physiology passes over this subject most superficially. Von

1 Fürth, O. von. dpud Panicle Physiologie der niederen Tiere. Jena,

Fischer, 1903. 8vo., xiv + 67

No. 441.] NOTES AND LITERATURE. 627

Fürth's very extensive compilation will therefore be a welcome

guide in this growing field of research. Although von Fiirth’s

volume marks a new departure, the book is arranged on extremely

conservative lines. After a brief introduction, it deals with the

chemistry of the blood, respiration, digestion, excretion, animal

poisons, secretion, etc., a series of heading that suggest at once the

sections of the old-fashioned human physiology. It seems really

remarkable that such a classification as this should have been

adopted, for if there is one lesson taught by comparative physiology

more clearly than any other, it is the non-essential character of the

blood. Large groups of the lower metazoa are complete organisms

and yet they are without this fluid. "Why then should the blood be

chosen as a means of introducing the student to the chemical physi-

ology of these lower forms? But aside from this traditional treat-

ment of the sections, the substance of these sections is refreshingly

modern, and with their excellent bibliographies they form admirable

summaries of many new fields of work. The exhaustiveness of the

treatment is well indicated by the subject digestion which covers

over a hundred pages and takes up in sequence digestion in the

protozoa, sponges, cnidaria, echinoderms, worms, molluscs, crustace-

ans, and other arthropods, devoting a chapter to each. Such a work

as this, despite its defects, must find its way to the hands of every

advanced student of animal physiology.

Mind in Nature.'— This little book is at bottom an argument for

a certain form of vitalism. The author, while admitting the value of

the chemico-physical descriptions of movements given by Loeb and

other investigators of similar interests and aims, insists that it is

impossible to account for those forms of movement which we usually

designate as action or conduct on this ground. He believes that

there is a gap in the physical series which must be bridged by some

such factor as the psychic if a complete description of action (Hand-

lung) is to be given.

'The work consists of a careful study of the forms of movement.

Reflexes are classified as :

I. Simple.

II. Complex 1 E NE a. Homometachronous.

ee b. Heterometachronous,

! Driesch, Hans. Die “Seele” als elementarer Naturfaktor. Studien über die

Bewegungen der Organismen. Leipzig, Englemann, 1903. 8vo, vi-97 pp.

628 THE AMERICAN NATURALIST. (Vor. XXXVII.

The synchronous reflexes are such complex movements of multiple

phases as exhibit a rhythm; the metachronous are chain reflexes

in which each step serves as a stimulus for the next, of these

the homometachronous are coórdinated, the heterometachronous

uncoórdinated.

In his discussion of instinct the author states that only simple

stimuli can initiate instinctive movements. By a simple stimulus he

means something which is essentially an elementary nature quality ;

such, for example, are light, motion, heat. The simple stimulus he

contrasts with the individualized stimulus, which is appropriate for a

certain specialized type of sense organ. In this discussion much

credit is given to Loeb for his analyses of instincts.

Attempts are made to get at the meaning of the concepts of neural

centres, spontaneity, autonomy, etc. Driesch thinks that the present

tendency to do away with the concept of brain centre is as far from

being desirable as is the uncritical acceptance of the old notion of

such centres.

The chapters on directed movements (taxes), reflexes, instincts,

and brain centres serve merely as an introduction to the author's

real subject, Activity (Handlung). The analysis of reflexes does

not furnish the information necessary for the understanding of

action, for in the latter there are characteristics which are not

found in the simpler forms of movement. The criteria of action

which Driesch presents are the “Individuality of Association " and

the * Historical Basis of Reaction." Volitional action differs from

directed or reflex movement in that it is infinitely variable; it is not

a matter of certain elements of stimulus and response in definite

and unchanging relation, but of practical unpredictability. Loeb

makes the great mistake of supposing that all movements of the

organism can be described in terms of the factors which are common

to reflexes. Now, in the opinion of Driesch this is impossible, since

in action there is *autonomy"; we therefore have to take into

account the associational facts, and in as much as the subjective as

such cannot be material for the biologist it is necessary to objectify

this factor. For Driesch the objective element which enables one

to give a description of action is the “ psychoid."

The book well deserves the attention of biologists who are interested

in the relations of their science to chemistry, physics, and psychology.

The chances are that few will be able to agree with the work as a

whole, but this makes it all the more valuable. It is of interest to

note that we have here another biological discussion which is

No. 441] NOTES AND LITERATURE. 629

avowedly unmetaphysical. The felt-need to say in the preface that

a scientific book is unmetaphysical is good evidence of the increasing

interest in philosophic problems among biologists. Furthermore, the

author who begins by assuring us that he is not going to be meta-

physical usually plunges at once into a discussion of metaphysical

problems with a naïveté which delights the technical philosopher. :

The ever increasing interest in the morphology of concepts is evi-

denced by Driesch in his attention to the meanings of the funda-

mental concepts with which he has to deal. However unsatisfactory

his general conclusions may be to the majority of his readers, he

has succeeded in pointing out certain problems which are worthy of

attention.

ROBERT YERKES.

The Biogen Hypothesis. !— Chiefly for the purpose of establishing

a clear working hypothesis as to the inner changes of the living cell

Verworn has attempted to make more precise the biogen hypothesis

based on the investigations of Hermann, Pfliiger, Ehrlich, Allen and

others, and to show the wide application of this to the active proc-

esses of cells. Biogen molecules, according to Verworn, occur in the

cytoplasm, not in the nucleus of the cell. Unlike albumen molecules,

they are ordinarily very labile. The nucleus, though containing none

of them, gives out material essential to their changes. The cyto-

plasm contains in addition to the biogen molecules reserve food mate-

rials and oxygen, the latter in weak combination. In hunger the

reserve food of the cell is first used and then certain biogen mole-

cules are sacrificed to others. To make good such loss food is appro-

priated and is made available to the biogen molecules through the

action of the enzymes. The stimulation of protoplasm consists in

changing its biogen molecules from a state of high lability to one of

low lability, a change brought on by oxidation. The recovery to the

state of high lability is an assimilative process that requires time, and

is represented by the refractive period in many operations during

which stimulation is impossible. Thus the stimulability of a mass of

protoplasm is a measure of the completions of the assimilative proc-

esses which repair the effects of stimulation so far as the biogen mole-

cules are concerned. The hypothesis thus affords a more or less

complete history for protoplasmic metabolism.

'Verworn, M. Die Biogenhypothesis. Jena, Fischer, 1903, Svo. vi + 114 pp.

630 THE AMERICAN NATURALIST. [Vor. XXXVII.

ZOOLOGY.

Animal Classification.! — Teachers of elementary zoólogy but more

particularly students of this subject are often given to crave a simple

classification of animals, and to fill this want Professor Wilder has

prepared a synopsis of the chief animal groups. The author has

disarmed criticism by his avowal that schemes of this nature are

mainly personal, but even in such outlines it is fair to expect consis-

tency and freedom from obvious error. Presumably the part on

vertebrates should be best written and yet by a strange coincidence

the Vertebrata (p. 39) is the only type to which no general definition

is given, and the classes of its gnathostome division are numbered

one to six with the omission of four. There is no reason to suppose

that the beginner would ever rightly determine, by the artificial key

at the end of the book, the groups to which such forms as the

bilateral sea-urchins and holothurians belong, for by the tables these

must come under “ Structure radiate.” The statement that follows

this, * Parts in 2s” would be a stumbling block to any one who knew

what bilaterality was. On the whole this key is so very artificial

that it is best used when one knows beforehand where the animal

belongs. Defects of the kind pointed out, while of no great impor-

tance to the advanced student, are serious matters with the beginner,

and render the tables much less useful than they should be.

Hertwig’s Manual of Zodlogy.’

Hertwig's Lehrbuch, the best German elementary text-book in

zoology, has been until recently accessible to the English-reading

student only through a partial and imperfect translation. Kingsley's *

new edition in English based on the fifth German edition will there-

fore be welcomed by many. A cursory examination of the new

volume shows that the American edition is likely to repeat the suc-

cess of its German forerunner. The translating is remarkably well

done and the general form of the book excellent. Here and there

exception may be taken to the course chosen by the translator. It

would have been better to have used exclusively the English term

cecology, which is coming to have a definite meaning, rather than the

! Wilder, H. H. A Synopsis of Animal Classification. New York, Henry Holt

& Co., 1902. gy 57 Pp-

? Hertwig, A Manual of Zoology. Vagos and Edited by J. S.

Kingsley, rs De Fifth German Edition. w York. Henry Holt & Com-

pany, 1902, 8vo., xii + 704 pp., 672 text moe dh

No 440] NOTES AND LITERATURE. 631

two terms cecology and biology (pp. 4 and 57), which, though

synonyms in German, are far from equivalents in English. The

choice of the form for proper names, if not of great importance,

would lead in English rather to Vesalius than Vesal (p. 12), a matter

in which the reader is given his choice with Galen and Galenius (p.

12). In discussing animal temperature poikilothermous, idiothermous

and homoiothermous are used without good reason, so far as we know,

for the more usual pcecilothermal, idiothermal, and homothermal.

The fact that the volume in its several editions has passed from one

century to another has led to some confusion which should have

been cleared up in editing; thus while we are correctly told (p. 17)

that the cell theory is of the “last century” and that the name Pro-

tozoa was given “in the century just closed” (p. 186), the “ Origin

of Species” is described (p. 24) as a “scientific work of this century.”

The proofreading has been unusually close; on page 13, line 28,

unbiassed is preceded by a useless dash and on page 435 Cumbarus

stands for Cambarus. The presswork and illustrations are as a rule

good, though many of the newly introduced, original figures, particu-

larly the half-tones, are too faintly printed. The defects that have

been pointed out are insignificant compared with the good qualities

of the volume, which deserves immediate acceptance as the best

general text-book of zoólogy for the majority of American colleges.

The Neurone Theory and its Adherents.' — Since the promul-

gation of the doctrine of the neurone by Waldeyer in 1891 numerous

general estimates of this theory have been advanced by almost all

the more noted workers in neurology. These expressions of opinion

have almost invariably come from advocates of the theory and have

been the means of introducing at most only slight modifications of

the general doctrine. Up to the present no single considerable pub-

lication has been devoted to a thorough review of the body of evidence

brought forward by the neuronists and to a radical and well directed

attack on their position. Nissl’s Weuronenlehre is such a publica-

tion.

The first chapter of this work takes up briefly Waldeyer's original

conception of the neurone and the modifications that during the last

ten years this has undergone. In deciding what the essentials of

the neurone theory are Nissl makes one of the clearest and most

pe issl, F. Die Neuronenlehre und ihre Anhänger. Ein Beitrag zur Lösung

Problems der Bersiehungen zwischen Nervenzellen, Faser und Grau. Jena,

nlp te 1903, Svo., vi + 478 2 Taf

632 THE AMERICAN NATURALIST. (Vor. XXXVII.

justifiable statements of the subject that has appeared. The neurone

theory is in essence the application of the cell theory to the complete

interpretation of nervous structures, in that the nerve fibres and the

neuropile are to be regarded as outgrowths and integral parts of ner-

vous cells whose bodies are represented by ganglion cells. Thus the

question of contact or continuity among neurones is set aside as

secondary and the real core of the matter is reached by the declara-

tion just given.

Following the introductory chapter come eleven others devoted

each to the exposition and rigorous criticism of the views of some

well-known neurologist; among the investigators whose opinions are

here analyzed are Edinger, Hoche, von Lenhossék, Van Gehuchten,

Ramón y Cajal, Kólliker, Verworn, and His. ‘The line of criticism

which pervades this part of the book consists in pointing out the

fallaciousness of the Golgi method and the failure on the part of the

neuronists to appreciate the full significance of the neuropile. The

Golgi method is notorious for incompleteness in its impregnations and

yet observations based upon it have been used again and again in

support of the idea that the neuropile is at least physiologically sep-

arable into discrete portions referable to given neurones. Since we

know so very little about the structure of the neuropile it would seem,

as Nissl rightly urges, that to pass it over simply as a terminal out-

growth of the neurone, or to ignore it almost entirely, as Verworn

does, is wholly unjustifiable. This treatment is all the more repre-

hensible because there is good reason to believe that the neuropile

may be the most important physiological element in the whole nerv-

ous mechanism.

The concluding chapters, eight in number, serve to develop Nissl's

own views as to the structure of the nervous system. These are

based largely upon the work of Apáthy and Bethe and centre chiefly

about the neuropile. The fibrillar network of the central gray, the

invasion of ganglion cells by the neurofibrillae, and the relation of

these to the pericellular Golgi network are discussed in much detail.

The scheme of nervous mechanism that Nissl constructs from

recorded facts is certainly in many particulars inconsistent with the

neurone doctrine. This doctrine was a happy suggestion as to the

relations of cells and fibres, but subsequent work on the nervous

system has shown that these elements are quite secondary and that

the real nervous material is the neurofibrilla. Since the neurone

theory does not touch these and since we know so little about their

anatomy and nothing whatever about their development, speculation

No. 441.] NOTES AND LITERATURE. 633

should be abandoned together with the insufficient neurone theory,

and facts concerning the neurofibrille should be sought. This in

general is Nissl’s position and it will probably carry to the mind of

the neuronist the conviction that if this is a fair example of what the

neurone theory will have to meet, that theory is still very safe.

Notes. — The earliest stages in the development of the teeth in

selachians have been investigated by Laaser (Jena. Zeitschr. f. Natur-

wissench, Bd. 37, pp. 551—578), who finds that in embryos of Spinax,

Acanthias, and Mustelus of three to four centimetres in length, a den-

tal ridge is formed by a thickening of the epithelium of the jaws.

The ridges are formed earlier in the lower jaws of Spinax and Acan-

thias and in the upper jaw of Mustelus. Teeth develop not only in

the dental ridges but also in the adjacent epithelium where in their

early stages they are indistinguishable from placoid scales. The

first hard part formed is the dentine, the enamel being entirely absent

at these early stages.

Professor Bastian (London, Williams & Margate. Pt. II. 1902,

pp. 63-147, pls.) presents in a second installment much additional

evidence in favor of his views on heterogenesis. Thus he believes

he has shown that vorticellae may be produced from a pellicle largely

composed of spirilla, that amoebae may be made to segment and

their parts be converted into ciliate infusoria, that the entire egg of

the rotifer Hydatina can be transformed into a ciliate infusorian

Otostoma, etc. The paper is illustrated by photographic reproduc-

tions but even these cannot shake the conviction of many zoólogists,

that because of the methods used something is probably wrong with

the observations recorded in the text.

Dr. J. Anglas has published as number 17 of the biological series

* Scientia” a clear account of the changes undergone by the tis-

sues during the internal metamorphosis of insects. The histogenesis

of early development is first taken up, then the process of histolysis,

and finally the reconstructive processes. The book contains a final

chapter on the causes of internal metamorphosis.

The origin and classification of leucocytes and a very readable

discussion of the theories of their relations to health and disease

have been published in the biological series of “ Scientia” numbers

15 and 16 by Dr. J. Levaditi.

Fischer ( Yena. Zeitschr. f. Natur wissensch, Bd. 37, pp. 691—726) has

634 THE AMERICAN NATURALIST. [Vor. XXXVII.

made a thorough-going embryological study of the carpus and tarsus of

the problematic mammal Hyrax. Since in the embryo the hind foot

shows traces of the first and fifth digits, the extremities of Hyrax

point to derivation from a primitive form with five digits. "The

embryonic carpus contains two centralia like the embryonic carpus

of the turtle. Traces of both prepollex and prehallux were found.

The carpus and tarsus of Hyrax must have been derived from a more

primitive form than Phenacodus. Since they show as many affinities

to the rodents as to the fossil ungulates, Hyrax has probably been

derived from some form in which these two types were united, the

Toxodontia, or possibly the more primitive Tillodontia.

The growth of micro-photography has been so rapid that the A B C

of the subject has been issued in a handy volume by W. H. Walmsley

(N. Y., Tennant & Ward, 1903. iv—155 pp., 13 pls.). Chapters are

devoted to the microscope, the camera, illumination, negative making

and printing. The experience of an expert, the high quality of

whose work is attested by the illustrations that accompany the

volume, is given freely to the beginner.

BOTANY.

Setchell and Gardners N. W. Alge.'— This is a careful and

thorough account of the marine Algz of the Pacific coast of America

from Cape Flattery north to the Arctic Ocean, and of the fresh water

species found near the shore through the same range, the Diatomacez

and Desmidiacez excepted. The information hitherto accessible has

been scattered through many books and papers in various languages,

and this is now brought together, but covers only the smaller part of

the present work, the rest being now presented by the authors for the

first time. This is specially the case as to the fresh water Algz, in

regard to which very little indeed is on record previous to this work.

Every species mentioned by previous writers is included in this list,

even if the authors consider the determination as unreliable, or that

! Setchell, W. A. and Gardner, N. L. Alga of Northwestern America. Univ.

Cal. Publications, Botany, Vol. 1, pp. 165-418; Pl. XVII-XXVII. Berkeley,

March 31, 1903.

No. 441] NOTES AND LITERATURE. 635

the plant in question is to be included under another name here ; this

makes the total number of species to be credited to the Flora some-

what uncertain; but leaving out about 5o forms, which may be con-

sidered as erroneously or uncertainly reported, the following species

or named varieties and forms will approximately represent the extent

of the Flora.

Fresh water. Marine.

Cyanophyceaz 99 26

Chlorophycez 65 76

Phaeophycez I 147

Rhodophycez 9 214

Total 174 463

This is really a much richer list than any one had before supposed

probable; the proportion of Cyanophycez is exceptionally large, com-

paring well with the same order in regions which have long been

studied by resident botanists. Dr. Setchell is well known as a spe-

cialist on the Cyanophycez, and in the expedition along the coast of

Alaska recognized many forms which would probably be overlooked

by most collectors. The Laminariacez are also well represented,

and to Dr. Setchell is due the clear presentation and arrangement

of these perplexing plants.

It is interesting to compare the Flora of the northwest coast with

that of the northeast coast of America. Comparatively few marine

species are common, but the proportion increases as we go north,

and the common species are mostly found also in northern Europe,

indicating a common arctic origin for all the high northern floras,

apart from this element there are a few cosmopolitan species, common

to both sides of the continent. A few species are common to the

Flora of eastern Asia, and a few are common to the European Flora

but not found on the west side of the Atlantic; the remaining species,

about half of the whole, in the case of the red Algæ more than half,

are, as far as known, limited to the Pacific coast of the United States.

As regards the fresh water Algae, the case is quite different; nearly

all the species are cosmopolitan, some in all latitudes, some in tem-

perate regions only; very few are limited to this region ; it is inter-

esting to note that most of the cosmopolitan marine species are of

the Cyanophycee and Chlorophycez, orders more largely fresh

water than marine. It is probably accidental that the four species

of Characez, all European, are reported from Alaska only, in the

extreme North.

636 THE AMERICAN NATURALIST. [Vor. XXXVII.

In the introduction the authors divide the entire west coast into

four quite well marked regions of algal growth; the Tropical, the

North Subtropical, the North Temperate and the Boreal; with the

suggestion that further study may make it necessary to divide

the latter into an Upper and a Lower Boreal. The approximate

boundaries are Magdalena Bay, Lower California; Point Concep-

tion, California, and Puget Sound. The present work includes

such of the Temperate element as appears in Puget Sound, and the

whole of the Boreal. The subtropical families Valoniacezee and

Dictyotacez are each represented by a single species. If the divi-

sion of the Boreal into upper and lower is adopted, each of these

regions corresponds to a range of surface temperature, there being a

variation of 5° C. as we pass from one to another; each region

having approximately a difference of 5? C. between the maximum

and the minimum. The division between the upper and the lower

Boreal has an isochryme of 5? C. and an isothere of 10? C. the

southern limit of the North Temperate having 20° C. and 25° C.

respectively.

As is to be expected from the latitude, the great Laminariacez are

the most conspicuous element of the Flora; this region probably

exceeds all others in the gigantic size of the individuals and the

variety of forms of this family. The genus Alaria is represented by

eighteen species and forms, one of which, 4. fistulosa, has a blade

reaching a length of 25 meters. The eighteen different genera of

Laminariacez form a very rich representation of the family. Litho-

. thamnion and the allied genera are well represented, comparing fav-

orably with other northern regions; while the jointed Corallinacez

have many forms, contrasting strongly with the single species found

on our Northeast coast.

While many individual collectors at various points have contributed

to this work, the greater part of the material on which it is founded

was obtained by the expedition from the University of California, in

the summer of 1899, on which Professor Setchell was accompanied by

W. L. Jepson, L. E. Hunt and A. A. Lawson; while it is certain that

additions will be made to the list by future explorers, the general

character of the marine Flora may be considered as fairly well estab-

lished.

The arrangement follows the system of Engler & Prantl in the

main; as to nomenclature, a very conservative course has been fol-

lowed, generic names long in use being retained, no effort having

been made to replace them by earlier but neglected or abandoned

No. 441] NOTES AND LITERATURE. 637

names; nor have specific names been changed unless the change was

unavoidable. “We have preferred to devote our time to the study

of the plant itself” the authors say, and certainly if the choice had

to be made, they have chosen wisely. We have an ample supply of

botanical literature, affixing the author's name to new binomials,

representing plants that the author would never recognize if he met

them. The authors of this work know their plants thoroughly, and

those who enjoy juggling with names, can do it at their leisure.

In the matter of specific limitations, there is quite a tendency to

broaden out a species, and give form names to what others would

consider autonomous species; not less than 142 “forme” being

named in this work, some representing former species, some being

newly distinguished. Laminaria, Alaria and Fucus give good exam-

ples of this practice; but perhaps the most striking are in Corallina

and Amphiroa; here the disappearance of former species is quite

startling. Two new genera, Whidbeyella and Collinsiella are pro-

posed, and nine new species; the authors propose nothing as a vari-

ety, recognizing the term only as used by other authors. Descrip-

tions of new species and forms are full and clear; there are eleven

good plates; the type and paper are excellent. Specimens are

referred to by collectors or exsiccatae members; there is a good

index, and a very full list of the literature of the subject. Exact

localities are given in almost every case, and there is an alphabetical

list of all the localities mentioned, with full indication of the latitude,

longitude, etc., of each ; this novel feature is contributed by Professor

George Davidson of the University of California.

No work of such general importance to this department of Ameri-

can botany has appeared since Harvey's Nereis Boreali-Americana,

fifty years ago; and while undoubtedly much will be added by the

subsequent studies of the active botanists who are doing such good

work on the west coast, it is unlikely that there will ever be any one

contribution that will contain as much new information as does this.

The authors deserve the thanks of all students of Alga.

FRANK S. CoLLiNs.

Notes.— The Proceedings of the. Society for the Promotion of Agri-

cultural Science, for the 24th meeting, contain the following articles of

botanical interest: Jones and Sprague, “ Plum Blight caused by the

Pear Blight Organism "; Saunders, “Some Results of Cross Ferti-

lizing,” and “Decrease in Vitality of Grain by Age”; Fernow,

“The Significance of the Farmer’s Woodlot” ; Pammel and Lum-

638 THE AMERICAN NATURALIST. [Vor. XXXVII.

mis, “The Germination of Weed Seed," and “Germination of

Maize"; Lummis, * Effect of Coal Tar, Coal Oil, Gasoline, Benzine

and Kerosene on Germination of Maize”; and Lazenby, “ Com-

position and waste of Fruits and Nuts.”

The American Botanist for May contains the following articles:

Dobbins, “Lycopodiums of the Green Mountains”; Gilbert, “A

New Fern from Bermuda [Asplenium muticum)”; Ryon, “ Poison

Ivy and its Effect”; and Barrett, “Deciduous Tropical Trees.” —

The editor’s “Botany for Beginners,” and a series of notes, con-

stitute a prominent feature of the number.

A new journal, Annali di Botanica, under the direction of Professor

Pirotta of Rome, has been started. ‘The first number, dated May 15,

contains articles on the development of the seed of Cynomorium,

description of a new Euphorbia, Æ. Va/liniana; a study of the

influence of climate and location on the structure of plants in the

Mediterranean region ; a study of the origin and differentiation of the

primary vascular elements of the roots of Monocotyledons, and notes

on Gherardo Cibo's herbarium and on a recently unearthed addition

to the herbaria of Liberato Sabbati.

The Botanical Gazette for June contains the following articles:

Sargent, “Crategus in Northeastern Illinois" ; Stevens, F. L. and

A. C., “Mitosis of the primary Nucleus in Synchytrium decipiens” ;

Bergen, “The Macchie of the Neapolitan Coast Region”; Butters,

* A Minnesota Species of Tuber " ; West, * A new botanical Research

Laboratory in the Tropics”; and, Ashe, “New or Little-Known

Woody Plants."

The Bulletin of the Torrey Botanical Club for June contains the

following articles: Harper, “ Botanical Explorations in Georgia

during the Summer of 1901, II Noteworthy species”; Bush, “A

list of the Ferns of Texas”; Eaton, Zsoefes riparia Canadensis and

I. Dodgei.”

The Journal of Mycology for May, with portrait of S. M. Tracy for

frontispiece, contains the following articles: Blasdale, “A Rust of

the Cultivated Snapdragon ” ; Morgan, * A new species of Sirothe-

cium”; Seymour, “ A Series of Specimens Illustrating North Ameri-

can Ustilaginee "; Morgan, “ Dictyosteliee or Acrasiee ” ; Murrill,

* Historical Review of the Genera of Polyporacez "; Durand, “ The

genus Sarcosoma in North America”; Ellis and Kellerman, * Two

new Species of Cercospora” ; Kellerman, * Another much-named

No. 441] NOTES AND LITERATURE. 639

Fungus”; “ Puccinia laterifes an Aut-Eu-Puccinia”; “ Alternate

Form of Aecidium hibisciatum”; “Ohio Fungi, Fascicle VII";

“Index to North American mycology ”; “Notes from mycological

literature, V ”; and editor’s notes.

The 18th volume of the Zransactions of the Kansas Academy of

Science contains the following articles of botanical interest: Sayre,

* Loco Weed"; Gould, * Notes on the Trees, Shrubs, and Vines in

the Southern Part of the Cherokee Nation”; Garrett, “A pro-

visional List of the Uredinez of Bourbon County, Kansas”; and

Smyth, “ Preliminary’ List of medicinal and economic Plants in

Kansas.”

The eleventh Annual Report of the Ohio State Academy of Science

contains short papers or abstracts on a number of botanical subjects.

The Popular Science Monthly for July contains the following

articles of botanical interest: Cook, “ Evolution, Cytology and Men-

del’s Laws”; and Zirngiebel, * The Preservation of Wild Flowers.

The Plant World for June contains the following articles : Safford,

* Extracts from the note-book of a naturalist on the Island of Guam,

VII"; Beattie, “Indian Hemp as an ornamental”; Williams, “A

collecting Trip to Bolivia”; Orcutt, “Uses of Cacti”; Mansfield,

“ [Osmunda regalis|” ; and Shear, “ Fungi on old Logs and Stumps.”

Rhodora for June contains the following articles: “ The identity of

tris Hookeri and the Asian Z. setosa” ; Sargent, “ Recently recog-

nized species of Crategus in Eastern Canada and New England,

IV”; Bissell, “A new station for Dentaria maxima”; Harvey,

“ Splachnum ampullaceum”; Evans, “Preliminary lists of New

England plants, XI, Hepatice”; Bissell, “ Galium erectum and

Asperula galioides in America”; Fernald, “Some variations of

Triglochin maritima”; and, Robinson, “A hitherto undescribed

Pipewort from New Jersey [Eriocaulon Par&eri ]."

Torreya for June contains the following articles: Cockerell,

* Notes on New Mexico Oaks" ; Earle, ^A Key to the North Ameri-

can Species of Panus"; Harper, “A new Arabis from Georgia” ;

Gleason, “A second Illinois Station for Phacelia Covillei” ; Harper,

“ Lycopodium cernuum in Georgia”; and, “ Britton, A new pecies

of Urera [ U. magna]. ”

Zoe for May contains the following articles: Brandegee “ Flora of

the Providence Mountains,” “Vegetation of the Colorado Desert,”

640 THE AMERICAN NATURALIST. [Vor. XXXVII.

" Notes and New species of Lower California plants," and ‘ Notes on

Papaveracee.” The number closes with a facetious review of an

entertaining recent publication on California botany.

As the opening number of Volume VIII of the Contributions from

the United States National Herbarium, Dr. Rose publishes a third

part of his “Studies of Mexican and Central American Plants,”

marked by his usual critical acumen. It is to be hoped that in the

various readjustments of the Government publication facilities, these

Contributions from the National Herbarium may not be neglected.

Part VI of Captain J. D. Smith's “ Enumeratio plantarum Guate-

malensium necnon Salvadorensium Hondurensium Nicaraguensium

Costaricensium," recently issued, consists of 87 octavo pages of

herbarium label records of recently collected Central American

plants. i

Fascicle 3, completing the 3rd volume, of Urban’s Symbole Antil-

/an@, issued in May, contains descriptions of miscellaneous genera

and species, by Urban, accounts of mosses, by Brotherus, Burmanni-

aceæ, by Urban, Ficus, by Warburg, Cruciferæ, by Schulz, and Sela-

ginellæ by Hieronymus.

A most valuable scientific treatise on the Bermudas, with an

extensive bibliography, by Professor Verrill, forms the second part

of the centennial volume, Volume XI, of the Transactions of the Con-

necticut Academy of Arts and Sciences, which is very fully illustrated

by text cuts and plates.

The economic grasses and forage plants of Idaho are the subject

of a paper, by Henderson, published as Bulletin No. 38 of the

Agricultural Experiment Station of the University of Idaho.

Notes on Faulkland Island plants, collected by Vallentin, are con-

tained in the Memoirs and Proceedings of the Manchester Literary

and Philosophical Society, vol. XLVII, pt. 3.

A lecture on the spring flora of Table Mountain, at the Cape of

Good Hope, by Engler, is issued as Appendix II to the Nożizblatt of

the Berlin Botanical Garden, under date of April rst.

Volume II, Fascicle 4, of Coste's 77oré descriptive et illustrée de la

France is devoted to a continuation of the Composite.

A revision of Chironia, by Schoch, is distributed as no. 19 of the

No. 441] NOTES AND LITERATURE. 641

Mitteilungen aus dem Botanischen Museum der Universität Zürich,

from the Beihefte zum Botanischen Centralblatt.

The species of Crataegus occurring about Rochester, New York,

are discussed by Sargent in a number of the Proceedings of the

Rochester Academy of Science, issued in June.

The species of Crataegus of Northeastern Wisconsin are discussed

by Schuette in the Proceedings of the Biological Society of Washington

of June 25th.

A preliminary paper on a natural arrangement of the species of

Ribes, by Janczewski, is issued as an extract from the Buletin inter-

national de l'Académie des Sciences de Cracovie, Classe des Sciences

mathématiques et naturelles, for May.

A new Solidago from the Yukon region is described by Gandoger

in the Buletin de la Société Botanique de France, issued on the 25th

May.

An interesting economic study of the species of Orobanche found

in the United States, by Garman, is published as Bulletin No. r05 of

the Kentucky Agricultural Experiment Station.

An excellent photogram of Yucca glauca accompanies an article on

the use of Yuccas for planting on sand dunes, in Arboriculture for

June.

Fletcher publishes, in the O//aza Naturalist, for June, some notes

on teratological specimens of Trillium grandiflorum.

An illustrated article on the Redwood is published in Forestry and

Irrigation for June.

Dr. Davis' paper on Oogenesis in Saprolegnia, printed in the

Botanical Gazette, has been included also in the Decennial Publica-

tions of the University of Chicago, and issued in separate form under

date of March 1, 1903.

A monograph of the Uredinez of Umbelliferze, by Lindroth, has

been separately issued from Volume 22 of the Acta Societatis pro

Fauna et Flora Fennica.

An addition to our knowledge of the fungus which occurs in the

caryopsis of Lolium, and to which the poisonous properties of

the latter have been attributed, is contributed, by Freeman, to

the Philosophical Transactions of the Royal Society of London,

Volume CXCVI.

642 THE AMERICAN NATURALIST. (VoL. XXXVII.

An attractive little treatise on the poisonous mushrooms of Europe,

with particular reference to France, accompanied by a wall chart

illustrating eight deadly or dangerous species, by Octave Grosjean, is

published by the author at Saint-Hilaire, near Roulans, France.

The root rot of the sugar-cane forms the subject of a thick quarto

volume, by Kammerling, published by van Ingen, of Scerabaia, Java.

From experiments conducted with the colon bacillus, Professor

Jordan concludes, in a paper on The Self-purification of streams,

reprinted from Volume X of the Decennia! Publications of the Univer-

sity of Chicago, that the enteric bacteria disappear almost completely

in less than 150 miles in a river like the Illinois.

A set of photomicrographs, accompanied by explanatory text,

illustrating the effects of Pseudomonas campestris on the turnip, are

published by E. F. Smith as Buletin No. 29 of the Bureau of Plant

Industry of the United States Department of Agriculture.

Sydow's Monographia Uredinearum, in its third fascicle, reaches

No. 879 of the species of Puccinia.

From an article by Leffmann, in the June Journal of the Franklin

Institute, it appears that Agar-Agar is considerably used in certain

grades of ice cream and jelly, and its use is said to be capable of

easy detection by means of Arachnodiscus and other diatoms which

are always found on it, even in prepared food articles.

Several important fern papers are contained in the Annals of

Botany for June.

The dehiscence of the sporangium of pteridophytes is further dis-

cussed by Steinbrinck in the Berichte der deutschen Botanischen Gesell-

schaft, issued May 27th.

The relations of the leaf bundles of Conifers to the thickening of

the stem are discussed by Tison in Volume II of the Mémoires de la

Société Linnéenne de Normandie, which also contains an article by

Lignier on the fruit of JW//iamsonia gigas and the Bennettitales.

The /rish Naturalist has recently been giving considerable space

to a discussion of the leaf markings of Arum maculatum.

Observations on the digestion of proteids with papain, by Mendel

and Underhill, are published in Vol. XI, part 1, of the Zransactions

of the Connecticut Academy of Arts and Sciences, recently distributed.

No. 441] NOTES AND LITERATURE. 643

The formation of oxalic acid in green plants is discussed by Benecke

in Heft 5, Abt. I., of the Botanische Zeitung.

Professor Davis considers the evolution of sex in plants in the

Popular Science Monthly for February.

The upper temperature limits of life are discussed by Professor

Setchell in Science of June 12.

An account of a new myrmecophilous plant, A/acaranga triloba, by

Smith, is published in ZZe Mew Phytologist of May 30th, which also

contains a number of other interesting morphological and ecological

papers,

A practical lecture on The use of timber by railroads and its rela-

tion to Forestry, by von Schrenk, is published in the Oficial Pro-

ceedings of the New York Railroad Club, for May, and is followed by

an address by Professor Fernow on railroad interests in forest

supplies, and a discussion of the subject.

A discussion of the Seasoning of timber, by von Schrenk and

Hill, constitutes Buletin No. 4r of the Bureau of Forestry of the

Department of Agriculture.

Economic seedling studies of Lilium harrisii, by Oliver, are pub-

lished as Bulletin No. 39 of the Bureau of Plant Industry of the

United States Department of Agriculture.

Economic notes on Ldgeworthia papyrifera, Aralia cordata and

Eutrema wasabi, by Fairchild, form Bulletin No. 42 of the Bureau of

Plant Industry of the United States Department of Agriculture.

Professor Halsted’s report on the botanical department of the New

Jersey Agricultural College Experiment Station, for 1902, contains a

number of articles on plant breeding and selection, the behavior of

mutilated seedlings, and parasitic fungi.

A study ef Northwestern apples, by Hansen, constitutes Bu/detin

76 of the South Dakota Experiment Station.

The teaching of botany in secondary schools is discussed by

several writers in the Journal of Applied Microscopy and Laboratory

Methods for June.

An account of the Department of Botany of Columbia University

and its relation to the New York Botanical Garden, by Underwood,

is published in the Columbia University Quarterly for June.

644 THE AMERICAN NATURALIST. (VoL. XXXVII.

According to a note by Hemsley in ature of May 21, the Kew

herbarium is now estimated to contain considerably more than

2,000,000 specimens, attached to 1,300,000 sheets, — and its greatest

value is qualitative rather than quantitative.

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BAILEY, FLORENCE M. Handbook of Birds of the Western eny States

ending the Great Plains, Great Basin, Pacific Slope, and Lower Rio Grande

alley. Boston, Houghton, Mifflin & Co., 1902. 8vo, xc + 512 pp., 33 ae -

text figs. $3.50.— BECK, C. and ANDREWS, H. Photographic Lens

Simple Treatise. London, R. & J. Beck, 1903. 8vo, 288 pp., 155 figs. mon

AGNES M. Problems in di xu London, Adam & Charles Black, 1903.

8vo, xvi + 567 pp., 81 illustr. $6.00. — HANN, J. Handbook of Climatology.

art I. General Climatology. "Translated from the German with additional Ref-

erences and Notes by R. De C. Ward. ew York, Macmillan, 1903. 8vo, xiv +

437 pp. 12 figs. $3.00.— HENSHALL, J. A. Bass, Pike, Perch sad Orken. The

American Sportsman's pep edited by TEE Whitney. New York, Macmil-

lan, 1903. 8vo, xxi 4 20 hae AM $2.00. — HoLpEN, E. S. The

Sciences. A Reading an i Children. Boston, Ginn & Co., 1903. 8vo, x +

224 pp., 198 figs. — LE DANTEC, ^ Traité de Biologie. Paris, Felix Alcan,

1903. 8vo, 553 pp., 101 figs. — LIVINGSTON, B. E. The Rôle of Diffusion and

Asmotic Pressurein Plants. University of Chicago ee Publications. Sec-

ond Seri Vol. viii. >. University of Chicago Press, 1903. VO,

xiii + 149 pp. $1.5c. — NEUMEIS’ R. Bet E M über das Wesen der

Lebenserscheinungen. ae bsg za zum Begriff des Protoplasmas. Jena, Fisher,

1903. 8vo, 107 p .2— Noyes, W. A. A Text Book of Organic Chem-

istry. New York, Bae Holt & x 4 1903. 8vo, xvii + 534 pp., 24 figs. —

PIERCE, G. J. A Text-Book of Plant Physiology. New York, Henry Holt and

Co., 1903. 8vo, vi J- 291 pp., 221 figs. — PLATE, L. Ueber die Bedentung des

Darwin "schen ae Ee und Probleme der Art ee Zweite ver-

merhete Auflage. Lei Engelmann, 1903. 8vo, viii + 247 p S., §. —

oe Y ., BISHOP, us B. and VAN DvkE, T. S. The b rod Fam-

New York, Macmillan, 1903. — 8vo, ix + 598 pp., 20 pls. American Sports-

man's Library. $2.00.—- WHITING, C. C. Walks in New ao pine London

and New York, John Lane, 1903. 8vo, 301 pp., 24 illustr. — WILLIAMS, R. P.

Chemical TPR a for Class Room and Home Study. Boston, Ginn "Ya 4 1903.

8vo, p

Feet E. The Soy Bean. Bull. R.I. Agr. Exp. Sta. No. 92, pp. 119-

127, 3 figs. — Banos, O. Description * a New Neotoma from Mexico. Proc.

Biol. Soc. Wash. Vol. xvi, pp. 89-90. — BARBOUR, T. A New Species of 2

Lizard from Sarawak, Borneo. ic. d Soe. deo jog xvi, pp. 59-60. —

BARBOUR, T. Two New Species of Chamzleo Z. Soc. Wash. pm

xvi, pp. 61- 62. — BARTSCH, P. A New Land shell "i a Proc. Biol.

Soc. Wash. Vol. xvi, pp. 103-104. — BassLER, R. S. The Structural Features

of the Bryozoan Genus re tee with Descriptions of Species from the Cin-

cinnatian Group. Proc. U. S. Natl. Mus. Vol. xxvi, pp. ig F 20-25.—

Bean, T. H. Catalogue of É TE of New York. Buli State Mus.

Zoos No. 9, 784 pp. — BE: .H. Notice of a S sisi d f ishes Made

. H. Brimley in Cane aay a Bollings Creek, North Carolina, with

645

646 THE AMERICAN NATURALIST. [VoL. XXXVII.

Descriptions of New Species of oe (N. Brimleyi). Prec. U. S. Natl. Mus.

Vol. xxvi, pp. 913-914. — BEARDSLEY, A. E. The Destruction of Trout Fry ie

Hydra. Bull. U. S. Fish. Comm. 1902. pp. 157-160. eben J.E. Rev

sion of the Crustacea of the Genus Lepidopa. Proz. U. S. Natl, Mus. Vol.

xxvi, pp. 889-895. — Buscx, A. A Revision of the American Moths of the Fam-

ily Gelechidz, with Descriptions of New Species. Proc. U. S. Natl. Mus. Vol.

pal. pp. 767- ei igs ic etnies F. W. Bush Fruits. Bull. R. S. Agr.

Exp. Sta., No. 91. 89-116, 4 figs. —Cary, M. A New Reithrodontomys

from Western Nébusko: Proc. Biol. Soc. Wits. Vol. xvi, pp. 53-54. — CAW-

DELL, A. N. ‘Notes on Orthoptera from Colorado, New Mexico, Arizona and

Texas with Descriptions of New Species. Proc. U. S. Natl. Mus ol. xxvi,

' pp- 775-809, pl. 55. — CAWDEL E ke N. The Phasmidae or Walkingsticks of the

United States. Proc. U. S . Mus. Nol. i, pp. 863-885, pls. 56-59. —

CLARK, H. L| Ihe Short- Nouthed Snake (kaise brachystoma Cope) in

Southern pes Proc. Biol. . Wash. Vol. xvi, pp. 83-89. — CORBETT, L.

C. and Davis K. C. Vegetable Fasea in the gapping Glades. Bull. Agr.

Exp. Sta. We a., No. 81. pp. 355-386, 10 figs. — Davis, W. M. The Stream

Contest along the Blue Ride. Bill. Geog. Soc. Pind Vol. iii, No. 5, p

213-244, 4 pls. — Dickson, C. W. Note of the Condition of Platinum in is

Nickel-Copper Ores from Pin: Amer. Journ. Sci. XV, pp. 13

DixoN, R. B. and KROEBER, A. L. The Native Dinni of California.

Amer. Anthrop. N. S. ` Vol. v, pp. 26, 9 charts. — Doran, E. W. The Ver-

nacular Names of Birds. 77e Auk. va. xx, pp. 38-42. — Durcnuer, B. H.

Mammals of Mt. Katahdin, Maine. Proc. Biol. Soc. Wash. Vol, xvi, pp. 63-72.

a DYAR, H. G. ist of North American aa and Key to the Litera-

ture of this Order by Insects. Bull. U. S. Natl. Mus - §2. xix + 723 pp. —

EAKLE, ‘A. E. Palacheite. Univ. Cal. Publ., Bull. iue Geol. Vol. iii, No. 9,

pp. 231-236, pl. 20. — EAsTON, B. S. The Constructive eats oss of Group-

Theory. Pubi. Uv. Pa. Miitheniation No. 2: pp. VW. E

Biologicai Reconnoissance in the Vicinity of Flathead Lake. ^ Univ. Mon-

tana, No. 10. pp. 91-182, pls. 17-46. — FELT, E. P. Crude Petrolium as an

Insecticide. Proc. 33d Ann. Meeting Soc. Promotion Agr. Sci. - 10. — FELT,

;rapevine Root Worm. Bull. N. Y. State Mus., Entomology No. 16.

pp. 49-34, 6 pls. — FELT, E. P. Eighteenth Report of the State San slo,

on the Injurious and other Insects of the State of New York. Bull. N. V. State

Mus., Entomology, No. 17. pp. 89-193, 6 pls. — FENNEMAN, N. M. On the

Lakes of Southeastern Wisco. Bull. Wis. Geol. & Nat. Hist. Surv., No. 8.

xv + 178 pp., 36 pls., 9 oo eru — FiGGINs, J.D. Some Food Birds of the

Eskimos of North Western Greenland. Proc. Linn. Soc. N. Y., 1900-1902. p

61—65. — FINLAY, G. The Granite Area of Vermont. Rept. Vt. State Geol.,

1902. pp. 46-50, 8 pis. — FISHER, R. T., von SCHRENK, H. and Hopkins, A. D.

The somit U. S. Dept. Agr. iiie d isis Bull. No. 38. pp. 40, 11

pls., 4 text-figs. — FISHER, W. Procelstoma from the Leeward

Islands vagis ant Group. Proc. U. e Pin Mus. Vol. xxvi, PP. 559-563. —

FOLEY, Conservative Lumbering at Sewanee, Tennessee. U. S. Dept. Agr.,

Bur: Forestry, Bull. No. 39. 36 pp., 12 pall pe — GILL, T. Note on the Fish

Genera Named Macrodon. Proc. U. S. t. Mus. Vol. x xxvi, pp. 101 5-1016. ais

Git, T. On the Relations of the a of the Family Lampiride or sons

Proc. U. S. Natl. Mus. Vol. xxvi, pp. 915-924, 3 figs.— GRANT, M. The

No. 441] PUBLICATIONS RECEIVED. 647

bou. From Seventh Ann. Rept. N. Y. Zool. Soc. 24 pp., 34 figs., 1 map. —

GRAVES, H. S. and FisHer, R. T. The d A Handbook for Owners of

bir o sup in Southern New England. sieh Dept. Agr., Bureau of Forestry,

Bull, 89 pp., 30 figs. — HALL, ges Practicability of Forest Planting

in the Unie — Yearbook U. S. ete Agr. for 1902. pp. 132-144, 5 pls. —

Hay o New Species of Fossil Turtles from Oregon. Univ. Cal. Publ.,

Full. 2 oe Vol. iii, No. 10, pp. 237-241, 5 figs. — Hay,

Small Collection of Crustaceans Potes the Island of Cuba. Proc. U. S. Natl.

vi, pp. fete 435. — HELME, A. H. Notes on the wee of

tine sland N PO roc. Linn. Soc. N. Y., 1900- pp- 19-30. ERRERA,

A.L. Le Rôle cM des Substances itii dans les Phéonine

SR Mem. Soc. Antonio Alzate, Nol. xiii, pp. 337-348. — HERRICK, C.

J. A Note on the Significance of the Size of Nerve Fibers in Fishes. ds.

Comp. Neurol. Vol. xii, pp. 329-334. — HERTY, C. H. A New Method of

Turpentine Satie rding. U.S. Dept. Agr., ses sa Forests ry, Bull. No. 40. 34

pp., 15 pls. — HINE, J. S. Tabanidæ of € o State Univ. Bull., Ser.

7, No.9. 57 pp., 2 pls. — IJ1MA, I. Se in i oscila Contribution

Ill. Your. Col Sci. Imp. Univ. Tokyo. Vol. xviii, pp. 124, 8 pls. — JORDAN,

D. S. and Fow , H. W. A Review of the Silanes Fishes of Japan. Proc.

^ S. Natl. Mas. AA xxvi, pp. 897-911, 2 figs. — JorDan, D. S. and FOWLER,

W. A Review of the Elasmobranchiate byst of Japan. Proc. U. S. Nail.

vedi Vol. xxxi, pp. 593-674, pls. 26-27. — JORDAN, D. S. and FowLEn, H. W.

A Review of the iiy iesu Fishes of Japan. "ue 0, S. Natl. Mus. Vol. xxvi,

,D. S. and FowLkn, H- W. A Review of the

Dragonets (Callionymidae) a Related Fishes of the Waters of Japan. Proc. U.

S. Natl. Mus. Vol. , PP. 939-959, 9 figs. — JORDAN, D. S. and Starks, E.

C. A Review of the ipn MUN Fishes of Japan. Proc. U. S. Natl. Mus.

Vol. xxvi, = 525-544, 3 figs. — JouNsoN, D. W. On Some Jurassic Fossils

from Durango, Mexico. Amer. Geol. Vol. xxx, pp. 370-371. — KEMPF, J. F.

Igneus Ro p and Circulating Waters as Factors in T Trans.

Amer. "ae Mining Engineers, 1902. 16 pp.— KNIGHT, A. P. Saw dust «n

Fish I Trans. Canad. Inst. Vol. vii, pp. 1-42,

4 d eek, N,

pner an Oligoclase-Corundum Rock near Spanish Peak, California. ae

Cal Pubi, Bull Dept. Geol. Vol. iii, No. 8, pp. 219-229. — LEHMANN, A.

Sur la Nature de l'activité des Nerfs. Bull. Acad. Roy. Danemark., 1903. pp.

205-233. — Lucas, F. A. Notes on the Osteology and PRA of the Fos-

sil Birds of the Genera Hesperonis, Hargeria, Baptornis an t Proc.

S. Natl. Mus. Vol. xxvi, pp. 545-556. — Lyon, M. W. eakop on the

Number oF Young of the Lasiurine Bats. Proc. U. S. Natl. Mus. Vol. xxvi, -

425-426, pl. 17. — McGREGOR, R. C. On Birds from Luzon, rtgni Masbat

Ticao, phe Culion, Cagayan and Polawan. Zu. Philippine Mus., 12

pp. — McGrecor, R. C. Notes on a Small Collection of Birds from dis Island

of Maui, Hawaii. Zhe Condor. Vol. iv, No. 3, pp- 59-62. — MCGREGOR, R.

C. A List of Birds Collected in Norton Sound, Alaska. Zhe Condor. Vol. iv,

pp. 135-144, 2 figs. — McMunnicH, J. P. Note on the Sea i Sagartia

Paguri Verril. Proc. U. S. Natl. Mus. Vol. xxvi, pp. 427-428. — RIAM, C.

H. Eight New Mammals from the Pase States. Proc. Biol. Soc. ges Vol.

xvi, pp. 73-78. —MERRIAM, C. H. ur New Mammals, Including a New Genus

(Teonopus), from Mexico. Prec. Fut Soc. Wash. Vol. xvi, pp. 79-82. — MER-

648 THE AMERICAN NATURALIST. [Vor. XXXVII.

RIAM J. C. New Ichthyosaria from the Upper Triassic of California. Univ.

Cal. Publ. Bull. Dept. Geol. Vol. iii, No. 12, pp. 249-263. pls. 21-24.— MONACO,

ALBERT PRINCE de. La Quatriéme Campagne Scientifique de la Princesse

Alice II. Compt. Rend. Acad. Sci. Paris. Tom. CXXXVI, pp. 211-215, I

NEEDHAM, J. G. A New "Genus and dped of Dragonfly from Brazil. Proc.

Biol. Soc. Wash. Vol. xvi, pp. 55-58. — OLMSTED, F. E. Tests on the Physical

Properties of ‘Timber. Yearbook U.S. Dept. Agr. for 1902. pp. 533-539, 2 pls.—

PRENTISS, D. em Description of an Extinct Misk from the Shell Heaps of the

ps Coast. roc. U.S. Natl. Mus. Vol. » pp. 887-888, 1 fig. — PRICE,

WwW ssa cd of Forestry upon the L eri dicun Yearbook U. S, Dept.

Agr. agg 1902. pp. 309-312, 3 pls. — RICHMOND, C. W. Birds Collected by es

bbott on the Coast and Islands of Northwest Sumatra. Proc. U. S. Natl.

je Vol. xxvi, pp. 485-524. — RICHARD, J. Sur l'état actuel du Musée

anographique de Monaco et sur les travaux qui s'y mt Campagne

Men du yacht Princesse-Alice en 1902. Bull. . Zool. France. Ann.

1903. pp. 57-79. — Ro A. Some New aile Species of kg

from the Coal Measures. (Wiss Univ. Sci. Bull. Vol. i, pp. 269-275, pl. 1

ROOSEVELT, T., MCCORMICK, R. L. and PiNcHoT, G. Forestry and the ^» um-

d — E S. Dept. Agr., Bureau of Forestry, caer No. 25. -X4. pp. —

Y, J. e Mammals of Westchester County, Proc. Linn. Soc.

is pp. 31-60. — SAUERWEIN, C. L’ uae Bordeaux

Société eke du Golfe de Gascogne, 1903. pp. 37. — SCHALLER,

'T. Minerals from Leona Heights, Alameda Co., California. saei Cal. Publ.,

Dept. Geol. Bull. Vol. iii, No. 7, pp. 191-217, pl. 19. — SCH H. von

Seasoning of Timber. U. S. Dept. Agr., Bureau of Forestry iw o. 41. 48

pp., 18 pls. — ScHUETTE, J. H. The Vie d ha dpi Wisconsin.

Proc. Biol. Soc. Wash. Vol. xvi, pp. 91-98. — SEWARD, .and ARBER, E. A.

N. Les Nipadites des Couches Eocénes de la dicus Mowe. Mus. as Hist.

Nat. Belg. Tom. II, 16 pp., 3 pls. — SHIMEK, B. The Loess of Natchez, Miss.

Amer. Geol. Vol. xxx, pp. 279-297, pls. 1c-16. — SuiMER, H. W. tro-

viste E DARE of gré Dikes of Grand Isle, Vermont. Rept. Vt. State pin

1902. 17 5-185, map. — SINCLAIR, W. J. New Tortoise from the Auri

ous Gerds of California. Univ. Cal. Publ., Bull. Dept. Geol. Vol iii, No. 11,

pp. 243-248, 2 figs. — SMrrH, H. M., and HARRON, L. G. Breeding Habits of

the Yellow Cat-Fish. Bul. U. S. Fish Com. 1902. pp. 149-154. sir eitis

T. R. R. Amphipoda from Costa Rica. Proc. U. S. Natl. Mus. Vol. xxvi, pp-

925-931, pls. 60-61. — STEJNEGER, L. Rediscovery of one of Holbrook's Sala-

manders. Proc. U. S. Natl. Mus. Vol. xxvi, pp. 557-558. — STEUART, J. H.

and ATWOOD, H. Poultry Experiments. Bull. Agr. Exp. Sta. W. Va. No. 33-

pp- 443-465, 3 pls. — WASHBURN, F. L. Seventh Annual Report of the Ento-

mologist of the State Experiment Station of the University of Minnesota. 73 pp-;

62 figs. — WESENBERG-LUND,C. Sur l'existence d'une faune relicte dans le Jac

de Furesó. Bull. Acad. Roy. Danemark, Ann. 1902. pp. 257-303, 2 m

YERKES, R. M. The Instincts, Habits and Reactions +i ‘ber rog. ead

Psychological Studies. Vol. i, pp. 579-638, 9 figs. — Yerkes, R. M. Habit

Formation in the Green Crab, Carcinus granulatus. , Biol. Bull. Vol. iii, pp.

1-244. — YERKES, R. M. and HucaiNs, G. E. Habit Formation in the Craw-

i Cambarus affinis. Harvard Psychological Studies. Vol. i, pp. 565-577. 4

figs.

No. 441. PUBLICATIONS RECEIVED. 6

Boletim de Museu Paraense. Vol. iii, Nos. 3-4, 1902. — Brooklyn Institute of

Arts and Sci Children’s dri Bulletin, No. 6, Mcl. — Bulletin of The

Fohns pM pees ıl. Vol. xiv, Nos. 146-148, May-July. — Comision de Par-

asitologia Agricola [Mexico]. Las Plagas de la Agricultura, Entrega. 6-8. —

Memorias do Museu Goeldi (Museo Paraense de Historia Naturale rom

iii. — New Century. Vol. vi, Nos. 30-31. — La Nuova Notarizia. Ser Apr.

1903. — AVut-Grower, The. Vol. 1, No. 8, March. — Papoose, The. Bae

May. — Proceedings of the Natural Science Association of Staten Island. Vol. vii,

No. 2 Vol. viii, Nos. um — Quarterly septi ds of Books Reviewed in

ing American Paks Vol. 1, No. 2, Jan.-J]une, 1902. 100 pp. —

"unm Chilena de Historia poe Ano. vii, NA i. zc Seil Science. Vol.

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NATURALIS

A MONTHLY JOURNAL |

DEVOTED TO THE NATURAL SCIENCES

Co ge eae

Past-Offics,

Sei

The American Naturalist.

ASSOCIATE EDITORS:

“Js A. ALLEN, PH.D., American Museum of Natural Hi: gees ne York.

5 = = E. A. Tipp PH.D., Johns Hopkins University, B.

— — WILLIAM S. BAYLEY, Pu.D., Colby University, RU

thac

d Univ Dey inbi

-A ; . National Museum, Washington.

D. S. JORDAN, LL.D., Stanford University.

CHARLES A. KOFOID, PH.D., University z Cali dis Berkeley.

J. G. NEED TAM, erp. Lake Forest Univ

THE

AMERICAN NATURALIST.

VoL. XXXVII. October, 190}. No. 442.

ADAPTATION TO AQUATIC, ARBOREAL, FOS-

SORIAL AND CURSORIAL HABITS

IN MAMMALS.’

I. AQUATIC ADAPTATIONS.

RAYMOND C. OSBURN.

THERE seems to be no doubt whatever that all mammals were

originally terrestrial animals. However, either the abundance

of food occurring in the water or the competition existing on the

land has from time to time led or driven many species to an

! In the American Naturalist of May, 1902, I published an article entitled ** The

Law of Adaptive Radiation," a development of the idea of divergent evolution

as applied to the larger and smaller groups of mammals. There was considered

first, general adaptive radiation including the radiation of marsupials and the six

independent radiations of the Placentalia, second the law of /oca/ adaptive radiation

and finally the bearing of adaptive radiation on Cuvier's /aw of correlation.

Pursuing this general idea of adaptive radiation it appeared desirable to reéxamin

and compare the mammals as to the adaptations of different kinds which arise

independently in different groups, in other words the imd adaptations.

A number of advanced students of the evolution of mammals undertook this

comparison and the results were so interesting and in many cases so novel that

they appeared worthy of publication in the American Naturalist. They form the

basis of the three or four articles which the Aafwra/ist will publish d

HENRY FAIRFIELD OSBOR

July, 1903.

651

652 THE AMERICAN NATURALIST. [Vor. XXXVII.

aquatic life. The following list, though making no pretensions

to completeness, will serve to indicate how great and varied a

number of forms have become, either from choice or necessity, :

more or less aquatic.

Cetacea, the entire order.

Sirenia, the entire order.

Carnivora,

Pinnipedia, the entire suborder.

Fissipedia, `

Lutra, the various species of otters.

Enhydris (Latax), the sea-otter.

Putorius, the mink and sumpfotter.

Rodentia, many scattered examples, as —

Myocastor (Myopotamus).

Hydrochoerus, the capybara.

Hydromys, the Australian water-rat.

Hydrochilus.

Microtus (Arvicola) the water-vole.

Ichthyomys.

Castor, the beavers.

Fiber and Neofiber, the muskrats.

Insectivora, many scattered examples, as —

Myogale, the desman.

Crossopus (Sorex) the water-shrew.

Neosorex, the American water-shrew

Chimarrogale.

Nectogale.

Potamogale.

Limnogale.

Ungulata,

Artiodactyla,

Hippopotamus.

Marsupiala,

Chironectes, the water-opossum.

Monotremata,

Ornithorhynchus.

Besides these there might be mentioned among extinct forms

the Zeuglodontidz, thoroughly aquatic animals doubtfully related

No. 442.] HABITS IN MAMMALS. 653

to the Cetacea, or according to some authors, to the Pinnipedia ;

the oreodont Merycochoerus, probably semi-aquatic, the creodont

Patriofelis, a possible ancestor of the Pinnipedia, and the extinct

rhinoceros, JZetamynodon planifrons Scott and Osborn, which,

though related to modern species which are not aquatic, had the

shorter nasals, more dorsal external nares and more dorsally

curved ribs which mark many aquatic forms, as will be shown

later.

That many forms have but recently become aquatic is certain

in such as have undergone little or no modification in structure,

while others must have assumed aquatic life in very remote

times if the amount of the adaptation to the environment is any

criterion. Kiikenthal points out that the amount of adaptation

depends upon the length of time during which the influence of

the water has been operating and upon the amount of connection :

retained with the land, and that we can assign the relationships

of aquatic mammals with certainty in proportion to the time that

has elapsed since they were separated from their terrestrial

relatives. For example, we know definitely the relationships of

the forms that have recently taken up the aquatic habit, such as

the otters, muskrats, etc., of the Pinnipedia we know that they

are Carnivora, of the Sirenia we know almost nothing (according

to various authors they have been held to show relationships

with the Ungulata, which view the evidence largely favors, the

Cetacea, or to form an entirely distinct stem.), and as to the

Cetacea we are absolutely in the dark. Kiikenthal even main-

tains that the latter group is diphyletic from quite different

sources and that the supposed relationships between the Odon- -

toceti and the Mystacoceti are merely parallelisms due to simi-

larity of environment. The reason for this is to be found in the

profound modification resulting from life in the water which

affects internal as well as external structures and leads to par-

allelisms in many structures in forms not genetically related.

For convenience in treatment, the various adaptations may for

the most part be roughly classified in three groups as follows : —

I. Adaptations connected with the general body form including

those of the head, trunk and tail regions. II. Those affecting

the limbs. III. Those affecting the integument.

654 THE AMERICAN NATURALIST. [Vor. XXXVII.

I. The tendency of the body to take on a **fish-like " form

is too well known to need any discussion, as it goes without

saying that such a form is best adapted to progress in the water.

It is most marked in such animals as are most aquatic, as the

Cetacea, Sirenia and Pinnipedia, and to a less extent in other

forms as Enhydris, Potamogale, etc. The anterior part of the

body tends to become more rigid and concentrated, especially in

the cervical region, while the posterior part of the body becomes

more mobile for purposes of propulsion.

In the head there is found in the Cetacea and the extinct

zeuglodonts a lengthening of the face with a shortening of the

cranium. In the most extreme cases the head is nearly one

third the length of the body. In the Sirenia the face is some-

what elongate, but not excessively so, while in the Pinnipedia it

is never very elongate and may be quite short, while the cranium

is broad and flat. The length of the head is conditioned so

largely by the length of the jaws that it would seem that its

shape is not a result merely of life in the water, so much as

of adaptation to certain kinds of aquatic food. The secondary

simplification of the teeth which takes place in all truly aquatic

mammals must also be connected with food conditions. In the

Mystacoceti the teeth are never functional but are present only

in the embryo and are absorbed before birth and replaced by

whalebone. As the food consists of very minute forms the

adaptation is evident. In the Odontoceti or toothed whales the

teeth are purely raptorial in character, simple and fang-like and

often retroverted, and admirably adapted for the capture of the

food, which in most cases consists of cephalopods, crustaceans

and fishes. The number of teeth may be greatly increased as

in Globiocephalus where the total number may be over roo, and

even twice that number may be found in Delphinus and Inia, or

the number may be greatly reduced as in Monodon and Ziphius,

or the teeth may be vestigial as in Hyperoodon. In the extinct

zeuglodonts the teeth had not become so simplified, as they still

possessed two roots and a crenulated crown. This **zeuglo-

dont” condition of the teeth is found at the present time in the

majority of the Pinnipedia. Inthe herbivorous Sirenia the teeth

when present are of the bilophodont type, undoubtedly adaptive

No. 442.] HABITS IN MAMMALS. 655

to the food, which consists of aquatic plants. In the recently

extinct genus Rhytina the teeth are absent and are replaced by

horny plates. In the manatee they are secondarily increased in

number. Along with the simplification of the dentition is found

occurring a great simplification of the jaws, particularly the

lower which tends toward the loss of all prominences for the

attachment of muscles, so that in the Cetacea the coronoid

process is often greatly reduced and the angle as well The

articulation with the squamosal becomes loose and simplified, and

the symphysis does not ankylose except in a few cases such as

Platanista. Even in some of the. Pinnipedia there occurs a

noticeable weakening of the jaw and looseness of the symphysis.

However, in the walrus whose food consists chiefly of bivalve

molluscs, the teeth are adapted to crushing the shells and the

jaw is remarkably heavy and strong and in the adult the sym-

physis is thoroughly ankylosed. The crushing jaw of the

Sirenia is also ankylosed. In general the pterygoid processes

also tend to become reduced, and there is a tendency toward a

looseness of articulation in all the bones of the head, particu-

larly in the Cetacea.

The shifting of the external nares from a terminal into a more

dorsal position is an adaptation to breathing at the surface of the

water. This is accomplished by a shortening up of the nasal

bones, which in the Cetacea become merely vestiges on the

anterior surface of the frontals. In the Cetacea the opening is

so shifted as to lie quite on top of the head, while in the Sirenia

and Pinnipedia also the shifting is quite noticeable. It is a note-

worthy fact that the true seals, Phocidze, and the dugong, Halicore,

which give other indications of a longer life in the water than

the eared seals, Otariidae, and the manatee, Trichechus, respec-

tively, have the external nares also more dorsal. The hippo-

potamus and the extinct rhinoceros, Metamynodon planifrons,

also show a somewhat dorsal position of the nares. In many

cases the external narial opening can be closed to exclude the

water, as in Sirenia and Cetacea. In the Odontoceti the two

external nares fuse into a single opening before reaching the

surface, a condition paralleled by some of the crocodiles.

The internal nares are also shifted backward to bring the

656 THE AMERICAN NATURALIST. [Vor. XXXVII.

opening more nearly over the epiglottis, and along with this in

the Cetacea is found an intranarial epiglottis! formed by the

prolongation upward of the epiglottis and the arytenoid cartilage

into the nostril, forming an independent continuous passage from

the exterior to the lungs. These are undoubtedly adaptations

to the capture of food in connection with conditions of respiration

at the surface of the water and permit of food being held in the

mouth or even of being swallowed without interfering with

respiration. In the Cetacea, naturally, the process is carried

farthest, and here the extreme is found in such forms as Globi-

ocephalus in which the postnarial opening is secondarily bridged

over by the pterygoids and the palatines are entirely excluded

from the anterior border. In Delphinapterus the palatines take

part for a small space in the formation of the anterior border of

the opening, and from this condition we find successive steps to

that found in the less modified forms, as the Pinnipedia, where

the whole border is formed by the palatines. The palatines are

posteriorly elongated as the first step in the backward shifting of

the internal nares, as shown by the Pinnipedia, Ornithorhynchus,

etc. The reduction of the salivary glands is also to be noted.

As the function of the saliva is chiefly a mechanical one con-

nected with deglutition the reduction of the glands in forms tak-

ing their food in the water is easily explained.

The loss of the external ears is another noticeable result of

aquatic life. In the Cetacea, Sirenia and Phocidz the external

ear is lost entirely, and among the eared seals, Otariidze, it is

found in various stages of reduction. The opening of the ear is |

often valvular so that it can be closed when in the water, and

this condition occurs even among those forms which are only

semi-aquatic, as in Crossopus and Neosorex. There isa tendency

among certain forms, also, toward the arrangement of the ears,

eyes and external nares in one plane near the top of the head so

that all may come into use at once without exposing very much

1 Possibly this is to be looked upon as the persistence of a larval structure, as

Howes has found an intranarial epiglottis in the young of a number of the more

primitive mammals having a forced lactation by means of mammary muscles, and

it is perhaps more than a mere coincidence that the whales have also a forced

lactation.

No. 442.] HABITS IN MAMMALS. 657

of the head. This is most marked in the hippopotamus but is

seen also in the capybara and beaver and other forms. In such

forms the éye-sockets may be quite prominent.

The shortening of the neck is another manifestation of the ten-

dency to take on a **fish-like" form, in connection with the ques-

tion of locomotion. In all truly aquatic forms this is noticeable.

It is brought about by a great shortening up of the cervical

vertebrze, and in extreme cases the loss of a vertebra (manatee) or

the fusion of some or all of the cervical vertebrae may take place

(most Cetacea). The occipital condyles also tend to become

flattened out and the odontoid process is reduced. The final

result of this process is an almost complete lack of motion

between the head and trunk, a condition finding its parallel in

the fishes, ichthyosaurs and other truly aquatic forms. This

loss of motion in the cervical region is more than compensated

for, however, by the greatly increased power of motion attained

by the more posterior portion of the body. Here the interverte-

bral connections are simplified and the vertebral column rendered

more mobile, since for the purpose of swimming, mobility of a

certain sort in the posterior part of the body is most useful.

The zygapophyses are progressively reduced and lost posteriorly

in the Sirenia and Cetacea, and other processes such as ana-

pophyses are entirely lost. Also the pleurapophyses or ribs of

the sacral region are lost as the pelvis loses its connection with

the sacrum (Sirenia and Cetacea). The spinous processes tend

generally to reduction, as in the cervical and anterior dorsal

region there is no need for the strong supporting muscles and

ligaments of terrestrial forms, and connections for leaping mus-

cles are lost posteriorly. The spinous processes of the poste-

rior body region and anterior caudal region seem to be second-

arily elongated, probably in connection with the up and down

motion of the tail in swimming, in many Cetacea, and the chevron

bones of the anterior caudal region in some forms are also elon-

gated. The centra of the vertebrae become amphiplatyan in the

Sirenia and Cetacea throughout most of the column, and the

intervertebral cartilages become thicker especially posteriorly.

The epiphyses, also, tend to unite at a very late period.

In all truly aquatic mammals the thorax takes on a character-

658 THE AMERICAN NATURALIST. [VoL. XXXVII.

istic cylindrical form, there being little or none of the lateral com-

pression such as is common among land forms, and this seems

to be the first step in the enlargement of the chest capacity, as

it is found in Pinnipedia as well as in the Sirenia and Cetacea.

The ribs at first tend to become highly arched dorsally and then

to move upward in their point of attachment from the centra to

the transverse processes. The beginning of this process is

found in the Pinnipedia and its culmination is seen in the whale-

bone whales, Balaenoidea, where all the ribs are attached only

to the transverse processes of the vertebrae. Possibly this is of

service in equilibration as the lungs can take a more dorsal posi-

tion. Accompanying these changes the diaphragm becomes

much more oblique and much more strongly muscular, undoubt-

edly giving greater control over the chest capacity in the peculiar

conditions of respiration necessarily accompanying aquatic life.

Perhaps the most striking external adaptation to aquatic life

is the assumption of **fins" for use in swimming. Many of the

Cetacea have developed a fleshy dorsal fin which undoubtedly

serves the same purpose as the similar organ among the fishes

and ichthyosaurs. The Sirenia and Cetacea have a large

expanded caudal fin supported by a dense framework of connec-

tive tissue and used as a propeller in swimming. This organ

differs from that of the fishes in being expanded laterally instead

of vertically, and this arrangement of the fin permits the animal

to rise to the surface more quickly for air and to dive again as

readily, and it accounts for the peculiar undulatory motion so

noticeable in these animals when they are swimming at the sur-

face. The flukes of the tail are said to be capable of a some-

what rotary sweep like the blades of a screw propeller at each

stroke of the tail. It is a noteworthy fact that nearly all

. aquatic mammals have this dorso-ventral flattening of the tail,

the only exceptions being Potamogale, Myogale and the musk-

rats, Fiber and Neofiber which have the tail expanded vertically

after the manner of a salamander. Potamogale is said to swim

like a newt with the legs folded against the body, and to be an

exceedingly rapid swimmer. The tail is so large and strong and

functions so completely as a propeller that the limbs have not

been modified even to the extent of webbed toes although the

No. 442] HABITS IN MAMMALS. 659

animal is quite aquatic in its habits. In Chimarrogale, Necto-

gale and Crossopus there is a stiff fringe of hairs which may serve

the same purpose as the compression of the tail. The caudal

vertebra are flattened laterally in Ornithorhynchus, Castor,

Halicore, and some Cetacea.

II. In the limbs, which at first play such a part in locomo-

tion in the water, we should expect to find marked adaptation.

Naturally those forms which move about occasionally upon the

land have the limbs less modified than those which have become

more completely aquatic, as progress upon the land calls for an

entirely different form of limb from that which is most useful in

the water. In some animals semi-aquatic habits have been

assumed while as yet no modification of the limb has taken

place. Thus in the water-vole, Microtus amphibius, there is no

trace of a web connecting the toes. Some other forms, as the

muskrats, have only rudiments of webs at the base of the toes,

while the minks have the toes partially webbed. The beavers

and the water-opossum, Chironectes, have the hind feet large

and fully webbed, while in the otters, the sea-otter and the duck-

bill, Ornithorhynchus, both feet are webbed to the claws. In

the more completely aquatic Pinnipedia the membranes, sup-

ported by connective tissues, reach beyond the digits, but the

claws are still present. In the Sirenia the whole forearm becomes

enclosed in the membrane and the last vestiges of the claws are

seen. The manatee retains slight vestiges of three nails,

(T. inunguis questionably), but the dugong and the extinct

Steller's sea-cow, Rhytina stelleri, lack the nails entirely. The

whales reach the climax of modification, as all of the limb out-

side of the body becomes enveloped, and the nails are entirely

lost (Leboucq has described the nails in the embryo). The

limb now becomes simply a balancing organ and the transition

from an ambulatory to a natatory limb is complete. Kiikenthal

aptly says, “In exact ratio to the adaptation to aquatic life do

we find the membranes just indicated or uniting the digits or

enveloping them entirely.” In lieu of webbing there is some-

times developed a stiff fringe of bristly hairs on the margins of

the feet as in Crossopus. This may be carried still farther by

the addition of a fringe of hairs on the sides of all the toes, as

660 THE AMERICAN NATURALIST. [Vor. XXXVII.

in Chimarrogale, or in addition to the fringes the foot may be

widened by disc-like pads and at the same time be webbed as in

Nectogale and Myogale.

Another transformation due to life in the water is found in

the shortening of the arm and forearm and lengthening of

the digits. This begins in the Pinnipedia where the arm is con-

siderably reduced in length though it is still serviceable to some

extent in locomotion and in most cases capable of supporting

the weight of the body. In the Sirenia and Cetacea, where

progress upon the land has been entirely given up, the arm is

still more reduced and in the latter group may be entirely with-

drawn into the body wall, only the hand remaining outside to

form the fin. In both these groups, but in the latter especially,

the arm no longer functions as a propelling organ but serves, as

in most fishes, merely as a balancing organ, the greatly devel-

oped tail furnishing the motive power. The extreme of adapta-

tion in the hand is reached in the addition of extra phalanges in

the digits, hyperphalangy, and the addition of an extra digit,

hyperdactyly, thus increasing the extent of the hand. Hyper-

phalangy is common among the Cetacea, where as many as

twelve phalanges may occur in a single digit, and even a greater

number than this in one species, Globiocephalus melas. It

apparently does not occur in other aquatic mammals except

occasionally an extra phalanx may be found in Sirenia, but a

close parallelism is seen in the ichthyosaurs and plesiosaurs.

Kükenthal accounts for hyperphalangy as a result of retarded

ossification and the formation of double epiphyses. These

epiphyses tend to ossify at later and later periods and finally

to become entirely separate bones forming the extra phalanges.

As compared with other theories that have been advanced to

account for hyperphalangy, this theory of Kiikenthal, strength-

ened by the array of facts which he brings to its support,

seems most reasonable and sufficient to account for the condi-

tions in the Mammalia at least. Hyperdactyly is not common

in the Mammalia but it is known to occur in some of the

Cetacea, e. g. Delphinapterus leucas, the white whale, by a

splitting of the fifth digit, as shown by Kükenthal and Leboucq.

In the ichthyosaurs the process went much farther, several sec-

ondary digits being formed.

No. 442.] HABITS IN MAMMALS. 661

Along with the change of function in the anterior limb occurs

the absence of clavicles, the reduction of the sternum and the

reduction and loss of articulation between the bones of the limb.

The distal elements, carpals and phalanges, tend to become

separated and imbedded in cartilage so that there is only a gen-

eral flexibility of the hand, but the humerus, radius and ulna

beco ne exceedingly short and lose all motion upon each other so

that there is not only no torsional motion of the radius and ulna

but also no motion in the elbow joint. As a final expression of

this the three bones often become ankylosed at the elbow in the

Cetacea. There is also a noticeable flattening of the bones (the

humerus is flattened distally only) and, connected with the change

in function of the limb, there is a reduction of prominences for

the attachment of certain muscles. Thus in the Sirenia and

Cetacea the deltoid crest and the olecranon process become very

much reduced. The entepicondylar foramen is also absent.

The spreading out of the ulna and radius permits the intermedium

to move up well between the distal ends of these bones in some

of the Cetacea, a condition not found in any other group of

mammals but paralleling the condition in Amphibia and many

reptiles. The elongated sickle-shaped hand of Globiocephalus

and some other cetaceans is paralleled by that of the Thalatto-

suchia and Ichthyosauria among the reptiles. j

In the hind limb the story is somewhat different, according to

whether or not the form in question possesses a tail fin. In the

Sirenia and Cetacea which have this organ there is no function

remaining for the hind limbs, as the balance can be preserved

by the anterior limbs and the tail furnishes the motive power, and

as a result the hind limbs are entirely lost, with the exception of

the merest vestiges of the skeletal structures which have lost

all connection with the vertebral column and are not at all visible

externally. The reduction process in the pelvis begins even in

the seals as the ilium never unites solidly with the sacrum as it

does in land forms. In all the Pinnipedia, a tail fin not being

present, the hinder limbs have gradually moved backward to

assume the function of a propeller and a motion somewhat similar

to the tail of the cetacean. In the Phocidz, the true seals, this

process has gone so far that the limbs have become quite bound

662 THE AMERICAN NATURALIST. [Vor. XXXVII.

up with the tail and are entirely useless for locomotion on the

land. It was this similarity in action and arrangement that led

Ryder in 1885 to derive the flukes of the tail in the Cetacea

and Sirenia from the hinder limbs, — a position no longer tenable,

as they are in no sense homologous. In the hair seals, Otariidee,

and walrus, Trichechide, the hinder limbs have not undergone

so much modification and are capable of being turned forward in

progression on the land, in which operation they are functional

to some extent. With the change from the ambulatory to the

natatory limb there comes about necessarily a great change in

the musculature of the limb.

III. The changes connected with the integument may be

noted as follows, — loss of hair, acquisition of blubber, loss of

the integumentary glands, smooth muscles and nerves of the

skin, and loss of dermal armature. The loss of hair is usually

not marked in those forms which spend only a portion of the

time in the water, but in the more completely aquatic forms

there is almost an entire absence of it. In the *hair seals,"

Otariida, which are the least aquatic of the group, there is a

fairly good coat of hair, and in the case of the fur-bearing species

this is intermingled with a dense coat of fine fur, but in the

walruses, Trichechidz, and “true seals," Phocidze, there remains

only short, appressed, coarse hair. In the hippopotamus and

the Sirenia there remains but very little hair in the adult,

and again in the Cetacea we find remaining usually only the

merest vestiges and sometimes these occur only in the embryo.

Kükenthal has pointed out that all these forms are distinctly

more hairy in the embryo than in the adult (except in the case

of the white whale, Delphinapterus, and the narwhal, Monodon,

which have lost all traces of hair even in the embryo), thus

showing their origin from forms that were more hairy. The

acquisition of blubber goes on pari passu with the loss of

hair, until in the Cetacea the blubber becomes extremely thick.

Kükenthal is responsible for the statement that in the seals

* hand and hand with the biological observation of the longer or

shorter time spent on land by the various species, we can deter-

mine the presence of a denser covering of hair or detect a

thinning of the coat, corresponding with the gradual increase of

No. 442.] HABITS IN MAMMALS 663

the layer of blubber." The reason for this is to be found in the

fact that hair is but a poor defense against the loss of heat when

in the water, while the layer of oil constituting the blubber

affords an excellent protection. Accompanying the loss of the

hair we naturally find also a reduction in the sebaceous glands,

smooth muscles and nerves of the skin. The sweat glands hre

also wanting in the Sirenia and Cetacea.

There are indications that the toothed whales, Odontoceti,

have been derived from forms possessing a dermal armature.

Kükenthal, to whom this observation is due has shown that in

Neomeris there remains in the adult considerable vestiges of

what must -be looked upon as a dermal armor. This has been

preserved usually only in those regions of the body where it may

be useful as a protection, as on the anterior margin of the flip-

pers, the anterior dorsal region and around the blow-hole, though

traces may occur on other parts of the body. The study of the

embryo shows that this is only a remnant of what was once a

much more extensive dermal armor. In the porpoises is found

the last appearance of this armor in the tubercles along the

dorsal fin. Here also they are more abundant in the embryo.

It is worthy of note in this connection that there have been

found with the extinct zeuglodons certain ossicles which indicate

a more extensive armor than is known to occur among recent

whales. The loss of armor is paralleled a number of times in

the marine reptiles. In the ichthyosaurs, the most aquatic

reptiles known, Fraas has shown that the dermal armor was

almost entirely lost, being retained only along the anterior

border of the fore limb, — the same position in which it occurs

in Neomeris.

Besides the adaptations already mentioned, it may be noted

that the bones of the truly aquatic forms are light and spongy,

particularly in the Cetacea, and in this group also the bones

become impregnated with oil. In the Sirenia the bones are

exceedingly dense and heavy, but in explanation of this it must

be remembered that these forms are not pelagic but live along

the shore in shallow water and find their food in the sea-weed

growing upon the bottom. The very heavy skeleton seems to

be an adaptation to bottom-feeding habits. In support of this

664 THE AMERICAN NATURALIST. (Vou. XXXVII.

view it may be added that the walrus, which is a bottom-feeding

form living chiefly upon bivalve molluscs, has the skeleton

noticeably heavier than that of any other of the Pinnipedia which

are generally piscivorous in habit.

The kidneys of most aquatic mammals are lobulated, Hippo-

potamus, Pinnipedia, Cetacea, but how this is to be explained by

aquatic life is not clear.

The testes are retained within the abdomen in the Cetacea,

Sirenia and the true seals, Phocide. In the less aquatic hair

seals, Otariidz, they are scrotal as in the majority of mammals.

Retia mirabilia, anastomoses of smaller arteries and veins, are

abundant!y developed in the Sirenia and Cetacea. These cause

a slowing down of the blood stream and it has been suggested

that this is connected with the oxidation of the blood in these

forms that breathe infrequently.

In the foregoing enumeration of adaptations the writer has

attempted to include only those that seem to be a result of

aquatic life, but in certain instances these may be open to

question. For example, Beddard has been inclined to question

the loss of hair in the Cetacea and Sirenia as due to aquatic life,

holding out the suggestion in the case of the whales that they

have probably been derived from armored forms in which the

hair was already lacking. That the Odontoceti have probably

been derived in this way is true and it is also true that they have

less hair than other marine forms, but it is equally true that

hairs have been found, at least in the embryo, in all but a few

species and in all cases these hairs are degenerate or vestigial in

nature. The inference seems plain that the ancestors of these

forms had these hairs better developed. As to the Mystacoceti

and the Sirenia, which are almost equally devoid of hair, there is

not the slightest evidence that they have been derived from

armored forms. On the contrary it has been shown that in the

embryo these forms are distinctly more hairy. This evidence

. taken in addition to the progressive degeneration of the coat

observed in the Pinnipedia makes the reduction of the hair by

aquatic life strongly probable.

In general it may be said for any character that when the

same tendency is observed in two such widely separated groups

No. 442.] HABITS IN MAMMALS. 665

as the Sirenia and Cetacea (and the latter group is almost

certainly diphyletic) it would seem difficult to explain on any

other ground than similarity of environment. When a third

group, as the Pinnipedia, shows the same tendency the matter

becomes almost a certainty.

COLUMBIA UNIVERSITY LABORATORY

New York, May 18, 1903.

BIBLIOGRAPHY.

BEDDARD, F.

A My of Whales. London.

Maicrestis. Cambridge Nat. Hist., Vol. 10. London.

'82. Monograph of the Insectivora. London.

FLOWER, W. H.

*70. An Introduction to the Osteology of the Mammalia. London.

FLOWER, W. H. and LYDECKER, P.

'91. An Introduction to the Study of Mammals Living and Extinct.

London

Guz, T. , r

'73. The Genetic Relations of the Cetaceans and the Methods involved

in Discovery, Amer. Nat. Vol. 3.

KÜKENTHAL, W.

'90. Ueber die Anpassung von Saügethieren an das Leben im Wasser.

Zool. Jahrb., Bd. 5

THOMPSON, D'Arcy, W.

'90. On the Systematic Position of Zeuglodon. Studies Mus. Zool.

Univ. College Dundee, Vol. 1.

WIEDERSHEIM, R.

Vergleichende Anatomie der Wirhelthiere. Fünfte Auflage Jena.

Wik isos, S. W.

On Certain Homoplastic Reps pen Air-Breathing Verte-

brates. Kansas Univ. Sci. Bull.,

WOODWARD, A. S.

'98. Outlines of Vertebrate Palaeontology. London.

RovanL NATURAL History, MAMMALS. Vols. 1-3.

STANDARD NATURAL History, MAMMALS. Vol. 5.

AMITOSIS IN THE EGG FOLLICLE CELLS

OF IHE CRICKET;

EDWIN G. CONKLIN.

Tuar type of nuclear division known as amitosis or direct divi-

sion which was once supposed to be universal is now known to

occur so exceptionally that all teachers of cytology will welcome,

I think, the announcement of its existence in great beauty and

profusion in a common animal of wide distribution. It is this

consideration which leads me to publish the observations here

recorded which were first made more than six years ago.

It has been long known that one of the most favorable objects

for the study of amitosis is to be found in the egg follicle cells

of certain insects (cf. Carnoy '85, Korschelt '86, Preusse

'95); in most if not all of these cases, however, it occurs only

occasionally and must be searched for among many cells which

do not show it. However in the case of the common crickets,

Gryllus pennsylvanicus, abbreviatus and domesticus, every follicle

cell in the enlarged portion of the ovarial tubes shows some stage

or other in the process of direct nuclear division, and these cells

are so easily prepared and they show every step of the division

with such diagrammatic clearness that they cannot fail to become

favorite objects for class demonstration.

A few words as to the structure of the ovarial tubes of the

cricket and as to the best methods of preparing them for the

demonstration of amitosis may not be out of place. If a mature

female cricket, which can readily be distinguished from the

male by the presence of the long median ovipositor, be torn in

two it will be seen that a large part of the abdomen is occupied

by the ovarial tubes which are arranged in two masses, one on

either side of the body ; these tubes are attached behind to the

right and left oviducts and in front, by long slender filaments to

! From the Zoólogical Laboratory of the University of Pennsylvania.

667

668 THE AMERICAN NATURALIST. [Vor. XXXVII.

the walls of the dorsal vessel. In fixing these masses of egg

tubes it is advisable to tease them apart, otherwise the fixing

fluids may not penetrate to the interior of the masses or the

tubes may be difficult to isolate afterwards. Almost any modern

fixing fluid will give good results, though I have had most suc-

cess with picro-acetic (Boveri), picro-sulphuric (Kleinenberg)

and Zenker’s fluid. After hardening in alcohol the ovarial tubes

may be stained from five to ten minutes in picro-haematoxylin !

and then mounted entire; such tubes show beautifully not only

the follicle cells in various stages of division but also the egg

cells in different stages of growth. However, for the accurate

study of the amitosis it is necessary to remove portions of the

follicle by means of needles; this can readily be done after the

tubes have been. stained and dehydrated and while they are in

the clearing fluid. These pieces of the follicle can then be

mounted in balsam and, if desirable, can be studied under an

immersion lens, though the nuclei are so large that all the

details of the division can be made out with a magnification of

from 125 to 300 diameters. It is advisable to double stain those

tubes from which the epithelium is to be stripped, in picro-

haematoxylin followed by a weak solution of eosin, or with acid

fuchsin and methyl green (Auerbach's formula) in order to bring

out the sharp contrast between the chromatin and the nucleoli.

Each ovarial tube of the cricket consists of a number of sec-

tions, all of which except the first contain ova in various stages

of growth. These sections are, Fig. 1, (7) the terminal filament

a thread of considerable length but of small diameter which

serves to attach the tube to the walls of the dorsal vessel; (2)

an enlarged section, the terminal chamber, which contains ovo-

gonia and young ovocytes which are not arranged in a linear

series; (3) a varying number of small ovocytes or eggs which

are arranged in linear series but are not completely separated

from one another by constrictions of the tube; (4) a section in

which the egg cells are separated by deep constrictions, the epi-

thelial cells growing all the way through the tube and thus form-

! Delafield's mess nd

Distilled wate

Kleinenberg's Moo dd (strong)

10 CC.

40 cc.

to drops.

No. 442.] EGG FOLLICLE .CELLS OF CRICKET.

ing partitions between the eggs; in this

section all the eggs are of approximately

the same diameter, but the lower ones

are much longer than the upper ones;

(5) a section consisting of two or three

eggs, each succeeding one being about

four times the volume of the preceding

and all being enormously larger than

any of the ova in the other sections of

the tube. Finally each of these tubes

is attached to one of the two oviducts,

through which the ripe ova escape.

In all sections of the tube the egg

cells are covered by epithelial cells; in

the second and third sections these cells

are relatively large and their nuclei are

far apart; in the fourth section they are

small and the nuclei are closely crowded

together, at the same time many of the

nuclei come to lie beneath the surface

layer and some of them seem to be

completely isolated in the yolk, to the

formation of which they probably con-

tribute. In the second, third and fourth

sections of the tube the cells increase

rapidly by division, the nuclei always

dividing by mitosis; in these sections I

have never seen a case of amitosis. The

ovarial tube is not as the name might

lead one to believe a tube composed of

follicle cells through which the ova

Fic. 1.— Ovarial tube of est I, terminal filament;

2, ovarial chamber; 3, section in which individual eggs

are not separated by follicle iib: 4, section in which

eggs are completely separated by the ingrowth of

follicle cells ; 5, last section of tube containing two or

t large ova; in this section all nuclei of follicle

cells are in process of amitotic division. X 62. y.

ASU),

i Foss

ae. 2

669

670 THE AMERICAN NATURALIST. (VoL. XXXVII.

descend from one section to another ; on the other hand in all

the sections below the ovarial chamber each ovum is perma-

nently surrounded by its own follicle cells which descend with

the ovum and increase in number with its growth in size. The

extent to which these follicle cells multiply can be estimated by

comparing their number in the fourth section with that in the

second, which would indicate that the increase cannot be less

than a hundred fold. In the fifth section mitosis rarely if ever

occurs, but here all the nuclei are found to be in some stage

of amitosis. In spite of the fact that all of the follicle cells of

this section are found in process of division the increase in the

number of the cells and nuclei in the successive segments of

this section is not great, not nearly as great as in the preceding

sections. Coincidently with the great growth of the egg cells

in this section the follicle cells and their nuclei grow rapidly in

size though they do not increase rapidly in number ; this would

indicate that the process of amitosis is here a very slow one,

which may, perhaps, account for the fact that it is to be seen in

all the nuclei.

Figures 2 and 3 represent portions of the follicle from the

region of the ovarial chamber and just below it; both show char-

acteristic mitoses but no evidence of amitosis in any of the cells.

Fic. 2.—Fragment of epithelium from the ovarial chamber (section 2) showing mitosis.

900.

Fic. 3.— Fragment of epithelium from section 3 of the egg tube, showing mitosis. X goo.

Figures 4-8 however taken from the greatly enlarged portion

(fifth section) of the tube show every nucleus in process of amito-

tic division. In this division, as is shown in all the figures named

but particularly well in Fig. 5, the nucleolus first elongates and

No. 442.] EGG FOLLICLE CELLS OF CRICKET. 671

then becomes constricted in the middle. The elongation and

constriction of the entire nu-

cleus follows after that of the

nucleolus. After the nucleolus

has completely divided into two

the nucleus also divides, but

although two separate daughter

nuclei are frequently found in

these follicle cells I have never

seen any indication of a division

of the cell body. After the

first amitotic division of the

nucleus the nucleolus in many

cases elongates again and divides Fre. 4.—Fragment of follicular epithelium

: è ° from the first egg in the fifth section of

in an axis at right angles to that he egg tube. X 333.

of its preceding division so that

two nucleoli are present in each of the daughter nuclei, Fig.

5. The daughter nuclei may also become elongated and even

constricted in the middle, but I have never seen them com-

Fic. 5.—Follicular epithelium from the second egg in the fifth section of the egg tube.

X 333- :

pletely divided into four nuclei within the single cell. This

division of the nucleolus and nucleus is almost always an equal

one and it usually occurs in a plane parallel with the surface

of the epithelium.

It is interesting to note that this case of amitosis almost

67 2 THE AMERICAN NATURALIST. [Vor. XXXVII.

exactly corresponds to the type described by Remak in '55.

He maintained that “cell division proceeds from the centre

toward the periphery. It begins with the division of the nucleo-

lus, is continued by simple constriction and division of the nucleus

and is completed by division of the cell body and membrane "

(Wilson, :00, p. 63). For a score of years after Remak's work

this was supposed to be the prevalent if not universal type of cell

division. Then it gradually came to be recognized that karyoki-

nesis or mitosis was the usual form of nuclear division and that

amitosis was comparatively rare, in fact its very existence was

Fic. 6.—Follicular epithelium from the third and largest egg in the fifth section of the

egg tube i

called in question. In particular the type of Remak which

begins with the division of the nucleolus was found to be most

unusual, being as Wilson says (:00, p. 115) *one of the rarest

forms of cell division (!).” Such preliminary divisions of the

nucleolus have been described by Carnoy (’85) in the egg follicle

cells of the mole-cricket, Gryllotalpa, and also in Lithobius and

in Geotrupes, by Wheeler ('89) in the follicle cells of Blatta, by

Hoyer ('90) in the intestinal cells of the nematode, Rhabdo-

nema, by Korschelt ('95) in the intestine of the annelid,

No. 442] EGG FOLLICLE CELLS OF CRICKET. 673

Orphryotrocha, by de Bruyne (’97) in the follicle cells of several

insects and by Montgomery ('98) in the peritoneal cells of

Polydora. In none of these cases however is it found so

abundantly and so plainly as in the follicle cells of the cricket.

I have examined the follicle cells of a number of insects and am

surprised to find how infrequent amitosis is in most of them.

In the mole cricket, as in all the species of true crickets which I

have examined it occurs in great profusion, but in the case of

the former the division of the nucleus is brought about by a deep

constriction on one side only of the nucleus. In the grasshopper

the nuclei are spherical and not bilobed and rarely show any indi-

cation of amitosis and the same is true of several other insects

which I have examined.

In most cases of amitosis the nucleolus does not divide and a

Fic. 7.—Section through the ae ROE AEE and underlying chorion of the smallest egg

in the eed section of the e

Fic. 8.— Section through the poa pubes of the largest egg in the ovarium tube.

X 333-

regular division of the nucleolus into two equal parts, preceding

an equal division of the nucleus such as is found in the cricket

is an occurrence of such rarity as to render it unusually interest-

ing. Another peculiar feature of these divisions is that the

nucleolus is always surrounded by a clear faintly-staining area

free from chromatin. This area is probably not the result of

shrinkage since it is found after all of the best methods of fixa-

tion which show no traces of shrinkage in other parts. This

clear zone around the nucleolus elongates with the elongation

674 THE AMERICAN NATURALIST. [Vor. XXXVII.

of the nucleolus and after the division of the latter it also

divides (Fig. 5). In the elongation, constriction and division of

the nucleolus into equal parts and in the presence of this clear

peripheral layer which also elongates and divides, the nucleolus

in the follicle cells of the cricket is very unlike ordinary nucleoli,

while in both of these respects it resembles an intranuclear cen-

trosome or *centro-nucleolus." Whether this resemblance is

merely a superficial one without significance (analogy) or a fun-

damental likeness (homology) cannot be affirmed without a more

extensive study of the structures in question, particularly of the

fate of the centrosomes which are found in the mitotic divisions

in the upper portions of the tube. If it should turn out that

these nucleoli are really comparable to centrosomes their peculiar

structure and form of division would find a ready explanation.

The biological significance of amitosis in this case cannot be

a matter of much doubt. These amitotic divisions are found

only in the terminal segments of the ovarial tubes at which time

the follicle cells are actively secreting the chorion (Fig. 7).

After the formatión of the chorion and before the egg is laid

these cells completely degenerate and disappear. A follicle cell

which has once divided by amitosis never again divides mitoti-

cally ; in fact no mitoses are ever found in this terminal section

of the tube. These facts all indicate that the amitotic division

is, in this case, one of the last functions of these cells and that

it is therefore an accompaniment of cellular senescence and

decay.

UNIVERSITY OF PENNSYLVANIA.

No. 442] EGG FOLLICLE CELLS OF CRICKET. 675

BIBLIOGRAPHY.

BRYNE, C. DE.

'97. * Les Cellules Doubles." Verhandl. anat. Gesell.

CARNOY, J. B

'85. La cytodieress chez les Arthopodes. La Cellule, Tom. 1.

Hoyer, H.

'90. Ueber ein für des Studium der “direction” Zelltheilung vor-

züglich geeignetes Objekt. Anat. Anzeiger., Jahrg. 5

KORSCHELT,

'89. Relves zur Morphologie und Physiologie des Zellkernes. Zool.

'95. Ueber Kerntheilung, Eireifung und Befruchtung bei Ophryotrocha

puerilis. Zeit. wiss. Zool., Bd. 40.

MONTGOMERY, T. H.

'98. Comparative Cytological Studies. Journ. Morph., Vol. 15.

PREUSSE,

'95. Ueber die amitotische Kerntheilung in den Ovarien der Hemip-

teren. Zeit. wiss. Zool, Bd. 59. _

REMAK,

'50-55. Untersuchungen über die Entwicklung der Wirhelthiere.

Berlin.

WHEELER, W. M

*89. The mbryslony of Blatta germanica and Doryphora decemlineata.

Vol. 3.

Jour. Morph., V

WILson, E. B.

00 The Cell in Development and Inheritance. New York.

NEW SPECIES OF PLANTS FROM THE

MATAWAN FORMATION.

EDWARD W. BERRY.

In A recent Bulletin of the New York Botanical Garden! I

have enumerated sixty-seven species of plants from the Mata-

wan formation (Mid-Cretaceous) describing . fourteen species as

new to science. Additional collections from near Cliffwood,

New Jersey, the only locality within the formation where recog-

nizable plant remains have been found, disclose numerous addi-

tions to this Flora, among which the following species have been

singled out as new ; and it has seemed best to publish them in

advance of more extended treatment which might be long

delayed. The remains are all from the lower portion of the

Matawan formation known as the Crosswicks Clays. The types

will be deposited in the paleobotanical collection of the New

York Botanical Garden.

Confervites dubius sp. nov. Fig. 9.

Remains referable to this genus have not heretofore been

recorded in this country, and as their microscopical characters

are obliterated, it cannot be certain that they are algal in nature

and not the macerated fibres of some higher plant. The dispo-

sition of the remains which are flexuous and interlaced would

indicate the former view, and as a well marked type of vegetable

remains they deserve a place in the flora of the Matawan forma-

tion. A number of foreign species have been referred to this

genus,? comparisons with which would be useless in view of the

unsatisfactory nature of the remains. Judging from the figured

! Vol. 3. No. 9. Sept.,

*Schimper, Pal. Végét. RE Tom I, p. 154, lists eleven, mostly — species,

all from European localities.

677

678 THE AMERICAN NATURALIST. (VoL. XXXVII.

Fics. 1-3, Gleichenia saundersii; 4, Pinus PONTS 5 and 6, Viburnum hollickii;

an yi Myrica heerit; Q, Cou vites dubius. Fig. 3 enlarged, all others four-fifths

natural size.

No. 442.] PLANTS FROM MATAWAN FORMATION. 679

specimen ' our form is practically identical with that of Confer-

vites aguensis Deb. & Ett., from the lower Senonian of West-

phalia.

Gleichenia saundersii sp. nov. Fig. 1-3.

In the flora of the Matawan formation I called especial atten-

tion to the total absence of ferns in that formation. Recent

collections contain three characteristic fragments of what I

regard as a new species of Gleichenia, adding another to the

considerable list of types common to the floras of the mid-

cretaceous Atlantic coastal plain and that of Greenland. The

larger fragments are each about 3 cm. long and 7 mm. wide and

the smaller is 16 mm. long and 3 mm. wide. Pinnules broadly

falcate-ovate, entire, bluntly pointed ; attached by a wide base,

about as wide as the pinnule is long ; length 4.5 mm. ultimately

becoming much smaller, each with a stout mid vein which sends

_off alternately on each side rather thick veins to the margin,

those running distad are all simple except the basal one which

is sometimes forked, those running proximad are usually once

forked ; texture thick and coriaceous.

Three species are found in the underlying Raritan formation,

Gleichenia giesekiana Heer is larger with longer narrower pin-

nules, Gleichenia micromera Heer is much smaller with narrow

linear right-angled pinnules, and the widespread G/ezcAenmia

stppet Heer has narrower pinnules with more rounded apices.?

The only other coastal plain species is the fragment which

Hollick3 refers to Gleichenia gracilis Heer, which is about the

same size as the Cliffwood fern but has the pinnules more acute

and runcinate. This reference of Hollick’s was only provisional

as he did not wish to found a species on so small a fragment.

His specimen is quite distinct from the Matawan form and also

seems to be distinct from Heer's type; unfortunately the vena-

tion cannot be made out. The numerous Greenland species

1 Hos. & v. d. Marck, Palaeont. 26: 177. pl. 36,f 135. 1880.

? Since the above was written small, poorly preserved fragments of the latter

Species have been detected in the Matawan formation by the writer.

nn. N. Y. Acad. Sci. Vol. 11, 1898, p. 57, pl. 5 f. 3.

680 THE AMERICAN NATURALIST. [Vor. XXXVII.

have usually smaller pinnules; there is some similarity with

Gleichenia acutiloba Heer, the venation being much the same,

but the pinnules are smaller and more acute. The species is

named for its discoverer Mr. B. H. Saunders of Clifton, N. J.,

who has collected a number of valuable specimens in the Clay-

Marls.

The genus Gleichenia is a most interesting one. In the

living flora it has about twenty-five species, widely distributed

throughout the tropics of both hemispheres, subtropical eastern

Asia, and the humid regions of the southern zone. The fossil

species are equally numerous and widespread. Aside from those

forms from the Paleozoic and older Mesozoic which have been

referred to the Gleicheniacez, which reference is not altogether

conclusive, as they probably represent synthetic forms from

which the later species may have been derived, the genus

enjoyed a wide adaptive radiation during the lower and middle

Cretaceous. In the lower Cretaceous (Kome) of Greenland

Heer has described fifteen species of Gleichenia, only one of

which ranges as far south as the Potomac formation; one other

occurs in the lower Cretaceous of the continental interior. By

the mid-cretaceous several of these Greenland species had become

wide-spread, identical species occurring in such widely separated

localities as Europe and Kansas, or Europe, Greenland and New

Jersey; four of the Greenland species find their way south

along the Atlantic coastal plain and three reach Europe. Two

species occur in the Laramie after which the species disappear.

With the gradual refrigeration of Tertiary climates the Gleiche-

niaceze moved southward, the only American fossil species of that

age being the doubtful GZezcZemia obscura Kn. from the late

Tertiary (Esmeralda formation) of Nevada. This southern

movement probably continued until Glacial times sending the

Gleicheniacez into the West Indies, along the Andes into

South America, along the eastern Asiatic coast, and across

southern Europe into Africa.

Pinus mattewanensis sp. nov. Fig. 4.

The well characterized impression of a single winged seed is

among the material from Cliffwood, N. J. Length 13.25 mm.;

No. 442.] PLANTS FROM MATAWAN FORMATION. 681

greatest width 10.5 mm. ; veins .75 mm. apart; proximal margin

nearly straight; apex broadly truncate. Newberry! figures

fragmentary winged seeds together with leaves from the under-

lying Raritan clays at South Amboy, N. J., and Hollick? figures

leaf fragments and a single seed (Fig. 19) from Tottenville,

Staten Island, in approximately synchronous strata, but no

remains referable to this genus have hitherto been found in the

Matawan formation. The specimen is unaccompanied by leaves

although poorly characterized remains of the latter are found in

the same formation. I have been unable to refer the seed to

any described species, none of which are nearly so wide, and

the same may be said of the existing species with which it has

been compared. In outline it is approached by the seeds of

some of our western species of Picea and it is also very similar

to the seeds of Cedrus deodara Loud.

In the living gymnospermous Flora Pinus is a dominant genus

with about seventy species widely distributed throughout the

northern hemisphere, thirty-nine of these occurring within the

limits of the United States. The fossil species are likewise

numerous (though many are of uncertain value) ranging from

the older Mesozoic upward. The genus becomes greatly

developed in the later Cretaceous and early Tertiary of the

Arctic regions, Greenland furnishing nine Cretaceous and six

Eocene (?) species and Spitzbergen four Cretaceous and twelve

Eocene (?) species, some of them widespread ; thus Pinus paleo-

strobus (Ett. Heer ranges from Switzerland and Tyrol to

Florissant, Colorado, occurring in the Baltic Tertiary, in Green-

land, Grinnell Land and Spitzbergen. Pinus quenstedti Heer is

likewise cosmopolitan, occurring in the Cenomanian of Moravia,

Silesia, and Bohemia, in Spitzbergen, in the Dakota group of

Kansas, and in the Montana formation of Wyoming. Pinus

(Cyclopitus) nordenskioldi Heer ranges from the Rhetic beds of

Norway and Spitzbergen to the Kootanie of British Columbia.

! U. S. Geol. Survey, Monograph 26, 1896, p. 48, pl. 9, figs. 17, 18.

2 Trans. N. Y. Acad. Sct. Vol. 12, 1892, p. 4, pl. 1, figs. 13, 19, 20, 22.

682 THE AMERICAN NATURALIST. [Vor. XXXVII.

Myrica heerii sp. nov. Figs. 7, 8.

This is one of the finest specimens that I have found in the

Matawan formation, consisting of a twig and four attached

leaves which are complete except for their apical portions ; the

block of clay in which they were found contains several more of

these leaves which cannot be uncovered without destroying the

specimen figured. The remains indicate a lanceolate leaf 13-14

cm. long by 2.7 cm. in greatest breadth ; base tapering, narrow,

gently incurved; the larger leaves have wide and regular,

strongly undulate, almost toothed margins; two of the leaves

are only slightly over one third the size of the larger and have

entire margins; petioles comparatively long and moderately

stout ; leaf substance thick, with apparently immersed venation,

as only a few secondaries can be made out on the impression of

the under side of the largest leaf; they leave the midrib at a

wide angle and are nearly straight almost to the margin where

they fork at a wide angle; their ultimate disposition cannot be

made out. The general similarity of size, shape and margin ally

these leaves to Myrica. Seven species occur in the underlying

Raritan clays, all of which are much smaller except Myrica emar-

ginata Heer which approaches our smaller leaves in size; it is

emarginate however and has entire margins and more ascending

secondaries. Three species have been found in the Staten

Island Cretaceous, one of which Myrica hollicki Ward is even

larger than our leaf, which it resembles greatly except that the

margin is more dentate. This species (%o%icki) might be con-

sidered ancestral to the widespread Tertiary Myrica banksiefolia

Unger which in turn was considered by Lesquereux as the

possible ancestor of the living Myrica californica Cham. of the

Western United States. The latter might easily be the descend-

ant of our Matawan leaf, some specimens seen by me are iden-

tical except for their slightly smaller size; other specimens are

more dentate; no other living species that I have seen so

nearly approaches the Matawan species in the character of the

undulations of the margin, although nearly all of the Myricaceze

are very variable in this respect, the same species often having

entire, or undulate, or dentate leaves on the same twig; this is

No. 442.] PLANTS FROM MATAWAN FORMATION. 683

particularly so in our common Myrica cerifera Linn. which

might also be derived from the Matawan species. It is at least

related and except for its somewhat smaller size and the irregu-

lar nature of the marginal characters it is strictly comparable.

The Cliffwood leaf is also very similar to Myrica (?) trifoliata

Newb." in size, shape and margin, but is longer petioled and not

trifoliate. Among the ten species found in the Dakota group

are several approaching ours in size, the one most similar,

Myrica aspera Lesq.? has the same texture, size and margin;

the base of the former is however narrower, the venation more

obsolete, the midrib more slender, and the petiole considerably

longer if we may judge from Lesquereux’s figure which appears

to be that of a sessile leaf. This species has also been men-

tioned as the possible ancestor of the living Myrica cerifera Linn.

Myrica torreyi Lesq. is the possible descendant of Myrica

heerit in the Montana and Laramie formations, with more

pronounced marginal teeth. A number of unrelated Cretaceous

leaves are somewhat similar in form and margin, as for instance

Ilex borealis Heer, Eleodendron speciosum Lesq., and Rhus

powelliana Lesq., but all differ in other particulars. Another

species from the Dakota group which resembles our leaf is

referred by Lesquereux to the somewhat smaller leaved species

Proteoides acuta Heer. Like the Matawan leaf this also contains

smaller entire margined leaves, but the tip is more extended, the

undulations of the margin are wider, and the base indicates that

the leaves were sessile.

Viburnum hollickii sp. nov. Figs. 5, 6.

Viburnum whymperi Heer, Knowlton, Bull. U. S. Geol. Surv.

163. pl. 19, f. 3. 1900 (mon pl. 17 f. 1, and pl. 18 f. 1).

The remains consist of the major portions of several leaves

indicating an ovate leaf between nine and ten centimeters in

length by 5 cm. in greatest breadth ; apex and base acute; basal

secondaries opposite, long, ascending in a nearly straight line

from the decurrent base, giving off three or four tertiaries on

! Undistributed plates, X/V. f. 2. from Dakota group of New Mexico.

2 U. S. Geol. Survey, Monograph 17, 1892, pl. 2, f. 2.

684 THE AMERICAN NATURALIST. [Vor. XXXVII.

the outside which run to the teeth of the margin; angle of

divergence acute, about 30°; a considerable interval to the next

pair of secondaries which leave the midrib at an angle of about

40° and ascend in a slight curve to the margin ; the three or

four succeeding secondaries on each side become more and

more ascending and are unbranched; margin for the basal

third entire, upper two-thirds shallow-toothed ; nervilles per-

current at right angles to the secondaries; midrib thin and

straight. I have tentatively included under this species a leaf

from the Montana formation which Knowlton (/oc. cit.) doubt-

fully refers to Viburnum whympert Heer, a Tertiary species

from which it manifestly differs. It differs from our type in

the lower secondaries being sub-opposite and supra-basilar,

otherwise it is markedly similar The genus Viburnum has

been heretofore unrepresented in the flora of the ancient Atlan-

tic coastal plain, the species Viburnum integrifolia referred by

Newberry to this genus being an entirely different leaf. Three

species occur in the Patoot beds of Greenland and abundant

remains are found in the western interior from the Dakota

group upward. Among the twelve species and varieties from

the Dakota group only one, Viburnum sphenophyllum Kn.

resembles the Matawan leaf. From this species ours differs in

its larger size, more ovate outline, lesser number of secondaries,

which are also more ascending and slightly more curved; the

margin is less prominently dentate and the basal third is entire ;

a pair of secondaries leaves.the base at an acute angle; these

are wanting in sfhenophyllum where all the secondaries are

parallel, the first pair leaving the midrib 3 mm. above its base

at an angle just twice as wide as in our type. In the existing

flora the nearest analogues of Viburnum hollickii are to be

found among the occasional simple leaves of the normally lobed

species such as Viburnum acerifolium Linn. and Viburnum opu-

/us Linn. The normally simple leaved forms have numerous

parallel secondaries and more toothed margins although the

outline is quite similar in Viburnum cassinoides Linn., Vibur-

num lentago Linn., and Viburnum nudum Linn.

Passaic, N. J., July 12, 1903.

SOME REMARKS ON THE FOSSIL FISHES

OF MOUNT LEBANON, SYRIA.

O: P. HAY.

IT 15 sometimes the good fortune of the archaeologist to carry

his excavations into the site of some long ago forgotten village

or city, and there to unearth the relics of its former inhabitants.

From these remains, perhaps scanty and broken, he essays to

determine the manners and customs of the people, their religion,

the grade of their civilization, the nature of their intercourse

with the neighboring tribes, and perhaps to learn what ancient

practices yet persisted and what new ones were coming into

vogue.

To the palaontologist the earth's crust, in its breadth and

thickness, is a burial ground from which he may exhume the

remains of the animals and plants that once lived on its surface

or in its waters. The words of Bryant, spoken of the races of

men, may truthfully be applied to other living things,

‘All that tread

The globe are but a handful to the tribes

That slumber in its bosom.”

But there are spots where the carcasses are sown thicker and

have been better preserved than elsewhere ; and to such places

the scientific birds of prey, who seek for, and must usually be

satisfied with, fragmentary bones, and imprints of skeletons, and

scattered scales and teeth, are gathered together; and, fed on

such booty, they have visions of the swarms of animals, fat, sapid,

and comely, that once populated the earth.

The Cretaceous period is one of great interest to the palceon-

tologist. It was apart of that long period which has been well

called the Mesozoic ; it was the closing third of the medieval age

of the world’s history. During the Cretaceous, vast advances

were made in the extinction of ancient forms of life and in the

685

686 THE AMERICAN NATURALIST. [Vor. XXXVII.

introduction of the present order of things; so that, with the

ushering in of the Tertiary, the world had become utterly trans-

formed from the condition it had at the beginning of the Creta-

ceous.

In the present paper we are to consider the fishes of the upper

Cretaceous period, especially those found in Mount Lebanon,

Syria.

The fishes of the Upper Cretaceous come to us principally

from four regions far removed from one another. One of these

is in western Kansas, another in the south of England, a third

in Westphalia, and the fourth in Syria. Our knowledge of the

fish-bearing strata of Kansas is of comparatively recent date ;

the other beds have long been known. Davis, who has written

an important paper on the fossil fishes of Mount Lebanon, tells

us that they were known to Herodotus, 450 years B. C. Trav-

ellers within the six hundred years preceding the nineteenth

century often expressed their astonishment at finding such per-

fect resemblances of fishes when the fissile rocks of that region

were split open. Accurate and scientific descriptions of these

remains have been presented only since the beginning of the

nineteenth century. Blainville, Agassiz, Pictet, Humbert,

Heckel, Davis, and A. S. Woodward have been the principal

writers on the subject.

While fossil fishes seem to have been found in several locali-

ties in Syria, two have become especially famous, Sahel Alma

and Hakel. The former is a village about eleven miles north-

east of Beirut ; Hakel is situated about twenty-three miles some-

what northeast of Beirut, and six miles from Jebeil, the ancient

Byblus.

Recently a third locality has been explored. This is near a

village called Hajula, situated about six miles south of Hakel.

About two years ago, at the instance of Rev. D. Stuart Dodge,

of New York City, this locality was visited by Alfred Ely Day,

professor of geology in the Protestant Syrian College at Beirut ;

and this gentleman succeeded in making a large collection of

fossils, especially of fishes. He collected many fishes at Hakel

also; but he was not permitted to work at the classical locality,

Sahel Alma. A large part of Professor Day’s collection was

No. 442.] FOSSIL FISHES OF MOUNT LEBANON. 687

presented by Rev. D. Stuart Dodge, in behalf of the Protestant

Syrian College, to the American Museum of Natural History,

in New York, and the present writer has had the opportunity

of studying and describing them. An account of the interesting

things in this collection, illustrated by fourteen plates, has

recently been issued by the Museum (Bull. Amer. Mus. Nat.

Hist, Vol. 19, pp. 395-452, pls. 24-37).

From the fish-beds at Sahel Alma there have been described

about sixty species of fishes; from Hakel, fifty species; and

now there come from Hajula thirty-four species. Doubtless

further collecting at Hajula wil much increase its number.

The fishes found at Sahel Alma belong, in general, to the

same genera as those at Hakel and Hajula; but, of its sixty odd

species, probably not one is found at the other localities. On

the other hand, twenty-one species that have been secured at

Hajula occur also at Hakel. These data make it quite certain

that the beds at Sahel Alma are on a different level from those

at Hakel and Hajula; while those at the latter places are on the

same, or nearly the same, horizon. Opinions have differed as

to which are older, the fish-beds at Sahel Alma or those at

Hakel; but it is apparently the view of the best modern author-

ities that those at Hakel are more ancient. This opinion

appears to be supported by the character of the fishes in each.

From a study of the fishes taken at Hajula the writer has con-

cluded that the beds containing them belong to a slightly more

recent time than that of the beds at Hakel.

As to the position of the fish-bearing strata of Mount Lebanon

in the geological scale, all authors now agree that they belong

to the Upper Cretaceous. The epochs of this division, as rec-

ognized in Europe, are, beginning with the lowest beds, the

Cenomanian, the Turonian, the Senonian, and the Danian.

Some authorities have assigned the fishbeds of Mount Lebanon

to the Turonian, others to the Senonian. A. S. Woodward,

in the earlier pages of the first volume of his splendid work,

Catalogue of Fossil Fishes, credits the Mount Lebanon fishes to

the Turonian, but in the later pages they are said to belong to

the Senonian. In the third volume they are more definitely

referred to the Upper Senonian; but in the fourth volume they

688 THE AMERICAN NATURALIST. [Vor. XXXVII.

are cited simply as coming from the Upper Cretaceous. The

deposits from which the fossil fishes of Westphalia are exhumed

are regarded as Senonian. Roemer (Zeitschr. deutsch. geol.

Gesell. vol. 6, 1854, p. 201) assigns these beds to the older

Senonian. Lepsius (Geol. Deutschlands, vol. 1, p. 177) and

Credner (Elem. Geol., p. 637) regard them as belonging to the

Upper Senonian. A comparison of the genera and species of

fishes from Mount Lebanon with those from Westphalia has

convinced the present writer that the horizons of the two groups

of beds are practically the same, and that the Mount Lebanon

fishes, therefore, belong to the Upper Senonian. Hence these

fishes lived near the close of Cretaceous times; and we are

enabled to observe the advances which this group of animals

had made during this long age.

At the beginning of the Cretaceous the shark-like animals

were few, so far as the record shows, and these mostly of now

extinct genera. There were doubtless species of Hexanchus

(Notidanus), a decadent genus represented now by the cow

sharks; for we know that species lived during the Jurassic.

There was a member or two of the Heterodontide (Cestra-

ciontidz), and possibly the Lamnidae were represented. No rays

or sawfishes are known from the Lower Cretaceous, but there

must have been species of Rhinobatus, since they had previously

existed and the genus is yet on the earth. But with the open-

ing of the Upper Cretaceous, in the Cenomanian, the Lamnidze

and the Scylliidze were well established ; while in the Senonian

beds at Sahel Alma there were three or four species of each

family. At Hakel only a single species of shark, Otodus sulca-

tus, has been found ; at Hajula, no shark,

Of the rays there are interesting forms at all three of the

Mount Lebanon localities. A. S. Woodward has described

from Sahel Alma the rostrum of a primitive sawfish, Sc/eror-

hynchus atavus, the rostral teeth of which are not in sockets

and which, as they are followed backward, graduate into the

shagreen scales of the side of the head! In the collection from

Hajula the writer has found three new species, one represented

by a considerable portion of a rostrum ; another, by a complete

rostrum and a considerable part of the head; and a third, by

No. 442.] FOSSIL FISHES OF MOUNT LEBANON. 659

much of the rostrum, the head and most of the pectoral fins.

There is likewise the trunk of a species of the genus present.

These specimens show that Sclerorhynchus belongs to the Pris-

tide, and further, that at that far away day the sawfishes

resembled closely those of our time, except in the mode of

attachment of their rostral teeth. Between that time and the

Eocene, these modified shagreen scales found deeper and deeper

lodgement in the cartilages of the rostrum, attained greater size,

and became the efficient weapons that we find them to-day.

The new Sclerorhynchus which is represented by the rostrum

alone has been called S. sentus, because on one flat side of the

rostrum there are two rows of short spines. The species repre-

sented by the complete rostrum has been named S. solomonts,

in recognition of the interest of the great Israelitish king in

natural history (i Kings, iv, 33). The third species is to be

known as S. %iram, in honor of the friend and ally of Solomon.

Of the genus Rhinobatus three species have already been

described from Sahel Alma and one from Hakel. A new one,

R. eretes, is found in the collection from Hajula. Likewise a

new ray comes form Hajula, and this has been named aya

whitfieldi, in honor of Prof. R. P. Whitfield, of the American

Museum. It is surprising to observe how little these rays have

changed since those Senonian times.

Of the true fishes, Pisces, we find in the Mount Lebanon fish

beds but few representatives of the old families of the Jurassic.

None has been obtained at Sahel Alma, and this speaks for the

later time of these beds. From Hakel there have been known

four species of pycnodonts; and now from Hajula there is

secured a new one, Coccodus insignis. It is represented by

many specimens, some of them quite perfect ; and is character-

ized by having a short and compressed occipital spine. It like-

wise presents clear evidences of pectoral and ventral fins. Hakel

furnishes a new and strange fish which possibly belongs to the

Belonorhynchide, and which has been named Séenoprotome

hamata. The head, as it is shown on the matrix, resembles in

miniature that of the African antelope, Bubalis, even to the horns.

The horns, or spines, are probably connected with the operculum.

The tip of each is very sharp, and just below this there is a

690 THE AMERICAN NATURALIST. (Vor. XXXVII.

barb, like that of a fishhook. There seem to have been no

vertebrz, and the body has been covered with bony plates.

The Macrosemiidz and Oligopleuride, scantily represented at

Hakel, have not yet been found at either Sahel Alma or Hajula.

Some of the fishes of the Upper Cretaceous had already acquired

many of the characters which are found in our modern forms.

In the majority of them the skeleton was extensively ossified,

the vertebrze were as we find them today, the fins had lost their

fulcra, and in many cases the rays had become spinous. A

number of the families that still persist had already come into

existence, and this is true of a few genera. The Elopidz, best

known by the tarpon of our southern waters, are recognized in

eight species at Sahel Alma, three at Hakel, and one at Hajula.

None of the species begins to attain the size of the tarpon.

The Ichthyodectide, a family close to the Chirocentridz, is

thought to be represented at Hakel and Hajula by a species

which Woodward calls /chthyodectes Lbanicus, but which the

present writer refers to a new genus, Eubiodectes, that is,

a biter that gets a good living. It grew to a size somewhat

greater than that of the shad.

Ctenothrissa is a genus which stands as the type of the

Ctenothrisside. It possessed many of the structures that are

greatly in vogue in our own day among the denizens of the deep.

It had serrated scales, and the ventral fins had been moved for-

ward to a position just below the pectorals. The writer has

described a new and beautiful species, Crenothrissa signifer,

from Hajula. It is characterized by the possession of very high

dorsal and anal fins ; and the ventral rays likewise were greatly

elongated.

The Clupeidz, so abounding in genera and species in the salt

and fresh waters of our time, were present in force during the

Senonian ; but only one of the genera of that time has survived.

Two species of Scombroclupea were exceedingly abundant at

both Hakel and Hajula. The specimens are nearly always

found in a contorted position. Pseudoberyx is known by three

species from Hakel. They are short-bodied, high, and com-

pressed fishes, with large and pectinated scales. Diplomystus

brevissimus is another small, elevated fish from Hakel and

No. 442). FOSSIL FISHES OF MOUNT LEBANON. 691

Hajula, and it must have swarmed in those waters. Another

species of the genus has been described by Cope from the

Upper Cretaceous of Brazil; several species, by Leidy and Cope

from the Eocene of North America; and another, from the

Oligocene of England, by Newton. To-day species are living

in the fresh waters of Chili and New South Wales.

The Dercetidee form an extinct family of elongated fishes,

most of whose members lived during the Senonian, but some of

which flourished during the Turonian. Two species of Lepto-

trachelus have been described from Sahel Alma, one from Hakel,

and now another is sent to us from Hajula. This is a species

very eel-like in form, and has been called Z. sezpeutiuus. One

species of the genus has been described by Cope from the Upper

Cretaceous about Yankton, South Dakota, and with it another

related form, Zrzenaspis virgulatus.

The Enchodontidee were numerous during the whole of the

Upper Cretaceous. A. S. Woodward regards them as hav-

ing as their nearest living allies the deep-sea families Odon-

tostomida and Aleposauride. The enchodonts were undoubt-

edly predaceous fishes, having long fang-like teeth in their jaws

and a huge gape. Five species come from Hakel, four from

Hajula, and three from Sahel Alma.

Another interesting family of fishes which have come down

the ages to our day from probably the Lower Cretaceous is the

Myctophidz, usually called the Scopelide. In our time they

are fishes which live in the open sea, many of them being inhab-

itants of the deep waters, and many possessing phosphorescent

organs. Figures and descriptions of many of the living forms

may be found in Goode and Bean's Oceanic Ichthyology. At

Hakel there have been collected eight species ; at Hajula, seven,

and at Sahel Alma, eight. While most of the species found at

Hajula occur also at Hakel, none found at these places have yet

been collected at Sahel Alma. Four new species come from

Hajula, two of which come likewise from Hakel; and one new

and rather small species, Osmeroides ornatus, also comes from

Hakel. Osmeroides is used here in the sense in which it

was originally employed by Agassiz and in place of the latter

proposed name, Sardinioides. In the place of Osmeroides,

692 THE AMERICAN NATURALIST. (Vor. XXXVII.

applied to a genus of Elopidz, the present writer prefers to use

the name Holcolepis. How much confusion might have been

avoided had the latter name been employed for the elopid genus

as soon as it was discovered that Osmeroides lewestensis was

not congeneric with O. monasteri! Specimens of a Nematonotus

of unusually large size and having a greatly elongated first dorsal

ray are referred to JV. /ongzspinus (Davis).

Eels are proverbial for their lubricity, but many of them were

caught in the net formed by the sediments of the Senonian

ocean. Davis had already in 1887 described from Hakel a little

species which he called Anguilla hakelensis, but which Wood-

ward refers to his genus Urenchelys. This author has likewise

been so fortunate as to find another species of the genus in the

Turonian of England. It is the oldest known eel. He describes

also a third species from Sahel Alma. The species of the genus

are shown to have about one hundred vertebra and a caudal fin

which is distinct from both the dorsal fin and the anal. The

present writer has named a fourth species from a specimen col-

lected at Hajula.

Besides these representatives of the family Anguillide, the

writer has found two species which present most of the char-

acters of Urenchelys, but which are peculiar in possessing well-

developed ventral fins, a new feature in eels. This character

brings the A podes into closer relation with the other bony fishes.

The species are regarded as forming a new genus, Anguillavus.

the type of a new. family, Anguillavide. One of the species,

represented by a complete skeleton six inches long, from Hakel,

and a fragment from Hajula, has been named, in honor of a

worthy woman, Anguillavus bathshebe. A larger species from

Hajula has been christened 4. guadripinnis. Besides ventral

fins, this species seems to have had a row of enlarged scales on

each side of the body, perhaps along the lateral line.

Still another eel hails from Hakel. It has been very slender,

much elongated, and apparently without a fin of any description ;

but the feature which most strikes our attention is the structure

of the vertebra. Throughout the length of the body, from the

head to the tail each vertebra resembles, not the ones

adjoining it, but the second one behind it or in front of it. This

No. 442.] FOSSIL FISHES OF MOUNT LEBANON. 693

condition appears to be much like that found in the tail of

Amia, and the writer explains it in the same way. The species

is called Enchelion montium, the type of the new family

Encheliidz.

The remaining fishes are regarded as belonging to the order

Actinopterygii, or Percomorpha. In these the dorsal and anal

fins are usually wholly or partly spinous and the ventral fins are

brought forward to beneath the pectorals. The order contains

the most highly developed fishes. The Berycidz, yet represented

by some marine and mostly deep-sea forms, appear to have been

very abundant during the Upper Cretaceous. A single species

is known from Hakel, two from Hajula, and a dozen from Sahel

Alma. These data appear to argue that the latter locality is at

a higher level than either of the others. A new species of

Pycnosterinx, P. /evispinosus, is described by the author from

Hajula. It is a small, compressed species, about an inch and a

half in length, deeper than long, and with a steep front.

Three species of the genus Omosoma are referred provisionally

by Woodward to the Stromateide, a family represented on our

Atlantic coast by the butter-fishes and harvest-fishes. The

three species are found at Sahel Alma only. Other genera sup-

posed to belong to the same family are described from Westphalia

and the south of England.

From Hakel and Hajula there have been obtained three species

of Aipichtys, a genus placed by Woodward in the family Caran-

gidz. One of these species is a new one from Hajula.

The Cretaceous fishes of the families just mentioned, the

Berycidz, the Stromateidae, and the Carangidze, with their spiny

fins, their thin and often ctenoid scales, their complete vertebrze,

their thoroughly ossified skeletons, and their thoracic ventrals,

form a strong contrast with the Semionotidze, the Pycnodontidze,

and even the Chirocentridz, which formed the greater part of

the fish-fauna at the beginning of the Cretaceous period.

It is remarkable that no vertebrates, except fishes, have been

found at Mount Lebanon. In the collections examined by the

writer, not a scrap of any reptile has been detected. Ichthyo-

saurs had indeed reached the evening time of their existence.

There were doubtless plesiosaurs swimming the deep, and we

694 THE AMERICAN NATURALIST. Vor. XXXVII.

might expect to find some trace of them in the Mount Lebanon

rocks. There were certainly mosasaurs haunting the shores and

venturing out on the waves. We should hardly expect to meet

with remains of dinosaurs, for these either stalked about on the

land. or wallowed in the swamps of lakes and rivers. Unless the

conditions were peculiar we might anticipate finding bones of

turtles and crocodiles, possibly of pterodactyls.

It appears probable that the deposits of the Mount Lebanon

district were laid down in deep water and at a considerable dis-

tance from any shores. The fineness of the materials of the

rocks favors this view. The large numbers of Myctophidze and

Berycide found there, seem to indicate that the depths were

considerable. The presence of so many sharks and rays is not

opposed to this view, as may be learned by an examination of

the lists of fishes in Goode and Bean's Oceanic Ichthyology.

In case the deposits were made in deep water and at a con-

siderable distance from land, few reptiles would be entombed in

them. Most of these animals probably found it more profitable

and more comfortable to remain near the shore.

Dana regarded the Niobrara deposits of Kansas, which have

furnished so many fossil fishes, as nearly equivalent to the Turo- -

nian of Europe. This was the opinion held by Cope also. A

comparison of the Kansas fishes with those of the Turonian

and Senonian confirms this opinion. Comparatively few of the

genera of Kansas fishes are found in the Senonian of Europe or

Syria. On the other hand, about a dozen genera from Kansas

are represented in the Old World Turonian.

In 1878 (Bull U. S. Geol. Surv., vol. 4, p. 67), Cope described

seven species of fishes which Hayden had collected in what is

now South Dakota. No definite statement was made regarding

the locality; but, written on the blocks of soft limestone bear-

ing the types, the present writer has found the word * Yankton.”

This no doubt indicates approximately the locality whence the

fishes were derived. Of these fishes, two species belong to

Leptotrachelus and one to Triæaspis, a genus close to Lepto-

trachelus. Of other known species of the latter genus, all are

found in the Senonian, although one occurs in the Turonian

also. One species from Yankton belongs to Spaniodon. The

No. 442.) FOSSIL FISHES OF MOUNT LEBANON. 695

other three species of the genus are from Sahel Alma. Three

of the species from Yankton belong to Leptosomus. Two

other species of Leptosomus are known from Westphalia and

two from Mount Lebanon. Cope supposed that these fishes

were obtained in the Niobrara deposits; but, judging trom their

close relationships with Mount Lebanon forms, it appears highly

probable that they came from a higher horizon, probably the

Pierre. It is a matter of considerable importance that some

geologist in that region should explore these beds. There

appear to be great numbers of the fishes, since on one block

about six inches square there are three of Cope’s types and

one on the opposite side. The reader will find a figure of this

block on Plate 5, of volume 19 of the Bulletin of the American

Museum of Natural History.

AMERICAN MUSEUM OF NATURAL HISTORY.

vew York, May 30, 1903.

ON THE OSTEOLOGY AND SYSTEMATIC

POSITION OF THE KINGFISHERS.

(HALCYONES.)

R: W. SHURFELDT.

BY FAR one of the most useful contributions to the recent

literature of ornithology is the work entitled “A Hand-List of

the Genera and Species of Birds,’ by R. Bowdler Sharpe,

Three volumes have already been issued, while the fourth and

last one is now passing through the press.

This work presents us with a hand-list of the species and sub-

species of the birds of the world brought fully up to date. The

“ Systematic Index” is practically a classification of the class

Aves, and in fact is Sharpe’s taxonomy of birds, and is there-

fore one of great value and worthy of our closest study. It

also takes into consideration the fossil forms of birds, and the

distribution of all forms. Among other Orders enumerated,

we find in Volume II of the * Hand-List" Order XXIX,—

the Coraciiformes, which is subdivided into fourteen sub-

orders created to contain a great many different kinds, and very

differently affined, birds. These are as follows :— (I) Steator-

nithes, (II) Podargi, (III) Leptosomati, (IV) Coraciz, (V) Halcy-

ones, (VI) Bucerotes, (VII) Upupz, (VIII) Meropes, (IX)

Momoti, (X) Todi, (XI) Caprimulgi, (XII) Cypseli, (XIII)

Trochili, and (XIV) Colii. These suborders are duly divided

into their families and other minor divisions. There is no

intention of discussing in full this arrangement here, and the

sequence of the suborders is given only to show the position

assigned the Halcyones or the kingfishers, the group which is

the subject of the present contribution. It is now more than

thirty years since Sharpe published his famous monograph on

the kingfishers, and it remains a classic in the literature of

ornithology. In it the family Alcedinidae was divided into

nineteen genera, created to contain the 125 species described

697

698 THE AMERICAN NATURALIST. [Vor. XXXVII.

in the memoir. Of these a subfamily division was made, the

line being drawn between the insectivorous Daceloninz, with

14 genera and 84 species, and the piscivorous Alcedininz, con-

taining the balance of the group. Now as will be seen, the

Halcyones are placed between the Coracize on the one hand,

and the Bucerotes on the other, a long ways removed from such

forms as the Galbulide or jacamars, the Buccones, and the

ground cuckoos, (Geococcyx), birds that appear in widely sepa-

rated and entirely different orders.

The Halcyones in the * Hand-List " we find still to be divided

into the two subfamilies Alcedininze and the Daceloninz, the

first still containing five genera, and the last by an increase of

one, now containing fifteen. The number of species, however,

have been increased from the 125 enumerated in 1870 to about

200, or in other words there have been about 75 species of

Kingfishers described within the last thirty-two years. A

knowledge of their anatomy, however, has by no means kept

pace with this remarkable discovery of new and undescribed

forms. The habits of the various kinds of kingfishers are

described with greater or less detail in Sharpe's monograph,

as well as the plumage and external characters, and as highly

important as this is for an understanding of their affinities, it

cannot be properly touched upon in this paper which deals with

the osteology. North America is extremely poor in king-

fishers, as we find but two species and a subspecies of the

genus Ceryle (C. a/cyon, C. torquata, C. a. septentrionalis), given

in the last A. O. U. * Check-List." Australia, Africa, and the

East Indies claim the greatest number of forms, but their dis-

tribution is extremely unequal, when taken as a whole, in so far

as other countries are concerned.

In studying the osteology of such species as I have been able

to obtain, I have taken special care to keep before me the geo-

graphical distribution of the family, the remarkable variations of

the plumage, the beaks, and particularly the feet and other

external structural characters; also similar data of the species

representing other families of birds which avian anatomists from

time to time have considered near relatives of the Halcyones.

I have also had in mind, during my researches, the notes of the

No. 442] OSTEOLOGY OF THE KINGFISHERS. 699

various species, their habits, and their nidology, all of which has

been done in order to avoid any biased opinion I might other-

wise arrive at on any particular point, where the osteology alone

had come to influence me in the formation of an opinion.

It is eighteen years ago since I have been engaged with

the osteology of the kingfishers, when I published a brief illus-

trated memoir on the “Osteology of Ceryle alcyon,” which

appeared in the Journal of Anatomy, (Vol. 18, London, 1884,

pp. 279-294), and yet it can be said with great truth that we

stand much in need of a thorough investigation of the general

structure of the Halcyones. Avian taxonomers and anatomists

are not unanimous on the systematic position of the kingfishers,

and still less so on their relationships with other groups of birds.

Some thirty years ago Cunningham contributed a brief notice

of some of the anatomical points of the kingfishers (Proc. Zool.

Soc., 1870, p. 280), and for various views upon the taxonomy of

the Halcyones and their affinities one should consult Wallace,

(Ann. Nat. History, ser. 2, Vol. 18, pp. 201-205): Eyton

(Contrib. Ornithology, 1850, p. 80.): Huxley (Proc. Zool. Soc.

1867, p. 467.): Coues (* Key to N. Amer. Birds,” rev. ed. p.

469, 1884.): Fürbringer (Untersuchungen z. Morph. u Syst. der

Vögel. Amsterdam, 1888, pp. 1555-1567): and the Mon-

ograph of Sharpe already cited.

According to Huxley's classification, the kingfishers (Alcedi-

nidz) belong to the Desmognathz, they forming a family

of a third group out of four of the division designated as the

Coc pha. He associates with them the Bucerotide,

Upupidze, Meropida, Momotidze, and the Coracidz ; and he also

believed that they approached the Pelargomorphze in their struc-

tural characters.

Newton tells us with great truth that “it is to be regretted

that hitherto no light has been shed by palzontologists on this

interesting subject, for the only fossil referred to the neigh-

borhood of the Family is the Hadcyornis toliapicus of Owen

(Br. Foss. Mamm. and Birds, p. 554) from the Eocene of

Sheppey — the very specimen said to have been previously

placed by Kénig (/con. foss. sectiles, Fig. 153) in the genus

Larus. (Dict. of Birds. 1893, Pt.2, pp. 488-489). In the

700 THE AMERICAN NATURALIST. [Vor. XXXVII.

same excellent work, and in the same article (** Kingfisher "),

Newton further states that “the common Kingfisher of Europe

is the representative of a well-marked family of birds, the 4&edi-

nide or Halcyonide of ornithologists, which is considered by

some authorities to be closely related to the Aucerotide ; but

the affinity can scarcely be said as yet to be proved ; and to the

present writer there seems to be at least some ground for believ-

ing that a nearer alliance is to be found in the Ga/bxZde,

Momotide, Meropide, and perhaps some other families —

though all may possibly be discovered to belong to one and the

same larger group."

The Halcyoniformes of Fürbringer form one of the suborders

of his order Coraconithes, and he divides them into three groups

(gens) vzz., the Halcyones, the Bucerotes, and the Meropes.

In the first-named we find but one Family (sensu latiori), the

Alcedinide, and this he divides into two others (sensu stric-

tori) the Halcyonide, and the Alcedinide. The Bucerotes

contain the Bucerotida and the Upupide, while the Meropes

contain only the single family Meropide.

In his Manual of North American Birds Robert Ridgway

places the family Alcedinidee, together with the Cuculidz, Tro-

gonidz and Momotidz in an Order Coccyges, but in differentiat-

ing the kingfishers gives a very slender array of characters, and

no structural ones. (Phila. 1887, pp. 27 1-279.)

It is an interesting fact, although we have as yet but little

knowledge of the habits of the jacamars (Galbulidz), that at

least some of the South American species secure their insect

food after the manner of some of the Daceloninz, and that fur-

ther they make their nesting holes in marl-banks, thus agreeing

with the nidification of some of the kingfishers (Ceryle).

Newton is not alone among ornithologists in the belief that the

kingfishers are-in some way more or less related to the jacamars

(Galbulidz) and the bee-eaters (Meropida). I believe it was

the naturalist Lesson who named one genus of jacamars, Jaca-

maralcyon, and another Jacamerops.

It is perfectly safe to say at the present writing that there

are no two avian classifiers who agree exactly on the question

of the relationships of the kingfishers, while on the other

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 791

hand, with others there exists the greatest possible variance

in the matter of opinion on this subject. No two classifiers

of birds would today agree as to the place of the Halcyones

in the system, or would award them the same place in any

scheme of classification. A unanimity 'of opinion can only be

arrived at upon this point when the morphology and complete

life-histories of all the forms are known, and have been com-

pared and intercompared. As it is we are a long way from any

such decision, and in fact we know very little of the anatomy of

any of the kingfishers, the jacamars, the bee-eaters, the cuckoos,

the trogans, the hornbills, the toucans, the hoopoes, and a dozen

other families more or less related. This being the case, I

must believe that any little contribution to the anatomy of any

of the birds in question, will in time prove to be useful, and it

is therefore with less hesitation that I bring forward here what

little I have accomplished in the osteology of the Halcyones,

not that I think that any part of it will completely settle any

particular taxonomical point at issue, but that it may help to do

so, when our knowledge in these matters becomes wider than it

is at the present time.

In studying the osteology of the Halcyones I have carefully

examined the skeletons of the following species of birds, and

compared the characters they have presented with great thorough-

ness and detail.

1. Ceryle alcyon, belted kingfisher. North America. |

2. Ceryle a. septentrionalis, texas kingfisher. Texas to

Panama.

3. Alcedo ispida, Europe and many parts of the east.

4. Geococcyx californianus, road-runner. Mexico and parts

of the western United States.

5. Various cuckoos of the genera Coccyzus, Cuculus, Croto-

phaga, and others.

6. Dacelo gigas, Australia (skull).

. Several of the Meropide.

. Steatornis caripensis, Northern S. Amer. and Trinidad.

Trogan (several species).

Bucerotidz (several species).

Nyetiornis amicta, Borneo.

^ O0 ^u

702 THE AMERICAN NATURALIST. [Vor. XXXVII.

12. Galbulida.

13. Trochili, many species.

I4. Cypseli, various species.

15. Momotus.

Also incidentally the representative of several other groups,

as the woodpeckers, Caprimulgi, and the toucans.

I am indebted to the United States National Museum for the

loan of some of the material used in the preparation of the pres-

ent memoir, and to that institution my thanks are due, as they

are to Mr. F. A. Lucas for the loan of the skeleton of Acedo

ispida from his own collection. For the trogan skeletons I

have pleasure in thanking Dr. Sclater, late Secretary of the

Zoólogical Society of London; and the skeletons of Geococcyx,

Mr. Herbert Brown of Yuma, Arizona. A number of the swifts

and goatsuckers were kindly sent me by Mr. Jno. H. Sage of

Portland, Ct., Mr. Gerrit S. Miller, Jr., of the U. S. National

Museum, and Dr. A. K. Fisher of the U. S. Dept. of Agricul-

ture. Many humming-birds and much of the other material has

been supplied me by Messers F. Stephens, H. K. Coale, Luther

N. Rossiter, H. W. Henshaw, J. G. Parker, E. M. Hasbrouck,

and others,— to each and all of whom I desire here to return my

thanks. Not a few of these specimens, now types of descrip-

tions, are in the collections of the British Museum, and the

Royal College of Surgeons of England.

With this preliminary introduction I am now prepared to

pass to the consideration of the comparative osteology of the

kingfishers (Alcedinidz) employing primarily for this purpose

the skeletons of specimens of Cery/e alcyon; Ceryle cabanisi and

Alcedo ispida. These I will not only endeavor to carefully inter-

compare, but in turn, compare them with the skeletons of the

various species and material set forth in the above list.

In my conclusions I shall have something to say regarding the

relationships of the Halcyones to other groups of birds.

The Skull. — By referring to the figures, herewith presented,

illustrating the skeleton of our common kingfisher (C. alcyon),

it will be noted that the superior osseous mandible of Ceryle is

considerably longer than the remaining part of the skull, being in

fact a three-sided pyramid, with a broad base and sharp-pointed

No. 442] OSTEOLOGY OF THE KINGFISHERS. 703

apex. A very perfect cranio-facial hinge joins this structure to

the cranium, and the tomial edges are sharp and slightly raised

above the level of the nether aspect of this mandible. This

skull is strongly desmognathous, and the rhinal chambers are

quite filled in by the spongy mass of the bulky maxillo-palatines.

Through this the parial subtubular narial passages run, and they

terminate externally upon either side, as rather elongated,

broadly spindle-shaped narial openings. Just to the rear of the

posterior border of either one of these latter there is to be found

a small circular foramen leading into the nasal passages just

referred to, a character seen also in C. cabanisi. The external

narial apertures are separated by a complete, rather thick septum

narium, while in Adedo ispida this partition is pierced by an

oval foramen, at about its centre, and in this species the fora-

mina described in the last sentence are absent.

The culmen, which forms one angle of this mandibular pyra-

mid, is rounded ; while the side that forms a good share of the

roof of the mouth is flat, marked its entire length by a small,

deep, median groove, that exhibits impressed venations branch-

ing from it upon either side. At the hinder end of this groove

a pair of small foramina are seen (Fig. 2, 3).

We find this mandible in Ceryle, although having a very

solid appearance from being closed in on all sides as it is, to be

extremely light, having internally very much the same structure

as in the hornbills, only rather coarser.

There is one other feature we notice on the superior aspect

of the mandible in Alcedo that is absent in Ceryle; this is a

pretty-well marked groove, leading on either side from the

anterior margin of the nostril back to the maxillary. It is shown

in the drawing of this view of the skull here figured, from a

specimen kindly lent me by Mr. F. A. Lucas of the United

States National Museum (Fig. 1, 5, #.).

Regarding the skull of Ceryle from above, we find the supe-

rior margins of the orbits sharp and regular, and separated from

each other by the smooth, rather broad surface of the frontal

region. This is slightly indented longitudinally by a shallow

median groove that traverses the cranium from the transverse

fronto-maxillary line, through the parietal portion. Here the

794 THE AMERICAN NATURALIST. [Vor. XXXVII.

surface is raised, on either side, into smooth rounded domes,

that are less marked in C. cabanisi, and that are bounded behind

by the prominent and projecting temporal fossa. These latter

are divided behind by a sharp median ridge (see Fig. 2). All

these features, though present in Alcedo, are far less noticeable,

while the median ridge tends to merge into the surrounding

surfaces. This is completely effected in the skull of such a

form, for instance, as Geococcyx, where the temporal fossa are

A B

Fic. r.— Pata or (A) and inferior (B) views * n skull of Acedo ispida; 7, lacrymal; n

palatine; £, inter-naso-maxillary groo sega and f, the maxillary. Natura

size, dd Shufeldt, Jour. anat. Phys. "Vol. Xu

well separated, and the median ridge has become a broad sur-

face, indistinguishable from the general superficies of the cranial

vault. Holding a mid-position between these two conditions,

we find an example in the skull of Coccyzus americanus, where

the fossæ approach each other again.

Ceryle has its occipito-basicranial region circumscribed, and

its plane faces posteriorly and only slightly downwards, much

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 795

as we see it in the skulls of some herons. The subcircular

foramen magnum is of fair size, and the condyle comparatively

small. Anterior to it the basitemporal space is limited, while

the usual groups of foramina are to be seen upon either hand.

There is also a groove leading up on either side of the foramen

magnum, that terminates in a minute opening on the not very

elevated supraoccipital prominence. The superior limiting mar-

gin of the occipital area in Ceryle is very sharp, while in Alcedo

and Geococcyx, where the same characters first enumerated are

present, this boundary is more rounded.

Viewing the skull of either C. alcyon or C. cabanisi laterally (Fig.

2), the unusually deep temporal fossa is again brought to our

notice, occupying much of the space behind, terminating only at

the superior margin of the large and very open ear cavity, and

allowing but just enough surface for the lodgment of the head

of the quadrate.

It has above it a flattened squamosal process, which is sepa-

rated by a shallow concavity, still more anteriorly, from a feebly

pronounced post-frontal projection. The ali-sphenoidal wall

bulges forward in the form of an elevated dome, a feature

characteristic also of Alcedo. Above, the thin and horizontal

frontal affords the orbit an ample roof, this bone being carried

forward to join with the extensive surface of the superior por-

tion of the large /acrymal.

Viewed from above, this latter bone (Fig. 2) presents an

oblong surface with slightly rounded angles. Its infero-anterior

margin makes a close articulation with the nasal in front and

the frontal above, rather more than one third being devoted to

the first and the remainder to the latter bone. There is thrown

down from beneath this superior oblong plate of the lacrymal

another plate, the connection being a somewhat constricted

neck, which forms the greater part of the anterior wall of the

orbit. This plate is highly pneumatic, bulbous, and has a

smooth surface with a light spongy interior; its shape is square,

with rounded angles. Its outer portion below rests upon the

horizontally expanded maxillary, while its inner and upper angle

has wedged into it a small oblong process that is developed from

the ethmoid: it is the pars plana. Above, they embrace a

706 . THE AMERICAN. NATURALIST. (Vor. XXXVII.

large subelliptical foramen, through which, in life, the nasal

nerve and vessels pass. This inferior plate of the lacrymal is

much larger than the superior, and is roughly placed at right

angles to it.

Agreeing with C. alcyon and C. cabanisi, we find in Aledo

ispida, the form of the lacrymal and its articulations, sub-

stantially the same. It differs principally in being thicker from

before, backwards, and the pars plana, which in this kingfisher

is triangular, meets it about the middle of the inner border

rather than at the superior and inner angle, as it does in

Ceryle. In Geococcyx, the superior plate has moved down on

the margin of the orbit nearer the maxillary, articulating almost

exclusively with the nasal. This change nearly does away with

any distinction between an inferior and superior plate, the two

having run together in nearly the same plane.

The ethmoid here develops a very large pars plana, and the

lacrymal is so twisted to pass down in front of it, the former really

forming the anterior wall of the orbit. In Coccyzus americanus,

the pars plana, is very extensive, and forms the entire inter-

orbito-rhinal partition, while the lacrymal barely articulates with

it, it being a light bone, having very much the form we found

in Colinus.

One large vacuity is found in the interorbital septum in

Ceryle alcyon, as shown in Figure 2. This has the appearance

above of being divided in two, by a very narrow isthmus of

bone, in Alcedo, but we find this deception due to the large

foramina for the exit of the nasal nerves from the brain-case in

this bird, these openings being exceedingly small in Ceryle.

To return to the latter we find that the quadrate presents

little or nothing that is peculiar. Its orbital process is sharp

and spine-like, projecting into the orbital space, though over-

shadowed by the bulging wall of the alisphenoid above. This

process of the quadrate is remarkably stumpy and short in Alcedo,

while it is broad and flat in Geococcyx and Coccyzus, where it

almost rests against the posterior wall of the orbit.

Upon the inferior views of the skull of Ceryle, we observe the

anterior extremities of the palatines to be broad, horizontally flat-

tened and thin plates. They are separated from each other by

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 707

an interval of about two millimeters, and merge beyond into

the bony roof of the mouth already described. The interval

between their anterior ends is continued backwards to a point

well within the articulation of the heads that articulate with the

pterygoids. Near their inner margins, posteriorly, quite a large

foramen exists in each, which is a constant character. Opposite

the pars plana of the ethmoid in Ceryle, each palatine throws up

a triangular plate of bone to project freely into the rhinal space.

The postero-external angles of these bones are truncate, a feeble

spine being produced near the middle in Cery/e alcyon, a character

better marked in C. cabanisi while in Aledo ispida this character

is developed as a spine-like spur, and is very conspicuous.

This feature is also present in Dacelo gigas, while in Merops

the long and slender palatines are devoid of any postero-external

elongations.!

In Ceryle the inner margins of the posterior moieties of the

palatines always fuse together, and this fusion is carried back as

far as the pterygoidal heads, which it includes.

In my specimens of Coccyzus americanus there are no spine-like

projections from the postero-external angles of the palatines, but

the angles in question are distinctly defined, differing in this

respect from Cuculus canorus, where “the palatines are rounded

off postero-externally " (Huxley). This is the case also in the

skulls I have at hand of Geococcyx, but Huxley found them to

be distinctly indicated in the specimens he examined of this bird,

so here this character may vary perhaps with the age of the

individual.

I find no vomer in any species of Ceryle, and in this it agrees

with others of the group, for we know **in Kingfishers and hoo-

poes there is no vomer." è Geococcyx has a small vomer. Again,

in our kingfishers, the pterygoids are very long and straight,

their distal extremities having on their outer sides little horizon-

tally flattened and projecting triangular processes, inconspicuously

developed in some specimens, and entirely absent in Alcedo,

Geococcyx, and Coccyzus. None of the forms just mentioned

! See Fig. 29, Huxley's * Classification of Birds, etc." Proc. Zoöl. Soc. 1867,

P- 447-

2 Parker, W. K. and Bettany, G. T. Morphology of the Skull, p. 264.

708 THE AMERICAN NATURALIST. |. (Vor. XXXVII.

possess basi-pterygoid processes, and the pterygoids in the

ground cuckoo are very short in comparison with the great

length of the skull, being somewhat shorter than they are in

Alcedo.

In Ceryle the maxillaries are long and broad, being horizon-

tally expanded plates passing, on either side, into the exten-

sively developed maxillo-palatine masses anteriorly, which nearly

1G. 2.—1.— Right l i l of C.

nf, nasal — " nasal septum; g, quadrate; Ag,

maxillary.— Superior view of the same skull, lower ma

ew of ^s oo skull, owe mandible removed.—4. Superior

ble of Ceryle alcyon,— Left pelvic limb of Ceryle alcyon : f. fe

fibula; Z, psec ek P ‘hatluciat metatarsal ; 4, phalanx of hallux ;

—ó. The o-metatarsus, »eZ, in front view, from same limb.— »

i from

erat

branchial. Allfigures somewhat less than natural s x a in the same propor-

tion. After Shufeldt, Jour. Anat. & Physiol. Vol.

No. 442] OSTEOLOGY OF THE KINGFISHERS. 709

fill up the forward part of the rhinal chamber (Fig. 2). In

Alcedo this expansion is not particularly noticeable, while in

other respects the arrangement of their parts is the same.

Alcedo also differs from the various species of Ceryle in the

manner in which the quadrato-jugal bar articulates with the quad-

rate. In Ceryle it is received. into an articulating socket on the

outer side of the bone in question, while in Alcedo it meets it

much more anteriorly, though somewhat lateraliy.

Passing to the base of the skull, again, we must note how

small is the anterior aperture of the Eustachian tubes, and

observe the sharpened lower border of the sphenoidal rostrum.

The form of the lower mandible is well shown in figures 1

and 4. It does not essentially differ in Acedo zspida. Both of

these birds sometimes have the ramal vacuity spanned across

with a thin plate of bone, which may be pierced by a minute

foramen. Sometimes in C. alcyon the ramal vacuity is nearly:

fitted in by this plate. Among the cuckoos this bone is very

much more like the general form as we find it in the passerine

types.

In Coccyzus a large ramal vacuity exists, and the internal

angular processes are long and pointed, curving upwards, while

rudimentary posterior angular processes here commence to

make their appearance. In Ceryle the articular cups are deep;

the mandible is pneumatic ; the inturned articular processes are

short and blunt, while the hinder ones are truncated. The bone

is V-shaped with a very deep symphysis.

Nothing worthy of particular note rewards our examination of

the internal aspect of the brain-case. The usual sclerotal plates

are present in the eyes, and proportionate in size with other parts.

The arrangement of the ear-cell is very simple; the delicate

bony tie beams found in its interior, in so many birds, is here

replaced by solid bone, pierced only by the necessary openings.

The hyoid apparatus (Fig. 2, 7) departs very markedly from

ordinary birds. This is seen principally in the broad first basi-

branchial (4%), with its slender, connate second basi-branchial

(b'bh) reduced in this bird to a mere spine of no great length.

The epi-branchials are very much shortened, and extremely

delicate in structure, being tipped behind as usual with cartilage.

710 THE AMERICAN NATURALIST. [Vor. XXXVII.

Behind, the glossohyal (gZ) is broad and quadrilateral in outline,

the cerato-hyals being scarcely discernible at its posterior and

outer angles (ch).

The Axial Skeleton— The vertebrae in Ceryle, when com-

pared with many other birds of about the same size, are large, in

comparison, with prominent processes. We find nothing to par-

ticularly distinguish the atlas. The plate closing in the neural

canal of this segment above is oblong in outline, with a minute

spine at each outer and posterior angle. The body is thick from

before, backwards, so the shallow cup for the occipital condyle

is never perforate, as it is in many birds. A large neural spine

is found on the axis, and the diapophyses are elevated. Situated

somewhat posteriorly, a neural spine, smaller than that of the

axis, is found on the third vertebra, and this process diminishes

in size as we proceed backwards, to disappear entirely on the

ninth vertebra. The twelfth has a small one again, becomes

larger still in the thirteenth, and in the next of the series appears

very much like the elevated quadrate plates as seen in the dor-

sals. In the third vertebra the foramen found in the plate

between the pre-and postzygapophyses, as a common avian

characteristic, is here sometimes scarcely perceptible. But in

this vertebra two other features arise — the cervical extremity of

the vertebral canal, with minute parapophyses projecting from it

on either side, and, secondly, the appearance of an hypapophysis

beneath. In some of the leading cervical vertebrz after the

third, usually the fourth, fifth and sixth, there is a bridge of

bone, on either side, connecting the posterior margin of a para-

pophysis with the antero-external base of the corresponding

postzygapophysis. This bridge becomes absorbed behind, in the

sixth and may be in the seventh and eighth vertebra, and then

projects from the parapophysis simply as a spine-like process.

The vertebral canal persists through the cervical chain to include

the twelfth vertebra; in the thirteenth it is closed in by a very

delicate little rib, consisting of but scarcely anything more than

head, neck and tubercle. In the fourth vertebra the hypapophy-

sis is but feebly developed, while the parapophyses are much

stronger; these latter disappear in the twelfth vertebra. The

carotid canal traverses the inferior aspects of the centre of the

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 711

fifth to the ninth vertebra inclusive ; while in the tenth, eleventh

and twelfth a median single plate reproduces the hypapophysis

once more. This process is three-pronged in the thirteenth ;

bifurcate, with nearly horizontal limbs, in the fourteenth, which

latter decrease in size in the next, although the process length-

ens, — and thus it continues, as a rule, throughout the dorsal

series, being shorter only upon thelast two. In some specimens

of C. alcyon I have seen this hypapophysis quite rudimentary

upon the last dorsal vertebra, and in a specimen of C. cabanisi

(U. S. National Museum) I find these spines present, though

they are not very long, upon the two leading vertebra of the

sacrum. There is also at my hand a disarticulated skeleton of

C. alcyon, and in these two individuals, this one and the afore-

said skeleton of C. cabanisi, I find after careful count, rather a

remarkable difference in their vertebra. In the skeleton of C.

alcyon there are nineteen free vertebrze from the occiput to the

pelvis, whereas in the skeleton of C. cabanisi, there are but

eighteen. Again, in the latter we find the first pair of free

riblets, (cervical ribs) on the ¢wed/fth vertebra; a stronger pair

on the thirteenth ; and a free pair again on the fourteenth ; the

last ones have epipleural appendages upon them. Now in the

skeleton of C. alcyon, the first free cervical riblets occur upon

the ¢hirteenth vertebra ; a stronger pair is found upon the four-

teenth; and finally, a free pair, with epipleural appendages,

upon the fifteenth. In my paper upon the osteology of C.

alcyon in the Journal of Anatomy (London, 1884) I see that

the specimen whose skeleton I there describe, had the same

arrangement as the skeleton of the individual now before me

belonging to the collections of the U. S. National Museum.

That there is a variability in the number of vertebrae appears

from what follows ;— for I find that in my paper, first men-

tioned, I say of the dorsal series of ribs and vertebra, in C.

alcyon that “The four dorsal vertebrz, with their ribs connect-

ing with the sternum, have nothing very peculiar to mark

them. They freely articulate with each other, and develop

stumpy metapophyses on their transverse processes. ; The unci-

form projections are not anchylosed with the ribs. Two

pairs of ribs are suspended from beneath the ilia, belonging

712 THE AMERICAN NATURALIST. (VoL. XXXVII.

to the antecedent vertebrae of the so-called “sacrum.” The

first pair have small unciform processes, their haemapophyses

articulating with these bones on the last dorsals in the usual

manner; the last pair, which are very delicate in structure, vary

exceedingly in length, and terminate in free extremities." Now

the just-quoted description does not tally with what I find in the

skeleton of C. alcyon from the U. S. National Museum, for in

the first place in the sternum of that individual there are fous

articular facettes upon the right costal border and but ż%ree

upon the Zz/z The fault here, however, or this discrepancy

seems to be due to some difference in arrangement at the fore

end of the series, as the /c/7 costal process is longer, and has a

suspicious looking spine anchylosed to it, at about the point

where the leading costal rib ought to have had its facet. The

total number of ribs in these two specimens of C. alcyon may,

however, have been the same, while a difference only existed as

to the number that connected with the sternum by costal ribs.

This will not apply, though, to what we find in C. cabanisi, for

here, although there are four dorsal pairs of ribs, supplied by

the 15th, 16th, 17th and 18th vertebrae, only the three leading

pairs connect with the sternum by means of haemapophyses ;

the last pair failing thus to connect. Then, finally, there is but

one pair of “sacral ribs," and their hemapophyses are still

shorter than the pair on the last dorsal ribs, and these sacral

ribs are likewise without epipleural appendages, being the only

pair that do lack them in this species. The arrangement of the

ribs in birds cannot always be relied upon nor even that the

same species always have the same number; but I cannot well

account for the difference in the number of the cervico-dorsal

vertebrae in these two kingfishers. The method of anchylosis

of the pelvic bones with the sacrum sometimes has something to

do with it, but apparently not here, for as near as I can count in

the pelves of these adult birds, there appear to be /qwe/ve ver-

tebrz in the pelvic sacrum of C. alcyon, and but eleven in C.

cabanisi. This evidently does not help, for to satisfactorily

account for the difference in question, the count should stand

the other way. i

Aside from the number of vertebrz in the pelvic sacrum, how-

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 713

ever, the pelves in these two species of kingfishers are very

much alike. In both, the pelvis is broad and shallow. The ilia

are widely separated from each other throughout their extent,

and, anteriorly, the sacral crista is low and inconspicuous.

Interdiapophysial parial foramina occur the entire length of the

pelvis, being very small, or sometimes only closed in at an area

constituting the central portion of the bone. Individual speci-

mens differ much in this respect, they being nearly absent in

Fio: 3 — Pelvis and coccygeal vertebræ seen from nite in Ceryle alcyon. —2. Left

liesi view of the shoulder girdle ys Cer. ryle alcyon ; bones a -~ ed as in iife; s, scap-

a; le. £ we. "

onn le aicy and shows very

y shaped sternum of this iortchar projects beyond

the body of the bone. — Left lateral view of the sternum of aie alcyon. Figures all

drawn by the author, eat size after Shufeldt, Jour. . & Physiol., Vol. 18,

1884.

[e]

L-

some (see Fig. 3). The superior surface of either ilium, anteri-

orly, is concaved, not extensive, looks upwards and outwards, and

is especially characterized by a sharp, backward-turned process

upon its outer margin. Viewed laterally, the pelvis appears

very shallow ; the pubis being a curved, slender rod of nearly uni-

form calibre that does not come in contact with the ischium

above, for its entire length. It projects somewhat posteriorly.

Upon this aspect a notch, triangular in outline, is seen to exist

between ischium and ilium at their posterior borders. Antitro-

714 THE AMERICAN NATURALIST. [Vor. XXXVII.

chanter is small, but the foraminal apertures, the acetabulum,

the obturator and ischiadic apertures, seen upon this aspect, are

of an average size.

Seen upon ventral aspect, we are to note, that usually but a

single pair of parapophyses are sent out as supporting braces

opposite the acetabule. The sacrum on this aspect exhibits,

along its middle portion, a medio-longitudinal groove that is very

striking even upon superficial examination; the centrum of the

anterior sacral vertebra is very deep and sharp. Specimens of

C. alcyon show three anterior sacral vertebrae that throw out

lateral processes against the nether aspects of the iliac walls.

C. cabanisi shows four. 'The pelvic basin is broad and capa-

cious in both these kingfishers, being more so in the former spe-

cies than it is in the latter.

The tail-vertebree are seven in number, not including the

rather small pygostyle. The three anterior ones usually have

long and slender diapophyses, while these processes in the last

four are broad and become gradually shorter as we proceed

posteriorly. "We also note that the last three caudal vertebra

develop bifurcated hypapophyses below, a character likewise

enjoyed by the pygostyle.

The sternum in the representatives of the genus Ceryle is

rather a curiously formed bone, and not only differently shaped

from the sterna of’ ordinary birds, but differs to some extent

among the several species. So far as I know, it is invariably

four-notched in this group of kingfishers, the xiphoidal prolonga-

tions being dilated at their hinder ends, more especially the outer

ones. Four facets for the haemapophyses of the dorsal ribs are

found on the superior aspect of each costal border, and beyond

the anterior ones prominent costal processes arise.!

The sternal carina is somewhat shallow behind, gradually

becoming much broader in front, where it protrudes considerably

beyond the body of the bone (see Fig. 3). In C. alcyon its

anterior margin is straight, while in C. cabanisi it is markedly

concaved; and in both species it extends the full length of the

sternum. Another difference is seen in the manubrial process,

1th lready stated that i alcyon examined, there were but

three of debe ions on the left costal Prec qug M the usual four upon the right.

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 715

for in C. alcyon it may be said to be almost entirely aborted,

whereas in C. cabanisi the supero-anterior angle of the project-

ing keel rears into quite a respectable manubrium.

The grooves for the coracoids do not meet in the median

plane by at least a millimeter or more in the larger species,

while we often find a pneumatic toramen present immediately in

front, upon the antero-superior edge of the projecting sternal keel.

Both these kingfishers, too, seem to possess a small, circular

foramen of this character, in the median line, on the thoracic

aspect of the body, just within the anterior coracoidal border.

Our belted kingfisher also has minute apertures of this nature at

the base of the interhaemapophysial pits along either costal

border.:

Coming next to consider the bones of the shoulder girdle

(Fig. 3) we find the scapula in C. alcyon to be rather broad, of

nearly uniform width throughout, being obliquely truncated from

within outwards for its posterior third. It has a conspicuous

process, which, when the bones of this arch are articulated,

extends in the direction of the head of the corresponding cora-

coid, having the furcula resting against its mesial aspect. Cery/e

cabanisi has comparatively a narrower scapula, and its posterior

end is more distinctly bent outwards. In it the coracoid

! Before passing to the consideration of the shoulder girdle, and closing my

description of the vertebral column and its attached bones, it is well to note an inter-

esting point in the vertebræ of Ceryle which I believe was firs t semiopemted by

W. K. Parker in his paper “On the Osteology of Steatornis caripensi (Proc

Zool. Soc. of Lond. Apr. 2, 1889; p. 175). In comparing the skeleton a Stentor:

nis with our kingfisher (a specimen of which,I had sent him), Parker says “In

a New-World Kingfisher (Cery/e alcyon) the "hinder dorsal centra make a great

approach to those of Steatornis, without, however, being opisthoccelous.” And

again upon the same page. “The sacral vertebre [of Steatornis] and the whole

pelvis are very much like siat of Ceryle alcyon,— the Kingfisher whose dorsals

show a tendency to the celous character, and have deep, concave-sided

dorsal centra, with long, eit ‘diated, inferior spines.

I have personally made this same compa rison, and can confirm the points

above noted by Parker, especially in the pelves of these birds, is the resemblance

striking. . Steatornis, however, lacks the peculiar ma upon the outer free

margin of either ilium, seen in the kingfisher, and upon ventral aspect it would

act as braces opposite the acetabulæ, as they are in Ceryle. For the remainder

of the skeleton in these two forms, the resemblance ends entirely, beyond the

points just cited.

7 16 THE AMERICAN NATURALIST. [Vor. XXXVII.

process, just described, does not reach the coracoid when the

bones are assembled zz sítu. The coracoid is considerably

dilated at its sternal end, with raised facet there on its posterior

aspect for sternal articulation. Its shaft is straight, not stout,

subcylindrical in form, and rather long. At its anterior end we

note a large glenoidal facet, and the usual inwardly-crooked,

tuberous head. A spiculiform, clavicular process, is to be seen

at some considerable distance below this, upon the mesial aspect

of the shaft. ‘his is equally well-marked in C. cabanisi, a

species having a coracoid after the pattern of that bone in the

typical picivorous kingfishers.

Our belted kingfisher has an os furcula of the broad, — very

broad U-shaped model — without a semblance of a hypoclei-

dium. The bone is slender below, but the clavicular limbs

gradually become broad, and much compressed laterally, as we

proceed in the direction of either head. So that, viewed as a

whole, either clavicular limb may be said to be almost blade-like,

with the free end, when articulated zw sztw, reaching back along

the antero-mesial aspect of the corresponding scapular for some

little distance. Ceryle cabanisi is peculiar in having a process

developed upon the superior margin of either clavicular head,

which, passing upwards and backwards, articulates with the

tuberous head of the corresponding coracoid at a point situated

at its antero-mesial aspect. I find no os humero-scapulare in any

of these birds."

The Appendicular Skeleton— In the pectoral limb the humerus

is the only bone possessed of pneumaticity ; — the pneumatic

foramen being single, very Open, and fills the entire base of the

fossa. Its surrounding margins in C. cabanisi are thickened,

and the fossa is markedly shallow. Proportionately, this bone is

very long, — being but two thirteenths shorter than the bones of

With respect to the method of articulation of the os furcula in C. alcyon, I

find that in my above-quoted paper upon the osteology of that species I said that

“In the articulated skeleton (Army Medical Museum, Washi ington, Section Com-

parative Anatomy, No. (55) it rests against the anterior border of the sternal

keel at the decies of the middle and lower third."

us n with caution, as it may not be invariably the case, and the

skeletons in the idus referred to, are by no means to be Me upon. In

C. cabanisi it does not thus rest upon the anterior carinal bord

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 717

the antibrachium. Its radial crest is but moderately developed,

and a deep valley is sculpt 'twixt ulnar tuberosity and humeral

head. In other respects this bone presents the usual characters

found among birds generally.

The osseous tubercles for the quill-butts of the secondaries,

found in many of the class along the shaft of the ulna, are here

absent or very faintly perceptible in some specimens. Both

bones of the antibrachium in the well-developed limb of these

kingfishers are as we usually find them in ordinary birds. C.

alcyon upon the superior aspect of the distal end of the radius,

exhibits a single, deep groove for the guidance of tendons. The

two carpal segments are well apart from each other, thus

affording quite an extensive share of the head of the metacarpus

for articular surface for the ulna. I find no small sesamoids,

either at the wrist or at the elbow. The index metacarpal, the

main shaft of the carpo-metacarpus, develops at the proximal

extremity of the shaft, the palmar aspect, the flattened process

which is seen in other groups of birds. It nearly reaches across

to the border of the adjacent middle metacarpal, and this latter

bone is distally longer than the metacarpal of the index, pro-

jecting slightly beyond it. The expanded portion of the proxi-

mal phalanx of index finger is non-perforated and well-developed.

One phalanx is allotted to pollex digit, two to index, and one

to medius, claws and spurs being absent in the manus of Ceryle.

Formerly, I believed the pelvic limb in the kingfishers was

entirely non-pneumatic, but by examining far more extensive

material, I am convinced that the femur may be pneumatic in

many individuals. The National Museum specimen (No. 18749)

of C. alcyon is peculiar in this respect, for the right femur

undoubtedly enjoys this condition, whereas the fellow of the

opposite would, I think, be taken for a non-pneumatic bone. For

its remaining characters we are to note, that although the tro-

chanter projects somewhat anteriorly, it does not rise above the

summit of the shaft. This latter is straight and cylindrical,

being but very feebly marked by ridges or lines for muscular

attachment. An ordinary avian patella is present.

On the proximal and anterior aspect of the tibio-tarsus the

pro- and ectocnemial ridges are but feebly produced: they are

718 THE AMERICAN NATURALIST. [Vov. XXXVII.

best marked in C. cabanisi. An evident feebleness stamps the

bones of the leg, that is when we came to compare them in size

with other long bones of the skeleton. Kingfishers having weak

pelvic limbs, we naturally find the skeleton of the parts likewise

weak. Nowhere is this better seen in the limb of this bird, or ,

birds of this group, than in the tarso-metatarsus and foot-skeleton.

At the distal extremity of the tibio-tarsus of C. a/cyon we find

the antero-osseous bridge for the confinement of certain tendons

to be situated very low on the shaft, indeed, it occurs immedi-

ately above the rather prominent condyles.

The fibula has but a slender spine of bone below the fibular

ridge of the tibio-tarsus, and in some specimens I find even this

missing. When this latter condition exists, C. a/cyon has as

short a fibula as any bird of its size, with which I am at present

acquainted.

The tarso-metatarsus is less than a third as long as the shaft

of the tibia (Fig. 6), the rather large free metatarsal for hallux

thus being compelled to take a position in articulation very near

the middle of the shaft of this bone. The **hypotarsus" is large

in proportion, and exhibits a median groove, posteriorly, for the

passage of tendons, and immediately anterior to it a single cylin-

drical perforation for the same purpose. Distally, the tarso-

metatarsus, has three well-developed trochlez, either of the

lateral ones being larger than the bigger one in the middle. The

foramen for the anterior tibial artery is present, while at the

proximal end of this bone in front we note a small protuberance

for the insertion of the tibialis anticus muscle, as well as two

small antero-posterior perforating foramina, placed close to each

other side by side.

The digits of the foot possess the normal number of phalanges

(2, 3, 4, 5), as they occur in the class. The basal joint of hal-

lux is somewhat laterally expanded at its proximal end. With

respect tothe joints of the other toes, the three anterior ones,

we note that the proximal joints have an articulatory movement

nearly in the same subhorizontal plane, while careful examina-

1 This corrects a statement I made on this head in my paper ** On the Oste-

ology of Cery/e alcyon, quoted above. There are a few other slight inaccuracies I

have taken the opportunity to correct in the same manner.

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 719

tion of the opposed surfaces of the bones composing the outer

and middle toes, show the effect of their being so long strapped

together in a common podothecal sheath, in the evident compres-

sion of the ridges and elevations usually found on these phalangeal

bones in avian feet where the digits are free.

SYNOPSIS OF THE PRINCIPAL OSTEOLOGICAL CHARACTERS

OF THE GENUS CERYLE OF THE FAMILY OF THE

KINGFISHERS.

Superior mandible once and a half as long again as the

cranium ; tapering and of a pyramidal form, flat upon its

under side.

—

2. Septum narium complete.

3. Cranio-facial hinge quite movable (this is especially the

case in the dried skull, after the quadrate and lacrymals

have been removed). !

4. Lacrymal large, with a superior and inferior portion.

5. Pars plana very small.

6. Large vacuity in interorbital septum.

7. Maxillaries very broad, horizontally flattened plates. |

8. Maxillo-palatines large, subspongy, fused in middle line,

nearly filling post-rhinal chamber.

9. Palatines broad, horizontal, with the postpalatine parts fused

together in the middle line. A large foramen pierces either

postpalatine.

10. Pterygoids long and straight.

11. Crotaphyte fosse deeply sculpt, and only separated by a

crest posteriorly.

12. Lower margin of sphenoidal rostrum sharp.

13. Lower mandible V-shaped; long deep symphysis ; articular

cated behind, with stumpy inturned processes.

cups deep, trun ;

basibranchial

Hyoidean arches peculiar, with very broad

and cerato-hyal parts. ;

Nineteen cervico-dorsal vertebra in C. akyon, and only

—_

THE AMERICAN NATURALIST. [Vor. XXXVII.

eighteen in C. cabanisi, with a variable arrangement of the

ribs.!

Vertebra comparatively large; long hypapophysial proc-

esses characterize the ultimate cervicals and the dorsals.

Centra of dorsal region laterally compressed, subopistho-

coelous (Parker). Pneumatic. Neural canal large in mid-

division. Carotid-hypapophysial canal open. Seven caudal

vertebre and a pygostyle.

Pelvis broad and shallow; ilia widely separated from sacral

crista, with a peculiar process on the outer free margin of

either bone. Postpubic element long and slender, projects

posteriorly, and is not in contact with ischium above. One

pair of vertebral ae thrown out as braces opposite

the acetabulae.

Sternum 4-notched behind, with carina projecting in front.

Manubrium either entirely aborted or much reduced. Pneu-

matic.

Os furcula broadly U-shaped, slender below, broad above,

where in C. cabanisi a peculiar process is developed extend-

ing to head of coracoid on either side. Coracoids long, and

moderately slender. Scapula with conspicuous coraco-

clavicular process, the blade of the bone being narrow,

longly truncated behind, and in C. cabanisi turned outwards

for its posterior third.

Pectoral limb well-developed: humerus pneumatic: long.

Process present on proximal end of index metacarpal, pal-

mar aspect. Blade of proximal phalanx of index digit

entire.

Lower part of pelvic limb feebly developed. Femur may

or may not be pneumatic. Patella present. Fibula weak.

Tarso-metatarsus very short, with its hypotarsus once

pierced and grooved for tendons. Podal joints run 2, 3, 4,

5 respectively, feeble, and in the anterior toes somewhat

laterally compressed. `

1 This character needs further investigation, and it should be either confirmed

or disproved

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 721

NEGATIVE CHARACTERS.

(Ceryle.)

1. Vomer absent.

2. No basipterygoid processes.

3. No hypocleidium on os furcula.

4. Os humero-scapulare absent.

5. No prepubis on pelvis.

6. Claws not present on digits of manus (?)

7. Aside from patellæ, sesamoids not found in the limbs.

FURTHER OSTEOLOGICAL COMPARISONS, WITH NOTES ON THE

POSITION OF THE KINGFISHERS.

My opinion upon the systematic position of the kingfishers

is but a tentative one, for I have not examined all the material I

. could wish in order that I might render it more positive. There

are many forms of kingfishers in the world, and they vary much

among themselves in their organizations. The anatomy of these

various species should be fully comprehended, and with this

knowledge at our command we should fully investigate the

structural economy of many forms of birds that we now sus-

pect of being alcedine affines, —— and it will only be when this

is done that we will gain any correct idea of the subject.

As the present memoir shows, I have carefully looked into

the skeletology of our two species of Ceryle. Further, I have

examined in connection with other extensive materials a skeleton

of Aledo ispida, and a skull of Dacelo gigas. Skeletons of

Coccyzus, Crotophaga, and Geococcyx have also been thoroughly

compared, and I have compared skeletons of a jacamar, of

Diplopterus, and one of the Meropida from Borneo, vzz.:—

Nyctiornis amictus. Also, I have studied some of the skeletons

of various Bucerotidz and the Trogones, as stated.

We find the skeletons of the extremes of the insectivorous

Daceloninze and the piscivorous Alcedininz to differ consid-

erably in their characters, and I am of the opinion that it will

be through a study of the osteology of the Daceloninz that

722 THE AMERICAN NATURALIST. [Vor. XXXVII.

we will eventually come to a knowledge of the kinships of the

kingfishers with other groups of birds.

To me, it is not difficult to see the reason for Huxley's

remarks when he said “ A/cedo and Dacelo repeat the structure

observed in Geococcyx, with minor modifications, and that ‘the

Alcedinide [approach] the Pelargomorphæ ” (Proc. Zool. Soc.

1867, pp. 447 and 467).

Already I have committed myself to the belief that the Hal-

cyones are most nearly related to the Galbulide. Still, I think

so, but I likewise think that the osteological resemblances will

best be seen when we come to compare the skeletons of certain

jacamars with the more aberrant Daceloninz.

The specimen of Jacamar (sp.?) now at my hand shows in

its skull some characters that agree pretty well with the corre-

sponding ones in the skull of Cery/e cabanisi, but as a whole the

Jacamar's skull agrees equally well, and in some respects better,

with the skull of Geococcyx. It differs from both in having an

entire orbital septum ; in having a remarkably long post-frontal

process, with a complete abortion of the squamosal process ; and

in the supraorbital prominence being so conspicuous. Its pars

plana and lacrymal most nearly approach Geococcyx, but its

lacrymal is peculiar in the great definition of its external notch.

The osseous mandibles are most like the kingfisher's, while

its palatines and its maxillo-palatines again remind us most of

the same bones in Geococcyx.

It seems to lack a vomer, and this would again suggest Ceryle

or perhaps some other more nearly related Kingfisher.?

With respect to the sternum, we remember that the fore-part

of the carina in Ceryle alcyon and Ceryle cabanisi differ con-

siderably in their form. Now, the fore-part of the sternum, the

carina of the same, and the costal processes in this Jacamar,

more nearly agree with what we see in these parts in C. cabanisi

than do the same parts agree between the two kingfishers just

! [ can still see this, notwithstanding the fact that the postero-external angles

of the palatines in Geococcyx are zo£ produced as processes, as they are in Alcedo

and Dacelo, and as Huxley thought they were.

2 There are some points about the skull of this Jacamar that call to my mind

the skull of certain of the Trochili.

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 723

mentioned. But in the jacamar the two notches upon either

side of the keel are notably profound, and the xiphoidal pro-

longations very slender ; the inner ones not being in the least

dilated at their hinder tips, while very handsome dilations

characterize the ends of the outer pair. Now to make the ster-

num of C. cabanisi agree with the sternum of the Jacamar, we

would simply have to deepen its notches to a similar extent, and

the two bones would then be very considerably alike. Again

the fore-part of the pelvis in the jacamar is quite after the

order of the kingfisher, but the resemblance is lost when we see

in the former how peculiarly the ilia and ischia are produced

behind ; but here, too, this appears to be but an exaggeration of

what we really see in Ceryle alcyon. Aside from the zygodactyle

feet of the jacamar, we could easily select some very good

characters, that seem to indicate that in the remainder of the

trunk skeleton, and the skeleton of the limbs of this bird there

is much more than a mere resemblance to the corresponding

parts in the skeletons of our two North American kingfishers.

When we come to compare our skeleton of Nyctiornis amicta

as representing the Meropida with the skeletons of the two

species of Ceryle we have under consideration, we meet with

still other characters that suggest inter-relationship for these two

families, while Nyctiornis has some peculiar characters of its

own. In principle, the arrangement of the bones of the palatal

aspect of the skull, agree in Geococcyx, Ceryle and Nyctiornis,

and I may say the resemblances are distributed with the most

perplexing singularity.

On the whole the superior osseous mandible in Nyctiornis is

more like what we see in Geococcyx than in either of our king-

fishers, while the lower jaw possesses characters common to

both, and is peculiar in not having even the vestige of a ramal

vacuity, wherein it disagrees with both.

The lacrymal in Nyctiornis is very similar to that hone as it

occurs in Geococcyx, but it is narrower, and not quite so tuberous ;

the exceedingly small pars plana, however, agrees with Ceryle,

and leaves in this bee-eater a great vacuity between lacrymal

and pars plana, which opens into the rhinal chamber. Its inter-

orbital septum is thick and practically entire; this agrees with

724 THE AMERICAN NATURALIST. [Vor. XXXVII.

the Jacamar, and differs with both the cuckoo and the king-

fisher. The form of the occipital region of the cranium and the

deep and sharply-defined crotophyte fossae of Nyctiornis agree

in many respects with Ceryle, but when we come to compare the

pterygoids and quadrates of our Bornean bee-eater with those

bones as they occur in Ceryle cabanisi, we are confronted with

decided differences. In the kingfisher the orbital process of a

quadrate is almost a hair-like process, while it is broad and strong

in Nyctiornis. Again, in the kingfisher the proximal end of the

pterygoid articulates with the mesial aspect of the inner mandib-

ular facet of the quadrate ; in Nyctiornis it articulates with a

facet situated directly between the orbital process of the quad-

rate and its mandibular condyle. The mandibular sesamoid in

the bee-eater is unusually large, is moulded upon the quadrate,

and articulates with a special facet upon the angular extremity

of the jaw.

The post-palatines of Nyctiornis fuse together in the middle

line, and with the small, rod-like vomer of this bird, while the

postero-external angles of the palatines are angulated without

possessing a very distinct process. The maxillo-palatines are

large and co-ossified across the middle space as in Geococcyx,

and they practically agree in these two species. At the lateral

aspect of the skull in the bee-eater we observe the post-frontal

and squamosal processes to be rather small, and the zygomatic

bar transversely compressed, and vertically rather broad. We

may add here that the skeleton of the hyoidean arches of the

kingfishers is peculiar, and is quite different from what we see

in the Meropida. I have not examined it in the jacamars.

Passing to the remainder of the skeleton we find many points

of agreement between C. cabanisi and Nyctiornis. Theip ver-

tebral columns and ribs are essentially very much alike, and this

is strikingly the case with the pelves of these two birds. In

their limbs and shoulder-girdles they also agree very well indeed,

but we find in Nyctiornis, on the other hand, a most remarkable

sternum. This bone has a manubrial process quite like what

we see in the sternum of C. cabanisi, but it differs in having an

osseous mesial bridge thrown across from its posterior aspect to

the anterior border of the body of the sternum, and this bridge,

No. 442.] OSTEOLOGY OF THE KINGFISHERS. 725

above, develops two lateral processes, one upon either side, while

the coracoidal grooves are below it. Posteriorly, this sternum

is even more peculiar, for externally its xiphoidal portion shows

two very deep lateral notches, with slender processes to their outer

sides which have large dilated hinder ends, while internally, that

is upon either side of the sternal keel, we find an elongated,

sub-elliptical fenestra. Either one of these fenestra have the

appearance of being formed by the hinder end of what would

have been the internal xiphoidal process, fusing with the pos-

terior point of the mid-xiphoidal prolongation.

There are two points, that should be mentioned, perhaps, in

the shoulder-girdle of Nyctiornis before closing my brief com- -

parative remarks upon its skeleton, and one is the conspicuous

expansion of the sternal end of either coracoid; and the other,

the fact that the outer aspect of either clavicular head has a

modified, jutting shoulder upon it for articulation with the

anterior aspect of the tuberous head of the corresponding cora-

coid.

There is evidently a hetero-kinship among Coccyzes, Halcyones,

Meropidz, Galbulida and undoubtedly other groups of birds,

but we will know little of the nearness or the remoteness of

these several affinities until the entire morphology and life

histories of the several groups are worked out.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS AND DEATHS.

EDUCATIONAL GIFTS.

Alleghany College, $20,000 from Mrs. H. F. Ballantyne.

Armour Institute $150,000 from J. O. Armour.

Barnard College, $1,000,000 from Mrs. E. M. Anderson for the IE

of land.

Beloit College, $50,000 from Andrew Carnegie for library building.

Carthage (IIl.) College, a conditional gift of $145,000 from Henry Denhart.

Cleveland, Ohio, $250,000 from Andrew Carnegie for library purposes.

Colby College, $46,000 from various sources.

Columbia University, $2,000,000 from J. Pulitzer for a school of Journalism,

$15,000 for scholarships.

Field Columbian Museum, a large sum (variously stated) from Marshall

Field. .

Hamilton College, $100,000 from Andrew Carnegie.

Harvard University, $50,000 from James Stillman for the hospital.

Illinois College, about $75,000 by the will of Dr. Hiram K. Jones.

Johns Hopkins University, $60,000 by the will of Mrs. Harriet Lane Johnston.

Kingfisher (Oklahoma) College, $25,000 from Andrew Carnegie.

Patterson, N. J., $130,000 for library building from Mrs. Mary E. Ryle.

Rush Medical School, $1,000,000 from various sources.

Syracuse University, one third the residuary estate of Mrs. Caroline S. Reid.

University of California, $500 from Mrs. A. S. Halliday for the library.

Vassar College, $13,000 and a new organ from various donors.

Wellesley College, $25,000 from Robert C. Billings.

Wesleyan University, $2000 by the will of Mrs. Henry F. Ackley.

Westwood, Mass., the greater part of the estate of Mr. H. O. Peabody, val-

ued at $1,000,000, for a girls’ school.

Williams College, $25,000 from Francis L. Stetson.

Winter Park (Fla.) College, $50,000 from Andrew Carnegie.

Yale University, $10,000 by the will of the late Walter D. Pitkins; $5,000

by the will of Mary P. Eakin.

APPOINTMENTS.

Carlos Ameghino, head of the division of paleontology, geology and

anthropology in the National Museum at Buenos Aires.— Dr. Rufus M.

727

728 THE AMERICAN NATURALIST. (VoL. XXXVII.

Bagg, professor of mineralogy and geology in the New Mexico Mining

School.— Charles J. Brand, assistant curator of botany in the Field Colum-

bian Museum.— Dr. E. Bresslau, docent for zoólogy in the University at

Strassburg.— William S. Bullard, assistant in zoólogy in the University of

Illinois.— Dr. S. M. Coulter, assistant professor of botany in Washington

University.— M. Danniel, professor of agricultural botany at Rennes.—

Dr. A. F. Dickson, professor of anatony in the college at Dublin.— Dr. J.

E. Duerden, acting assistant professor of zodlogy in the University of

Michigan.— Miss Ida Evans, instructor in biology in Rockwood College.—

Dr. A. C. Eycleshymer, director of the anatomical department of the Uni-

versity of St. Louis.— Nevin M. Fenneman, assistant professor of geology

in the University of Wisconsin.— Dr. Hans Fitting, docent of botany in

the University at Tübingen.— Russell D. George of Iowa University, pro-

fessor of geology in the University of Colorado.— Dr. Carl Wilhelm Genthe,

assistant professor of Natural History in Trinity College.— Gay M. Hamil-

ton, instructor in geology in the New Mexico School of Mines.— Dr. H.

Spencer Harrison, demonstrator of biology in University College, Cardiff.—

Emery R. Hayhurst, assistant in physiology in the University of Illinois.—

Dr. Ludwig Hecke, professor extraordinary of plant diseases in the Vienna

Agricultural School.— Dr. William A. Hilton, assistant in histology and

mbryology at Cornell University.— Dr. A. Hrdlicka, assistant curator in

the anthropological division of the U. S. National Museum. — Dr. B. F.

ingsbury, assistant professor of embryology in Cornell University.— C.

K. Leith, assistant professor of structural and economic geology in the

University of Wisconsin.— Dr. A. G. Leonard, state geologist of North

Dakota.— Dr. J. G. McCarthy, assistant professor of anatomy at McGill

University.— Dr. J. J. R. McLeod, professor of physiology in the Western

Reserve University.— Dr. Alphonse G. Malaquin, associate professor of

zoólogy in Lille.— Dr. Page May, lecturer on the physiology of the nerv-

ous system at University College, London. — Dr. T. H. Montgomery, Jr.,

professor of zoólogy in the University of Texas. Dr. J. P. Murlin,

Cornell University.— Amos W. Peters, instructor in zoology in the Univer-

sity of Illinois.— Dr. Joseph Felix Pompeckj, professor extraordinary of

paleontology in the University at Munich.— Dr. Raymond H. Pond, pro-

Dr. John L. Sheldon, professor of bacteriology in the University of West

Virginia.— Dr. Sievers, professor of geography in the University at

Giessen.— Dr. J. Rollin Slonaker, assistant professor of neurology in the

Leland Stanford University.— Ralph E. Smith, professor of botany in the

University of California. Dr. Johannes Sobotta, pr

: : : ofessor extraordinary

of anatomy in. the University at Wiirzburg.— M. E. Stickney, instructor

No. 442.] GIFTS, APPOINTMENTS, RETIREMENTS. 729

in botany Denison University— Michael X. Sullivan, instructor in chem-

ical physiology in Brown University.— Carl B. Tames, instructor in biology

in Western Reserve University.— Dr. Johannes Thiele, custodian of the

zodlogical museum of the University of Berlin.— Henry L. Ward, custo-

dian of the Milwaukee Public Museum, for five years.— D. G. Winter,

assistant in histology and embryology in Cornell University.

RETIREMENTS.

F. W. Cragin, from the chair of geology in Colorado College.— Sir

James Hector, from the directorship of the Geological Survey of New

Zealand.— W. W. Stockberger, from the position of instructor in botany in

Denison University.

DEATHS.

Andeas Allescher, mycologist, in Munich, April 10, aged 75.— William

Talbot Aviline, connected with the British Geographical Survey for many

years, May 12, aged 81.— Clarence Bartlett, Director of the Zoological gar-

dens in London.— Dr. J. Blum, zoologist, in Frankfurt a. M., April 24, aged

70.— Luigi Bombicci, professor of mineralogy in the University at Bologna,

May 17, aged 70.— Paul du Chaillu, African Explorer, in St. Petersburg,

April 3o, aged 68.— Dr. A. H. Chester professor of chemistry and mineral-

ogy in Rutgers College, April 13, aged 60.— Frangois Crépin, director of

the botanical garden at Brussels.— Charles Darrah, student of cactacez, in

Heaton, England.— Leonardo Fea, entomologist and assistant in the Civic

Karrer, geologist and secretary 5

aged 78.— Dr. A. Kaufmann, student of Ostracoda, in Basel, in March.—

Dr. W. C. Knight, professor of geology in the University of Wyoming, July

8.— Dr. Nickolaus Alexis Kubilin, geologist and former director of the

730 THE AMERICAN NATURALIST. [Vor. XXXVII.

Department of Mines of Russia, in St. Petersburg, April 23, aged 72.—

Prof. J. V. B. Laborde, director of the physiological Laboratory of the

Faculty of Medicine in Paris, April 8, aged 72.— J. Peter Lesley,

for many years geologist of the State of Pennsylvania, at Milton, Mass.,

June 1, aged 83.— Rev. Thomas Ansell Marshall, student of Hymenoptera,

at Ojaccio, Corsica, April 11.— Dr. Frederick V. August Meinert, long the

editor of the * Entomologiske Meddelelser," in Copenhagen, Mar. 3, aged

o.— Dr. Emmanuel Munk, professor extraordinary of physiology in the

University at Berlin, August 1, aged 51.— Dr. C. L. J. X. de la Vullée

Poussin, professor of mineralogy in the University at Louvain, Mar. 15,

aged 76.— Mr. Renard, professor of mineralogy in the University of Genth,

aged 60 years.— Heinrich, Baron Schilling von Canstatt, entomologist, in

Friedrichshafen, May 18, aged 53.— Dr. Heinrich Schurtz, ethnologist and

assistant in the city museum in Bremen, May 4.— Dr. Max Westermaier,

professor of botany in the University of Fribourg, Switzerland, May 1,

aged 50. me

(No. 441 was mailed October ro, 1903.)

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VOL. XXXVII, No. 443 NOVEMBER, 1903

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL

DEVOTED TO THE NATURAL SCIENCES

IN THEIR WIDEST SENSE

CONTENTS

I. Adaptations to Aquatic, Arboreal, Fossorial and Cursorial Habits in Mam-

mals. IL Arboreal Adaptations oy . LOUIS I. DUBLIN 73

Il. Mutation in Plants . - > DR. D. T. MACDOUGAL

III. Distribution of the DEEE Fishes of Mexico. DR. 5. E. MEEK

731

771

IV. Ezamination of Organic Remains in Postglacial Deposits

DR. P. OLSSON-SEFFER 785

799

800

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VI. Correspondence

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o ‘scientific Ce ees of the — |

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THE

AMERICAN NATURALIST.

Vor. XXXVII. November, 1903. No. 443.

ADAPTATIONS TO AQUATIC, ARBOREAL, FOS-

SORIAL AND CURSORIAL HABITS

IN MAMMALS.

II. ARBOREAL ADAPTATIONS.

LOUIS I. DUBLIN.

IN THE struggle for existence it is apparent that single forms

and whole groups of forms would independently become modified

to a life off the ground. Very often only by such an adaptation

could small defenseless animals save themselves from the attacks

of larger and speedier carnivores. In addition, there is the ques-

tion of food. The larger animals to whom the bulk of terres-

trial food naturally goes are virtually absent from the trees.

We accordingly find a multitude of animals that have made this

region their abode where, freed from their enemies and with an

abundance of food they have prospered.

PARTIAL LIST OF ARBOREAL MAMMALS.

Order MARSUPIALIA,

Family Didelphyidz (all except Chironectes).

* Phalangeridz

* — Macropodidze (Dendrolagus only).

* — Dasyuridz (Dasyurus and Phascologale only).

73!

732 THE AMERICAN NATURALIST. [Vor. XXXVII.

Order EDENTATA.

Family Bradypodide, Myrmecophagidz (only Tamandua and

Cycloturus).

Order UNGULATA.

Sub-order Hyracoidea (Dendrohyrax).

Order CARNIVORA.

Family Felide (many partly, the Jaguar alone truly arboreal).

* Viverridz (the Fossa, Viverra and Arctictis).

* Procyonidae (Procyon, comin Bassariscus, Nasua

and Bassaricyon).

Family Mustelidz (the Martens and Helectis).

Family Ursidz (the Brown Bear).

Order RODENTIA.

— Anomaluride.

Sciuridz (Sciurius).

* — Lophiomyidz.

* Myoxide.

* . Hystricidee (only the American sub-family Synethe-

rina).

Order INSECTIVORA.

Family Tupaiidæ.

* . Erinaceidæ (Gymnura only).

* — Galeopithecidze.

Order CHEIROPTERA.

Order PRIMATES.

(All except Homo and the Baboons.)

It will be observed from this list that with the exception of

the Monotremata, the Cetacea and the Sirenia all the mamma-

lian orders have arboreal representatives. Thus of the six

No. 443.] HABITS IN MAMMALS 733

existing marsupial families two are completely arboreal while

arboreal forms are found in one or more representatives of each

of the remaining families. Among those forms that are not

arboreal there still persists a considerable number of vestigial

structures and conditions in the pes all pointing unmistakably to

a previous arboreal life. In like manner among the edentate

sloths, many of the smaller Carnivora, Rodentia and Insec-

tivora and finally the Cheiroptera and in large part the pri-

mates have become arboreal.

This adaptation however is probably a secondary one, acquired

independently by the different orders. We should therefore

expect diverse forms of the adaptation to exist. Here we shall

distinguish the following main types :

-. I. Partially arboreal. These are still capable of terrestrial

life. Here belong the majority of the carnivores, insectivores,

and rodents, and Dendrohyrax.

II. Strictly arboreal. This contains the remaining forms

and is divisible into three sub-groups.

(a) Modified for running on branches.—Arboreal marsupials

and lemurs. |

(b) Modified for suspension from branches.—Sloths and bats.

marsupial type.— Remaining arboreal primates.

It is clear that this classification expresses corresponding dif-

ferences in foot structure. In the first group the pes is little

different from the typical terrestrial running foot. The pha-

langes have, as in the raccoons, become much elongated and the

soles are often naked. In some cases a distinct plantigrady has

replaced the previous digitigrady.

It is in the second group that the greatest modification has

occurred. In the first subdivision (a) the foot has become an

almost perfect grasping organ; the hallux being opposable ; the

second and third digits have reduced and united ; the fourth toe

is greatly elongated. There is also a distinct regression of the

claws; for as the foot becomes more and more prehensile in

structure the nail is no longer indispensable and is lost (Dollo).

In the second sub-division (b) of the second group, the manus

and pes have become much elongated and centrally strengthened

734 THE AMERICAN NATURALIST. [Vor. XXXVII.

and the nails have been modified into hooks by means of which

the body is kept in suspension. The number of digits is reduced

to two in Cholepus and three in Bradypus. The carpal and

tarsal elements are laterally compressed and there is some anas-

tomosis. This forms a more compact centre of resistance, while

the proximal bones develop a more or less complete ball and

socket joint in connection with the distal ends of the radius and

tibia, to permit a more perfect rotation.

In the last sub-division (c) both the manus and pes have

become grasping organs. The hallux or pollex, or both, are

generally opposable. Many modifications occur in the pes very

similar to those already described for the marsupials.

But in spite of these differences in main type there are

developed certain important characters which distinguish arbo-

real forms as a group from related terrestrial and aquatic types.

These like responses to the same conditions are to be observed

in what are otherwise most diverse forms. These characters are

the following :

I. The tail is often prehensile and, as in some of the Cebidze,

naked at the tip being a sort of fifth arm with which the animal

can move from branch to branch. Frequently correlated with

this adaptation is the loss of the thumb.

2. Ectodermal spines are often developed. These may

occur on the root of the tail as in the Anomaluride, on the

shoulder or on the feet as in Gymnura and some of the Anthro-

poidea. In all these cases the spines are climbing organs.

3. The limbs are much elongated. This elongation may

occur in different segments in different forms. In the swinging

apes, it is the fore-arm rather than the hand which is elongated.

In the tree-sloths all the limb segments except the compressed

carpalia and tarsalia and proximal phalanges are lengthened, the

very long remaining phalanges and the claws forming a hook for

suspension. In other forms the tarsals are greatly lengthened

as in Tarsius, Galago and other lemurs. These elongations are

obviously connected with the climbing and leaping habits of these

forms.

4. The hallux or pollex, or both, are generally opposable.

This gives the hand or foot a stronger hold on the branches and

No. 443.] HABITS IN MAMMALS 735

is perhaps the most important element in the arboreal limb. It

disappears however when the animal moves in suspension as in

the sloths.

5. The clavicle and scapula are well developed. These give

strength to the fore extremities and thus increase the climbing

power. It is interesting to observe that, as occurs in the Hystric-

idze, the clavicles will be developed in one arboreal form while a

terrestrial member of the same family will have vestigial clavicles

or none at all. Together these two bones strengthen the pec-

toral arch “in the transverse direction; that is, against lateral

strains of pulling and pushing, which came almost entirely from

the use of the anterior limbs (Cope).”

6. The ilium is broadened in some forms, particularly in

Anthropoidea and the tree sloths. This adaptation is for the

support of the viscera. In the edentates the pubis is directed

posteriorly.

7. In arboreal forms the ribs and chest are powerfully

developed as compared with the conditions in their non-arboreal

relatives.

8. The number of the dorsolumbar vertebra is often

increased. It is in the tree sloths among the Edentata that

the greatest elongation occurs. In the two-toed Cholepus the

number is twenty-seven, and twenty-five in the species didactylus

and hoffmanni respectively, while the number typical for the other

forms of the order is about nineteen. In the three-toed Brady-

pus the number is the typical nineteen. Curiously enough. it is

the cervical region which is here elongated there being nine cer-

vical vertebrae instead of six or seven as in the remaining Eden-

tata. While one form has specialized itself to firm suspension

the other has more or less sacrificed this character for a perhaps

more valuable one — the mobile neck. Among the Rodentia

where the typical number of dorsolumbars is nineteen, Capromys

which is arboreal possesses twenty-three. Hyrax and Dendro-

hyrax have thirty and twenty-eight respectively ; fully six more

than that prevailing among the terrestrial ungulates.

If inverse evidence can be of any value, it is known that in

the human species, ancestrally adapted to arboreal life, there is

a tendency toward the shortening of the back ; there being gen-

136 THE AMERICAN NATURALIST. [Vor. XXXVII.

erally, one less vertebra in man than in the still arboreal apes.

On the other hand among the marsupials where typically arbo-

real forms prevail the number is constant for the group — nine-

teen. This may be due to the fact as Dollo has shown that the

terrestrial forms have but very lately modified themselves to this

mode of life — the whole group of marsupials having been at one

time arboreal. In like manner the number in the carnivores is

constant (twenty). In this group the arboreal forms have but

lately diverged from their terrestrial relatives. In the Insec-

tivora there is also no difference of any significance.

Among isolated adaptations may be mentioned the modified

feet of Hyrax and Dendrohyrax. As described by Dobson

these animals are enabled to climb perpendicular walls and trees

without the use of claws; nor is there an opposable hallux or

pollex. The thickly padded tuberculated soles are drawn up

by certain flexor muscles thus leaving a vacuum by means of

which the animal retains its hold. In the Cercolabide there are

in addition to other arboreal characters such as spines, tubercles

on the soles which may serve as in Hyrax.

BIBLIOGRAPHY.

ALLEN, H.

°93. A Monograph of the Bats of North America. Bull. U. S. Natl.

Mus. No. 43.

BEDDARD, F. E.

:02. Mammalia. Cambridge Nat. Hist. Vol. to.

GOPE, E: D.

':89. The Mechanical Causes of the Development of the Hard Parts

jin the Mammalia. Journ. Morph., Vol. 3

DoBSsON, G. E. :

‘76. On Peculiar Structures in the Feet of Certain Species of Mam-

mals which enable them to walk on Smooth Perpendicular Surfaces.

Proc. Zool. Soc. 1876. London.

DorLo, L.

'98. Les ancéires des Marsupiaax étaient-ils arboricoles? Trav. Stat.

Zool. Wimereux, Tom. 7.

FLowER, W. H. and LYDEKKER, R.

'91. An Introduction to the Study of the Mammals Living and

Extinct.

MUYBRIDGE, E.

"77. Animal Locomotion.

MUTATION IN PLANTS:!

D. T. MACDOUGAL.

Ir is presumably safe to say that all students of natural his-

tory agree in the opinion that living matter has qualities at the

present time that it did not originally, or always possess, and

furthermore it is universally conceded that protoplasm is under-

going such development that it is constantly acquiring new prop-

erties, and taking form in an increasing number of types, kinds,

or species of organisms as a consequence. In other words living

matter is increasing the number of its qualities, multiplying the

number of forms in which these qualities are variously grouped,

and at the same time undergoing such differentiation that an

increasing complexity is the general tendency of the organic

world. These facts once realized the biologist finds himself

confronted with two stupendous interrogatories. By what

method is the ‘general development and differentiation of

organisms brought about as expressed in the formation or

origin of new species, and secondly what are the general fac-

tors which shape this progression? The amount of mere dis-

cussion ensuing from the presentation of conflicting views

brought out by these questions, in comparison with the total

scientific effort to obtain positive evidence upon the points

involved is appalling to contemplate. Happily the biological

world is becoming intolerant of wrangling and speculative con-

tentions, and has earnestly set about finding the facts that will

afford an adequate and satisfactory solution to the main problems.

The cult of the study of statistical variations may be regarded as

one expression of this newly assumed attitude, while the devious,

intricate and oft-times labyrinthine ways of cytological investi-

1The general discussion of the mutation theory embodied in this paper,

together with an exhibition of the seedlings of CEnothera was given before he

Zoological Seminar of Columbia University, April 23, 1903. The comparisons

between the mutants were not completed until August, 1903.

737

738 THE AMERICAN NATURALIST. [Vor. XXXVII.

gations have, or should have, their chief purpose in the discovery

of the physical mechanism of heredity.

The terms discontinuous variation, or mutation in connection

with the study of inheritance, descent, and the origin of species

may be taken to mean the autonomous physiological processes

by which one or more individuals of a species give rise to off-

spring which exhibit qualities, or groupings of qualities not pos-

sessed by their immediate ancestors and not previously exhibited

by the individuals comprised in the parent species (progressive |

mutation), or by which one or more individuals give rise to indi-

viduals lacking qualities or groupings of qualities exhibited by

the ancestral forms (retrogressive and degressive mutation).

These aberrant individuals or mutants may transmit their char-

acters to their offspring in such a manner as to give rise to a

new line of descent constituting the origin of a new type by

mutation.

The number of freaks, sports, bud-variations, and specimens

of plants with abnormal forms and sizes of leaves, stems, and

flowers, some of them highly teratological, to which attention

has been called by various writers in botanical periodicals under

the designation of mutants makes necessary the emphasis of

the fact that observations on a single individual, or a single gen-

eration of individuals are of but little value in distinguishing

fluctuating variations from mutations. Results worth a moment's

consideration may be obtained only by the most careful exclusion

of the possible effects of disease, of animal or plant parasites,

of hybridization, and by a careful analysis of the phylogenetic

value of the divergences as tested by observations on successive

generations of living forms. It is in this manner, and in this

manner alone, that discontinuous, saltatory variations may be

distinguished from, the results of common, fluctuating and indi-

vidual variability. Mutation rests in the main upon such sub-

stantive, discontinuous variations as the acquisition of new

characters, or the loss of old ones hitherto transmitted by the

parent type, or upon simultaneous alterations of both kinds.

These changes may be accompanied by, or may result in, the

masking of current qualities, or the unae and energizing

of latent qualities of the parent type.

No. 443.] MUTATION IN PLANTS. 739

The essential differences between the two processes appear to

have been originally set forth by Charles Darwin,' and are treated’

at length by deVries. The formal distinctions drawn by deVries

appear to need some slight modification and elaboration in order

to make them universally applicable. Thus he holds that con-

tinuous, or fluctuating variability occurs only in accordance with

Quetelet's laws, and that it involves only the number, size and

weight of organs, and does not include differences in qualities.

Cultural experiments of various kinds during the last few years

have given results in which the qualities as well as the number,

size and structure of organs have been materially altered, but

such induced variations or divergences were not transmissible.

This particular factor in distinguishing between fluctuating and

mutating variability therefore becomes a safe one, when it is

modified to make mutating variability include only newly

acquired and transmissible qualities. The presence of a plant

or an animal parasite may not only change the mechanical fea-

tures of an organ but may also cause most radical alterations

in its physiological properties. A single example of the latter

may be cited in the case of the common species of Euphorbia in

which the affected leaves alter their geotropic sensibility in such

manner that they change from diageotropism to apogeotropism.

Such variations are not transmissible however, and in this lies

the true test between mutation and fluctuating variation. A

still further distinction consists in the fact that mutations ensue

in the rudimentary state of the individual, while the alterations

in qualities induced by any of the above factors in fluctuating

variability may be caused in various stages of the development

of the individual, but in a rudimentary stage of the organs

concerned. Mutative alterations arise with the individual, are

not the direct result of external factors, and are perfectly

transmissible, while fluctuating variations may arise by the influ-

ence of external factors at various stages in the individual

development, and are not transmissible in their entirety.

Much of the confusion inevitable to any discussion of the

subject may be avoided if it is borne in mind that we habitually

1DeVries. Mautationstheorie, Bd., 2, 1903.

740 THE AMERICAN NATURALIST. (VoL. XXXVII.

deal with two different conceptions under the term species, one

based upon systematic and the other upon physiological, or

sexual affinities. The last named conception considers species

as phylogenetic groups embodying certain elementary characters

and showing certain capacities and habits, some of which may

not find expression in external form and structure. The sys-

tematic conception of species runs closely parallel to the above

and should finally express the actual blood relationship of all of

the forms in the vegetable kingdom. It is practically impos-

sible however, to take into account features not actually expressed

in some definite measurable structure, or which may not be deter-

mined by some rigid physical standard, and comply with taxo-

nomic methods. Thus numerous undoubted instances are known

of two or more groups of forms embodving separate lines of

descent, which, however, may not be separable by taxonomic

standards. The present discussion is of course concerned only

with the physiological conception of species, although as may be

seen by an examination of the features of the mutant forms

brought under consideration, these present anatomical character-

istics sufficient to warrant their recognition upon any taxonomic

basis.

The special purpose of the present paper is to consider dis-

continuous variation as a probable method of the origin of new

species, and to present the results of two season's observations

on the form, habit and behavior of some of the mutant forms

discovered by deVries seventeen years ago.

The observation and recording of marked examples of discon-

tinuous variation in lines of descent is as old as biological sci-

ence itself. Recently this procedure has been brought into the

focus of attention anew as the result of the deVries investiga-

tions, which tend to demonstrate that it is an important means

by which species come into existence. More than three cen-

turies ago (1690), Sprenger the apothecary of Heidelberg, who

had Chelidonium majus under cultivation, noted the sudden

appearance of a type with laciniate leaves in his garden. This

form which is also distinguished by other characteristics, was

found to be constant and self-maintenant in competition with the

parent type, and has remained distinct to the present day with-

No. 443.] MUTATION IN PLANTS. 741

out artificial selection, and no specimens have ever been seen

which could not be traced back to this original lot of individuals

in Heidelberg. The citation of a large number of equally well

or better authenticated instances of the sudden origin of types

is to be found in Korschinsky’s memoir to which reference is

made below.

The space at command does not permit even an outline of an

historical sketch of the views of the more prominent writers on

descent, concerning discontinuous variation as a means of origin

of species. It may be said, however, that Darwin attributed

some importance to “single variations” in his earlier writings

but seemed to relinquish this favorable view of the matter under

the pressure of criticism to which he was subjected in connec-

tion with all phases of his opinions on the origin of species.

Kolliker's theory as to the transmutability of egg elements as a

means of heterogenesis in 1864 will be recalled in this connec-

tion.

Dollo is credited by deVries with being the first to announce

definitely the conclusion that species might originate by muta-

tion (1893) (JMutatzonstheorie, Bd. 1: p. 46). . Bateson goes so

far as to say in his summary of The Material for the Study of

Discontinuous Variation (1894) that * It (The evidence of varia-

tion) suggests in brief that the discontinuity of species results from

the discontinuity of variation.

Korschinsky (1899) published a most valuable historical

account of the better authenticated instances of types sup-

posedly originating by discontinuous variation, and made a

comparison of the theories of natural selection and hetero-

genesis. The German reprint of his paper (//ora, 89, pp. 240-

363, 1901) is the completest yet published in citation of facts

and in review of pertinent literature, and it forms a logical his-

torical prelude to the observations of deVries.

The first well-guarded scientific observations of the origin of

new types as a result of discontinuous variation were made by

deVries, who by the expenditure of a great amount of labor

carried out an extensive series of experiments in the cultivation

of plants of the old Œnothera lamarckiana type. The general

facts obtained by him have been brought to notice repeatedly

742 THE AMERICAN NATURALIST. (Vor. XXXVII.

within the last three years and it will not be profitable to

rehearse the details at this time. Briefly stated deVries's inves-

tigations may be embodied in the following paragraphs.

I. Observations were chiefly concerned with a large num-

ber of plants growing wild and under cultivation, of the type of

(Enothera lamarckiana. The identity of the parent form was

found by comparison with the original description of the plant

made a century earlier, and by comparison with a type specimen

in the Muséum d'Histoire Naturelle in Paris collected in 1788.

The actual name of this plant in the revised nomenclature is a

matter of minor importance in the present connection.

2. Numbers of individuals of the parent type, as a result of

cross- and self-pollination indifferently, constructed seeds which

developed into independent forms, constant and self-maintenant,

which differed in habit, structure, stature, appearance and prop-

erties from the parent type.

3. The aberrant or mutant forms might be divided by

characters as sharp and numerous as most of the so-called

minor species of the systematist.

4. No forms intermediate between the mutants, or between

the mutants and the parent type were found.

5. That the mutant forms were really groups of phylogenetic

value was proven by their behavior when crossed with one another,

with the parent form, and with other species in the same genus.

The hybridization experiments with these forms has yielded

some exact evidence as to the preponderance of phylogenetically

older characters by reason of the fact that the mutants are

forms the exact ages of which are known. Of the crosses of

CE. lata and CE. nanella with the parent form, from a half to

three-fourths were found to be of the parent type, and the

remainder of the mutant type form. The crossing of mutants.

with each other produces a generation many of which show

reversionary characters. The mutation hybrids are constant in

succeeding generations. The separation of antagonistic charac-

ters in the first generations is weighty evidence in support of

the theory of elementary characters, and for the mutation theory.

6. The new types were either constant from the beginning,

or if weak, inconstant or perishing, showed no tendency to revert

No. 443.] MUTATION IN PLANTS. 743

to the parent type, and their MT or fixity might not be

increased by artificial selection.

7. Morethan one mutant might arise simultaneously from

the parent individuals.

8. Any one of the several mutants observed might originate

from several parent individuals simultaneously.

9. The mutant forms might arise from successive genera-

tions of the parent types.

10. The mutant forms might in turn give rise to new types

after their separation from the parent type.

The above statements rest directly upon observations of care-

fully conducted experimental cultures and admit of but little

argument as to interpretation. With this positive evidence at

hand questions at once arise as to the frequency, occurrence,

prevalence, exclusiveness, and as to the mechanism of discon-

tinuous variation as a method of origin of new species. When

we take up these points we at once enter a field of speculation

in which it may be seen there is opportunity for unlimited

argument, and in which with the bias to which most of us are

subject as a result of our training and investigations, it is diffi-

cult to maintain a purely judicial attitude. It will be profitable

to recall some of the more important facts bearing upon these

matters however.

First, as to the occurrence of discontinuous variations in plants

the following examples cited by Korschinsky will be illustrative :

Erythrina crista galli was introduced into cultivation in 1771 and

` no aberrant forms were seen until seventy-three years later: Bego-

nia semperflorens showed deviating forms only after fifty years :

Cyclamen persicum gave no unusual forms until after one hun-

dred and twenty years of observation: no mutations were

Observed in /pomoea purpurea in one hundred and twenty years.

De Vries observed many thousands of individuals of a hundred

species growing in the vicinity of Amsterdam in 1886 and 1887

and found mutations in only one, that one CEnothera lamarck-

tana. He points out that remains of plants of various species

found in mummy cases four thousand years old have been found

identical with living species in all recognizable characters. As

a result of a rough examination he also concludes that the

744 THE AMERICAN NATURALIST. [Vor. XXXVII.

elementary characters of any species of a higher plant may be

reckoned at a few thousand — about 6000 in CEnothera. If

Lord Kelvin's estimate of the period during which life has

existed on the earth is accepted it might be concluded that in a

general way the average interval separating mutable periods of

any plant must be several thousand years, although nothing

in the nature of the question may be taken to indicate anything

like uniformity in.the matter. Some writers have put forward

the conclusion that at least ten times the above named period,

or twenty-five hundred million years, would be necessary for the

derivation of the existing forms of plants and animals by natural

selection. It must be admitted that both ideas are valuable

chiefly as attractive examples of imaginative grasp rather than

as affording any real evidence in the matter.

It will be recalled that the various theories which have been

put forward to account for the origin of species have been held

by their authors and advocates to be mutually exclusive, and it

seems to have been, and is still taken for granted by the major-

ity of writers, that all organic forms, both plants and animals,

have arisen in the main by one simple method of biological pro-

cedure. The development of biological science has certainly

reached a stage where this a priori generalization may well be

abandoned. I can not say that a candid review of the mechan-

ism of protoplasm, or of the pertinent evidence, from any point

of view compels adherence to this ancient assumption.

The great amount of critical study that is being directed to

the study of hybrids and hybridization is widening the horizon of

this subject momentarily, and the result of our recently acquired

information leads us to conclude that species may originate by

crossing. In such instances the new types are due either to new

combinations of unit characters or to reversionary qualities, it

being necessary to keep in mind the fact that by such union of

two types no new characters are brought into existence. It must

be regarded as unsafe moreover to declare any plant a hybrid of

any other given forms unless the process of origin has been care-

fully followed. The fact remains that hybridization is a demon-

strated source of origin of species however, and it is becoming

more and more generally recognized that more than one method.

—

No. 443.] MUTATION IN PLANTS. 745

of procedure may have been followed in the development of the |

prevalent types of vegetal organism. This view of the subject |

has been thoroughly discussed by von Wettstein and need not

occupy our attention further at the present time (Bericht, deut.

Bot. Gesell., Bd. 13, p. 303, 1895).

DeVries concedes that species might originate by more than

one method, but he holds that natural selection may account for

neither the origin, nor the preservation and continuance of spe-

cies. He furthermore calls attention to the fact that Darwin

repeatedly asserted that characteristics or qualities were formed

very slowly but might disappear suddenly, or in other words that

retrogressive and digressive species formation might ensue by

discontinuous variation or mutation. (Mutationstheorie. Bd. 2:

661. 1903.)

It is necessary to point out that the use of the term natural

selection as applying in any sense to the origin of species by mu-

tation is wrong in view of the special meaning long attached to

that phrase. Natural selection implies constant and progressive

variation in one or many directions, the individuals distinguished

by the greatest improvements constituting the fittest and surviv-

ing from successive generations. The constant and repeated

survival of the fittest and most improved effecting in time such

an amount of departure from the original as to constitute.a new

type. The mutants which arise in discontinuous variability are

seen to depart in all directions from the original, but none of

these may be fitter than the parent type and may perish. It is

probable that many thousands of mutants come into existence

for every one that is capable of existence in competition with the

parent type. The repeated failure of the successive series of

mutants can in nowise affect the character of the later crops

of discontinuous derivatives, and hence the failure of the non-fit

and the endurance of the improved form are not dependent upon

natural selection. Every mutant that survives must not only be

suitable for its environment but must be of a structure and habit

that will enable it to compete successfully with existing types,

in comparison with which it is enormously weaker in numerical

strength. It must therefore gain a foothold at once, with but

little opportunity for adaptations of any kind. Every mutant is

746 THE AMERICAN NATURALIST. [Vor. XXXVII.

a possible species and the only selection which might be said to

act is that which determines the type able to live: this selec-

tion has nothing to do with the origin of the surviving form

however.

Thus of the sixteen mutants discovered by DeVries one had

already established itself when found, although seen to arise anew

from the parent type subsequently. Perhaps one or two of the

others might have succeeded in gaining a foothold, but the major-

ity of the new forms must have inevitably perished if subjected

to the ordinary competition of the prevailing meadow species.

As to the cause of mutation, and the mechanism of the proc-

ess but little except of a speculative nature may be offered.

Korschinsky assumes that heredity and variability are opposing

forces or tendencies which are ordinarily balanced. External

agencies such as successive seasons of good nutrition might

allow the tendency to variation to overcome the hereditary

stability and allow the origination of a new form as a result

of the unloosed, superfluous unbalanced energy. He supposes

that whatever the agencies may be that cooperate to bring

about the mutative condition, these forces act upon the develop-

ing embryo in the seed, although he hazards no guess at the

manner in which this might be accomplished (Zora, 89: 240,

1901). The above it may be noted is in direct contrast with

the proposal of Darwin that the development of new types is

more rapid when species are competing under adverse condi-

tions, or when the struggle for existence is fiercest.

So far as DeVries's theory of mutation is concerned it may be

said to be the logical outcome of, and to rest upon his hypothesis

of intracellular pangenesis. By this, protoplasm is taken to con-

sist of ideally minute pangens, which make up the living sub-

stance. The pangens and aggregations of pangens are the

bearers of the elementary characters of the species. Altera-

tions in the numerical relations of pangens are made to account

for fluctuating variability. The inactivity of pangens and groups

of these units would cause degressive or retrogressive mutation.

The formation of new characters in progressive mutations would

depend upon the development of new pangens, this process con-

stituting premutation. The formation of identical pangens in

separate species would account for parallel mutations.

No. 443.] MUTATION IN PLANTS. 747

But little definite evidence is at hand as to the time at which

the changes antecedent to mutation, constituting pre-mutation

occur, although certain stages of development may be designated,

previously to which they must come about. Mutations of the

higher plants are first apparent in the seedling but the actual

alterations or departure from the hereditary behavior must have

taken place at least as far back as the formation of the sexual

elements the union of which produced the embryo, and may

have occurred evén earlier. In any case the mutants are per-

fectly formed in the embryo and influence of any kind upon the

germinating seed may not alter their nature (see page 746). It

may be seen from the foregoing that the mutative processes may

be connected with either the vegetative body or the sexual ele-

ments, and may be found within the sporophyte, or be confined

to the gametophyte.

If the pre-mutative alterations occur in the vegetative proto-

plasts of a self-fertilized individual both gametes would presuma-

bly carry the same characters to the union. If, on the other

hand, premutation occur in one of the sexual elements, or if it

occur in the vegetative cells of species which are cross fertilized

only, the embryos formed would be the result of the union of one

mutant gamete and one of the regular inherited form. Inasense

such mutants might be considered as hybrids. This theoretical

aspect of the question seems to find a reflection in the behavior

of CE. lata, one of the mutants with pistillate flowers only. When

pollinated by the parent form, CE. damarckiana, it produces CE.

fata and GE. lamarckiana.

DeVries conjectures that the causes inducing mutation are

partly internal, and partly external to the organism. The state

of external factors necessary to the process probably occur only

at uncertain intervals, and is supposed to embrace a combination

of extremely favorable and unfavorable conditions. i

Probably no more profitable subject for research in the whole

realm of natural history offers itself to the investigator than the

problem of the causes which produce new species. The above

supposition deserves early attention from the experimentalist

since it is one that is comparatively easily capable of proof and

disproof.

748

THE AMERICAN

NATUR?

ILIST.

[Vor. XXXVII.

nothera

Lamarckiana

e Figs. 2 an

3-)

1G Ga ettes of seedlings two months, and five months of age-

‘Sic aborted from heiter ea. (Se

0. 443.] MUTATION IN PLANTES. 749

1G. 2. CEnothera rubrinervis. and five months of age. Photo-

graphed from herbarium sheets.

Seedlings two months,

(See Figs. 1 and 3.)

750 THE AMERICAN NATURALIST. [Vor. XXXVII.

Being desirous of testing the general facts of mutation as

illustrated by the behavior of the cenotheras under environ-

mental conditions different from those at Amsterdam, seeds of

CE. lamarckiana, Œ. rubrinervis, Œ. lata, Œ. nanella, Œ.

brevistylis, and CE. gigas were procured from Professor DeVries

and these were placed in soil in the propagating houses of the

New York Botanical Garden May 15th, 1902. Germination fol-

lowed in a few days, and a number of individuals ranging from

fifteen to forty of every species were pricked out and suitably

repotted from time to time. The cultures were examined three

to seven times per week except during February, 1902, and July,

1903. The amount of work necessary to make minute and exact

observations on all of the above forms being too great a demand

upon my time, chief attention was devoted to a comparison of

the parent type with rxdrinervis and nanella, two mutant forms.

In order to systematize the results general notes were made

continuously upon the habits of the growing plants and formal

comparisons were made at successive stages as follows :

First stage — July 11th, 1902. The plantlets were nearly

two months old and still retained the cotyledons.

Second stage — October 15th, 1902. A distinct tap root

had been formed and a rosette of leaves had been developed.

Third stage.— June 1st— roth, 1903. Adult rosettes had

been formed, and the smaller leaves which appear around the

base of the stems were apparent. Some flowering stems were

beginning to push up. :

Fourth stage.— August 1oth-15th, 1903. A number of

inflorescences had been produced and flowers were opening daily

in great profusion on some of the forms. Some of the inflores-

cences were enclosed in paper bags in order to secure pure seeds

by means of artificial transfer of pollen.

The more apparent anatomical differences among the forms

examined are shown quite strikingly by the series of photo-

graphs and drawings which illustrate this article.

The main fact to be kept in mind in regard to the parent

form is that it is a recognized and constant species, which has

not undergone noticeable alteration during the long period it

has been under exact observation. The seeds from which the

No. 443.] MUTATION IN PLANTS.

ysettes

of seedlings two months,

Fic. -(Enothera nanella. Re

AE from herbarium sheets. . Figs. 1 and 2

-I

and five months of age.

Lon]

752 THE AMERICAN NATURALIST. [VOL. XXXVII.

experimental material was derived were sown in a bed at s' Grave-

land near Amsterdam in 1875 and bad been allowed to spread

over an adjoining neglected field until in 1884 an area of 2800

square meters was covered. This material showed the presence

of a form so different from the parent type, when examined by

deVries in 1886, as to lead him to consider it as a new species,

and this mutant, QE. drevistylis, which did not arise again during

the observations, maintained itself in the same locality during a

period of twelve years, records of it having been made as late as

1898, and it is still cultivated among the other mutants grown

by deVries and myself. Other forms appeared during the

course of the next fourteen years as has been described in detail.

It was deemed advisable to make independent comparisons of

the plants grown in my own cultures with the type specimen

with which deVries identified his parent form, and to this end

Miss A. M. Vail made a visit to the herbarium of the Muséum

d' Histoire Naturelle in Paris, in May, 1903, at my request, and

also latera journey to Amsterdam and inspected the cultures

of Génothera under Professor deVriess own guidance. Miss

Vail has kindly prepared the following report on the matter :

“The parent form, Guothera lamarckiana Ser. was found by

deVries to agree in every particular with two specimens in the

Muséum d'Histoire Naturelle in Paris. These specimens con-

sist of, first: a plant cultivated in the Paris Garden that had

formed the basis of the original description of Ginothera grand-

iflera Lam. It bears a label indicating it as having been

included in the herbarium of Lamarck which was acquired by

the Museum in 1850. On the margin of the sheet in the hand-

writing of Poiret (the author of the section dealing with Gzze-

thera in Lamarck's Encyclopedia) is the following inscription

* CEnothera —(grandiflora)— nova spec. flores magni lutei, odore

grato, caulis 3 pedalis. This specimen is in flower only and

consists merely of the branched upper portion of the shoot with

numerous rather small leaves and conspicuously large typical

flowers. The second specimen comes from the collection of

Abbé Pourret that was contained in the collections of Dr.

1 For a brief general account of the experimental cultures, see MacDougal, The

Original of Species by Mutation. Torreya, Vol. 2, pp. 65-68, 81-84, 97-100, 1902.

No, 443.] MUTATION IN PLANTS. 753

Barbier inherited by the Museum in 1847. It is filed in a cover

with CE. biennis L., and bears that name on the sheet, a small

label inscribed with a series of prelinnean names, and another

with ‘ Onagra vulgaris Spach’ and ‘ GZuothera biennis Linné,

both apparently in Spach’s handwriting. This is the plant

referred to by de Vries as having been collected presumably by

Abbé Pourret in the Paris Garden during his visit in 1788.

The specimen represents an unbranched upper portion of a

shoot with numerous large well-developed leaves, partly mature

capsules and several flowers that are somewhat smaller than

those of the previously mentioned specimen. These two speci-

mens differ in no important particular. Tracings of them com-

pared with living plants grown in the New York Botanical

Garden from seeds sent by de Vries agree quite perfectly.

“A search through the herbarium of the Muséum d'Histoire

Naturelle and that of the New York Botanical Garden does not

bring to light any specimen of a wild North American plant that

can be referred to CE. /amarc&iana as it is now known and cul-

. tivated in Europe, nor does it seem to be known to collectors in

North America at the present day.

* Several specimens were found however, which might be con-

jectured as representing a North American plant from which

CE. lamarckiana might have been derived. One of them is a

plant collected by Michaux now preserved in the Muséum at

Paris, and cited by de Vries in the Mutationsteorie (Bd. 1:

316) and referred by him to a plant frequently cultivated in

Europe under the name of Enothera grandiflora Ait CE. suave-

olens Desf. but which he considers different from CE. /amarck-

tana. A tracing was also made of this plant which consists of

two specimens fastened on the same sheet upon which numerous

inscriptions bear witness to much diversity of opinion as to its

real identity. A small slip of paper bears in Michaux's hand-

writing * Znothera grandiflora, another (the customary label

of the Michauxian specimens) the inscription *CEnothera grandi-

flora Poiret Encycl, in the writing of that author of the

section dealing with Cxothera in Lamarck's Encyclopedia ;

beneath that * G2nothera suaveolens Hort. par.’ in the writing

of Desfontaines, and lastly * Onagra vulgaris grandiflora Spach.’

754 THE AMERICAN NATURALIST. [Vor. XXXVII.

in the writing of Spach. The larger of the two specimens con-

sists of a simple entire plant not fully developed, showing root,

leaves, flowers, and capsules, but no basal leaves. The other

specimen, which is smaller, is incomplete and fragmentary. A

comparison of the tracing of the larger specimen with material

in the herbarium of the New York Botanical Garden shows that

it is identical with a specimen under the name of Onagra biennis

grandiflora (Ait) Lindl, collected by E. S. and Mrs. Steel on

Stony Man Mountain, Luray, Virginia, August 15th, r9or.

The comparison also shows that the wild plant has undergone

no change of any kind during a period of over a century.

“The following memoranda and citations may be of interest

as throwing some light on the history of CE. lamarckiana previ-

ous to 1788. :

Linnzeus in his Species Plantarum says that Eunothera biennis

was brought from Virginia in 1614 and was then (1753) common

in Europe. In Hortus Cliffortianus (1737) he states on p. 144,

that it is a native of Virginia, having been brought from there

to Europe 120 years before and was at the time he wrote spon-

taneous and plentiful in the fields of Holland. In Hortus

Upsaliensis (p. 94. 1748) he gives the date of its introduction

as 1620, then declared it to be spontaneous in Belgium, Italy,

‘Gallia and Germania.’ So that from the middle of the 17th

century it was generally in cultivation in the botanical and horti-

` cultural establishments of Europe.

Referring to some of the prelinnean writers we find that

Tournefort in Just. ret. herb., on p. 302 (1700) enumerates nine

species of Onagra, the first four of which only are of interest

here, as follows :

(1.) Onagra latifolia. Lysimachia lutea, corniculata. C. B.

Pw. 245.

(2. Onagra latifolia, flore dilutiore. Lysimachia corniculata

non papposa, Virginiana, major, flore sulphureo. H. L. Bat.

(3. Onagra latifolia, floribus amplis. L ysimachia Virgintana,

altera, foliis latioribus, floribus luteis, majoribus. Cat. Alt-

dorf.

Onagra angustifolia. Lysimachia angustifolia, Canadensis,

corniculata H. R. Par. Lysimachia corniculata, lutea, Cana-

densis minor, seu angustifolia Mor. H. R. Bles.

(4.

No. 443] MUTATION IN PLANTS. 755

In the first of these references Caspar Bauhin in Pinax on p.

245 (1671) writes of an American evening primrose under the

name of Lysimachia lintea corniculata, as being a Virginian Lysi-

machia growing in the Garden at Padua in 1619 and adds that

it was a pleasing plant and easy to propagate from seed. The

second reference goes back to Hermann's Catalogus, 1687, where

on p. 396 he records a species of Virginian Lysimachia with

sulphur colored flowers as growing in the Garden at Leyden.

The third reference is to a plant with larger leaves and larger

flowers from the Altdorf Garden. In Jungermann's Catalogus

plantarum quae in horto Medico Altdorphino reperiuntur we read

that a Lysimachia lutea Fl. majoribus, odore Tabaci. and a (Lysi-

machia) Virginiana lutea Delphinium quorundum, were known in

the old Bavarian garden at Altdorf in 1635 and the statement is

again repeated in another Caza/egus in 1640. It was a suffi-

ciently remarkable plant for Tournefort to note especially in his

Institutiones, and it might be inferred that this large flowered

plant from Altdorf was the ancestor of Œnothera lamarckiana.

It would appear as if a form of what is generally claimed to be

CEnothera biennis L. with delicate sulphureous flowers grew in

the Leyden Garden and another with larger flowers in the gar-

den at Altdorf. Under the same name, Lysimachia corniculata,

an American evening primrose is said to have been growing in

the Messina Garden in 1640 and it was known in the Paris Gar-

den at about the same time or a little earlier and in 1653 in the

Copenhagen Garden. Morison also records it as occurring in

the Hortus Blesensis in 1669. This last reference is the one

quoted by Tournefort as his fourth species. Again under the

same name of ZL. corniculata Sherard speaks of it on p. 44 of his

Schola Botanica as growing in the Paris Garden in 1689 and,

presumably, descendents of the plants he saw were those col-

lected by Abbé Pourret a century or so after and later made the

type of the much discussed Œ. grandiflora Lem. = GE. lamarck-

tana Sen. The plant described by Linnzus in the Species

Plantarum was doubtless a composite species and it would be

particularly interesting in this connection to know just what he

meant by the plant described in the Hortus Cliffortianus as

being plentiful in the fields of Holland. A tracing of the speci-

756 THE AMERICAN NATURALIST. [Vor. XXXVII.

men which could be considered as the type of the plant described

by him in the Hortus Cliffortianus has been kindly furnished by

Dr. A. B. Rendle of the British Museum, and although the flow-

ers are somewhat smaller than those of the living plants of Œ.

lamarckiana as grown in the New York Botanical Garden nurs-

eries, yet the general characters are identical, notably that of

the entire or slightly emarginate petals. This character is cer-

tainly not typical of the wild weed-like CE. biennis of waste lands

£. s: 1

Fic. 4. h kii

ering stem. Photograph of living plant taken from directly above. (See Figs, 5 and 6.)

in North America to-day. In any case it seems extremely doubt-

ful that all these cultivated evening primroses should be referred

to so ungainly and unornamental a plant as CE. biennis.

Prof. deVries in an article on the introduction of CE. damarck-

tana in Holland (Ned. Kruidk. Arch. ser. 2, Vol. 6, p. 579,

1895) gives a long and detailed history of the ancestors of the

plants taken into cultivation for his experiments. T hey were

traced to plants escaped from cultivation and originally raised

from seed received from a seedsman of Erfurt, Germany. Prof.

No. 443.] MUTATION IN PLANTS. 757

deVries also states that QE. biennis and CE. muricata are found

in Holland, notably on the dunes,

It seems well established that a large flowered CEnothera was

seen in the Altdorf Garden in 1635, which is probably referable

to none other than /amarckiana. Later notes of its occurrence

are in existence, but the first definite record of the species was

FiG. 5.—GEnothera rubrinerv us. Adult

sette immediately sie the fume of the flowering stem.

Figs nd 6

ros

Photograph of iiving plant taken from directly above. (See I

in 1788. It has been found constant since this date, both in

; : "ld: its evolutionary procedure is

gardens and when running wild: its evolut 21 i

therefore none the less valuable as scientific evidence than as i

it were an indigenous wild growing species.

as present

CEnothera lamarckiana is a species which, so far as prese

758 THE AMERICAN NATURALIST. [Vor. XXXVII.

knowledge is concerned, has no exact duplicate in the native

flora of any region, and two probabilities are suggested as to its

origin: It may have been a native of a restricted range in “ Vir-

ginia" in which it has been exterminated by agricultural opera-

tions, and hence cannot be found at the present time: or it may

have arisen by some such sudden, and abrupt, discontinous vari-

ation, as that by which deVries’ mutants came into existence,

from CE. diennis in the gardens, at Padua, Altdorf or else-

where: at least no intermediate forms are known."

Only eleven specimens of hybrid seedlings derived from

CEnothera lata were brought to the adult stage, in my cultures

and of these but two conformed to the type of Œ. /a/a, the

remainder being the O. damarckiana form. (ŒE. lata does not

perfect its stamens but it is capable of being pollinated from the

parent. The offspring followed the laws governing parent and

mutant hybrids, which with deVries were found to consist of

18% to 20% of the mutant type and the remainder of the par-

ent. My own results agree with this. It is clear that this form

would not have survived beyond the season of its appearance as

it does not display any marked propagative capacity.

— CEnothera nanella originated in deVries's cultures in 1888

and has since been followed by him through fifteen seasons.

The qualities of this form separate it from the parent in such

manner that it might be considered as a variety by some sys-

tematists, although its behavior and physiological properties are

constant and very clearly distinguishable. In following out

the development of the plant during the eighteen months over

which my own observations extended it became evident that it

differs most widely from the parent in its earlier, and also in its

adult stages, being most like it in the full rosette stage. The

most apparent feature is its diminutive size, both in the young

plant and in the mature flowering shoot. The stem shows but

little capacity for branching and did not reach a height of more

than 20 to 25 cm. in my cultures, or about one fourth that of the

parent, which sends out numerous vigorous branches. The first

few leaves have very broad laminz with irregular apical portions,

.and are short petioled. Later leaves are more nearly like the

parent type but remain shorter petioled which has the effect of

No. 443.] MUTATION IN PLANTS. 759

making a denser more crowded rosette. The bases of the lam-

ina are almost cordate in some instances, and vary from oblong

ovate to ovate in outline, being sparingly toothed. The plants

established in the soil in the open air did not bloom until about

three weeks later than the parent and CE. rubrinervis. No

noticeable departure from the characteristics assigned this form

by deVries was found.

Seedlings of Enothera rubrinervis were seen to have nar-

rower leaves throughout from the earliest stages. The rosettes

were very closely appressed to the soil, and in this stage the

margins of the long petiolate leaves were inrolled, thus decreas-

Adult rosette immediately preceding the formation of flowering

FriG. 6.—GEnothera nanetla.

5 ‘igs. 4 and 5.)

tem. Photograph of living plant taken from directly above. (See

ing their apparent width. Attention is to be called here to the

fact that comparisons of leaf forms in plants of this kind

are permissible only between organs on corresponding por-

tions of shoots. The laminae were more bluntly toothed than

those of the parent type, and the midribs occasionally bore a

tinge of red, while the entire shoot including the leaves of the

upper part of the stem showed a tendency to the formation of

anthocyan. The physical qualities of the leaf were strikingly

different from those of the parent, perhaps the most notice able

feature being the great brittleness of the leaves and stems of

760 THE AMERICAN NATURALIST. (VoL. XXXVIEL

young plants, indicative of high turgidity and weak development

of mechanical and supporting tissues. Both of these characters

have been observed by deVries, who notes that the bundles of

Fic. 7.—CGEnothera lamarckian Adult pl : er beginning of openi Ei

ers. Photograph of living plant grown in the soil in the open air, and temporarily fixed

ina pot. (See Fig. 8.)

bast fibers of the flowering stems were composed of elements-

with thinner walls than those of the parent type.

The leaves of the full rosettes, were silvery white owing to

Fig. 8.

—( En.

.p!

&rown in the soil in the open air, and temporarily vies in a pot. (See Fig. 7.

No. 443.] MUTATION IN PLANTS. 761

the fact that the hairs on both surfaces were both longer and

more numerous than on the parent type. The average length

of the hairs on the upper surfaces was 35 as compared to 28

in the parent type, and cn the lower surfaces 42 as compared to

30. Theaverage number of stomata on a unit of area of the

upper surface of the leaves of rubrinervis was 37 as compared

othera rubrinervis. Adult plant two weeks after beginning of opening of flowers.

with 34 in Enotera lamarckiana. The brittleness character-

istic of the tissues of rubrinervis may be seen to extend even to

the hairs, since these structures are easily detachable from the

dried specimens, and hence giving rise to the conclusion that

rubrinervis is less densely pubescent than lamarckiana as given

in the systematic description below.

Photograph of living plant

762 THE AMERICAN NATURALIST. [Vor. XXXVII.

Still another major difference between the forms in general

habit is that of the method of branching and the growth of the

branches. In Œ. lamarckiana, the branches from the basal por-

tion of the shoot were of a length amounting to more than half

that of the shoot which is also true of CE. rubrinervis. The

upper branches of the former remain short and stout however,

while those of rubrinervis attain greater lengths which decrease

upwardly so that a plant may have a roughly globular outline.

The majority of the features in which the mutant departs

from the parent, as described above, are of a nature that would

equip the new form for living under more arid conditions than

the parent, although the actual endurance of rubrinervis to

decreased supply of. moisture was not tested. So far as this

single observation goes then, it is to be seen that the new char-

acters of mutants are harmonious in their adaptive relations.

CEnothera rubrinervis originated in deVries’ cultures in 1899,

and has also appeared by independent mutations since that time.

It has been found to be independent and self-maintenant in

competition with the parent form.

A large number of flower buds in both ruġrinervis and

lamarckiana were pierced by some insect, and the larvze coming

from the eggs deposited made great destruction, and also caused

the abnormal enlargement of the buds and capsules, which failed

to perfect seeds. :

De Vries has continued to find the recurrence of some of the

mutants in the successive crops of seedlings of Cnothera

lamarckiana indicative of the fact that the mutating period of

the parent has not yet been passed. No departures from the

parent type were found among the individuals which have come

into bloom up to this time in the New York Botanical Garden.

The leaves of the seedlings of CE. lamarckiana are easily dis-

tinguishable from those of /aza, nanella, and rubrinervis even in

the earlier stages, although not so easily separable from some of

the other forms such as drevistylis and leptocarpa according to

deVries. The earliest leaves were ovate, or round-ovate with

rounded apices, or sometimes slightly pointed. These leaves as

well as those formed at the age of five months were distinctly

petiolate but with the laminz relatively narrower. Adult basal

No. 443.] MUTATION IN PLANTS. 763

leaves of the rosette in the period immediately preceding flower-

ing were petiolate with the apices bluntly pointed and with broad

lamine. The margins of all of the earlier leaves were sparingly

but sharply toothed.

Plants set out early in May were blooming profusely early in

August. The basal branches coming out from the axils in or

near the rosettes were strong and vigorous but the upper branches

of the stem were short and offered a distinct contrast to the

longer, more slender branches of rubrinervis, with which it was

also contrasted by its denser foliage and larger more showy

flowers. Both stems and branches were thicker and heavier

than in rubrinervis.

After noting the great variance in behavior and appearance of

the parent and two mutants as described above, mature plants in

bloom, the dried material of the younger plants, and photographs

were submitted to Dr. J. K. Small, who had previously published

an arrangement of the American species, and who is familiar

with them in (Small, J. K. CEnothera and its Segregates. Bull.

Torr. Bot. Club. 23: 167—194, 1896.) the herbarium and in

the field. Dr. Small has kindly prepared the following statement

concerning three forms, which is given in full below :

The characteristics of GZuothera. lamarckiana and CE. rubri-

nervis as given by Dr. Small are set in parallel columns for con-

venience of comparison :

764 THE AMERICAN NATURALIST. [Vor. XXXVII.

Oenothera lamarckiana Ser.

I. Seedling about two months

old.— Leaves sparingly pubescent ;

blades ovate to suborbicular, the

larger about 2 cm. wide, obtuse or

rounded at the apex, each abruptly

narrowed into a petiole.

II. Seedlings 5 months old. —

Rosettes relatively dense: leaves

copiously fine-pubescent; blades

typically oblong, the larger ones

fully 3 cm. wide, quite approxi-

mately denticulate, obtuse, or some-

what apiculate at the apex, much

longer than the petioles.

III. Adult flant. — Plant very

stout and luxuriant, 0.5 to I m.

tall. Stem markedly channeled,

sparingly hirsute with rather

spreading hairs, nearly simple, or

with several relatively short as-

cending branches near the base,

and few very short ones above:

leaves very numerous, 2-2.5 dm.

long about the base of the stem ;

blades shallowly and often irregu-

larly toothed, those of the lower

cauline leaves broadly spatulate to

petiole, those of the upper cau-

line leaves oblong to oblong lanceo-

late, acute, or somewhat acuminate,

short-petioled: bracts subcordate

at the base: hypanthium 4.5-5.5

cm. long, about 8 mm. wide at the

mouth, prominently ridged : sepals

4-5 cm. long, longer than the tu-

bular portion of the hypanthium,

the free tips 8-10 mm. long: pet-

als firm 4—5 cm. long, emarginate :

anthers 13-15 mm. long: stigmas

5-6.5 mm. long. (See Figs. 1, 4,

7 and 9.)

Oenothera rubrinervis deVries.

I. Seedlings about 2 months old.

tana; blades elliptic, the larger

ones about 1.5 cm. wide, acute or

acutish at the apex, each gradually

narrowed into a petiole.

II. Seedlings 5 months old. —

Rosettes lax: leaves less densely

pubescent than in Oe. Lamarck-

tana ; blades spatulate to elliptic-

spatulate or oblong-spatulate, the

larger ones about 2.5. cm. wide,

remotely denticulate, acute, or ab-

ruptly pointed at the apex, about

as long as the petioles or shorter.

III. Adult plant.— Plant rel-

atively stout, less luxuriant than

Oe. Lamarckiana. Stem scarcely

channeled, hirsute, with rather as-

cending hairs, typically branched

throughout, the branches near the

base elongated, decumbent, the up-

per ones gradually shorter: leaves

numerous ; blades less prominently

toothed than in Oe. Lamarckiana,

those of the lower cauline leaves

spatulate to broadly oblong, obtuse

or acutish, each narrowed into a

relatively long petiole, those of

the upper cauline leaves elliptic-

oblong to oblong or oblong-lanceo-

late, acuminate, short-petioled :

bracts rounded or round-truncate

at the base: hypanthium 5.5 to 6

mouth, obscurely ridged: sepals

3 to 3.5 cm. long, shorter than

the tubular portion of the hypan-

thium, the free tips 5-6 mm. long:

petals tender, 3-3.5 cm. long,

notched: anthers 6-10 mm. long.

stigmas 7.5-10 mm. long. (See

Figs. 2, 5, 8 and 10.)

No. 443.] MUTATION IN PLANTS. 765

CEnothera nanella was taken by deVries to have a degree of

separation from the parent type that would lead it to be consid-

ered as a variety, a conclusion which is borne out by Dr. Small’s

description as given below:

I. Seedling about two months old. — Resembles that of Eno-

thera lamarckiana ; but the leaf-blades are less uniform, some of

them ovate or oval, others ovate and somewhat lobed near the

apex, others broadly ovate, or prominently apiculate.

II. Seedling 5 months old— Nearly like that of Œ. /amarc£-

zana; but leaves inclined to have longer petioles. —

III. Adult plant— Plant, stout and stocky in all parts, resem-

bling CE. famarckiana, but smaller, less than 3 dm. tall.

Stem obscurely channeled, hirsute with somewhat ascending

hairs, simple: leaves approximate, 7—12, 5 cm. long near the

„base of the stem; blades shallowly, often rather remotely, but

quite evenly toothed, those of the lower cauline leaves spatulate

to oblong, acute, or acutish, each narrowed into a semi-terete

petiole, those of the upper cauline leaves broadly oblong to

oblong-ovate, acute or slightly acuminate, nearly sessile: bracts

subcordate at the base: hypanthium 3-3.5 cm. long, about 5

mm. wide at the mouth, obscurely ridged: sepals 3-3.5 cm.

long, longer than the tubular portion of the hypanthium, the

free tips 5-6 mm. long: petals 3.5-4 cm. long, emarginate :

anthers 11-12 mm. long: stigmas 4-5 mm. long.

GENERAL SUMMARY.

Discontinuous variation as a possible method of origin of

species was considered by Charles Darwin in his studies of

plants and animals under domestication, and he concluded that

if new forms did arise in this way that they were not self-main-

tenant (1868). On the other hand Galton took the position that

the evolution of species is not necessarily by minute steps (1889),

but Dollo (according to deVries's, Mutationstheorie, Bd. 1, p. 46,

1901) was the first to accept discontinuous variation as the prev-

alent method of origin of species (1893). Bateson (1894)

brought together a large amount of evidence as to types which

have arisen in this manner, and a comprehensive summary of

Fic. 9. oe lamarckiana. A, leaf f

middle,

rom basal zra of adult rosette; B, leaf fro

from upper portion of rosette; D, leaf from middle of ain

stem; E i i dd lower part of inflorescence; F, jove with petals removed; G

petal. (See rie 10.)

Fic. 1 Fort pcnc rubrinervis. A, leaf from lower part of — E ^ ^- —

iddle portion, and C, leaf from upper portion of rosette ;

ering stem; Æ, bract from lower part of inflorescence; F, "ica with ipu removed;

G, petal. (See Fig. 9 7

768 THE AMERICAN NATURALIST. [Vor. XXXVII.

the principal evidence furnished by plants was made by Kors-

chinsky in 1899. Systematic observations upon the subject were

begun by deVries in 1886 and have been continued until the

present time. As a result of his investigations deVries formu-

lated his ** Mutationstheorie," which has appeared in book form,

the separate parts of which have been published in the period of

1901-1903. This hypothesis rests upon the theory of pangenesis

previously formulated by him.

The parent type, O. lamarckiana, from which deVries saw

mutant forms arise has been found constant in its characters in

cultivation in Europe and America and also when running wild.

This type is not identical with any known member of the Ameri-

can flora, and is most nearly allied to Oxagra biennis grandiflora

(Oenothera biennis grandiflora) from which it is suggested it

might have arisen by mutation.

The mutant derivatives of the parent form are found to be

constant in their characters, with no connecting or intergrading

forms, as illustrated by the cultures of the parent, CE. nanella

and C£. rubrinervis, in the New York Botanical Garden during

1902-1903. The mutants are clearly separable from the parent

and from each other both by physiological and taxonomic stand-

ards. Furthermore the specific character of the mutants was

borne out by their behavior when hybridized with one another.

It has become evident from the results so far accomplished

that the testing, study, proof or disproof of the theory of the

origin of species by mutation involves an actual examination of

lines of descent, and observations upon successive generations

of organisms of known genesis. In this manner only may

mutant forms be distinguished from hybrids, individuals with

aberrant non-transmissible characters and teratological formations.

The nature of the questions involved, and. the essentially mate-

rial character of the evidence to be considered is such that all

controversial discussions not supported by facts of this character

must be viewed with distrust. In no instance is this more

plainly apparent than in the recent treatment of the subject by

Vernon (Vernon, H. M. Variation in Animals and Plants. 1903).

This author says “ Hence it (Gnothera lamarckiana) is probably

a garden variety of (Enothera biennis (Evening Primrose), and

No. 443.] |. MUTATION IN PLANTS. 6c

799

may be a hybrid plant, whilst the mutations obtained by deVries

may be merely partial or. complete reversions to the original

ancestors of the plant." It is quite possible, and even probable

that CE. damarckiand may have been originally derived from the

same type as CE. biennis as noted above, but to designate it as

a “garden variety," and as such ineligible as: research material

is simple evasion. The plant in question has been under more

or less continuous observation for a hundred and fifteen years

during which period it has been constant in its characters, and

has shown no evidence by anatomical similarity or physiological

behavior of being anything but an independent species. With

what species could ézennis hybridize to produce /amarckiana?

The genus comprises a comparatively small number of types, all

natives of America, and none of which were available as a

hybrid mate to dzennis at the time of the origin of lamarckiana.

The conjecture in question is totally unsupported after the most

rigid search for evidence upon the matter.

Again to consider the mutants as reversions to the original

ancestors of damarckiana is impossible, since the mutant forms

exhibit qualities not possessed by any other known members of

the genus, including dennis.

The point raised by Bateson and Saunders (Reports to the

Evolution Committee. Royal Society. I. p. 153, London,

1902) that the pollen of /amarckiana contains deformed grains,

Which points to its origin by crossing, is without significance,

since the author has found that the stamens of plants of dzennzs

growing in the vicinity of. New York exhibit a much larger pro;

portion of deformed pollen than that of the specimens of /amar-

ckiana cultivated in the New York Botanical Garden.

It has been impossible so far to assign mutations to definite

causes, or to forecast the frequency, or occurrence of the phe-

nomenon. These phases of the subject constitute the most

important problems of the subject, which await investigation.

Theoretical evidence upon such a subject can have but limited

value, and conclusions of any satisfactory degree of finality md

be expected only from direct experimental research under cir-

cumstances in which the probability of error is reduced to a

minimum.

779 THE AMERICAN NATURALIST. [Vor. XXXVII.

So far as the origin of mutations is concerned, it seems well

decided that the premutative alterations in seed-plants ensue in

the vegetative and sexual cells previously to the formation of the

embryo in which they first appear, and that no environmental

disturbances may bring about the alterations in question by

direct action on the seedling.

It is not the purpose of this paper to discuss the various theo-

ries which have been put forward from time to time to account

for the origin of species, but to bring under consideration the

facts upon which the conclusions as to the origin of species by

discontinuous variation have been based by deVries. These

facts make inevitable the conclusion that new types of specific

rank, taxonomically separable, and physiologically distinct and

constant, without intergrading and connecting forms, have arisen

in CEnothera by discontinuous variation. That mutation is

the principal method of evolutionary procedure is mot proven.

That natural selection is universally prevalent is certainly dis-

proven: that natural selection or any other method is capable of

accounting for the existence of any single species has not been

proven with the finality offered by the evidence of discontinuous

variation. It may be said, therefore, that species have actually

been demonstrated to have arisen by mutation, some are known

to have arisen as the result of hybridization, and that evidence

has been accumulated which has been interpreted to demonstrate

the origin of species by natural selection, and by adaptation.

Nothing in the nature of living organisms demands that all spe-

cies should have originated in the same manner, or that one

simple, or single method of procedure should have been followed.

New YORK BOTANICAL GARDEN,

August, 1903.

DISTRIBUTION OF THE FRESH- WATER FISHES

OF MEXICO.

SETH EUGENE MEEK.

Mexico consists of a high plateau bordered on each side by a

narrow coastal] plain. It lies between the United States and

Central America, but is not separated from either by natural

boundary lines. The southern half of this country lies in the

torrid zone, the rest in the North Temperate. Its geographical

position, its elevation and diversity of climate make it, from a

biological standpoint, a most interesting country. The Rocky

Mountains extend into the northern United States as a single

range to the Yellowstone Park. Here is given off to the west

the Wasatch range, which extends south into Mexico as the

western range of the Sierra Madre. The Rocky Mountains

proper become in Mexico the eastern range of the Sierra Madre.

These two mountain ranges include a plateau, the elevation of

which varies from three to eight thousand feet above the sea.

This plateau is drained by four river systems :— the Colorado

river on the north and west, the Rio Grande, central and eastern

portion, the Rio Panuca and the Rio Lerma, the southern portion.

The southern end of this plateau is the beautiful valley in which

is built the City of Mexico, while the two mountain ranges cul-

minate in the famous peaks of Ixtaccihuatl and Popocatapetl.

The valley of Mexico though at one time it probably drained

into the Lerma now. comprises a drainage system of its own.

The great central plateau comprises the larger part of Mexico.

On the east and the west is a low narrow plain from which the

ascent to the plateau is steep. South of the valley of Mexico

the mountains extend as one range through Central America to

become the Andes in South America. The Mexican plateau in

general is a treeless plain, covered with only a slight vegetation.

The Yucca, the Mesquite, various species of Cacti, sage

brush, a few stunted cedars and the like, together with a sparse

growth of various species of grasses, comprise the larger part of

the vegetation of this region. During the rainy season anda

771

772 THE AMERICAN NATURALIST. [Vor. XXXVII.

short time after it, there is a luxuriant growth of vegetation: but

after a few months of exposure to the piercing rays of a tropical

sun the character of the country changes, and it assumes the

air of a parched desert. It is subject to a short rainy season

and a long dry one. The rivers which are large in the rainy

season become almost dry by the end of the long dry season.

Many of the lakes in northern Mexico become dry and the

streams which flow into them contain but little water except in

the upper part of their courses where they are fed by mountain

springs, and streams of this character contain but few species of

fish.

The study of any group of plants or animals in a country

like this is very interesting, but no group of living things pre-

sents a more interesting subject for the study of geographical

distribution than the fresh water fishes. Living as they do

in the water their only highways of travel are in the streams

and lakes and so their dispersion is largely governed by the for-

mation of our fresh water lakes and rivers and is therefore

intimately associated with the later chapters of the geological

history of the earth.

The two large rivers which reach Mexico from the north and

which have furnished highways by which Northern Mexico

became stocked with fishes are the Colorado and the Rio

Grande. The former flows into the Gulf of California, the lat-

ter into the Gulf of Mexico. In their upper courses these two

rivers are near each other, but their fishes are not the same.

The only fish common to both river basins is a small dace

(Rhinichthys dulcis) and this is also found in the head waters of

the Arkansas, the Missouri and the Columbia rivers. From the

Colorado river there are known 32 species of fishes which are

distributed in 18 genera and 5 families.! Of these 32 species

all but ro? are thus far known only from this basin. Nine of

! Catostomidze (Suckers) 9 species, Cyprinidz (Minnows) 18 species, Salmonidz

(Trout and White fishes) 2 species, Poecilide (Killifishes) 2 species, Cottide

(Blobs) 1 sp

ne ae pew (Girard), Rhinichthys dulcis (Girard), Agosia chrysogaster

Girard, eus oscula (Girard), Lepidomeda vittata Cope, Coregonus williamsoni

Gi urus (C

irar ope), Poecilia occidentalis (Baird & Girard), Cottus

PEE (Gi.

No. 443.] FRESH-WATER FISHES OF MEXICO.

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Map of Mexico showing fish faunal areas.

774 THE AMERICAN NATURALIST. [Vor. XXXVII.

these exceptions belong to other western streams, the other

species (AAznichthys dulcis) is found in all rivers whose sources

are in the Rockies. Of the 18 genera 4' are thus far known

only from the Colorado basin.

More than half of the Colorado fishes are minnows (Cyprin-

idæ), and of these the white salmon (Prychocheilus luctus Girard )

is the largest member of the family. In the Colorado River

specimens of this species are occasionally taken which reach a

weight of 80 pounds. The blob (Cottus punctulatus) is the

only spiny-rayed fish known from this basin.

Up to within the past year and a half very little was known

concerning the fishes of the Rio Yaqui, the largest river in

Northwestern Mexico. The few fishes previously taken in that

river indicated that its fauna was that of the Colorado. The

finding of a bull-head in this basin in 1896 seemed a little

strange and it was difficult to account for its presence there.

With these facts in mind, when I was collecting fishes in Chi-

huahua it was with no small amount of interest that I visited

Lago de Castillos which is a part of a small river basin between

the head waters of the Rio Conchos and the Rio Yaqui. At

Castillos I found only the Rio Grande chub. In the Yaqui I

also found this chub, and a number of species I had taken in

tributaries of the Rio Grande, at Chihuahua, and San Andres.

Of 14? species known from the Rio Yaqui, 9 (listed below in

bold type) have been taken in the Rio Grande basin, 2? have

been found no where else ; one of these (Gila minace) belongs

to a genus peculia to the Colorado river basin, the other

(Catostomus sonorensis) belongs to a genus well represented in

both the Colorado and Rio Grande. Of the remaining 3 species

2 (Agosta chrysogaster and Pacilia occidentalis) belong to the

Colorado river fauna. In the lower portion of the Rio Sonora

! Xyrauchen, Tiaroga, Meda, Plagopterus.

? Ameiurus pricei (Rutter), Pantoriens plebius (B. & G.), Catostomus bernardini

Girard, Campostoma ornatum Girard, Pimephales Fonfértur (Girard) Gila minacce

Meek, Leuciscus nigrescens rand. Notropis ornatus (Girard), Notropis lutrensis

(B. & G.) Agosia chrysogaster Girard, Salmo spilurus Cope, Cyprinodon elegans

(B. & G.) Pecilia occidentalis B. & G.

3 Catostomus bernardini and Gila minace.

No. 443] FRESH-WATER FISHES OF MEXICO. 775

3 or 4! species which belong to the Colorado river fauna have

been taken.

The presence of so many fishes from the Rio Grande basin

can be thus interpreted: — The head waters of the Rio Paphig-

ochic, a tributary

of the Rio Yaqui, lie east of the central

wet

wm ae i

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Map diving head waters of the Rio Yaqui and a western tributary of the Rio Grande.

range of the Sierra Madre mountains.

That portion of this

stream no doubt formerly had its outlet through Lago de Castil-

los into the Rio Conchos and in this way became stocked with

fishes from the Rio Grande? The fact that the fauna of the

! Catostomus bernardini Girard, PAychocheilus lucius Girard, Agosia oscula

(Girard), Pecilia occidentalis (Baird & Girard). m ; :

? A collection of fishes was made recently by the writer in the Rio Mezquital

at Durango. This collection has not yet been studied; it is however composed

largely of Rio Grande species.

776 THE AMERICAN NATURALIST. [Vor. XXXVII.

Rio Yaqui is so much like that of the isolated river basins in

northern Mexico rather strengthens this belief, though its ulti-

mate proof must depend on the geologist.

In Northern Chihuahua west of the Rio Grande and adjacent

to the head waters of the Gila river, the Rio Yaqui and the

Rio Conchos, there is a considerable area which is drained by

several small river systems, all of which have no outlets. Five

of these small basins have been examined as follows: the Rio

Carmen which drains into Lago de Patos, the Rio Santa Maria

which drains into Lago de Santa Maria, the Rio Casas Grande

which drains into Lago de Guzman, the Rio Castillos which

drains into Lago de Castillos, and a small stream at Sauz, in the

state of Chihuahua. During the rainy season the water collects

in the lowest portion of these valleys forming large lakes.

Most of these lakes become quite or entirely dry by the end of

the dry season, but there is always enough water in the upper

courses of the rivers which flow into them to sustain a consid-

erable number of fishes. All of these streams were at one time

a portion of the Rio Grande. These five basins have not been

equally explored, though it is likely that all have about the same

fauna. From these basins have been taken ro species ! of fishes.

Of these 10 species none has been recorded from the Gila

river or the Rio Sonora. All except 3 (Usted in bold type)

arereported from the headwaters of the Rio Yaqui in Chihua-

hua, and from the Rio Conchos. The three exceptions are

species closely related to Notropis /utrensis, an extremely vari-

able and widely distributed minnow, and which is abundant in

both of these rivers. One other minnow (Notropis ornatus) is

! Pantosteus plebius (B. & G.). Casas Grandes; Rio Carmen ; Sauz.

Compostoma ornatum Girard. Casas Grandes.

Pimephales confertus Girard. Casas Grandes; Santa Maria.

«ciscus nigrescens (Girard). Casas Grandes; Santa Maria; Carmen

Castillos; Sauz

Notropis tirida Girard. (Identification doubtful), Santa Maria.

Notropis santamariæ Evermann & Goldsborough. Santa Maria.

Notropis formosus Girard. Casas Grandes.

Nae lutrensis (B. & G.). Casas Grandes; Santa Maria; Carmen; Sauz.

eles elegans (B. & C.). „Casas Grandes ; Santa Maria; Carmen

busia affinis (Baird & Girard). Sauz.

No. 443] FRESH-WATER FISHES OF MEXICO. 777

common to both the Rio Conchos and Rio Yaqui, but at pres-

ent is not known from any of the four small river basins. No

other species than those here mentioned is known to be common

to the Rio Yaqui and Rio Conchos.

Aside from the five small river basins mentioned above there

Skiffia erme Meek, Q.

Skifia lerma M eek, d'.

1 no outlet to the sea, of

are in central Mexico several others witl

From a number of these

which the Rio Nazas is the largest.

no collections of fishes have been made, though their fishes so’

778 THE AMERICAN NATURALIST. [Vor. XXXVII.

far as known are that of the Rio Grande. From the Rio Nazas

are known 12! species of fishes, 6 of these (listed in bold type)

have been taken in the Rio Grande or its tributaries, all of the

others except Stypodon signifer and Characodon garmani, belong

to genera well represented in the Rio Grande. The genus

Stypodon is known only from the Rio Nazas, and Characodon is

a tropical genus. Every large lake or river, as a rule, contains

one or more species of fishes not found in other localities.

It is evident that the larger number of the Rio Grande fishes

have migrated directly or indirectly from the Mississippi valley,

23 of its 85 species being found in the Wabash in Indiana.

This fauna has crowded its way over the divide and has become

more firmly established in the Pacific coast streams of Sonora

than has the Colorado river fauna, and one species (Notropis

nigroteniatus) at least has gotten as far south as the Rio Balsas

in southern Mexico.

The southern portion of the Mexican plateau is drained by

two rivers: the one to the east, the San Juan del Rio, is a

small stream which flows into the Rio Panuco. The other,

the Lerma, flows into the Pacific. The Lerma is the longest

river in Mexico. The valley of Mexico was formerly a part of

the Lerma drainage system. The fish fauna of this region is

very different from that either to the north or the south. From

the area which includes the valley of Mexico,? the head waters

of the San Juan del Rio? and the Lerma basin, are known at

present 49 species of fishes, not one of which is known to occur

1 Ameiurus 77;c; Rutter, Carpiodes tumidus Girard, Pantosteus —

(Garman), Hybognathus int Garman, Stypfodon signifer G n, i

cus nigrescens (Girard), Leuciscus modesta (Garman), jig garmani

Jord. Ev., Rhinichthys simus Garman, Cyprinodon latifasciatus Garman,

Characodon garmani Jordan & Evermann, Etheostoma pottsii (Girard), Ethe-

ostoma australe Jordan.

2 The following is a list of the fishes known from the Valley of Mexico, those

printed in bold type are d to this region

Algansea tincella (C. & V.), Aztecula azteca (Woolman), Evarra eigenmani

Woolman, Evarra FE ABA C Meek, Girardinichthys innominatus Bleeker,

Zoogoneticus miniatus Meek, Skiffia variegata Meek, Chirostoma jordani Wool-

man, Chirostoma humboldtianum (C. & V.), Chirostoma estor Jordan.

? List of species known from the headwaters of the San Juan del Rio.

Algansea tincella, Aztecula mexicana, Goodea caliente.

No. 443.] FRESH-WATER FISHES OF MEXICO. 779

in any other river. These 49 species belong to 17! genera, 10

of which are peculiar to this region.

Of the genera found elsewhere, Characodon is represented

in southern Mexico, central America, and lower California.

Gambusia comprises a number of small viviparous fishes usually

inhabiting swamps and springs all the way from Southern

Illinois to Panama. The other five genera, Lampetra, Ameiurus,

Moxostoma, Notropis- and Hybopsis are northern genera, and

all except Notropis are not represented by any species farther

south than the Rio Lerma. Of the 49 species found in this

region, 33 belong to two families; 17 to Peeciliidee (the Killi-

fishes) and 16 to Atherinidz (the Silversides). It is curious to

note here that all of the killifishes are viviparous, yet only one

species, Gambusia infans Woolman, has the anal fin of the male

placed well forward and modified into an intromittent organ such

as is characteristic of Heterandia, Poecilia and the like. In the

other 15 species the anal fin of the male has its normal position

and size. It is slightly modified by the shortening of the first

five or six rays, and their slight separation from the rest of the

fin by a shallow notch. This modification was first noticed by

Günther in Characodon lateralis Gunther. It was also described

by Bean in Zoogoneticus robustus (Bean), and by Jordan and

Snyder in Goodea caliente J. &. S. but no significance was

attached to it. Just what part this fin plays in fertilizing the

eggs in the body of the female is not known, but it evidently

plays a porminent part in this operation.

I was fortunate to collect these fishes during the breeding

season and so their viviparity was easily proved. The largest

killifish known from the Lerma Basin reaches a length of 8 or

10 inches. The accompanying figures are made from a photo-

graph of the largest female of this species I was able to obtain.

It was purchased from a fisherman who did not suppose it would

find its way into a distant museum and this explains its rather

dilapidated appearance. The ovary consists of a membranous

! The genera in italics are peculiar to this region. ;

Lampetra r, Ameiurus r, Moxostoma 1, Algansea 4, Aztecula 3, Notropis 1,

Xystrosus 1, Evarra 2, Falcula 1, Hybopsis 1, Zoogoneticus 5, Gi : ys

1, Characodon 2, Chapalichthys 2, Gambusia r, Goodea 7, Chirostoma 16.

780 THE AMERICAN NATURALIST. [Vor. XXXVII.

sack with a number of infolded partitions. Removing a portion

of one side shows the ovary full of quite well developed young.

The little fishes in it are not arranged in any definite order.

The spawning time for these fishes is near the close of the

dry season. At this time the water is more concentrated, as is

also the food on which the young must feed. The aquatic

insects, crustaceans and small fishes which would feast on the

eggs if deposited then are also more concentrated, so that depos-

iting the eggs at this time would mean considerable destruction

to the species. As it is, the young are born in a well developed

stage, and have time to reach some size before the wet season

sets in. They are then perhaps in the best condition to become

widely distributed as the volume and area of water increases.

As the dry season approaches again, and small streams and

ponds become dry, many of these small fishes perish. They are,

however, present everywhere to establish themselves in every

body of water which may carry them through to the next rainy

season. 1

The gestation of many tropical fishes presents some strange

peculiarities. Some of the catfishes carry the eggs in the mouth

till hatched, while a few others are thought to be viviparous.

Viviparity among the tropical killifishes in general seems to be

the rule rather than the exception. It would seem that in the

tropical fresh waters of America, there is much more provision

made for the care of the young than in the cooler waters of the

Northern continent.

It was rather surprising to find such a large number of Chi-

rostoma in the Lerma basin; no other river in North America

indeed, has so large a proportion of its fishes belonging to a salt `

water! family. It is probable that this number will be consider-

ably increased when this basin is more thoroughly explored. I

had seen but a few specimens of Chirostoma before going to

Mexico, and so never had an opportunit y to study these fishes.

And while I was careful to pick up specimens of all species, yet

my unfamiliarity at that time with the group, no doubt, caused

me to overlook some species. Again there is a number of small

1The Chirostoma are the only fishes belonging to a salt water family found on

the Mexican plateau.

‘BZunod Sururejuoo Areao yw *( c /

782 THE AMERICAN NATURALIST. [Vor. XXXVII.

isolated lakes which have never been visited. It is known that

some of these lakes, as Patzcuaro and Zirahuen, have in them

one or more characteristic species and no doubt most of them

have. The Lerma river system is far from being thoroughly

explored, but apparently its fish fauna is quite as distinct and

characteristic as if it were an island in the sea.

All of the many beautiful lakes in this area, now isolated,

evidently at one time drained into the Lerma, and so became

stocked with fishes. It is often argued that fishes become estab-

lished in isolated lakes by their eggs being carried accidentally

by water birds. Although while these birds are feeding some

eggs of fishes might cling to their feathers or legs and be taken

to adjacent waters, yet I much doubt the dispersion of fishes in

this way. Were this method of dispersion at all common fishes

would surely have been found in Shoshone and Lewis lakes in

the Yellowstone Park; moreover in the isolated lakes in the

valley of the Lerma the viviparous fishes are about as evenly

distributed as are the egg laying ones.

The Rio Balsas is one of the largest rivers in Mexico. It is

southeast of the Lerma and drains about the same area; and

though these two rivers are so near each other, not a single spe-

cies is known to be common. But one species of the silverside

and two of killifishes, are known from the Balsas, and yet these

two families comprise nearly two thirds of the fishes of the Lerma

basin. The Balsas is far from being thoroughly explored ;

enough, however, has been done to indicate the nature of its

fauna and to indicate that it contains comparatively few species

of fishes.

In Mexico there are four quite distinct fish faunas, and though

they overlap at the borders, the map may fairly indicate where

each fauna prevails.’ The origin of these fish may be approxi-

mately given as follows: from the Colorado river 9, from the

Rio Grande 80, from the Lerma 49, and from Central America

about 108 ; total 246.

The fish fauna of northern Mexico is essentially that of the

Rocky Mountains and eastern United States, or that part of

the United States adjacent to Mexico. This eastern fauna has

1The fishes of the region marked unknown probably belong to the Rio Grande

fauna.

No. 443] | FRESH-WATER FISHES OF MEXICO. 783

crowded its way over the divide and has become even more firmly

established in the Pacific coast streams of- Sonora than has the

Rocky Mountain fauna.

The South and Central American faunas prevail largely as far

north as the City of Mexico. The few forms which extend far-

ther north apparently keep to the lowland streams ; especially

is this true on the Pacific side. The most northern representa-

tive of the South American fauna, one of the Cichlids,! is found

in Mazatlan. On the east coast this family has a representative

in Texas. The fauna of the Lerma, the only river basin exten-

sively studied, is quite distinct from either North or Central and

South America. This fauna is richer and more characteristic

than was formerly supposed.

Mexico in general is not a well watered country. Nearly all

of the small streams and many of the large ones become much

reduced in size by the end of the long dry season, and such

streams never sustain a large number of species of fishes. On

the Mexican plateau the largest and most important lakes are

found in the Lerma basin; Lake Chapala, the largest and the

only one which has a large river for outlet and inlet, sustains

the largest fish fauna. Patzcuaro, a large lake with no inlet or

outlet does not have so varied a fauna, but supports a large

number of individuals. In view of the fact that more species of

fishes belong to tropical Mexico than to a like area farther north

it seems strange that a great river like the Balsas which lies

wholly within the tropics should contain so few species. This

river is fed by many mountain springs, and even in the dry sea-

son contains an abundance of clear water. Collections of fishes

have been made at but three places in this river basin, and in

all only rt? species of fishes taken, a number much fewer than

one would expect.

! The name Mojarra is used for the Cichlids in Mexico, it is also much used on

the plateau for the larger Peecilide

? Istlarius balsanus Jordan & Sutyder Algansea sallei (Gunther), Notropis

ther), J rtragenepier es mexicanus Filippi, Gambusia gracilis

nyder, Chirostoma jordani Woolman

—

nigrolteniatns (Gun

Heckel, Pæcilia limantouri Jordan p

Melaniris balsanus Meek, d nasutus Gunther, Heros zstlanus Jordan

& Snyder, Awaous taisiaca (Li um

Algansea sallei and a. jordani are in my opinion wrongly ascri

in this river basin.

784 THE AMERICAN NATURALIST. [Vor. XXXVII.

From many lakes and rivers in Mexico no collections of fishes.

have been made. In conclusion I will say that since the fish

fauna of Mexico is far from being thoroughly explored, the

faunal areas as I have outlined them, and their origin and prob-

able lines of dispersion must be regarded as tentative.

FIELD COLUMBIAN MUSEUM,

Chicago, June, 1903.

EXAMINATION OF ORGANIC REMAINS IN

POSTGLACIAL DEPOSITS.

PEHR OLsson-SEFFER.

LiTTLE or no attention has been paid in America to the study

of fossil plants in the postglacial deposits. They do not offer

such a fascinating field to the investigator as the tertiary and

other older formations. They do not show a multitude of forms

of animal and vegetable life, beautifully preserved from the times

when the earth was young; only a few fragments of recent

types, difficult to determine and. mostly of a very diminutive

size, necessitating a constant use of the microscope. But they

are, nevertheless, interesting, especially to the student of descrip-

tive phytogeography, as recorders of the history of the vegeta-

tion, and to some extent as indicators of climatic conditions in

times gone by. In this respect the great importance of an inves-

tigation of, for instance, the formation of peat-bogs, cannot be

overrated, and in northern Europe this study has developed

during the last decades into a special science, called in Germany

Moorkunde. A name of a more international character, telma-

tology,! has been used by some authors,” and seems acceptable.

The Scandinavian countries, especially Sweden, have been the

center of this study, and consequently, the development of the

Scandinavian Flora and vegetation is better known at the pres-

ent day than that of any other part of the world. r

It is the purpose of this paper to give a brie

methods for collecting, preserving and examining the plant-

remains in recent deposits, as these methods are now generally

employed by paleobotanists, with a few additions from the

writer’s experience in the study of telmatology. In another

f review of the

! From TéApa = swamp or bog. ;

? Klinge, J., for example, nearly twenty years ago. Not having access to the lit-

erature, I cannot at, the time of writing. ascertain who proposed this name.

Lagerheim suggested (1902) a nàme derived from Havewos 7. ^. = combustible,

but both priority and suitability speak in favor of ‘Telmatology.

785

786 THE AMERICAN NATURALIST. [Vor. XXXVII.

place the development of these formations and their relation to

certain plant-communities will be treated.

Japetus Steenstrup of Copenhagen was the first to begin the

difficult task of identifying the organic remains in peat bogs and

similar deposits. After him Axel Blytt of Christiania, A. G.

Nathorst and Gunnar Andersson of Stockholm, Rutger Sernander

and Henrik Munthe of Upsala have been the principal workers

in this field. Many pupils of Andersson and Sernander have in

later years pursued the study in Germany, Russia and other

countries, and the literature on the subject is rapidly increasing.

The first paper on the method of examining fossil plants in

postglacial deposits was published by Andersson in 1:892.

Improvements on his method were made known in 1892, 1893

and 1896? Munthe gave (1894) a detailed account of biologi-

cal investigation of clays,s and Professor G. Lagerheim * recently

('02) related some new experiences with regard to the technique

of telmatological research.

All these papers are in the Swedish language and the writer

thinks he is justified in bringing the methods in question under

the notice of American paleobotanists and phytogeographers, as

a study of the evolution of the plant-covering based on paleonto-

logical testimony is likely to find adherents in the United States

and Canada, where postglacial deposits, so widely distributed

and covering immense areas, offer special advantages for this

line of research.

The principal kinds of recent deposits in which we meet with

fossil plants, are fresh water alluvium, lacustrine deposits and

peat bogs. Wherever these formations are developed, accumu-

lation of partially decomposed organic matter has been the most

important agent in their construction.

When this process of decomposition is proceeding in presence

! Om metoden för vüxtpaleontologiska undersökningar af torfmossar. Geolog.

Jören. fórh. Stockholm, vol. XIV, pt. 2, pp. 165-175.

2 Om slamning af torf, Zoc. cit. vol. XIV, pt. 6, pp. 506-508; Om metoden for

botanisk undersókning af olika torfslag. Swenska igitnr tidsk., 1893,

and Om konservering af kvartiira vüxtlámningar. Geo/eg.füren. fórA., vol. XVIII,

pt. 6, pp. 492-498.

3 Om biologisk undersökning af leror. Geol. fören. forh., XVI, pt. 1, pp. 17-28.

* Torfiekniiks notiser, Joc. cit., XXIV, pt. 6, pp. 407-411

No. 443.] ORGANIC REMAINS. 787

of an excess of water, humic acid and certain hydro-carbons are

formed, and it is to these substances the said deposits owe their

anti-septic properties, which make it possible for organic remains

to resist decay for a sufficiently long time to allow deposition of

the sediment, in which they are finally imbedded.

Trees falling into the water, branches, roots, leaves, seeds,

and other parts of plants are often in this way preserved, and

retain sometimes their shape, color and anatomical structure to

a surprising degree, so that there is no difficulty in discriminating

the distinct species. It is, however, only lignified and corky

tissues that are able to resist decomposing. All those organs

which have not cell-walls modified in this way, are liable to be

destroyed. Of leaves, for instance, only the epidermis and vas-

cular bundles remain, while mesophyll and similar tissues decay.

The fossil remains are therefore often quite different in

appearance from the plants that fell into the water, where they

were deposited. Among Salices that are found in post-glacial

deposits, species with hard leaves, as Salir aurita L., S. cinerea

L. and S. nigricans Sm. remain unaltered, both with regard to

form and consistency, although, of course, the color is changed ;

the nervation is very distinct. In the case of S. myrsinites L.

only the skeleton of the ribs is left. S. lanata L. and S. lap-

ponum L. are very difficult to recognize, because the character-

istic tomentum has disappeared, and instead, the nervation,

which in the living condition cannot be traced, is rendered very

conspicuous.

In beginning the study of telmatology one of the greatest

difficulties met with is the fact that there are, as yet, only a few

study collections accessible, and no complete works of reference

with excellent illustrations and descriptions such as are available

in other branches of paleontology. The student has usually to

prepare for himself the comparative material he wants.

By means of certain maceration processes the same effect can

be accomplished in a few minutes in the laboratory that required

Thus it can be also ascertained to some

ether a certain plant can be preserved

d clays, or if it will be com-

he influence of water

a long time in nature.

degree of probability, wh

in a fossil state in mud, peat, an

pletely decomposed when subjected to t

and other agencies in the deposits.

788 THE AMERICAN NATURALIST. (VoL. XXXVII.

For this purpose the plant is boiled in Schultze’s maceration

mixture, which consists, as every botanist knows, of potassium

chlorate and nitric acid. Leaves, seeds and other parts of the

plants, which are usually found fossil, soon acquire the same

dark-brown color that is so characteristic for peat, and it is

almost impossible to distinguish these preparations from the real

fossils. Plants, however, which are almost instantly destroyed

by this strong reagent, never occur in the said deposits. It

can, therefore, be taken for granted, if the tissues are destroyed

within a minute or two, that the result would have been the

same in water, but if only bleached, or in a lesser degree mace-

rated, it can be supposed that the organ would have resisted

decomposition. °

These macerated objects can then be mounted and preserved

in the way usually adopted for microscopic preparations. Every

student of fossils in postglacial deposits should in this way secure

the material needed for comparison.

The collecting of fossils consists partly of field work and

partly of operations in the laboratory. For the former purpose

the student should be supplied with the following tools. A

small steel spade, about 20 cm. in length and 14 cm. in width,

with a handle like that on a mason’s trowel, and with sharp

edges, for cutting purposes ; a pointed knife with a blade of at

least 14 cm. in length ; a pair of forceps, a soft camel's-hair brush ;

a white china plate ; and a pocket microscope. ‘Further, a num-

ber of flat-bottomed test-tubes of different sizes: 60 x 18 mm.,

50x 16 mm, and 40 x 12 mm being the most suitable sizes ;

strong, wide-mouthed glass bottles, 80 x 40 mm., and some glass

jars of about 12 cc. capacity.

If collecting is done in deposits more or less petrified and

hard, such as calcareous sinter or tuff, the usual tools of a

geologist are needed.

Sometimes it will be found impossible to remove fossils found

in loose sand deposits, because they are too brittle, and in such

cases it is advisable to fix the sand particles together with water

glass, as silicate of potassium or sodium, readily and completely .

soluble in water generally are called: Although the fossils pre-

served in this way lose their color, and if not prepared most

No. 443.] ORGANIC REMAINS. 789

carefully will break, this method, nevertheless, has many advan-

tages ; and Andersson ' recommends always to be supplied with a

bottle of soluble glass when collecting in sand- and clay-deposits.

If care be taken to let the preparations dry slowly, the result

will often be surprisingly good.

At the places chosen for taking the samples of peat or similar

soft deposits, vertical holes are dug to the desired depth, the

cutting being trimmed with the sharp spade, care being taken.

not to disturb the succession of strata, or to get any recent plant

fragments mixed into the mass. Careful notes of the freshly

cut layers should be taken immediately, before the peat begins

to darken through the influence of the air. Samples, ro cc. in

volume, should, in general, be taken at intervals of 50 cm.

throughout the profile. Wherever any marked differences in

the soil are observed, separate samples should be secured. The

depth of every sample must be measured and noted on the labels

and in the field book, as well as any observations regarding the

consistency, color, odor, and other characteristics of the respective

strata from which samples are taken. These samples are pre-

served either in jars or in clean canvas bags, and later examined

in the laboratory.

The collector should also search for fossils on the spot.

this purpose the white plate is filled with water to the rim, and

slices cut out from the stratum to be examined are carefully

broken into pieces and washed, and any seeds or other remains

removed with the brush and forceps, and preserved. This

examination is facilitated if the peat is placed for some time in a

diluted potassium or sodium lye, which must, however, be care-

fully washed away afterwards. Lagerheim’s oxalic acid method,

which will be described later, is still better for the purpose.

Series of samples are taken on different places of the bog,

usually in a line across the deepest part of the formation so as

to give a section of the basin, in which it has developed. The

number of profiles to be opened depends naturally on the extent.

and topography of the formation, but from three to five profiles

For

! Andersson, G. Om senglaciala och postglaciala aflagringar i mellersta Norr-

land. Geol. fören. fórh., vol. xvi, pt. 6, p. 550

790 THE AMERICAN NATURALIST, (Vor. XXXVII.

between the centrum and the shore are sufficient in most cases

for a bog of, say, 10 acres.

The collecting often has to be done under great difficulties on

account of the swampy character of the peat, which is often of

so loose a consistency, that it is impossible to open a hole to any

depth. In this case an earth-auger or soil-sampler, has to be

employed. Of these instruments there are many kinds in use.

One of the best for peat sampling purposes, that has come under

the observation of the writer, was described in 1894 by A. G.

Kellgren.*

His peat-auger consists of a steel pipe 1.5 m. in length and

about 4 cm. in diameter. The

accompanying illustration (Fig. 1)

€ shows how the auger is arranged.

The lower end of the pipe is closed

with a piston which is pointed at

the apex, and can be lowered and

raised in the pipe with a steel rod,

managed from the upper end.

If the sample to be taken is from

a compact peat, the auger is sunk

to the required depth, the piston is.

drawn back into the pipe, and then

. the auger lowered for about 10 cm.

4 The lower end of the pipe will thus

: be filled with the earth which the

auger retains, when withdrawn. In

order to secure the samples in a nat-

ural state, the first 10 cm. of pipe

$id. r0 PARE at the end of the auger is split in

two halves, and these are secured

by hinges on one side, and fixed to the main stem of the pipe

with a screw arrangement which holds them together. When

the sample has been obtained, this 10 cm. end of the pipe con-

taining it is unscrewed and opened, thus allowing the sample to

be removed intact.

1 En ny konstruktion af mossborr. Geol. fören. fórA., vol. xvi, pt. 4, pp. 372—

374- :

No. 443.] ORGANIC REMAINS. 791

When sampling is to be done in very loose or almost liquid

sediments, the piston is pushed below the pipe, and when the

end of this is filled with the mass, the piston is drawn back to

its former position, which secures the sample.

The end of the pipe, which comes in contact with the sample

should be kept scrupulously clean and free from rust. When

the piston is withdrawn into the pipe, the cutting through the

sediment, is, of course, done by the sides of the pipe as the

auger is lowered. It is, therefore, essential that this part of the

pipe be made of the best steel, so that the sides can be ground

to a knife-edge and kept in that condition. The pitch of the

screw must be low, and the boring should always be done very

slowly. If samples are wanted from greater depths, the handle

of the auger is shifted and new lengths of pipe and steel-rod are

added.

The ulmic and humic substances, or those chemical compounds

to which the peat owes its peculiar character, are developed in

the presence of water and when dried are subject to molecular

alterations, by reason of which they lose their ability to re-absorb

water. There is no reagent known, as yet, that can restore to

dry peat its original properties. The usual means employed in

microscopy for causing swelling do not give satisfactory results.

Experiments with lactic acid have also failed.

All collections from peat and other moist deposits have, there-

fore, to be kept in some preserving fluid, and must not be

allowed to dry, because this would considerably increase the

difficulties of determination, and in some cases even make identi-

fication impossible. With collections from fossiliferous clays and

sand deposits this is not absolutely necessary, but desirable.

Früh has shown through experiments that the ulmic and humic

substances are immune for bacteria and fungi, and by reason of

this they are almost completely absent from the peat-water,

which can be used for some time as a preserving medium.

Alcohol is generally employed, but samples of peat can be kept

ina fresh state covered with the swamp-water in air-

ly disinfected with carbon disulphide

In case the samples have to be transported for some distance,

the vessels containing them should be well filled with the pre-

serving liquid so as. tó prevent unnecessary shaking.

for years

tight vessels, if previous

792 THE AMERICAN NATURALIST. (VOL: XXXVII.

Dried peat can to a certain degree be made suitable for exam-

ination, if boiled in water for three hours and afterwards satu-

rated with 5 fer cent. ammonia water for 48 hours. It should

then be subjected to the same treatment as fresh peat, when

prepared for examination. Fossil seeds and fruits, which have

been allowed to dry, can be restored to their original shape and

volume by the influence of a weak (2-3 fer cent.) ammonia

solution.

When peat has been under the influence of air for some time,

it darkens, and the more this change of color proceeds, the more

difficult will it be to find and determine the fossil remains. If

the water contains iron in solution, as is often the case, the

samples will, in a very short time, be almost black, which consid-

erably lessens the possibility of a successful botanical exami-

nation.

In order to restore the original color to the fossils it is there-

fore necessary to let them undergo a bleaching process. This is

effected in several ways. The oldest method, employed by

Schróter! (1883), was to use Schultze's mixture for bleaching,

as neither ammonia, potassium hydroxide, or calcium hypochlorite

gave good results. This reagent certainly makes the dark-brown

and opaque plant-remains from the peat transparent, so that ner-

vation, cells, etc., can be studied, but usually acts too strong and

often destroys the objects.

At present, Gunnar Andersson’s nitric acid treatment is the

method most used. According to this method the peat samples

are put for 24—30 hours into commercial nitric acid diluted with

twice as much water. In the phytogeographical laboratory of the

University of Upsala, where the writer first studied telmatology

under the guidance of Dr. Sernander, a solution of one part

nitric.acid (65 per cent.) and 3 parts water was used for mace-

rating peat of loose texture, and one third acid when the samples

were compact. From 12 to 16 hours treatment according to my

experience, is sufficient in most cases for bleaching the peat, so

that the fossils can be washed out.

The advantages of this method are certainly very great : all

! Die Flora der Eiszeit. Zürich, 1883, page 21.

No. 443.] ORGANIC REMAINS. 793

clays, whether calcareous or not, disintegrate, and the samples

of the usually tough and oily mass from the strata lying under

the peat proper swell and are macerated. The dark volor is

bleached, the fossils are filled with gas-bubbles and float on the

surface of the fluid, so that they can easily be collected.

But the method also has its drawbacks. Nitric acid of the

strength required is liable to act with more or less damaging

results on the organic tissues, and thus make the fossils more

friable still than they were before. Certain minute microscop-

ical remains are usually totally destroyed.

The process of bleaching should, of course, be done under a

hood or similar device to get rid of the fumes of the acid. In

case the examinations are done in the field one is confronted

with the additional difficulties of transporting the acid.

Lagerheims's method of bleaching with oxalic acid is undoubt-

edly an improvement, because no injurious fumes are developed,

the fossils are not affected, and the acid is in a solid form, and

consequently easy to handle and transport. From the fact that

oxalic acid is able to decolorize organic iron compounds Lagerheim

concluded that it would be a good reagent for bleaching peat,

especially when it contained iron in solution and had darkened

in the air. Acting on this suggestion he found that pieces of

peat immersed in a 3 fer cent. solution of oxalic acid, almost

instantly lose their dark color, which changes to brown. For the

bleaching process a glass vessel is most suitable, and if this is

exposed to daylight, or still better, to sunlight, the brown color

fades gradually, until, after a few hours, the peat mass is ready

for washing.

The influence of light is explained by the fact, already

observed by Downes and Blunt (1879), that solutions of oxalic

acid evolve carbon dioxide when exposed to the action of light.

Other catalyzing agents are, for instance, salts of iron, which

‘usually are present in peat. To the writer's knowledge the

composition of these iron compounds that cause the dark coloring

of peat has not yet been ascertained. Lagerheim is inclined to

. think that we have to do with some organic iron compound.

Peroxide of hydrogen is formed! in the process of oxidation

! Richardson, A.: The action of light on oxalic acid. Proceedings Chem. Soc.

London, 1894, (137); 88.

794 THE AMERICAN NATURALIST. [Vor. XXXVII.

of the oxalic acid solution, which probably takes place aécording

to following reaction :

C,O.H, + O, = 2CO, + H:O, and, it is perhaps this peroxide

of hydrogen that, in combination with some other compounds,

effects the bleaching.

According to Richardson, the total amount of hydrogen per-

oxide formed in the solution increases with the concentration of

the acid, while at the same time the proportion of peroxide to

acid formed decomposed decreases simultaneously, and since the.

described action of the head occurs with greater rapidity if con-

siderably diluted, only a very weak solution should be used.

If the fossils, especially leaves, are wanted almost colorless,

the following method of bleaching is recommended. A solution

(not too strong) of potassium permanganate, is employed where

they are allowed to lie for some time, and then transferred

directly into the oxalic acid solution.

To extricate fossils from calcareous peat it is necessary to

remove the carbonate of lime, and this is best done with hydro-

chloric acid. If, however, the material contains lime in a small

degree only, application of the acid will result in effervescence,

which causes the decomposition and penetration to take place

very slowly and unevenly. In order to prevent this, the peat-

particles are thoroughly saturated with strong alcohol, and the

hydrochloric acid is applied afterwards. The separation will

now proceed easily and uniformly, and the gas-bubbles are

bursting so soon, that no undesirable foaming is caused. Should -

this occur, the mass is again treated with alcohol. The sepa-

rated peat-material can then be preserved in the alcoholic cal-

cium-chloride liquid.

Whatever method is employed for bleaching, this process has

to be done very carefully, so as to prevent the fossils from being

destroyed by the acids. The next step is the “slumming,” or

washing of the macerated mass. For this purpose there are

different devices for slumming vessels. These are all constructed

on the plan of creating a rising current of water through the

mass, which is poured over a sieve of brass netting with meshes

not smaller than 1.5 mm. in diameter.

! Hydrogen peroxide alone does not bleach peat.

No. 443.] ORGANIC REMALNS. 795

A good arrangement is to have a porcelain vessel fitted with

two sieves, the upper one with meshes of about 2 mm. and the

lower r.5 mm. The sieves are placed about 5 cm. apart, and

two currents of water, the velocity of which can be regulated by

cocks, should be used, one under each net. Figure 2 shows a

contrivance made on these principles and used by the writer

with good success. The residue of the slumming need not be

examined, if portions of the sample have been reserved for micro-

scopic examinations for spores, pollen, alge, bryozoa, molluscs,

rhizopods, cirripeds and other Crustacea, fragments of echino-

derms and insects, and

other minute animal

remains. In order to

get a more complete

collection of these

fossils the slumming

water, which, of course,

has previously been ex-

amined and found free

from diatoms, should

be allowed to pass

through a silk net as

Shown in Fig. 2.

The slumming is

comparatively easy if '

the material is some- = Pectore

what sandy, but when

Sticky or miry, the

mass has to be stirred and sometimes broken by the hands of

the operator.

With regard to the slumming and preparation of clay s for =

study of diatoms or for mechanical analysis, the technical details

have so often been described that we need not go into them

here. : : :

When the fossils are ready for preservation twigs, pieces a

bark and wood, cones, nuts, rhizomes of grasses, and other larger

fossils are usually kept in 40 fer cent. alcohol or a a I-2 per

cent. solution of formalin. Seeds and fruits are preserved in

Fic. 2.— Slumming vessel.

796 THE AMERICAN NATURALIST. [Vor. XXXVII.

alcohol or in sterilized water. In the latter case, the glass tubes

are immediately sealed with melted paraffin, which acts both as

a cork and as an isolating medium. Larger leaves are best pre-

served in a mixture of 2 parts of glycerine, 1 part carbolic acid,

and 7 parts water. Remains of delicate mosses and small leaves,

parasitic fungi, and algz should be preserved in Canada balsam

like ordinary microscopic preparations.

Andersson recommends another plan of preserving, which has

its advantages. After being dehydrated in alcohol, the fossils

are transferred to a 30 fer cent. solution of benzin-alcohol,

thence to a 70 fer cent. solution, afterwards, to pure benzin, and

subsequently to a saturated solution of naphthalin in benzin.

The fossils are kept here for some time, until the fluid has well

penetrated. When the objects are drying the benzin evaporates,

and the surface is covered with small crystals of naphthalin.

These gradually evaporate and the object will remain almost

entirely unaltered. No shriveling is observed, and the contrac-

tion is estimated to be only 1 fer cent. This treatment can be

employed well for preserving larger objects.

When examining and mounting minute and fragile fossils, it

will be found convenient to do the bleaching on the object-slide.

The material is then. washed in water in order to remove all the

acid, and afterwards placed in alcohol until all gas-bubbles have

disappeared. The washing should be repeated in absolute alcohol,

and when the objects are thoroughly dehydrated they are trans-

ferred to a mixture of'equal parts of xylol (or toluol) and absolute

alcohol, subsequently to pure alcohol for a moment, and by this

time they are ready for mounting in Canada balsam.

The fossils are usually rather brittle, so that when sections are

wanted the razor and freehand cutting will be found unsatisfac-

tory and imbedding in paraffin and the microtome have to be

employed as for histological work.

For an exhaustive investigation, the statistical method will be

useful to follow, and from the proposed size of samples 10 cc., a

fair idea can be obtained of the quantity of fossils in a certain

stratum. i

To get a clear conception of the history of the vegetation of a

place, it is not, however, sufficient to examine the fossils in the

No. 443.] ORGANIC REMAINS. 797

deposits. The topography of the neighborhood has to be care-

fully studied, and the existing vegetation investigated, especially

with regard to composition and relation of the various plant-com-

munities. But also ecological conditions have to be observed in

this connection, because in some cases they are of considerable

help in interpreting the successive evolutional phases of the

vegetation.

LELAND STANFORD JR. UNIVERSITY,

DEPARTMENT OF SYSTEMATIC BOTANY.

NOTES AND LITERATURE.

EXPLORATION.

Hatcher’s Narrative of the Princeton Patagonia Expedition.

— In a quarto of twelve chapters,’ illustrated by fifty large heliotype

plates and a map, Mr. Hatcher gives an account of three expeditions

to Patagonia after fossil vertebrates, adding at the end a chapter on

the geography of the region.

This is by far the best description of travel in Patagonia that has

been written. Mr. Hatcher’s observations are keen and accurate,

while his judgment is expressed in a fair manner, based on notes

made during a period of three years experience, in which time they

have been corrected and verified. The matter is of such absorbing

interest that one’s attention is held throughout. Here we see the

trained naturalist at work, and gain some idea of the hardships and

difficulties entailed and the joys experienced in exploring a com-

paratively unknown land.

There are many well-noted observations on the physical features

of the country, animal and vegetable life, geology, etc., which make

this book a compendium of interesting information regarding the

little known land of Patagonia. The heliotypes in several instances

do not clearly represent the subject illustrated, and it seems possible

that another method of reproduction might have been used to

advantage.

In the chapter entitled “ Geography " the author treats the great

physical divisions of the surface, describes the rivers, indicates the

origin of the numerous lakes by pointing out their relation to the.

present inland waters of the channels, and reasons out in a convinc-

ing manner the changes through which this part of the continent

. has passed from the earliest times. The different tribes of Indians,

their habits and mode of life are accurately described. The last

chapter is on the resources of the country.

In speaking of lizards, p. 84, Mr. Hatcher says

pampas, a great variety of small lizards of varying size, shape

and color, but no snakes." This observation on lizards should have

: €... over the

rsity Expeditions to Patagonia, 1896-1899.

! Reports of the Princeton Unive

i Geography of Southern Patagonia. Prince-

Narrative of the Expeditions

ton, The University, 1903. 4to. Xxvi-314 PP- 5! pls., map.

799

8oo THE AMERICAN NATURALIST. [Vor. XXXVII

been confined to that part of Patagonia north of the Rio Santa Cruz,

for this river forms the natural southern boundary line for lizards as

well as of armadillos though a few have been scattered south of it

by man. In describing the Guanaco on page 271, he says: ‘Their

presence in Fuego, to which island the rhea, puma and deer have

not gained access, is but an illustration of their superior powers of

self distribution." The distribution of guanaco on Tierra del Fuego

is far more probably attributable to the agency of man, for since

time immemorial the channel Indians have plied between Patagonia

and Fuego in their canoes and might papiy have introduced these

animals.

BARNUM BROWN.

ZOÖLOGY.

A Summary of the Coccidze.! — The new “ Catalogue of the Coccide

of the World” by Mrs. M. E. Fernald, just published by the Massa-

chusetts Agricultural Experiment Station, will be of immense value

s dw of these insects. For the first time since Signoret’s

* Essai " appeared, nearly thirty years ago, the species are catalogued

with full bibliographical references. The preparation of the work

has been a tremendous task, involving a search through the scattered

literature published in every part of the world, and in all sorts of lan-

guages. ‘There are few places where it could have been attempted,

and few people who would have had the courage and perseverance

to carry it out.

In the catalogue, 1449 species of Coccide are recognized as valid.

The time since 1758, when the tenth edition of the “Systema

Nature " appeared, may, so far as the Coccide are concerned, be

divided into four periods. The first is from 1758 to 1799, during

which 38 species were described. The second, from 1800 to 1850,

saw the description of 57 valid species. ‘lhe third, in which scienti-

fic coccidology really began, culminated in Signoret's famous work

which covered the whole subject as then understood. In this period,

! Fernald, Maria E. A Catalogue of ihe Coccidz of the World. Special Bu/-

Jetin Mass. Agr. Exp. Sta. No. 88, pp. 3

No. 443.] NOTES AND LITERATURE. Sol

from 1850 to 1875, 145 species now held valid were published. At

the end of the period, only 240 species were known, as against 1449 -

in 1903. ‘The last period, from 1876 to 1903, saw the publication of

no less than 1209 species, and no doubt the next will make known a

far larger number. ‘The beginning of the fourth period was coinci-

dent with the first labors of Maskell of New Zealand, whose writings

extended over many years, and made known the rich coccid-faune

of New Zealand and Australia, treating also of a good many species

from other regions. To-day, the workers are more numerous than

ever before, but still not sufficiently so to deal with the material

which might readily be obtained. The coccid-faunz of Cuba and

the Philippines, for example, are practically unknown, though there

is no doubt that they are rich and interesting.

The genera recognized in the list number 168, distributed in sub-

families as follows: — Diaspinze, 34; Coccinz, 57; Tachardiinz, 2 ;

Dactylopiinze, 54; Conchaspinz, 1; Phenacoleachiinz, 1; Orthe-

Zinz, 3; Margarodine, 5; Monophlebinz, 11. I have thought it

worth while to prepare the following summary, in which the genera

are all listed according to their distribution, the number of species

being given after each generic name. In doing this, I have made

use of my own knowledge of the introduction of species into localities

by human means, and of the fact of certain localities being errone-

ously cited. The purpose has been to throw light on the natural

distribution of the genera.

(1.) Genera which are cosmopolitan or nearly so. :

Palxococcus, r1. Asterolecanium, 27. Eriococcus, 6o.

Icerya, 16. . Lecaniodiaspis, 17. Phenacoccus, 36.

Coccus, 29. Pulvinaria, 6o. Pseudococcus, roo.

Chionaspis, 59. Aspidiotus, 77. Lepidosaphes, 62.

Some of these, such as Aspidiotus, Coccus and Chionaspis, are

assemblages of more or less discordant elements, and will no doubt

eventually be subdivided. On the other hand, the species of Lecani- -

odiaspis, Asterolecanium and Eriococcus are certainly closely allied ;

and in Lecaniodiaspis there are undoubtedly native species in South

Africa, Arizona and Japan, which are as nearly alike as they could

very well be without being the same. When we consider the limited

means of travel possessed by the Coccidz, and the long period which

must have elapsed during this wide migration, the persistence of type

is something remarkable. The monophlebid genera Icerya and Pa-

leococcus show the same sort of thing; and it is noteworthy that

802. THE AMERICAN NATURALIST. [Vor. XXXVII.

Palæococcus is one of the few genera found fossil. Icerya is want- -

ing in the colder parts of the Palæarctic and nearctic regions, being

essentially a tropical and subtropical type.

(2.) Genera which are very widely distributed, but not

cosmopolitan.

Margarodes, 10. Europe, Africa, America.

Orthezia, 19. Palzarctic, Nearctic, Neotropical.

Rhizococcus, 14. Europe, New Zealand, Australia, Texas; but

the species may not be strictly congeneric.

Spharococcus, 1g. Australia, Japan, North America. The

species may not be truly congeneric.

Ceroputo, 7. Europe; North America, extending into the Neo-

tropical region.

Ripersia, 37. Holarctic, New Zealand, Australia, and doubt-

fully congeneric species in India and Trinidad.

Tachardia, 24. Cosmotropical.

Lichtensia, 12. Holarctic and Neotropical, but mainly the lat-

ter, if the species are really congeneric.

Ceroplastes, 62. Very abundant in the Neotropical region,

extending into the warmer parts of the United States; Medi-

terranean region to South Africa; a few species in Asia and

Australia, those in the latter country probably introduced.

Saissetia, 20. Cosmotropical.

Phenacaspis, 15. North America, Asia, S. Africa, Australia.

Chrysomphalus, 31. America, Australia, Asia. Numerous in

Mexico.

Targionia, 18. Holarctic, Australia, India.

The doubt expressed as to the species of certain genera properly

belonging together rests on the fact that the generic characters seem

more or less artificial or arbitrary, and may include species which

have independently come to have certain peculiarities. Thus, Rhi-

zococcus is essentially an Eriococcus without a sac; but we do not

know that the sac might not be independently lost in different parts

of the world. The only way to settle these matters is by the study

of all stages and both sexes of numerous species.

(3-) Genera having few species very widely separated.

Tessarobelus, 2. One in New Caledonia, one Panama. Prob-

ably they are not strictly congeneric.

No. 443.] NOTES AND LITERATURE. | 803

Llaveia, 7. Neotropical except one from the East Indies, which

may not be properly referred here.

Stigmacoccus, one in Brazil, one in India.

Ortheziola, 3. Two European; one in West Indies, but I sus-

pect introduced. '

Conchaspis, 3. Neotropical and Ceylon. No doubt spread by

man.

Cerococcus, three North American, one in India.

Solenophora, 9. North and South America and New Zealand,

in each case doubtless native.

Gossyparia, s. Australia, New Zealand, Europe. Perhaps not

all congenefic. :

Erium, 7. Australia and America. _

Rhizzcus, 4. Europe; one in West Indies, perhaps introduced.

Ripersiella, 4. Three in North America, one in New Zealand.

Takahashia, one in Mexico, one in Japan.

Protopulvinaria, one Neotropical, and Mr. Green has an unde-

scribed one in Ceylon. I suspect that the Neotropical one

was introduced from Asia.

Mallococcus, one in Brazil, one in China.

Ctenochiton, 15. New Zealand, Australia, and one in Mexico.

Cardiococcus, two in Australia, one in Mexico.

Inglisia, 8. New Zealand, Asia, Neotropical.

Ceroplastodes, s. North America, Australia, Asia.

Akermes, 9. Neotropical and Australia. Some are probably

not congeneric.

Paralecanium, 7. Australia, Asia, one in Brazil. :

Diaspis, 30. America, and several Old World species which

may not be strictly congeneric.

Poliaspis, 8. Australia, S. Africa, Japan.

Leucaspis, 8. Europe, Australia, Japan, Western

'The species are not all congeneric. ;

Cryptophyllaspis, 4. One in the United States, one in. Ceylon,

one in the Bismarck Archipelago, and one (accidentally

omitted from the list) in Madeira. I think that these are

probably not all really congeneric. :

Odonaspis, s. "Tropical Asia; one in Brazi

I think.

Gymnaspis, 3.

[ believe introduced in the latter.

Some of the above cases should afford satisfaction t

America.

|, surely introduced,

Asia, Australia and the Neotropical region;

o believers in

804 THE AMERICAN NATURALIST. [Vor. XXXVII.

Antarctica. It is to be remarked, however, that some of them may

appear in different light when we know more of the Coccidz of

tropical Asia. Nevertheless, it is hard not to think Ripersiella,

Akermes, Ctenochiton etc. significant of some southern route.

(4.) Holarctic Genera.

I use the term Holarctic as a convenience, but do not mean there-

by to abandon the Nearctic and Palzarctic as separate regions.

Xylococcus, 3.

Kermes, 28. On oaks. A species described from Australia

can hardly be congeneric.

Trionymus, 2. Exeretopus, 2.

Antonina, 7 (also China). Eriopeltis, 3.

Eulecanium, 71 (one in Brazil). Physokermes, 3.

Lecanopsis, 3. Aclerda, 7 (and Natal, one species

doubtfully congeneric).

(5.) Nearctic Genera.

Olliffiella, 1 (making a gall on oak). | Gymnococcus, 3.

Pseudophilippia, 1 (Florida). Philephedra, 1 (New Mexico).

'Toumeyella, 6. Comstockiella, 1 (on palms).

Pseudodiaspis, 2 (one in Mexico).

These are all southern genera, from the very borders of the Neo-

tropical, or evidently derived therefrom. "There is no characteristic

Nearctic genus, generally distributed throughout that region, and

not found elsewhere.

(6.) Neotropical Genera.

Porococcus, 2 (Mexico). Dactylopius, 3 (one enter-

: ing Nearctic).

Tectococcus, 1. Carpochloroides. 1.

Apiococcus, 4. Capulinia, 3.

Cryptokermes, 1. Termitococcus, 2.

Pulvinella, 1. Tectopulvinaria, 1.

Alichtensia, 1. Edwallia, 1.

Platinglisia, 1. Schizochlamidia, 1.

Pseudokermes, 2. Eucalymnatus, 4 (one may

Stictolecanium, 1. be native in Asia).

Mesolecanium, 12 (enters Sonoran). Megasaissetia, 1.

Neolecanium, 12 (enters Nearctic). Platysaissetia, 1.

Protodiaspis, 1. Xanthophthalma, 1.

No. 443.j NOTES AND LITERATURE. 805

Pinnaspis, 2 (also Old World, whence Morganella, 2 (widespread

probably introduced). - by human means).

Pseudischnaspis, 3. Pseudoparlatoria, 4.

Diaspidistis, 1.

It will be noticed that many of these are monotypic ; representing,

no doubt, particular excessively differentiated types, not illustrative

of any general tendency. Pinnaspis is so like the Old World Hemi-

chionaspis that I feel nearly sure that it is introduced into America.

One of its species is now widespread in both hemispheres.

(7.) New Zealand Genera.

Ccelostomidia, 5. Phenacoleachia, 1.

Eriochiton, 3 (one from India). Lecanochiton, 2.

Three of these appear to be genuinely isolated and very peculiar

types.

(8.) Australian Genera.

Frenchia, 2.

Apiomorpha, 34.

Opisthoscelis, 12.

Ascelis, 5.

Olliffia, 1.

Ourococcus, 3.

Lachnodius, 3.

Austrolichtensia, r.

Monophlebulus, 1.

Callipappus, 6.

Antecerococcus, 2.

Birchippia, 1.

Cylindrococcus, 4.

Sphzrococcopsis, 1.

Epicoccus, 1.

Pseudoripersia, 1. — .

Myxolecanium 1 (New Guinea).

Alecanopsis, 1.

Here we have a genuinely peculiar fauna, the gall making genera

Maskellia is the only Diaspine genus

being especially remarkable.

he neotropical list includes

in the list, it will be noted, whereas t

seven genera of this group.

(9. Genera of tropical Asia.

cies, doubtfully congeneric, are African).

Anomalococcus, 1.

Amorphococcus, 1.

Geococcus, 1.

Kermicus, 1.

Pseudopulvinaria, 1.

Vinsonia, t.

Monophlebus, 9 (three spe

Drosicha, 6 (also Australasia).

Walkeriana, 7 (also African).

Kuwania, 2 (one in Japan).

Cheetococcus, 1.

Ceronema, 3 (Australia to Japan).

Ericerus, 1

Howardia, 1 (original locality uncertain).

806 THE AMERICAN NATURALIST. [Vor. XXXVII.

Hemichionaspis, 1o (also Africa, etc).

Fiorinia, 25 (extended to Australia and New Zealand).

Pseudaonidia, 6 (also African). Aonidia, 12 (one is Palzarctic).

Greeniella, 1. Ischnaspis, 1 (origin uncertain).

It will be noticed that the faunz of tropical Asia and Africa have

elements in common, as might have been expected.

(10.) Genera of tropical Africa.

Lophococcus, 1. Cissococcus, r.

Halimococcus, 1 (also tropical Asia, undescribed species in Mr.

Green's possession).

Tylococcus, 1 (Madagascar). Lagosinia, 1.

Gascardia, 1 (Madagascar). Cryptinglisia, 1.

Stictococcus, 1. Selenaspidus, 1 (origin uncertain).

Some of these are very remarkable. The Coccid fauna of Africa

is as yet very little known, and it doubtless contains many wonderful

things.

(11.) Palearctic genera.

Gueriniella, 1. Pollinia, 2 (one is S. African.)

Newsteadia, 1. Nidularia, 1.

Pheenicococcus, 1 (Algeria, an offshoot from the Ethiopian fauna).

Puto, 1. Tetruüra, t.

Fonscolombia, 2. Cryptococcus, 1

Kuwanina, 1 (Japan). Spermococcus, r.

Luzulaspis, 1. Filippia, 1.

Chelonicoccus, 1 (doubtful genus). ^ Parafairmairia, 1.

Aulacaspis, 6 (extends to tropical Asia, etc.).

Parlatoria, 17 (extends to tropical Asia, etc.).

Syngenaspis, 1.

Epidiaspis, 1.

It would seem that Europe has more peculiar types than North

America.

T. D. A. COCKERELL.

Another Text-book of Entomology.' -*'To induce the student

to become acquainted, through personal observations in the field and

laboratory, with some of the important biological problems as pre-

‘Hunter, S. J. Elementary Studies in Insect. Life. Topeka, Kan. Crane

and Co., 1903. 12mo, xviii + 344 pp., 234 figs.

No. 443.] NOTES AND LITERATURE. 807

sented by insects,” is stated to be the object of this book. It aims

to present a course in study of insects, prepared from the biological

. point of view. Hence, the opening chapters deal with such general

themes as metamorphosis, senses and sense organs, coloration, social

life, habits and instincts, and the relations between flowers and

insects. Discussion of these themes occupies 119 pages. Then fol-

lows a chapter of 43 pages on economic entomology pure and simple,

and another of 39 pages on systematic entomology. In the latter

the orders are briefly discussed, with scant reference to either their

metamorphoses or other biologically interesting characteristics.

Another brief and heterogeneous chapter of 20 pages discusses

geographic distribution, the struggle for existence, parasitism, natu-

ral selection etc. As an academic discussion of the biology of

insects this part is singularly lacking in unity and coherence.

Part II is devoted to methods, equipment and laboratory exercises.

There are outlines for practical study of but two of the biological

themes discussed in part I: metamorphosis, and the habits of ants.

For the balance, there is but another outline for the study of grass-

hopper anatomy, and another key to the orders and principal families

of insects. ‘There are many illustrations — far too many in fact in

herein each part is figured for the student;

the laboratory outline, w

they

and while they appear to have been made from good originals,

are not well reproduced, and too often appear inky.

In so far as the study of the biological aspects of insect life is con-

cerned the profession is better than the performance. There is

considerable rambling discussion of biological themes (and this

appears to be all that is offered as new in the method of the book)

but, with the two exceptions noted above, there is no program set

before the student for the study of them. The lapse into anatomy

and the making of systematic determinations for the student's train-

ing is the more regrettable because there were many better laboratory

outlines for grasshopper dissection, and a few better keys, already in

existence. It may be, however, that the thin strata of insect biology,

anatomy, collection making, systematic determination and economic

procedure brought within the covers of a single book will afford light

digging for short course students.

The chief criticism of it as a laboratory book is that it is pedagogi-

cally weakest in those matters on which it lays greatest stress. In

this respect it is much like some other modern books of botany and

zoólogy, which introduce ecological and biological phenomena, and

get no farther than mere academic discussion of them. This endan-

808 THE AMERICAN NATURALIST. | [Vor. XXXVII.

gers a relapse to text book methods, and neglect of the first hand

study of the facts. Is it too much to expect of teachers who empha-

size the value of these subjects that they should guide the students

in the gathering, correlating and interpreting of biological data by

definite and practical methods that will insure the least waste of

time and energy? Shall we never get rid of thé spectacle of a

teacher and a score of students trying to peer at once into the same

worm hole? Shall we be forever content with merely demonstrating

biological phenomena ?

TJG N

Two Papers on Insect Wings.' — A remarkable American silk-

worm moth (Zz/ea polyphemus) with its right wings deviating from

the normal in both color pattern and venation, furnishes to Dr. G.

Enderlein a theme for the discussion of ontogenetic processes.

The important part of the paper is that embodying the results of a

search for ontogenetic processes in the light of which the aberration

might be explained. These remarks from the introduction are inter-

esting and set forth the author's point of view: “Experimental

investigations serve only as a means of varying ontogenetic processes

and also, therefore, of influencing, hindering or varying the com-

bined sum of inner mechanical forces by means of the outer mechan-

ical forces, in order to advance a little by means of the knowledge

of their reaction to these forces. By such means no new forms will

be brought to light ; for no noteworthy, sudden result can be induced

in phylogeny by the operation of external influences: phylogeny is

but the slowly progressing result of summarization; so we would

have to expect only phylogenetic forms which still exist somewhere

in ontogeny. Retarded developments (Hemmungsbildung gen), which

we may designate as reversions, are of equivalent value, whether we

think of the anlage in the egg or of a later outcome of ontogenetic

processes. But it should not be understood that all reversions have

existed as actual species, for there may be manifold. combinations of

phylogenetically younger and older component factors."

So the author sets about a very careful investigation of the struc-

ture of Saturnian and other Lepidopterous pupa. Incidentally he

makes some contributions to the knowledge of the nervous system

! Enderlein, G. Eine einseitige e aS bei Telea polyphemus vom

ontogenetischen Standpunkt: ein beitrag zur Kenntniss der peisea der

Schmetterlinge. Zool. Jahrb., Abt. für pole u. Ontogenie der Thiere. Vol.

1902, pp. 44, 3 pls.

No. 443.] NOTES AND LITERATURE. 809

and digestive and reproductive organs, but his main purpose and his

best results are in the study of the pupal tracheal system, and in the

relation of the same to the venation of the adult wing. Most of his

conclusions are abundantly evidenced, though some of them are not

entirely new: but one of them at least — his two systems of trachez

and veins, radial and median— is of very doubtful value: it rests

on altogether insufficient evidence. Lepidoptera alone with their

single pair of longitudinal trachea trunks, are too highly specialized

to show what was the primitive manner of grouping. In the light of

facts presented by other more generalized orders — facts that are set

forth in a paper that the author repeatedly cites ! — such grouping

seems little less arbitrary than that of Spuler (into * spreitentheil "

and *faltentheil ”) which Dr. Enderlein justly condemns.

He finds in the end that his aberrant moth represents in the con-

figuration of its venation from the ontogenetic point of view a pupal

stage preserved in adult life: from the phylogenetic point of view, a

one-sided reversion to a phylogenetically earlier stage, that is now

normal to no living Saturnian moth.

The second and more recent paper is by Tower,’ who makes a

large and timely contribution to the knowledge of the development

of the wings in Coleoptera. He calls attention to the great diversity

of larval form and habitat and habits, and tabulates the differences

in number of larval stages, length of larval life and time of first

appearance of larval wings, and then discusses in detail the three

types of early wing formation he finds in the order. His “simple

type,” which he finds to be “the dominant type of wing development

in beetles" is most interesting, because it bridges the gap in type of

wing development between the holo- and the hetero-metabola.

During the last larval stage the wing is directly evaginated down-

ward underneath the cuticle and is merely : uncovered by the last

moult. His “recessed type” in which the wing rudiment withdraws

slightly from the surface to the bottom of a hypodermal pocket, as in

Corethra, and his * enclosed type," in which the wing withdraws and

is shut in by the closure of the pocket, as in Lepidoptera generally,

are in beetles much less common.

He concludes that “the wings and spiracles arise in homologous

! Comstock and Needham, The wings of Insects. Amer. Nat., vols. 32 and

3» LA PE» . * EJ

n and development of the wings m Coleoptera.

2 Tower, W. L. The origi

Zool. Jahrb. Abt. für. Anat. u. Ontogeny der Thiere. Vol. t7, 1903, pp- 516-572,

Pls. 14-20.

810 THE AMERICAN NATURALIST. |. (Vor. XXXVII.

positions on the sides of the. segments, as determined by the position

of homodynamous muscles, and that the hind wings are derived

without much doubt from the degenerate spiracle disc of the meta-

thorax”: and adds, * All the evidence here presented concerning the

wings of Coleoptera and Hetermometabola is most positively opposed

to the theory of the origin of the wings of insects as dorsal backward

prolongations of the tergum."

'There is little new ground broken in this paper, but there is much

more careful tillage of a hitherto indifferently cultivated field.

G.N.

BOTANY.

A New Flora of the Southeastern United States. — A new

handbook of the flora of this region has long been a desideratum,

the data concerning the southern states being confined to antiquated

books, the latest of which, Chapman’s Fira, is a decade old, and to

scattered descriptions, collector's notes, and isolated specimens,

chiefly in the larger herbaria,— apart from one of which they could

hardly have been brought together satisfactorily. Dr. Small, who

is curator of the museums and herbarium of the New York Botani-

cal Garden, and who has personally made extensive trips over a con-

siderable part of the territory covered by the new Flora, consulting

Elliott’s herbarium at Charleston and the original Chapman herba-

rium at the New York Garden, is in an unequaled position to under-

take the preparation of such a handbook, and though his own

experience in the several years during which it has been going

through the press shows the rapidity with which such works call for

change when once launched, he has succeeded in placing before the

public a manual which, if of awkward form and size for field use, is

indispensable to every herbarium and botanical library in which the

North American flora receives attention.

! Small, J. K., Flora of the Southeastern United States being descriptions of the

Seed-plants, ferns, and fern-allies growing naturally in North Carolina, I

arolina, Georgia, Florida, irrita Alabama, Mississippi, Arkansas, Louisi

and the Indian Territory and in Oklahoma and Texas east of the One hu caveat

meridian. New York, published by the author, 1903. 8vo, xii + 1370 pp.

No. 443.] NOTES AND LITERATURE. 811

In all respects the treatment of the subject is modern. The phylo-

genetic arrangement of Engler and Prantl is essentially adopted; the

Neo-American practice in nomenclature is followed as consistently as

it is likely to be followed,— familiar generic names being added as

synonyms when discarded under this practice; and genera and spe-

cies have been subjected to the prevalent minute segregation. Opin-

ions may and do differ greatly as to the desirability of some of these

practices, and in the last-named respect Dr. Small is scarcely sur-

passed by any living botanist on this side of the ocean: but by what

appear to be well-made keys and terse contrasted descriptions he

makes reasonably clear his idea of the species that he names,— 6 364

in number, grouped under 1494 genera, pertaining to 236 families, of

62 orders.

Notes.— An unusually satisfactory series of plates showing autum-

nal coloration of foliage is contained in a paper on “ Tree planting on

Streets and Highways,” by W. F. Fox, published at Albany, by the

Forest, Fish and Game Commission of New York.

A capital treatise on woody plants in winter, illustrated by numer-

ous habit, bark and detail figures, by Schneider, has been issued

from the press of Gustav Fischer, Jena, under the title Dendrolo-

gische Winterstudien.

An account of the characteristics of some southern trees, by Emma

G. Cummings, is contained in Part I of the Zransactions of the

Massachusetts Horticuitural Society for 1903.

“A Primer of Forestry,” by Pinchot, is published as Farmers’

Bulletin No: 173 of the Departme tof Agriculture.

The value of oak leaves for forage is discussed by Mackie in

Bulletin No. 150 of the Agricultural Experiment Station of the

University of California.

A series of practical little Bulletins, somewhat comparable with

those issued by the Experiment Stations in this country, is being

distributed by the Biologische Abtheilung fir Land- und Forstwirth-

schaft, of the Kaiserlches Gesundheitsamt, of Berlin, under the title

“ Fligblatter.”

The forage conditions and problems of Eastern Washington, East

ern Oregon, Northeastern California and Northwestern Nevada are

812 THE AMERICAN NATURALIST. [Vor. XXXVII.

discussed by Griffiths in Buletin 38 of the Bureau of Plant Industry

of the United States Department of Agriculture.

The Yearbook of the United States Department of Agriculture, for

1902, recently issued, contains the usual variety of papers, primarily

of economic interest but a number of them botanically valuable.

* Loco and other poisonous plants of Montana" are discussed by

Blankinship in Bulletin No. 45 of the Montana Agricultural Expert-

ment Station.

The cultivation of sisal in Hawaii is the subject of a paper by

Conter, published, with illustrations, as Bulletin No. 4 of the Hawaii

Agricultural Experiment Station.

A very attractively gotten-up account of Luther Burbank and his

work in horticulture, by Wickson, has been reprinted from the Sunset

Magazine by the Southern Pacific Company of San Francisco.

The new ideals in the improvement of plants are discussed by

Bailey in an illustrated article in Country Life in America for July.

A comparison of hybrids with their parent forms, by de Vries, is

published in the Revue Générale de Botanigue, of June 15.

An article on plants as a factor in home adornment, by Corbett, is

reprinted from the Yearbook of the Department of Agriculture for

1902.

No accounts of school gardens, published in this country, are more

interesting or better illustrated than those contained in current vol-

umes of the Transactions of the Massachusetts Horticultural Societ y.

Dr. Grout, who a few years since wrote a little guide to the study

of mosses with the aid of a hand-lens only, has issued the first part

of a well printed and nicely illustrated book of larger scope under

the title Mosses with Hand-Lens and Microscope. His purpose is to

present a handbook of the more common mosses of the Northeastern

United States with the avoidance of unnecessary technicalities.

A new list of the “ Pteridophytes of Iowa," by Lyon, reprinted on

July 3rd from Minnesota Botanical Studies, contains the interesting

information that gametophytes of Botrychium obliquum and B. vir-

ginianum have been collected in that State, both species at Echo

Lake, and the last named also at Grand Marais.

As Part 3 of the current volume of Contributions from the United

No. 443] NOTES AND LITERATURE. 813

States National Herbarium, Mr. Maxon publishes a study of certain

Mexican and Guatemalan species of Polypodium.

A monograph of the Belgian species of Cladonia, by Agriet, con-

stitutes the third fascicle of Volume 4o of the Buletin de la Société

Royale de Botanique de Belgique, for the year r9or, issued in June

1903.

The bitter rot of Apples forms the subject of a paper by von

Schrenk and Spaulding, published as Bulletin 44 of the Bureau of

Plant Industry of the Department of Agriculture. The fungus com-

monly known as Glocosporium fructigenum, but in one of its forms

first named Septoria rufo-maculans by Berkeley, is here named G/o-

merella rufomaculans, the genus standing practically for Gnomoniop-

sis of Stoneman, but not of Berlese.

Diseases of the apple, pear, and quince are discussed in Bulletin

183 of the North Carolina Agricultural Experiment Station.

The Journal of the Royal Horticultural Society for April, as is usual

with that journal, contains a good many botanically interesting mat-

ters, among others a continuation of Cooke's * Fungoid pests of the

garden."

Two new diseases of the raspberry, cane blight and yellows, are

discussed in Popular Bulletin no. 226 of the Mew York Agricultural

Experiment Station, issued last December but dated December 1903.

Like preceding numbers, Professor Peck’s Report of the State

Botanist for 1902, published as Buletin 286 of the University of the

State of New York and issued in May, contains descriptions and fig-

ures of a considerable number of pileate fungi.

A helpful feature of the Ohio Mycological Bulletin consists in the

printing of accent marks over generic and specific names, — but

unfortunately the popular rather than the correct accentuation is

occasionally given.

A phalloid (probably /thyphadlus celebicus) is described by Fischer

in Mededeelingen van het Prafstation Oost-Java, 111, No. 46, as liv-

ing in close symbiosis or scarcely injurious parasitism on the roots of

the sugar cane, in Java.

Monascus purpureus and its systematic position, are considered, by

Ikeno, in the Berichte der Deutschen Botanischen n of June

24.

814 THE AMERICAN NATURALIST. [Vor. XXXVII.

In the Proceedings of the Academy of Natural Sciences of Philadel-

phia, LV, Part I, Keeley gives directions for the preparation of Dia-

toms for microscopical examination of their structure.

The American Botanist for June contains the following articles, all

popular : — Bradshaw, “Poppies”; Stillman, “A climbing Fern";

Barrett, “Odd Odors”; Dallas, “ Hints for Beginners in the Study

of Mushrooms”; [Clute], “ Botany for Beginners — III”; Field,

* New Jersey Tea”; and, Goetting, * Children's Names for Flowers."

The Atlantic Slope Naturalist, a new, popular, little journal, con-

tains in the July-August number a short note by Bessey on the “ Dis-

tribution of Forest Trees on the Nebraska Plains," and a note by

Rotzell on “The Smoking of Red-Willow Bark by the American

Aborigines."

. The botanical Bzhang till Kongi. Svenska Vetenskaps- ARademiens

Handlingar, Volume 28, recently published, contains a number of

important papers in various fields of botany.

The Aryologist for July contains the following articles: — Harris,

“Lichens — Sticta”; E. G. Britton, “West Indian Mosses in

Florida”; Renauld, “ Hypnum capillifolium Baileyi" ; Williams,

* Additional Mosses of the Upper Yukon River"; Grout, “The

Peristome, V"; and a continuation of reprints of Cardot and

Thériot’s * Mosses of Alaska” from the Proceedings of the Washing-

ton Academy of Sciences.

The Bulletin of the Southern California Academy of Sciences for

April-May contains the following botanical articles: Abrams, “ New

Southern California Plants”; Davidson, “New plant records for

Los Angeles County, —II.; Hasse, “ Additions to the Lichen-Flora

of Southern California”; Hasse, The Lichen-flora of San Clemente

Island”; and Hasse, “ Additions to the Lichen-flora of Southern

California, — II.”

The Bulletin of the Southern California Academy of Sciences for

June contains the following botanical articles : — Heller, “ Notes on

Plants from Middle Western California”; Davidson, “New Plant

Records for Los Angeles County, — III.” ; and Hasse, “ Additions

to the Lichen-flora of Southern California, — III.”

The Bulletin of the Torrey Botanical Club for July contains the

following articles — Nash, “A preliminary Enumeration of the

Grasses of Porto Rico"; True and Gies, “On the Physiological

No. 443.] NOTES AND LITERATURE. 815

Action of some of the Heavy Metals in Mixed Solutions" ; Rennert,

“The Phyllodes of Oxypolis filiformis, a swamp Xerophyte te: gad,

Fink, “ Some common Types of Lichen Formations.”

The Canadian Record of Science, Volume 9, No. 1, contains a sum-

mary of the “ Progress of Botany in the 19th Century,” by Campbell,

and a note by Emberson on the “Trees of Montreal Island.”

` The Journal of the New York Botanical Garden for July contains

an account by Professor Underwood of explorations in Jamaica and

an account of the Tree-fern house of the New York Garden.

The Plant World for July contains the following articles : — Saf-

ford, “Extracts from the Note-Book of a Naturalist on the Island of

Guam. — VIII"; Barrett, “ [Size of flowering Furcrza] " ; Waters,

“The resting period of Plants”; Schneck, “The Cross-bearing

Bignonia or Cross Vine”; George, “The Preservation of Native

Plants”; Bailey, “Plants of Universal Application "; Harper, “ The

Water Hyacinth in Georgia"; and Waters, [ Tipularia ]."

Rhodora for July contains the following articles: — Fernald,

“Chrysanthemum leucanthemum and the American White Weed” ;

Sargent, “Recently Recognized Species of Crataegus in Eastern

Canada and New England, — V.”; Robinson, * Preliminary Lists

of New England Plants,— XII. [Eriocaulacez, Phytolaccacez, Aizoa-

cee, Portulacacez, Caryophyllacez, Illecebraceze and Sarracenia-

cez]"; a short biographical sketch of Henry Griswold Jesup;

Flynn, “ A second Station for Cyperus Houghtoni in Vermont”; and

Morss, “ Clematis verticillaris in the Middlesex Fells."

contains the following articles: — Lloyd, “A

". Nash, “A new Aletris from

dovicas from the island. of St.

Torreya, for July,

new and cheap form of Auxanometer

Florida”; Cowell, “Two new Carlu

Kitts, W. I.”; Flynn, * Plants new to Vermont, found in Burlington

and vicinity”; Britton, “ A new Lippia from Porto Rico, and A new

Waltheria from the Bahamas”; and Harper, “ Elliottia racemosa

again.”

lementary text-books one of the best

r Round, published by the American

ell written book

Among the many recent e

is Andrews’ Botany all the Yea

Book Company, a well illustrated, well devised and w

for the secondary schools.

Volumes 2 and 6 of Asche

europäischen Flora are being currently published

numbered.

rson and Grebner’s Synopsis der Mittel-

in parts serially

816 THE AMERICAN NATURALIST. [Vor. XXXVII.

An analytical account of the higher groups, families and genera of

Mexican plants, by Conzatti, is in course of publication by the

Secretaria de Fomento of the City of Mexico, under the title “ Los

géneros vegetales mexicanos."

A dictionary catalogue, with annotation and illustrations, of the

economic plants of Porto Rico, by Cook and Collins, forms Part 2 of

the current volume of Contributions from the United States National

Herbarium.

A brief popular account of * Vegetation in Greenland,” with illus-

trations from herbarium material, is prre by W. E. Meehan in

Floral Life for July.

Part III of Cooke’s “Flora of the Presidency of Bombay,” com-

pleting the first volume, extends through the order Rubiacez.

As is customary with the Proceedings of the Linnean Society of New

South Wales, Part IV of Volume 27, recently issued, contains a

number of interesting papers referring to the Flora of Australia.

Volume LXII of the Natuurkundig Tijdschrift voor Nederlandsch

Zndié contains important papers by Koorders on the botany of the

Dutch Indies.

The result of evolutionary and physiological investigation of the

physiological róle of mineral nutrients in plants, by Loew, is pub-

lished as Bulletin 45 of the Bureau of Plant Industry, U. S. Depart-

ment of Agriculture.

A summary, by Pond, of MacDougal’s Influence of Light and

Darkness upon Growth and Development, is separately printed

from the Monthly Weather Review for April.

An extensive paper by Eberhardt on the respective influence of

dry and moist air on the form and structure of plants, is contained in

the Annales des Sciences Naturelles, Volume 18, Nos. 1—3, of the cur-

rent series.

The conclusion of Schulz’s monograph of the genus Cardamine is

contained in Engler’s Botanische Jahrbiicher, Volume 32, Heft 4.

Papers on Rubus and Cratzgus, by Ashe, constitute the larger

portion of Part I of the current volume of the /ournal of the

Elisha Mitchell Scientific Society:

An Arceuthobium of Tsuga in the Northwest is named Razou-

No. 443.] NOTES AND LITERATURE. 817

mofskya tsugensis by Rosendahl in Minnesota Botanical Studies of

July 3, 1903.

A brief economic article on Agaves, by Nelson, is reprinted from

the Yearbook of the Department of Agriculture for 1902.

Guerin publishes, in the Journal ad’ Agriculture Tropicale of June

30, an interesting account of the oil palms of Guatemala known as

Corozos, those of the Atlantic slope referred to .4//a/ea cohune and

those of the Pacific slope to EZezs melanocarpa.

An account of Japanese bamboos and their introduction into

America, well illustrated from photographs, is published by Fairchild

as Bulletin 43 of the Bureau of Plant Industry of the United States

Department of Agriculture.

A morphological and anatomical study of a hybrid of Agropyrum

violaceum and Elymus arenarius, by Gallee, is published in Botanisk

Tidsskrift, Volume 25, Heft 2.

No. 19 of Holm’s “Studies in the Cyperacez," dealing with the

genus Carex in Colorado, is published in the American Journal of

Science for July.

Biographic sketches of Bescherelle, giving a list of his publications,

are contained in No. 3-4 of the current volume of Buletin de la

Société Botanique de France.

A short biographical sketch of Micheli, with protrait, is published

in No. 8s of the Actes de la Société Helvétique des Sciences Naturelles.

CORRESPONDENCE.

To the Editor of the American Naturalist :

Sir: At the May meeting, this year, of the Philadelphia Acad-

emy of Natural Sciences, Miss Sarah P. Monks read a note on the

* Regeneration of the Body of a Starfish.” The brevity of the com-

munication and the inconspicuous place given it in the published

Proceedings of the meeting are not calculated to assure it the atten-

tion it deserves.

I quote from the report: “In studying regeneration in Phataria

(Linckia) fascialis she had cut arms at different distances from the

disk, and a number of the single rays produced new bodies. The

free ray made a new body and the rest of the starfish produced a

new ray .... In the photograph of a six-rayed Phataria, the cut ray

attached to the body shows a small ray sprouting, while the free ray

shows four new rays. This was cut July, 1902, and the photograph

taken February, 1903." EST

Miss Monks is to be congratulated on having at last produced

the experimental evidence demanded by the skepticism of recent

writers on the soundness of Haeckel's conclusion! reached long ago

that “jeder abgeloste Arm [of certain starfishes] reproducirt die

ganze Scheibe nebst den übrigen Armen."

I have been permitted by Miss Monks to examine all her speci-

mens bearing upon this subject, and have followed her experiments

with much interest and deem it but justice to her to say that in

reality she has the data for a considerably fuller presentation of the

question than would appear from the meager report which has

elicited these comments. It is to be sincerely hoped that a fuller,

well illustrated account of her observations may be published before

long.

WiLLIAM E. RITTER.

University of California,

Aug. 22, 1903.

! Die Kometenform der Seesterne und der Generationswechsel der Echinoder-

men. ZeitscA. wiss. zool., Bd. 30, 1878, p. 424.

818

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THE

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NATURALIST

A MONTHLY JOURNAL

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CONTENTS

Page

I, Adaptations to Aquatic, Arboreal, Fossorial and Cursorial Habits in Mam-

mals. III. Fossorial Adaptations ` H. W. SHIMER 819

IL On the Structure of the Pteraspidæ and baai.

PROFESSOR W. PATTEN 827

IIL An Out-Door Equipment for College Work in Biology

PROFESSOR J.G. NEEDHAM 867

IV. *Preliminary Report on the ‘‘Palolo” Worm of Samoa, Eunice viridis (Gray)

W, McM. WOODWORTH 875

V. Further Notes on the Habits of Autodax lugubris

PROFESSOR W. E. RITTER 883

VI. A Trip to the Truchas Peaks, New Mexico WILMATTE P.COCKERELL 887

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THE

AMERICAN NATURALIST.

VoL. XXXVII. December, 1903. No. 444.

ADAPTATIONS TO AQUATIC, ARBOREAL, FOS-

SORIAL AND CURSORIAL HABITS

IN MAMMALS.

III. FossoRiAL ADAPTATIONS.

H. W. SHIMER.

Tux purpose of the following article is to summarize a few of

the principal modifications in external shape and in the skeleton

independently acquired by mammals in different orders which

have become wholly or partially adapted to a life beneath the

surface of the ground. The highest specialization in this direc-

d, as we should expect, in those forms which secure

not only safety but also their food within the earth (e. g. the

moles). Such forms are completely fossorial. On the other

hand the procuring of food above ground and the use of the

burrow merely as a safe place in which to live and rear young

requires fewer fossorial modifications, and such forms may be

called semi-fossorial, although, naturally, between fossorial and

semi-fossorial no fixed line can be drawn. The following charac-

ters are best seen in the truly fossorial forms.

819

tion is foun

820 THE AMERICAN NATURALIST. [Vor. XXXVII.

A. EXTERNAL MODIFICATIONS.

I. Body more or less fusiform.— An obvious adaptation to

progression in such a dense medium as earth. In the common

mole (Condylura), for example, the body-diameter is greatest at

the shoulder, and diminishes gradually to a point at the nose.

In some fossorial forms, e. g. the wombat (Phascolomys) and

woodchuck (Arctomys), the body is very thick.

2. Eyes imperfectly developed or obsolete — Normally devel-

oped eyes are traditionally useless to an animal living in com-

plete darkness, and would be a continuous source of pain from

injury received in burrowing. The degree of degeneration

is no doubt partly dependent on the length of time which

has elapsed since the assumption of fossoria] habits, and on the

relative completeness of withdrawal from the upper air. In

the pocket gophers (Geomyidz) and Bathyergide the eyes are

small; in Spalax typhlus they are mere black specks among the

muscles (although retaining a relatively complete structure); in

the marsupial mole (Notoryctes typhlops) they are imperfectly

developed and functionless ; in Talpa they are vestigial; in the

Cape golden mole (Chrysochloris) the eyes are covered with

skin.

3. External ears, small, tending to disappear — External ears

impede burrowing especially as they are situated at the upper

and anterior part of the body where much friction would nat-

urally occur. Hence in the Geomyidz and in Lutra the exter-

nal ears are small, in the ratel (Mellivora) very minute and in

the Bathyergida they have become reduced to a mere ring

of skin around the auditory aperture, while in Notoryctes,

Chrysochloris and Talpa they are absent.

4. Limbs short and stout.— Since in a truly fossorial animal

strength to dig is of more importance than swiftness of progres-

sion on the surface of the ground the limbs are short and stout

(Ornithorhynchus, Echidna, Talpa, etc.). This of course does

not hold good for the majority of semi-fossorial forms, as in their

life above ground they need speed either to get food or to

escape enemies. Many of these, however, as our common wood-

No. 444.] HABITS IN MAMMALS S21

chuck (Arcfomys monax) and the wombat (Phascolomys), have

legs quite short and stout; while for example in the pig-footed

bandicoot (Cheropus castanotis) the legs are long and cursorial

in type. In the latter animal the external ears are also very

long.

5. Manus broad and stout, with long claws— The fore feet

and hind feet have undergone divergent specialization, since the

fore feet are used to loosen the earth, while the hind ones merely

throw it further backward. Hence the manus becomes broad and

stout with very strong claws. Inthe common mole (Condylura)

the manus is as broad as the whole body, and hence it can dig

with exceeding rapidity, excavating with one sweep of the arm

a place wide enough for entrance. In Echidna also the manus

is broad. The enlarged strong claws are well seen in Notoryctes,

Phascolomys, Taxidea and the Geomyide.

6. Pes modified to throw the loose earth backward— The

pes has the claws well developed although not nearly so strongly

as in the manus. Some animals have developed special adap-

tations for throwing back the loose earth. In Phascolomys the

second, third, and fourth digits are partly connected by skin.

This web is of course strongly developed in such swimming

forms as Chironectes and Scalops but it is also a great aid to

them in burrowing. Heterocephalus has the feet fringed with

stiff hairs, while the Octodontidz have long stiff hairs at the

roots of the claws. The hallux is at times vestigial as in

Phascolomys.

7. Tail usually short.— A tail seems to be a useless append-

age to an animal surrounded closely by earth so it becomes

quite short even in many semi-fossorial forms as in the hare,

ratel, and woodchuck. In the wombat, moles, etc., it is ves-

tigial. Oryzoryctes is an exception, for although fossorial it

has a comparatively long tail.

B. SKELETAL MODIFICATIONS.

I. Skull in top view triangular, apex forward.— The sub-

conic form of the skull is one of the obvious adaptations to pro-

gression through a dense medium. It is very well seen in

Condylura, Crossopus fodiens, etc.

$22 THE AMERICAN NATURALIST. [Vor. XXXVII.

2. Zygomatic arches not extended outside the greatest width

of the skull— All projections from the usual regularly shaped

skull become reduced. The zygomatic arches are very slender

(e. g. Condylura, Talpa, Erinaceus, Orycteropus).

3. Prenasal ossicle.— A small prenasal ossicle is developed

at the tip of the mesethmoid cartilage in Talpa as it is in the

pig (Sus), due doubtless to the same cause, that of using the

nose as an aid in digging.

4. Inctsor teeth chisel-shaped, projecting forward. — In some

forms the large incisors serve to keep earth out of the mouth,

in others they are used as an aid in digging. The wombat

(Phascolomys) for example has a pair of strong chisel-shaped

projecting lower incisors. Among the rodents Spalax typhlus

for example has the well developed lower incisors projecting

beyond the lower jaw; the bamboo rat (Rhizomys) uses its

incisors to aid in digging, in the Bathyergidæ the entrance of

earth is prevented by the upper incisors which extend in front

of the closed lips.

5. Cervical and lumbar vertebre more or less fused.— The

fusion of these vertebræ gives strength and firmness in pushing

(e. g. Notoryctes and the armadillos). It is possible that the

peculiar intercentral ossicles observed below the lumbar verte-

bree especially in Talpa and the Erinaceidæ may be of use

in strengthening the spinal column. Phascolomys and the

Dasypodidæ have the cervical vertebræ wide and depressed ; in

the latter several of them are commonly anchylosed. In Talpa,

the fourth, fifth and sixth cervicals are much lengthened and

overlap each other.

6. Transverse processes of lumbar vertebre very short —

Since in truly fossorial forms there is but little up and down or

from side to side movement in the lumbar region, the corre-

sponding muscles and their attachments, the transverse processes

are feebly developed (e. g. Erinaceus).

7. Sacrais fused— The main stress in pushing comes on

the sacrals; in the majority of fossorial forms (e. g. Condylura,

Lepus) they fuse even to a greater extent than in cursorial

animals, as no lateral or vertical displacements but only rigidity

are required of them.

No. 444.] HABITS IN MAMMALS 823

8. Sternum well developed — The anterior part of the trunk

requires rigidity, great strength and ample surfaces for the

attachment of the hypertrophied digging muscles. The individ-

ual bones tend to become short and broad and the processes

for muscular attachment prominent (e. g. armadillos). In the

moles change of position of the fore limb is correlated not only

with the broadening but especially with the elongation of the

presternum ; at the same time the clavicle is extremely broad,

and shortened so that the limbs may project as little as possible

from the sides of the body ; the limbs are but slightly shortened,

so that the leverage of the muscles is unaffected. In Chryso-

chloris this need is met by the invagination, as it were, of the

walls of the thorax for the reception of the arms, the ribs and

sternum being convex inward. The clavicle is usually, as in

Chrysochloris, curved backward from the sternum, so that the

shoulder may slope gradually forward and not be an angular

projection interfering with progress through the earth.

9. Bones of fore limb strong, tuberosities prominent— The

fore limbs being the principal organs for digging are well devel-

oped, while the tuberosities, ridges, etc., for the insertion of mus-

cles are very prominent. Sesamoid bones are frequently devel-

oped in the palms, as in Echidna where also the breadth of the

hand is increased by a radial sesamoid (os falciforme of Talpa).

The humerus is usually stout and broad as in Talpa, Phas-

colomys, Echidna, etc. with prominent deltoid and supinator

ridges. These are carried to an extreme in Talpa where the

deltoid ridge joins the inner tuberosity above. The olecranon

process is always strongly developed. A supracondylar foramen

is usually present.

10. llumand ischium rod-like gerade! to the vertebral column

and fused to the sacrum.— The hind limbs being mainly used for

pushing the body through the earth, their point of attachment

to the body must be firm and they must deliver the forward

thrust in a more or less horizontal plane. So we find that the

ilium is long and fused usually throughout its entire length to

the vertebral column ; this is especially true of the moles.

1I. Bones of hind limb not so strongly developed as those of

fore limb.— Besides pushing the body forward the hind limbs

824 THE AMERICAN NATURALIST. | [Vor. XXXVII.

are principally used for throwing back the loose earth. Although

not acquiring the excessive development of the fore limbs, the

femur usually has its trochanters well developed. In Erinaceus

it has a strong ridge below the third trochanter. Greater

strength is given by the partial anchylosis of the tibia and

fibula; in Chrysochloris they are welded at the lower end.

Strength is effected in the pes by the great development of

the calcaneum, which plays a prominent part in pushing the ani-

mal forward. In Talpa the pes has a large sesamoid bone at the

side of the tibia corresponding to the os falciforme of the manus,

but otherwise it exhibits none of the great modifications of the

manus.

C. PHYSIOLOGICAL MODIFICATIONS.

1. Hbernation.— In the temperate zones where the ground

is frozen during a portion of each year, fossorial mammals would

have difficulty in getting food. Especially is this the case with

those semi-fossorial forms such as the woodchuck which live on

soft succulent herbage. Such forms are thus compelled to spend

the winter in a long sleep (e. g. woodchucks, gophers, ham-

sters, etc.).

PARTIAL List or FOSSORIAL AND SEMI-FOSSORIAL MAMMALS.

Monotremata : Ornithorhynchus, Echidna ; Marsupialia: Phas-

colomys, Dasyurus, kangaroo rat, Bettongia leseuiri, pig-footed

bandicoot (Chæropus castanotis), marsupial mole (Notoryctes

typhlops); Edentata: Dasypodidæ, Orycteropus ; Insectivora :

Talpa, Condylura, Scalops, water shrew (Crossopus fodiens), des-

man (Myogale), Erinaceus, Oryzoryctes, Chrysochloris ; Roden-

tia: Lepus, Spermophilus, Cynomys, Arctomys, Geomyidæ,

Spalacidæ, Rhizomys, Octodontidæ, Ccelogenys, Viscacha

(Lagostomus trichodactylus), Bathyergidæ, Heterocephalus ;

Carnivora: Lutra, ratel (Mellivora), Javanese skunk (Mydaus),

American badger (Taxidea).

The anatomical conditions prende to progressive modifi-

cation along fossorial lines evidently include the following: reten-

No. 444.] HABITS IN MAMMALS. 825

tion of the primitive characters of small size, generalized (z. e.

short, more or less plantigrade, pentadactyl) limbs with moder-

ately sized claws and normal relations of the radius and ulna,

clavicle and muscles used in digging unreduced, face pointed

elongate, dentition adapted to food found in the earth. Hence

it is natural that the majority of fossorial forms should have

sprung from primitive and defenceless rodents, insectivores and

edentates and that the carnivores (especially those with special-

ized sectorial dentition) the ungulates (mostly herbivorous, and

cursorial) the primates (typically light limbed, light clawed,

short faced), have as a rule failed either to find protection

` from foes or abundant food by turning into the barred road of

fossorial modification.

BIBLIOGRAPHY.

BEDDARD, F.

: 02. icc Cambridge Nat. Hist. Vol. 10.

FLower, W. H. and LvDEKKER, R.

'91. An Introduction to the Study of the Mammals Living and Extinct.

ON THE STRUCTURE OF THE PTERASPIDZE

AND CEPHALASPID/ZE.

W. PATTEN.

I. Tur fact which first suggested to me the possibility of a

genetic relationship between the Vertebrata and Arthropoda was

the similarity in structure and development between the median

eyes of arthropods and the pineal eyes of vertebrates.

To test this idea a careful study of the brain, sense organs,

cranial nerves, nephridia and skeletogenous structures was made,

the results of which showed so clearly a fundamental similarity

between the structure and the arrangement of these parts in the

cephalothorax of certain arthropods and in the head of verte-

brates, as to justify the conclusion that the vertebrates were

derived from fully developed arthropod types, and that the solu-

tion of the various problems in the morphology of the vertebrate

head must be sought for in the evolution of the arthropod ceph-

alothorax.

For the advocate of the annelid theory of the origin of verte-

brates, or of any other theory that assumes the vertebrates to be

derived from soft bodied ancestors, an appeal to paleontology in

support of anatomical or embryological evidence is well nigh

hopeless. But for those who support the arthropod theory, such

an appeal is imperative because paleontology is not likely to

remain forever silent when both extremes of the series of hypo-

thetical annectant types could be preserved as fossils.

The fossil forms that at first sight seemed most completely to

bridge the gap between their respective types are the trilobites

and Merostommata, representing the arthropods, and the ostra-

coderms representing the vertebrates. It seemed probable that

a study of their remains, especially those of the ostracoderms,

whose structure presents so many interesting problems, might

furnish evidence for, or against, the supposed genetic relationship

. between these two groups. Moreover the fact that these ani-

827

828 THE AMERICAN NATURALIST. [Vor. XXXVII.

mals are among the oldest representatives of their respective

"types known, that they were contemporaneous and lived under

very similar conditions, and that they presented many obvious

external resemblances in form and mode of life, clearly invited

a more detailed comparison between them.

The evidence thus far obtained, justifies the belief that pale-

ontology will add its evidence to that of anatomy and embryology

in favor of the origin of vertebrates from arachnids.

From the following historical review it will be seen that while

it is now generally assumed that the ostracoderms have pro-

nounced affinities with the vertebrates, there is the greatest

diversity of opinion among leading authorities as to whether cer-

tain organs characteristic of the true vertebrates are present in 7

the ostracoderms or not, and there is also the greatest difference

of opinion in the interpretation of those structures which are

actually known to occur there. In many cases the despairing

admission is made, that if certain openings, or structures, are

not this, that, or the other, what can they be? This is not due

entirely to the imperfect preservation of the remains, because in

some cases they are beautifully preserved.

At the very outset, the obvious differences between the

ostracoderms and true vertebrates, and the resemblance between

them and the arthropods, was a subject for repeated comment.

In fact the resemblance was so striking that it led many of the

best earlier observers to describe certain merostomatous arthro-

pods as fishes and various ostracoderms as arthropods. More

recent authors while admitting that there is a superficial resem-

blance between these two great groups, explain this resemblance

as one due merely to mimicry or parallelism, and not to a genetic

relationship. Some authors even deny that the peculiar struc-

ture of the ostracoderms is an indication of their primitive

character, but regard them as either highly specialized, or as

degenerate offshoots from the class of typical fishes.

At first the discussion centered round the question whether

certain genera of the ostracoderms, especially Cyathaspis and

Pteraspis were arthropods or vertebrates. But whether or no

these genera, and consequently the whole group of the ostraco-

No. 444] PTERASPIDÆ AND CEPHALASPIDA. 829

derms, were annectant types uniting the true fishes with the

arthropods, and possessing some of the characters of both

classes, so far as I know, was not discussed by any writer till

I raised the question in 1889, in my first paper ** On the Origin

of Vertebrates.”

Huxley and Lankester answered the first question with great

emphasis in favor of the vertebrates, basing their conclusions

primarily on the microscopic structure of the head shields, the

presence of fish-like scales in Pteraspis, and of a caudal fin in

Cephalaspis. Their authoritative opinion settled the question

for the time being, and soon afterwards, the almost universal

belief in the derivation of vertebrates from soft bodied annelid

ancestors that would leave little or no trace behind, and the wide-

spread conviction that the growing science of embryology was

to be the final court of appeal in all broad phylogenetic prob-

lems, turned the attention of morphologists away from the pale-

ontological aspect of the problem.

The development of new lines of zoólogical research and the

failure of embryology to realize the over confident expectations

of its disciples, as well as the frequent and flagrant abuses of

embryological data, have produced within the last decade a spirit

of impatience, or of indifference, towards phylogenetic specula-

tions in general and a reaction set in, not only against the

annelid theory of the origin of vertebrates, but against all

theories that attempt to bridge this, the widest gap in the whole

organic kingdom, by a purely speculative use of embryological

data.

The association of such names as Hugh Miller, Louis Agassiz,

Huxley and Lankester with the early history of Pteraspis and

the Cephalaspidz adds greatly to the interest that has long

centered in this group.

In .Sz/urza, (London 1854, p. 252), Sir Roderick I. Murch-

ison speaking of CepAa/aspis agassizii says: "* This fish with its

large buckler-shaped head and its thin body, jointed somewhat

like a lobster, is perhaps the most remarkable example of a fish

of apparently so intermediate a character, that the detached por-

tions of its head when first found were supposed to belong to

Crustacea.”

830 THE AMERICAN NATURALIST. [Vor. XXXVII.

In a foot note Murchison adds: * Mr. Miller has requested

his readers to compare the head of Asaphus (now Phacops)

caudatus, a well-known Silurian trilobite, with that of C. Zye//zt,

to illustrate how the two orders of Crustaceans and Fishes seem

here to meet,— in the view of persons who have not mastered

the subject."

Eichwald says (54, p. 105): “It is very remarkable that this

colossal crab (Pterygotus) formerly regarded by L. Agassiz as a

fish....occurs in the dolomitic chalk of Rootziküll in Oesel,

together with another genus, Thyestes, standing between Crabs

and Fishes and resembling Bunodes and Cephalaspis."

Hugh Miller, the discoverer of Pterichthys, says (Old Red

Sandstone, p. 50), in comparing a trilobite with Cephalaspis

“The fish and the Crustacean are wonderfully alike.”

* They exhibit the points, ....at which the plated fish is linkéd

to the shelled TEER

Sir Roderick Murchison, when first shown specimens of

Pterichthys wrote regarding them that, “if not fishes, they more

clearly approach to crustaceans than to any other class.”

Again, “ They (Cephalaspis and Pterichthys) form the connect-

ing links between crustaceans and fishes.” Agassiz was at first

in doubt as to whether Pterichthys was a fish or a crustacean.

The following quotation illustrates the attitude of modern

paleontologists toward the ostracoderms. A. S. Woodward,

whose opinion on this subject is entitled to the greatest respect,

in his recent text-book of Paleontology ('98, p. 5) states that

* Nearly all the genera (of the Ostracodermi) mimic in a curi-

ous manner the contemporaneous Eurypterids”; and on p.

24 of the Introduction, that “The oldest Ostracoderms ..

sometimes claimed as the immediate allies of the crustacean or

arachnid Merostomata of the same period, are fundamentally

different from the latter in every character which admits of

detailed comparison; they are to be regarded merely as an

interesting example of mimetic resemblance between organisms

of two different grades adapted to live in the same way and

under precisely similar conditions."

Surely, no one knows either the precise conditions, under

which these forms lived, or the “way” they lived. It would cer-

No. 444] PTERASPIDE£ AND CEPHALASPIDA. 831

tainly be a very unusual thing if all the ostracoderms mimicked

animals so different in grade of organization according to Wood-

ward's view, as the Merostommata are. As a matter of fact,

there is no more reason for assuming that the ostracoderms mim-

icked the Merostommata, than that the Merostommata mimicked

the ostracoderms, and there would have been no obvious advan-

tage to any of them on either supposition.

Moreover the features in which the ostracoderms mimicked

the eurypterids are characteristic of a very extensive class and

are the very characters which are important in differentiating the

ostracoderms from the true fishes, such as, for example, the

small pointed body, large shield-shaped head with its peculiar

cornua, cephalic appendages, shell covered orbits, unusual char-

acter of the parts surrounding the mouth, and the minute

structure of the nearly continuous dermal armor. It is the

combination of all these characters that makes the resemblance

between the ostracoderms and Merostommata difficult to under-

stand on any other assumption than that of genetic relationship.

Clearly it is not convincing, or a final solution of the problem,

to say that these extensive resemblances between two great

classes of animals are due solely to either mimicry or parallelism.

II. Tue Preraspipa. Going back again to the older

writers, we shall see that much of their discussion having any

bearing on the position of the ostracoderms was on the struc-

ture and relations of the Pteraspide. But the fact that these

animals were the first fish-like animals to appear on the earth's

surface, and that they were found in some of the oldest fossil-

iferous rocks known, did not influence their views as to the nature

of these forms so much as one might have supposed. Although

certain parts of Pteraspis and Cyathaspis were at one time

thought to be the bones of a cuttle fish, or the shields of trilo-

bite-like crustaceans, a more careful study of their microscopic

structure, and a comparison with related forms, showed that they

belonged to a group of animals with unquestionable vertebrate

affinities. This fact seems to have shut off all further consider-

ation of their phylogenetic signification, for as soon as their ver-

tebrate affinities were once established they were pigeon-holed

832 THE AMERICAN NATURALIST. [Vou. XXXVII.

among the true fishes, and their existence practically ignored.

Although the pteraspids are now generally placed among the

true fishes, their head shields do not present a single recogniz-

able vertebrate character. The various surface markings have

been supposed to indicate the presence of median or lateral eyes,

olfactory, or auditory, organs, or gill openings, or the impres-

sions of gill pouches,— not because they show any particular

resemblance to corresponding organs in true vertebrates, but

because, as has been frequently said, there is nothing else with

which they can be compared. There are no traces in these

remains, which in general are fairly well preserved, of upper and

lower jaws, fins, notochord, or vertebral column.

The genus Pteraspis was first proposed by Rudolph Kner in

1847 to include the forms described in 1835 by Agassiz as

Cephalaspis lewisit, and C. /leydzi. Their appearance was so

unlike the ordinary fish remains that for a long time Kner did

not suspect that they had been already described by Agassiz

in his Pozssons Fossils.

From a study of their minute structure Kner believed them

to be the internal shells of cephalopods allied to Sepia.

In 1856, F. Roemer described a form closely related to C.

lloydii as Paleoteuthis, and referred it to the Sepiidz, but sug-

gested that the forms described by Kner were crustaceans

related to Dithyrocaris or Pterygotus. ;

In 1864, Lankester divided the Pteraspidz into the three

genera, Pteraspis, Cyathaspis and Scaphaspis. But in 1872,

Kunth described a shield of Cyathaspis, below which he found

one belonging to Lankester's genus Scaphaspis, and he rightly

concluded that the two shields belonged to the same animal.

He maintained that the lower shield bore the same relation to the

upper one that the tail plate of a rolled up trilobite does to its

head shield, and that between the two were a number of pieces

comparable with the segmental trunk plates of a trilobite. Other

plátes were present which Kunth regarded as locomotor organs,

or foot-jaws. From the above facts Kunth concluded that these

remains were not those of a fish but of an arthropod. In refer-

ring to Huxley's statement that there is no molluscan or crusta-

cean structure with which such remains could be for a moment

No. 444] PTERASPIDAX AND CEPHALASPIDA. 833

confounded, and to Kner's belief that Scaphaspis was the shell

of Sepia officinalis, Kunth adds “so schienen mir diese Ansichten

in verein mit unserem vorliegenden Stücks mir zu beweisen dass

wir es mit einer Crustacean Abtheilung von ganz eigenthümlicher

Schalstructur zu thun haben. Denn jedenfalls giebt es weder

einen Fisch noch eine Sepien Schulpe, die eine ühnliche Structur

wie die Schilder zeigte; wohl aber ist die Organization des gan-

zen Stückes beweisend fur Crustaceen Character " (p. 6).

Both Schmidt ('73, p. 33c) and von Alth (p. 47) agree with

Kunth that Scaphaspis is the ventral shield of Pteraspis, but

they deny that any of the remains described as Pteraspis, Cya-

thaspis or Scaphaspis are crustaceans, although no valid reasons

are given for doing so.

Huxley (58, p. 277) in reply to Agassiz, who had remarked

on the singular resemblance between the shell of C. Koydii and

that of crustaceans, and to Roemer’s and Kunth’s opinion

that Pteraspis was a crustacean, seems to have closed the dis-

cussion for the time with his oft quoted statement that ** No one

can, I think, hesitate in placing Pteraspis among Fishes. So far

from its structure having ‘no parallel among fishes,’ it has abso-

lutely no parallel in any other division of the animal kingdom.

I have never seen any molluscan or crustacean structure with

which it could be for a moment confounded."

Roemer accepts these statements apparently because they

came from Huxley, although he does not make an unconditional

surrender of his opinion, for he says “ Allerdings manche Anal-

ogie der aüsseren Form mit Crustacean-Formen dar bieten

wurde.”

In 1855, R. W. Banks in his paper on the Downton Sand-

stones, after commenting on the association in these beds of

Lingula cornea, Pterygotus and Pteraspis (Cyathaspis), made

the following observation, p. 98, **On the under side of the

sharp projections before referred to (on either side of the

rounded snout) are protuberances which seem to be projecting

horny eyes similar to those of Crustaceans.”

He remarks further on, that doubtful as it is whether the

buckler-like fossil remains above referred to belong to fishes or

to crustaceans, it is certain that they are closely allied to Cepha-

834 THE AMERICAN NATURALIST. [Vor. XXXVII.

laspis lloydii and C. lewisii. In a final note, it is announced

that Professor Huxley is now minutely examining their structure

to determine their true relationship either to the crustaceans or

to the fishes. When Huxley's paper appeared, although he

gave a very good description of the minute structure of the

shell of these animals and concluded that they are not crus-

taceans, he entirely ignored the existence of the eye tuber-

cles, although their presence afforded very weighty evidence

against his conclusion.

Lankester (768, p. 26) admitted the presence in Cyathaspis

of tubercles corresponding with similar tubercles in Pteraspis,

which are “produced by the supposed orbits”; but how a

vertebrate eye, or an “orbit,” could be preserved as a beauti-

fully rounded protuberance when all the other soft parts are

completely destroyed, is not discussed.

Lankester however, (73, p. 241), still maintained the validity

of his genera Scaphaspis and, in another article (73, p. 190)

makes the following statement: “It is to me à source of two-

fold regret that Dr. Kunth has perished in the Franco-Prussian

war, for not only have we thus lost a chance of obtaining addi-

tional knowledge of the Berlin Cyathaspis, but I shall be unable

to obtain from him the admission that his conclusion is not in

accordance with the facts." Lankester ('91) finally admits that

von Allth's discovery shows Scaphaspis to be the ventral shield

of Pteraspis, and thus we may assume that his “twofold regret "'

for Kunth's untimely death was in a measure mitigated.

Lankester attaches much importance to the presence of scales

on the anterior trunk region of Pteraspis, for these scale-like

structures are regarded as conclusive proof that the Pteraspidze

belong to the vertebrates. As Lankester says ('68, p. 18)

«All that is known as regards the scales of these Fishes is from

a single specimen found in the Cornstones of Herefordshire."

This specimen, he says elsewhere (73, p. 191) “Shows seven

rows of rhomboidal scales attached (not merely adjacent to) to a

portion of the head shield of Pteraspis. That these are true

scales, or lozenges of sculptured calcareous matter is absolutely

certain. It is also absolutely certain that the shield is pteras-

pidian and that the scales and shield belong to the same individ-

No. 444] PTERASPIDA! AND CEPHALASPIDZ.. 835

ual organism. The scales are fish-like. I know no Arthropod,

nor any other organism except a fish which possesses any struc-

ture even remotely representing them." “The shields of the

Chitonidz and Cerripedz are the only animal structures, except

the scales of a ganoid fish (with which they agree exactly) which

they could even vaguely suggest." “The form of this shield,

and its details as to apertures, processes, etc., agrees with the

view that it belongs to a fish most fully. It has not the

remotest suggestion of Crustacean affinities about it."

After commenting on the fact that the fossil in question was

marked with long parallel striz, and that the middle layer con-

tained the polygonal cavities he adds ('64, p. 195), * This

structure, which has no parallel among fishes, or, indeed, any

group of the animal kingdom, leaves no possibility of a doubt

that the specimen is a fragment of Pteraspis." .... Lankester

further maintains (68, p. 4) that by the discovery of these scales

* the piscine nature of these fossils was definitely set at rest."

These positive statements are somewhat contradictory and

would now be hardly warranted by the facts of the case. The

crustacean character of the shields has been repeatedly com-

mented on by competent observers. In his own monograph

(p. 61) he has described a fragment, possibly connected with

Cephalaspis which he names Kadlostrakon podura (Tolypelepis ?)

* on account of the resemblance to the well-known microscopic

markings of the scales of the insect Podura."

It is not true that there are no arthropods with structures

even remotely resembling these scales, because in Pterygotus the

entire body is covered with an ornamentation which bears an

astonishing resemblance to fish scales, so much so as to deceive

such a keen observer of fishes as Louis Agassiz. Moreover in

many trilobites and in the Ceratiocarina, Clarke, the surface

of the shell is ornamented with ridges and grooves not unlike

those of Pteraspis in external appearance.

Lankester probably would not have made these statements

had he kept Pterygotus in mind, or had he been acquainted with

the minute structure of the shield of Limulus.

Moreover all recent students of the shell of Pteraspis are

agreed that it is zo? “exactly ” like that of a ganoid fish, in fact

its microscopic structure is altogether of a different character.

836 THE AMERICAN NATURALIST. [VOL XXXVII.

But after all, the presence or absence of these scales in

Pteraspis has little weight, it seems to me, in answering the

real question at issue, namely whether Pteraspis in addition to

its vertebrate affinities has not also a genetic relationship with

the arthropods.

Lankester, Woodward, Traquair, Rohon and others agree in

denying the existence of

arthropod characters to

the pteraspids, appar-

ently because of the

abundant evidence now

available that Pteraspis

is related to Cephalaspis

whose ichthyic affinities

have rarely been ques-

tioned, rather than be-

cause the arthropod fea-

tures of Pteraspis have

been dispassionately

considered and found

wanting.

But within recent

years there seems to be

a growing tendency to

doubt the affinity be-

Pteraspis and

Cephalaspis. Reis pro-

tests against their union,

and apparently Traquair

is in doubt, treating

them together largely

as a matter of conven-

ience. Lankester in his

earlier monograph states

tween

Fic. r.— Cephalic buckler of Pteraspis, dorsal view,

after Lankester. The shape of the lat

ings, d. * fev been slightly modified; 7. ». the

supposed ral eye openings, LN the points

for edt of appendages

that “The Heterostraci

are associated at present

with the Osteostraci

because they are found in the same beds, because they have like

No. 444] PIERASPIDE AND CEPHALASPIDA. 837

Cephalaspis a large head shield, and because there is nothing else

with which to associate them." More recently he has said (97)

“There is absolutely no reason for regarding Cephalaspis as

„allied to Pteraspis, beyond that the two genera occur in the

same rocks, and still less for concluding that either has any

Fig. 2.—Cephalic buckler of Pteraspis, ventral view, showing the small oral

y filled with several pairs of plates. The large ventral plate,

s. c. is supposed to be cut away on one side showing the median eye pit

the paired muscle markings of the inner surface of the dorsal shield.

connection with Pterichthys." Zittel says, Vol. III, p. 147,

“ Mir scheinen die Beziehungen der Pteraspiden und Cephalaspi-

den nach Form und Structur so entfernt dass beide besser ms

838 THE AMERICAN NATURALIST. [Vor. XXXVII.

besondere Ordnungen betrachtet werden." He remarks further

on that while the Cephalaspide certainly appear to be ganoids,

the position of the Pteraspidae is very doubtful.

It seems to me that there can no longer be any doubt on

these points, since the Pteraspide and Tremataspidz are pretty

closely united with the Asterolepide by the similarity of their

oar-like appendages, and with the Cephalaspidze, by the similar-

ity of the median and lateral openings in Tremataspis, Thyestes

and Cephalaspis.

At present, of the genus Pteraspis, only a part of the cephalic

armor and a few scale-like structures belonging to the anterior

part of the trunk, are known. The head, Figs. 1 and 2, is arrow

shaped and covered by a dorsal and ventral shield, the latter,

the so-called Scaphaspis, consisting of a single ovate or oblong

piece, varying in different species from 14 to 34 in. in length.

The lateral edges of the ventral shield probably fitted closely

to the lateral edges of the dorsal one, leaving under the rostrum

a comparatively small opening in which the various mouth parts

were situated. ;

The boat-shaped dorsal shield is composed of seven portions,

marked off on the outer surface of the shield by furrows, and

on the inner surface by ridges. In young specimens, the ros-

trum and the central disc may be found separately. Each piece

has its surface ornamentation of ridges and furrows arranged in

concentric lines parallel with its margins. This fact, together

with other considerations, led Lankester to believe that each

piece ossified from a separate centre and that their complete

anchylosis occurred only in the adult.

Muscle Markings. —In 1872, A. Kunth described in Cyathaspis

integer a series of six “flache Höcher,” situated on the under

surface of the dorsal shield, which he regarded as indications

of segmentation, Fig. 4. Lankester, (73), describes similar

impressions on the shield of Cyathaspis banksii and believes

that in both cases they indicate the position of a series of

branchial chambers.

In Pteraspis also, Lankaster has described five narrow ridges,

with four broad shallow depressions between them, which radiate

No. 444.] PTERASPIDA AND CEPHALASPIDA. 53 9

from the centre of the inner surface of the dorsal shield, Fig. 2,

m.r. They are perhaps best marked in Preraspis crouchii and

P. rostratus.

These markings on the dorsal shield of the Pteraspide are, I

believe, best explained as indications of the original segmenta-

tion of the head, produced in part by the attachment of strong

segmental muscles extending vertically from the inner surface of

the dorsal shield, either to a cartilaginous cranium, or to a series

of gill-like or jaw-like segmental appendages on the ventral side.

They suggest the markings produced in this manner on the dor-

sal surface of the cephalothorax of Limulus, or the lobulations

on the dorsal shield of a trilobite head.!

Eyes.— There is a conspicuous pit on the inner surface of

the shield, appearing on the outer surface as a small tubercle,

which marks the position of the median eyes.

Lankester (70) shows in his Fig. 6, Pl. IV, three tubercles

near the median anterior part of the head of Cyathaspis, and the

same tubercles were figured by A. Kunth in 1872. In both

figures, the resemblance of the shields to the cephalothorax of

an arthropod is intensified by the shape and arrangement of

these three ocelli-like tubercles (Fig. 4).

The location of the lateral eyes of Pteraspis is supposed to be

indicated by two smooth tubercles, or in some cases by circular

openings, near the anterior margin of the shield, Z. ». It is hard

to understand how the usual type of vertebrate eye could be pre-

served in the form of these tubercles. If they are lateral eye

tubercles, they, like the compound eyes of arthropods, must have

been enclosed by a firm dome-like covering, continuous with the

outer layers of the shell. I have never seen any specimen of

Pteraspis in which these so-called lateral eye notches were actu-

ally covered by an extension of the outer shell layer, although

such a covering is found over the large median eyes of Cephalas-

pis and Tremataspis. It is possible that there are two pairs of

such notches, or openings, in Pteraspis, Cyathaspis and Tolypele-

pis, one serving for the lateral eyes and the other for the attach-

ment of swimming appendages similar to those of Tremataspis.

'Rohon has described a somewhat similar segmentation of the head in

Thyestes.

$40 THE AMERICAN NATURALIST. [Vor. XXXVII.

The projections sometimes seen in the so-called orbital notches

of Pteraspis, appear to be the remnants of appendages.

Dorso lateral openings.— In Pteraspis a large oval opening on

each side of the dorsal shield leads into a wide canal that runs

diagonally forward and downward into the interior of the head

(Figs. 1 and 3, 7. o.). Lankester ('68, p. 17) says, “It is very

difficult to find any explanation of these open excavated struc-

tures unless they be spiracles."

Although the margins of these openings are not well enough

preserved to show such important details as are seen in Trema-

taspis, their shape and position indicate that they may be homolo-

gous with the lateral openings

in the dorsal shield of Cepha-

laspis, Thyestis and Trema-

do.. taspis. Like the openings in

A Mh. iu uM estie

=< genera, they were probably

Fic, 3.— Cross section through the posterior covered by an extension of

t pane the outer shell layer, possibly

with its lateral openings is from an actual by loosely fitting polygonal

section. The ventral shield is restored.

plates. This may be the

reason why Claypole, (p. 566), found no trace of such openings

in Palaeaspis.

Sections of two different specimens (Fig. 3) show that there

is no extension of the inner layers of the shell over the opening,

like that seen in Tremataspis and Cephalaspis.

If this interpretation of the lateral openings in Pteraspis is

accepted, there is no reason for supposing that they have any-

thing to do with the gills, and the argument that Pteraspis is a

fish because it has spiracles, or gill openings, is eliminated.

Appendages.— Yn Paleaspis americana, Claypole ('92, p. 554),

finds indications of appendages consisting of small objects resem-

bling spines * margined with what appeared to be a flat fringe

around their smaller and pointed ends; ....the central spine-

like portion was covered with a layer of the same tissue as that

of which the shield was composed." He then adds that the

resemblance of the structure to a crossopterygian fin was obvi-

ous. ‘Two forms of fins appear to prevail, the one broad and

No.444] PTERASPIDA AND CEPEALASPIDE. 841

the other narrow, indicating that more than one pair belonged to

the same individual."

These facts are of great significance in view of Lindstróm's

discovery of the appendages in Cyathaspis, (Fig. 4) and of my

own discovery of ap-

pendagesinTrematas- maw

pis. Dean COS p.71) |

maintains that the

structures in question

do not have the sig-

nificance claimed for

them by Claypole,

and, that they do not

even belong to Palæ-

aspis but to some

elasmobranch. But

Claypole's observa-

tions seem to have

been made with great Fic. 4.— Restoration of the head shield of Cyathaspis,

1 1 fads ^ oar-like appendages, af. the three median

wales : and in view of eye pits, ef. and the pear muscle markings, ». m.

the importance that sometimes regarded as the impressions of gill pouches.

The eye pits and muscle markings are best seen in

must be attached to casts of the inner surface

them, if confirmed,

they deserve more careful consideration than they have hereto-

fore received.

III. THE CEPHALASPIDÆ are found in the upper Silurian and

in the Devonian formations, and range in size from a few inches

to possibly two feet or more in length.

Shape of Head.— In Cephalaspis, (Figs. 5 and 6), the typical

genus of the family, the head was completely enclosed in a con-

tinuous cephalic buckler or shield, which seen from above is

elipsoidal in outline, either rounded or pointed in front and

truncated behind. The lateral angles of the posterior margin

extend backwards to form the cornua, while the median posterior

portion forms a broad crest often armed with a prominent median

Spine.

Cross sections (Fig. 8 Æ) show that the head is thin and

842

THE AMERICAN NATURALIST. [Vor. XXXVII.

spoon-shaped, with its concave

surface facing downwards. It

rests on a flat, thick-walled rim

which gradually widens posteri-

orly till it forms the under side

of the broad triangular cornua.

The ventral wall of the shield

is thin, and is pressed closely

against the dorsal wall. Toward

the middle line it is gradually

reduced to a flexible membrane

m adt

which terminates abruptly, leav-

ing a large median opening in

which the soft parts of the head

were situated.

Eyes.— Near the middle of

the dorsal surface are the oval

openings for the eyes. Cross

sections of the orbits, and frag-

mentary casts of their outer sur-

face, show that the eyes were

nearly spherical, and that they

projected dome-like above the

surface of the head. They were

covered by a thin coating of

hard material continuous with

the outer layer of the shield.

The floor of the orbits is very

concave and consists of a basket

work of bony trabeculae formed

by an extension of the inner

eres. shell layer.

appendages, so-called *'*opercular flaps,

c, cornua; @ o. marginal organs; 7. ros-

perforation; e. eyes; f. v. t-orbital val-

ley; 4. f. dorsal fin; f. 5. fringing proc-

esses.

843

PHALASPID£..

AND

PTERASPIDAE

No. 444.]

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844 THE AMERICAN NATURALIST. [Vor. XXXVII.

The eyes, therefore, of Cephalaspis, like those of Tremataspis

and Bothriolepis were imbedded between two layers of the der-

mal armor, and in this respect agree with the condition which

prevails in both the median and the lateral eyes of Limulus (Fig.

Two depressions lie just in front of the orbits (the so-called

ant-orbital fosse (a. f.). They are separated by a median crest,

m m m — P À——— a iiir Pr

Fic. 7.— Diagrammatic cross section of trunk of Cephalaspis.

the summit of which is perforated by a narrow slit which leads

into the interior of the head.

On the inner surface of the shield, near the lateral margins,

are two very large oblong thickenings composed of a net-work

of osseous trabecula (Pl. I, Figs. 2, 3 and 4, and text figs., 5

and 6, d. o.). Another prominent mass of the same tissue lies in

the median line, just behind the orbits and just beneath the post-

orbital valley (5. c.).

These lateral thickenings have been obscurely indicated in

some of Lankester's figures, but I can find no reference to them

in the text. They are probably the “pair of great rounded

lobes meeting in the middle line" mentioned. by Woodward (p.

179, Part II), and the “pouch-like sensory organs” of Dean

In the specimeris here figured the lateral lobes do not meet

in the median line and I have not seen any specimens in which

they do. When the outer surface of the shield is preserved, it

No. 444] PVERASPIDAS AND CEPHALASPIDE. 845

shows a well defined opening corresponding in position and out-

line with the mass of trabeculz below. In some specimens, the

opening is covered by a special group of loose, irregular polygonal

plates, well shown in Pl. I. Fig. 3.

The mass of trabeculze below the post-orbital valley is conspic-

uous and is often very sharply outlined (Pl. I. Fig. 2). Itis

oblong, pointed in front, and broad and somewhat angular behind.

The post-orbital valley when seen from above, in well pre-

served specimens, presents a clearly defined oblong opening,

similar in shape to the underlying mass of trabeculz, and filled

with polygonal plates similar to those of the marginal openings.

In Thyestis, Fig. 9, I have found marginal and post-orbital

Fic. 8.—A.— Cross section of the cephalothorax of Limulus, showing arrangement of the

bony trabeculz in the median line, below lateral eyes and on the margins of the shield.

bo section is too far back to show the trabeculae below the median eyes. B.— Cross

tion through the head of Cephalaspis, showing orbits, lateral organs, and part of the

nd shield.

openings similar to those of Cephalaspis, but smaller and more

like those of Tremataspis. The covering polygonal plates, how-

ever, are absent.

There can be no doubt that the marginal and post-orbital

openings of Cephalaspis and Thyestis, and perhaps the marginal

openings of Pteraspis, are the same as those so well shown in

Tremataspis, and that in all these genera some important organ

of a sensory nature was lodged between the two layers of the

shield at these points. I have shown that the arrangement of

the marginal openings in Tremataspis is very suggestive of that

of the lateral eyes and dorsal organs of Limulus (:01, p. 7).

846 THE AMERICAN NATURALIST. [Vor. XXXVII.

The marginal cells of Eukeraspis Lank., seem to be a special

development of bony trabeculze similar to those below the lateral

openings, but more loosely arranged. They serve to unite the

dorsal and ventral laminze, and to give additional strength to the

rim of the shield. :

I have already pointed out (:01), that in Limulus there is a

system of supporting bony trabecule similar in structure and

arrangement to those in Cephalaspis. In Limulus, as in Cepha-

Fic. 9.— Head shield of Thyestes, X 4. Letters as in Figs. 5 and 6.

laspis, the principal masses of the trabeculz lie along the mar-

gins of the shield, in the cornua, and beneath the median and

lateral eyes.

Trunk Scales and Fins —The dorsal surface of the trunk is

covered with a single row of saddle-shaped, overlapping scales,

sometimes fused into larger groups. The ridge scales extend

on to, and support, the anterior margin of the dorsal fin, which

appears to be merely an expansion of the dorsal ridge.

No. 444] PTERASPID.E AND CEPHALASPIDA. 847

The dorsal fin must have been nearly immovable, as it is

sheathed with parallel rows of oblong scales which diminish in

size toward the free margin. The tail lobe is covered by similar

scales. It was diphycercle, not, as is usually stated, heterocercle.

In some cases I have seen indications that it terminated in a

long, banner-like filament.

The flanks are covered by two principal rows of scales, the

dorso-lateral one consisting of plates placed at a sharp angle with

the dorsal crest, and in some cases twice as numerous as the

crest scales (Pl. II, Fig. 7). The posterior edge of each scale

overlaps the anterior edge of the next following one.

The lateral trunk plates are much larger, and stand nearly

vertical. There is a semicircular incision on the posterior angle

of the ventral end of each scale, into which fits the head of the

fringing processes. The scales begin to break up into irregular

polygonal plates about opposite the cloaca (Pl. II. Fig. 6, and

text Fie s5).

The ventral surface of Cephalaspis murchisoni is flat and tri-

angular in contour. It widens out towards the ventral surface

of the head, and narrows toward a point a little distance behind

the cloaca. Its lateral boundaries are sharply defined by the

projecting fringing processes. It is covered with small scales

arranged in well defined rows, directed from either side diagonally

inwards and forwards (Pl. II, Fig. 5). In Cepkalaspis lyelt,

they are directed backwards.

The outermost ones of the ventral scales appear to be nearly

square and in some places (Fig. 6) present a distinct joint for the

articulation of the fringing processes. Next to them comes a

rather large oblong plate. The remaining scales in each row

diminish in width as they approach the median line, where they

are sometimes curiously crossed as though formed by imbricating

filaments.

Cloaca.— In one beautiful, heretofore undescribed specimen

in the British Museum (P. 8804 ? ) about two thirds of the dis-

tance from the anterior end of the trunk, there is a well defined

transverse slit that no doubt represents the cloaca (PI. II, Fig.

5). Its posterior lip is smooth and rounded, the anterior one is

more sharply defined, and forms the basal line of a triangular

area covered with minute rhomboidal scales.

848 THE AMERICAN NATURALIST. [Vor. XXXVII.

Back of the cloaca, the scales are arranged in obscure V-

shaped rows, gradually breaking up into the squarish plates seen

on the sides and ventral] surface of the root of the tail.

Mandibles.— In my paper before the Fifth International Con-

gress at Berlin, I referred to the presence of certain problem-

atical structures in the head of Cephalaspis. I stated that my

attention was first called to this subject several years ago by

one of Lankester's figures (Pl. X, Fig. 2) which seemed to sug-

gest the possibility of some kind of appendages on the under

side of the shield, in the mouth region. Lankester made no

reference in the text to the structure in question. On examin-

ing this specimen in the Edinburgh Museum of Arts and

Sciences (No. 182), two oval bodies were found, lying in about

the middle of the under side of the shield, just back of the

orbits, and conspicuous on account of their peculiar shape and

smooth shining surface.

Each body had a smooth under surface with fluted sides.

Their symmetrical form and arrangement shows beyond doubt

that they formed an organic part of the head.

The muscles moving these structures at first appeared to be

attached to the large, circular, muscle scar on the inner surface

of the dorsal shield, between the cornua and the posterior part

of the dorsal crest (Fig. 5, m. m.) but I am now inclined to

believe these marks were made by muscles moving the so-called

opercula.

It is extremely probable that similar mouth parts occur in

other ostracoderms. In Tremataspis, I have shown that the

oral plates so completely fill the opening in this region that

there is little room for the presence or action of upper and lower

jaws of the vertebrate type. The shape and arrangement of

these plates indicate that the large anterior pair in Tremataspis

represent the mandibles which like those of the arthropod type

must have moved to and from the median line when in use.

In Bothriolepis, we see even more clearly, as we shall describe

in more detail elsewhere, that the so-called mental plates were

mandibles of this kind. They are the only pair of plates present

that could serve as jaws, and their general contour, the long

No. 444] PTERASPIDAS AND CEPHALASPIDA. 849

muscle ridge on their inner surface and the thickened, inturned

median ends, that were probably armed with chitinous, or horny

sheaths, show that they acted against one another in the median

line, either as cutting, or crushing jaws. Their structure and

position show in the clearest manner that the mouth must have

been situated detween these mandible-like plates, not in front of

them. The presence of these remarkable structures around the

mouth of the ostracoderms shows more clearly perhaps than

anything else how wide the gap is between them and the true

fishes.

Appendages — Powrie was the first to call attention to the

paddle-shaped lappets, or so-called pectoral appendages of Cepha-

laspis. They were later described and figured by Lankester and

Powrie as ** Ellipsoidal expanses with some calcareous matter in

their structure which has caused them to be preserved.” They

are characterized by a kind of reticulate or areolate marking and

although they show no trace of fin rays, they were regarded as

a remarkable kind of pectoral fin, ** efficient in causing currents

of water to pass to the branchial organs."

Woodward has more recently examined some well preserved

specimens of C. murchisonii and makes the following statement

in regard to them: (Cat. B. Museum, p. 186). “A novel

point of much general interest is elucidated by the middle layer

of the shield, which is well preserved in several specimens."

* The present specimens prove distinctly that it extends back-

wards as a pair of postero-lateral * flaps’ beyond the rest of the

shield." ‘The outer layer is broken away, so that direct con-

tinuity can be observed between the appendage and the middle

layer.” ‘The structures are merely a portion of the shield

itself, divested of the outer and inner layers to insure flexi-

After quoting Lankester’s opinion of them, he adds, “ Some

connection with the gills has thus already been suspected and it

now seems most probable that the appendages in question

actually correspond to a pair of opercula, and may henceforth be

designated as such.”

I have studied these appendages in the collections of the.

$50 THE AMERICAN NATURALIST. [Vor. XXXVII.

British Museum and those at Edinburgh, but could find no

evidence that the folds are formed as extensions of the middle

layer of the shield. They appear to me to be covered by the

usual shell layers, but broken into small shell-like plates united

in such a manner as to allow some flexibility to the whole struc-

ture. They are not in my opinion specializations of the poste-

. rior lateral margin of the cephalic shield, or of the cornua, but

true appendages attached to the under side of the head. The

large circular muscle scars seen on the inner surface of the dor-

sal shield (Fig. 5, m. m.) probably served for the attachment of

muscles moving these appendages.

I cannot agree with Woodward and Lankester in regarding

these structures as opercula, for sections indicate that the sides

of the head were very thin, and that the most important cranial

organs were near the median line. Hence the so-called opercula,

standing so far back, and to one side, and when the animal was

at rest lying flat-wise against the bottom, were not in a position

to cover the gills, or to produce currents of water through them.

Traquair regards the lappet-like flaps of the Ccelolepidz as

pectoral fins. These “lappets”’ it seems to me are represented

in the Cephalaspidz, by the cornua. Traquair's position is a

difficult one to maintain, in that it compels him to look on the

cornua of Drepanaspis as fins “rendered utterly functionless

as fins by being enclosed in unyielding bony plates" (p. 846).

The true interpretation is to be obtained, I believe, by revers-

ing this order of events and assuming that the ridged cornua of

forms like Cephalaspis, Pteraspis, Drepanaspis, and others, are

homologous and the most primitive because they are most like

the cornua of their arthropodan ancestors. It is well known

that cornua like those of Cephalaspis are present in many

trilobites, and that in Limulus the two walls of the cornua

are united by bony trabecula produced by an elaborate develop-

ment of the inner shell layers, and suggestive of the reticulated

structures filling up the cornua of Cephalaspis.

According to Dean, ('95, p. 69) the large oar-like appendages

of Pterichthys are “lateral head angles produced and special-

ized " for locomotion.

If it is so easy in this instance to create de novo highly

No. 444] PTERASPIDA AND CEPHALASPIDA. 851

specialized appendages like those of Pterichthys, with their

necessarily complex arrangement of muscles and nerves, then

the elaborate discussions over the origin of the paired append-

ages of higher vertebrates would seem to be a waste of time.

But some such explanation as that offered by Dean is forced on

those who regard these animals as true fishes by the difficulty,

from their point of view, in explaining the presence of so many

different kinds of appendages in a vertebrate head, for it is

generally assumed that the appendages of Pterichthys are not

homologous with those of Cephalaspis, and that neither one nor

the other are homologous with true pectoral fins."

The Fringiug Processes. — Lankester showed long ago, in his

reconstruction of Cephalaspis, a fringe of peculiar plates along

the ventral margins of the trunk, which, although they produced

a most unusual appearance, have not attracted the attention

they deserve. Whatever their significance may be, there is

apparently nothing known in true fishes that is exactly compar-

able with them.

In one specimen belonging to the Powrie collection in the

Edinburgh Museum (No. 163) the body was badly crushed,

throwing the plates to one side where they lay flatwise and well

separated. On examination with a lens, some of the plates

appeared to consist of at least two joints, possibly three, the

distal one being the smallest. The surfaces of the plates were

covered with coarse spines. The details of this specimen were

not brought out by the photographs with sufficient clearness to

allow them to be reproduced.

In the beautiful Ledbury specimens of Cephalaspis murchisont,

described by Woodward, the fringes are clearly seen in various

positions, either from the sides or from below (PI. I, Figs. 1 and

5.) In most cases they form a series of regularly overlapping,

-oblong plates, apparently in their normal position on the ventral

margin of the trunk. Each plate appears to be three lobed, the

segmentation being indicated by the gently rounded outline of

each joint, as well as by the transverse lines that separate one

! Jaeckel (:02, p. 111) regards the *opercular flaps ” of Cephalaspis as homolo-

_gous with the dermal skeleton of a crossopterygian pectoral fin.

852 THE AMERICAN NATURALIST. (VoL. XXXVII.

joint from the other. The mode of articulation with the body

scales is not shown by any specimens of this species examined.

In the magnificent specimen originally described- by Agassiz,

(Pl. II, Fig. 6) the animal liesat full length in a natural position.

Here the fringing processes are seen edgewise, and have a differ-

ent shape and appearance from those of Cephalaspis murchisont.

They hang freely away from the trunk, in a nearly vertical

position, with their distal ends bending backwards in graceful

curves. Each process has a rounded head that fits into a cup-like

depression on the posterior ventral margin of the large dorso-

lateral trunk scales. Below the rounded head, the process is at

first quite slender, then somewhat abruptly thickened, and

finally reduced to a thin lamella with indistinct boundaries.

There are from twenty to thirty pairs, beginning just back of

the cephalic shield and gradually decreasing in size from that

point toward the tail end. The most posterior ones are reduced

to mere spines, or rhomboidal plates, loosely articulated to the

lateral trunk scales.

Finally in a small specimen in the Powrie collection at Edin-

burgh (No. 139), where the whole animal is seen in outline

from below, the fringing processes of both sides are shown

folded inwards and flattened against the ventral wall of the

body (Fig. 10). On the left side of the figure, one can count

about twenty processes.

The varying appearance of the processes is due in part to

their position and to the way in which they are exposed, and in

part to the fact that the plates RE to different species

differ considerably in structure.

In Cephalaspis murchisoni, the fringe plates are lobed and

overlap one another so that their flattened surfaces are directed

diagonally forward and outward. In Cephalaspis pagei, they

appear to have a similar shape and arrangement, but are armed

with coarse projecting spines that give them a decidedly arthro-

pod appearance. In Cephalaspis lyellii, the plates lie one

directly behind the other without overlapping, while the .promi-

nent articulating head, the narrow neck, the posterior swelling

and the thin, backwardly directed distal ends give them a unique

and characteristic appearance.

PTERASPIDA AND

No. 444.]

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5854 THE AMERICAN NATURALIST. (VoL. XXXVII.

There can be no doubt that the fringing processes projected

freely from the ventral margin, and that they were freely mov-

able forward and backward, and to and from the median line.

The conditions we have described are not less extraordinary

than the fact that such conspicuous structures should have

remained so long practically unnoticed. Lankester does not

discuss their possible signification in his text, but merely intro-

duces them into a very diagrammatic cross section as plates pro-

truding like bilge keels away from the body. Many of the

newer specimens of Cephalaspis murchisoni in the British

Museum, as well as some of the older type specimens, show in

the clearest manner that the fringing processes are articulated

to the ventral margins of the body and are not artificial folds

made by pressing together the margins of the dorsal and ventral

walls (Fig. 7).

That the whole group of ostracoderms was provided with a

series of fringing plates similar to those of Cephalaspis is very

probable, for a series of fringing plates are known to exist in

Tremataspis, and I have found indications of fringing processes

in the trunk of a fine specimen of Pterichthys preserved in the

geological collection of McGill University.

Morphology of Vertebrate Appendages.— Organs so widely

distributed in a primitive group of animals as the fringing proc-

esses are, must have great morphological significance. But

while there is little doubt that they are the antecedents of the

lateral fold of vertebrates, for no other structures so clearly

reproduce in size, position and function the hypothetical folds

from which the paired fins are supposed to take their origin,

that fact does not help us to determine the morphological

significance of the fringing plates themselves.

The fact that the fringing plates are marked with the same

surface ornamentation as the trunk scales shows they are not

comparable with internal fin rays. On the contrary, their orna-

mentation, shape, and mode of articulation indicate that they

are independent, segmental structures. It is difficult to inter-

pret such structures as anything else than appendages, having

the same significance as the rudimentary abdominal appendages

of arthropods.

No. 444] PTERASPIDE AND CEPHALASPIDA. 855

Assuming that to be the case, we may form some idea of their

probable mode of development by a comparison with those of

Limulus. Miss Hazen and the writer have shown that here

each abdominal appendage arises first as a fold of ectoderm,

into which grows a muscle bud that soon divides into two princi-

pal parts to form the adductor and abductor muscles. Mean-

time the nerve to the appendage appears and an axial core of

cartilage is formed which grows from the basal mesoderm

through the middle of the muscle cells, toward the apex of the

appendage.

As these processes agree in every essential particular with

those known to occur in a segment of the lateral fold of verte-

brates, there can be no serious objection, from an embryological

standpoint, to the interpretation of the lateral fold as a series of

fused abdominal appendages. Assuming then that the lateral

fold is formed, phyllogenetically, by the fusion of a series of seg-

mentally arranged, and independently movable structures, such

as the fringing processes of the ostracoderms, it is clear that

the oar-like cephalic appendages of the ostracoderms cannot be

regarded as specializations of either a lateral fold, or of gill arches

in the Gegenbaurian sense. On the contrary we must consider

the paired cephalic appendages, gill arches and fringing processes

as various modifications of one set of serially homologous struc-

tures, the pectoral and pelvic fins of modern vertebrates, being

a comparatively recent specialization of a partially fused series of

such appendages in the trunk region.

I assume, therefore, that the highly specialized condition of

the visceral arches and appendicular structures of modern fishes

arose through the modification of paired segmental appendages.

Even in the more remote ancestors, such as the Phyllopoda,

Trilobita, Phyllocarida and Merostommata. These appendages

varied greatly in form and function in different animals, and in

different regions of the body in the same animal.

In the ostracoderm type, we may assume that certain ones of

the anterior cephalo-thoracic appendages were retained as oar-

like swimming appendages. Two or three pairs were retained

about the mouth followed by several pairs of respiratory append-

ages of an unknown character.

856 THE AMERICAN NATURALIST. [Vor. XXXVII.

The trunk appendages were represented by the series of

fringing processes.

We may assume that the evolution of the true vertebrates

was accompanied by the fusion of the paired mouth parts into

unpaired upper and lower jaws, by the further specialization

of the gill pouches, the reduction of the free cephalic append-

ages to such embryonic structures as certain oral papilla or

tentacles, external gills, and the larval balancing organs seen

in certain amphibian larva, and by the fusion of the abdominal

appendages to form the lateral fold from which in turn arose

the pectoral and pelvic appendages.

The above interpretation of the origin of paired appendages

retains the strong points of both the gill arch, and the lateral

fold theories, without the weak points of either. It gives us

precisely what Gegenbaur claims has heretofore been lacking in

the lateral fold theory, namely: (1) a reason for the existence

of the primary fold ot ectoderm that initiates the formation of

the lateral fold; (2) a reason for the migration into it of seg-

mental detachments of muscle, nerve and cartilage; and (3) a

primary function for the lateral fold out of which a set of loco-

motor organs could be logically developed.

We may explain the presence in the ostracoderms of two or

more pairs of cephalic appendages that are not homologous with

one another or with the pectoral fins, without being forced to

assume that such highly specialized structures are nothing but

movable spines or cornua, or flexible flaps, without any known

antecedent function or significance. We may agree with

Gegenbaur that there is a certain homology between gill arches

and specialized portions of the lateral fold, without assuming the

extensive migrations of gill arches demanded by his theory, and

we may agree with Dohrn that structures homologous with gills,

or associated with them, extended far back into the trunk region

without assuming that true gill arches and visceral — were

present there.

IV. MODE or Lire OF THE Oereaconenes. — We may

form some idea of the mode of life of the Ostracodermata by

considering the shape of the body and its armor, the nature of

the appendages, and the position of the eyes and mouth.

No. 444.] PTERASPIDE AND CEPHALASPIDA. 857

That the Cephalaspidz were sluggish animals is indicated not

only by the clumsy shape, and large size of the heavily armored

head, but also by the absence of an axial skeleton, and by the

feeble development of the trunk and the dorsal and caudal fins.

The overlapping of the large scales on the dorsal surface of

the trunk in Cephalaspis, and the presence of minute ventral

scales, indicate some freedom of movement in a dorso-ventral

direction and a restriction of those lateral movements so essen-

tial to sustained swimming when well developed appendages for

that purpose are absent.

' The dorsal fin was short and low and covered with close set

scales that would allow but little freedom of movement.

The Cephalaspidz, therefore, were in all probability bottom

feeders. Any one that has watched Limuli, both old and young,

ploughing slowly through the soft mud and sand, leaving little

more than their median ocelli and lateral eyes exposed, could

hardly avoid the conclusion that many of the trilobites and Ceph-

alaspidæ, whose eyes are placed high up on the convex surface

of similar shovel-shaped heads, must have moved about on the

bottom in a similar manner.

But Limulus frequently crawls with considerable rapidity

over the surface of shallow bottoms, or turning on its back with

the aid of its caudal spine, even the largest and heaviest female

may leave the bottom and swim slowly away with legs, operculum

and gill covers beating the water in unison with oar-like strokes.

. The young larvz, especially in the trilobite stage, swim very

persistently in this inverted manner. It is also well known that

Branchipus, Apus and many copepods swim in this position,

and there can be but little pon that ns: trilobites and

merostommata did the same.

The Cephalaspidee were certainly disproportionately heavy at

the anterior end, so that any attempts to swim by movements

of the trunk alone would tend to push the head deeper into the

mud or sand, a tendency that would be increased by the sharp

downward slope of the anterior dorsal surface. It is also proba-

ble, judging from the shape of the head and trunk, that its

centre of gravity was situated above the mass centre, so that if

the animal did succeed in getting off the bottom, it would have a

858 THE AMERICAN NATURALIST. [Vor. XXXVII.

tendency to topple over and thus bring the ocular surface under-

neath. In this position, however, owing to the convexity of the

surface, now underneath, any forward movement would tend to

elevate the head and thus counteract the effect of its greater

weight and volume.

When the Cephalaspide, therefore, left the bottom they

probably turned over on to the ocular surface and made their

swimming excursions in much the same manner as the above

mentioned arthropods.

Whether the locomotion of the Cephalaspidz was aided by

the movements of branchial appendages concealed beneath the

arching under surface of the head, can only be determined by

the study of more perfect remains than have been as yet dis-

covered. Certainly the very strongly concave under surface

of the head indicates the presence there of some organs freely

movable and of considerable size, projecting ventrally and later-

ally from the middle region of the head.

What we have said in regard to mode of life of the Cepha-

laspide will apply, with slight modifications to other members

of the ostracoderms.

The Asterolepidz, with their powerful, but badly constructed

and impractical appendages and large centrally placed eyes, rep-

resent the extreme development of the free swimming type.

It is quite certain that in some genera the attachment of the

slender swimming appendages close to the ventral side, and the

very highly arched dorsal surface, must have made it extremely

difficult, if not impossible, for these animals to swim with the

ocular, or dorsal surface, uppermost.

The methods of locomotion in the ostracoderms iid the

arthropods thus have a double value for they show us that ani-

mals like Limulus, the Phyllopoda and probably the Trilobita and

the Merostommata, adopted when swimming a vertebrate posi-

tion with the neural side uppermost, and that the ostracoderms

must have frequently assumed the typical invertebrate position.

In Pteraspis, Cyathaspis and Tolypelepis, the lateral eyes are

probably absent, or were covered with such thick layers of the

shell as to render them nearly useless. The trioculate median

eye, however, is well developed and is represented in Pteraspis

No. 444.] PTERASPIDA AND CEPHALASPIDE. 859

by a small pit on the under side of the dorsal shield, in Tolypele-

pis by an inconspicuous smooth spot on the outer surface, and

in Cyathaspis by three obscure markings. These conditions

indicate a considerable degeneration of the visual organs in these

genera and must have profoundly modified their mode of life

toward one of less activity and greater simplicity. We have

here a condition approaching that of the lampreys, where the

lateral eyes may be nearly functionless, while the median ones

attain a degree of development that is hardly exceeded by that

in any other vertebrate.

In Cephalaspis, the mouth was almost certainly situated high

up on the vaulted under surface of the head and the character

of the mandibles, as well as the small space into which such

organs must have been crowded indicates that the mouth was

very small. Similar conditions prevailed in Tremataspis, Bothri-

olepis and other members of the ostracoderms. Under these

conditions and with their necessarily slow and clumsy move-

ments, the ostracoderms could not have been rapacious animals.

The position of their eyes, whether well developed or not, points

with equal decisiveness to the same conclusion, for whether

swimming or crawling, they could not see where, or when, to

seize their prey, because their eyes would always be behind their

own body. They must have been dependent on highly spe-

cialized gustatory, or tactile, organs situated near the mouth.

As a parasitic life for such animals is out of the question, it is

probable that they lived on the soft bodied animals or decom-

posing organic matter that could be exposed or forced into the

mouth as they slowly ploughed their way through the soft mud

or sand.

DARTMOUTH COLLEGE, HANOVER, N. H.

July 10, 1903.

EXPLANATION OF PLATES.

PLATE I.

Fic. 1.—Cephalaspis murchisoni, showing several lobed fringing processes, /7. British

useum.

Fic. 2. Piste of Cephalaspis. Outer layers of the shell are absent, showing outlines of

bony trabeculz below the lateral openings and the post-orbital vall

Fic. sc outline of the lateral openings filled with polygonal pines. British

1 th

Fic.

Hide Mohan.

Fic. 5.— Ventral uli of the trunk of Cephalaspis, showing the cloacal opening.

PLATE II.

Fic. 6.— Soret view of the trunk of Cephalaspis nes wert den the fringing processes --

their articulation with the lateral trunk scales. tish Mus

Fic. 7.— : Hind and anterior poc of the trunk o arabna MEE arrangement of es

trunk scales. British Muse p

: 24 1 L B f* 4% 2. pis R? e British Museum.

eq:

Fic. 8.— Side view

No.444] PTERASPIDÆ AND CEPHALASPIDA. 863

BIBLIOGRAPH Y.

ALTH, ALOIS V.

x Ueber die palaozoischen gebilde Podoliens, etc. Abhandl. k. k.

geol. Reichsanstalt Wien

'55. On the Tilestones, or Downton Sandstones. Proc. Geol Soc.,

London. Vol. 12

CLAYPOLE, E. W.

'92. On the American Pteraspidian Palewaspis. Quart. Jour. Geol. Soc.

Vol. 48

DEAN, BASHFORD.

'95. Fishes, Living and Fossil. London and New York.

Hux ey, T. H.

'58. On Cephalaspis and Pteraspis. Quart. Jour. Geol. Soc. Vol. 14.

HIEEN UM E. VON.

Die Grauwackenschichten von Liev. ünd Ehsthland. Bull. de. la

Soc. Imp. d. Naturalistes Moscou. T. 27, No. 1.

esp eoe e.

Ueber die Stammform der Wirbelthiere. Sitzungs-Berichten der

gesell. naturf. Freunde Berlin. No. 7.

JACKEL, O.

: Ueber Coccosteus, etc. "Site. Bericht. Gesell. natur. Freunde, Ber-

lin. = No. 4$.

"m RUDOLPH

Ueber die beiden Arten Vesp d lloydii and lewissii, Agassiz.

Haidinger's Naturwiss. Abh. Bd.

KUNTH, A.

'72. Ueber Pteraspis. Zeitschr, deutsch. geol. gesell. Bd. 24.

LANKESTER, E. Ray.

'68. A Monograph of the Fishes of the Old Red Sandstone of Britain.

Part I. The Cephalaspide. Paleontographical Society.

LANKESTER, E. Ray.

"JO. Part II. (concluded). The Cephalaspide.

LINDsTROM, G.

'95. On the Remains of a Cyathaspis from the Silurian Strata of Goth-

land. R. Swedish Acad. Science, Bd. 21, 1-3, Stockholm.

PATTEN, WM.

'89. On the Origin of Vertebrates from Arachnids. Quart. Jour. Micr.

Scd. Vol. 31, Part 3.

PATTEN, WM

93. On the Morphology and Physiology of the Brain and Sense Organs

of Limulus. Quart. Jour. Micr. Sci. Vol. 35, No. 137.

864 7HE AMERICAN NATURALIST. (Vou. XXXVII.

PATTEN, WM.

'7"&. On Structures Resembling Dermal Bones in Limulus. Anat. Anz.

Bd. 10, No. 14.

PATTEN, WM. and REDENBAUGH, W. A.

'99* = The Endocranium of Limulus, Apus, and Mygale. Jour. Morph.

Vol. 16, No. 1.

PATTEN, WM. and REDENBAUGH, W. A.

'99». Ibid. The Nervous System of Limulus Polyphemus.

PATTEN, WM. and HAZEN, A. P.

The Development of the Coxal Gland, Branchial Cartilages, and

Genital Ducts of Limulus. Jour. Morph. Vol. 16, No. 3.

PATTEN, WM.

:01. On the Origin of Vertebrates, with Special Reference to the

Ostracoderms. Address before the V. International Congress of

Zoologists, Berlin.

PATTEN, WM.

:03". On the Structure and Classification of the Tremataspide.

Mem. Acad. Imp. Sci. St. Pétersbourg. Vol. 13, No. 5.

PATTEN, WM.

03". On the Appendages of Troas Amer. Nat. Vol. 37,

No. 436.

PATTEN, WM.

The Structure of the Ostracoderms. Science. Vol. 17, No. 430.

ROREM FERDINAND

Palzoteuthis. S E PERE IV.

Rouwon, J. V.

'93. Die obersilurischen Fische von Oesel. I. Theil. Mém. Acad.

Imp. Sci. St. Pétersbourg. Tome 38. No.1

Rouon, J. V.

9 Die obersil. Fische von Oesel. II. Theil. Ibid. Tome 41, No. 5.

Ronon, J. V

'93». Zur Kenntniss der Tremataspiden. Bull. de l'Acad. Imp. de sc-

de St. Pétersbourg. Tome | :

Ronon, J. V.

'95. Die Segmentirung am Primordial-Cranium der Obersil Thyesti-

den. Verhandl Kaiserl Russ. Mineral. Gesell. St. Petersburg.

Bd. 33, No. 2.

Ronon, J. V.

'99. Ueber die Parietalorgane and Paraphysen. Sitzungsbr. d. Kónigl.

rage Gesellschaft der Wissenschaften.

SCHMIDT, F.

"13. iae Remarks on Pteraspis. Geol. Mag. Vol. 1o.

TRAQUAIR, R. H.

'88. On the Structure and Classification of the Asterolepide. Mag-

Nat. Hist. Vol.2

No. 444] PTERASPIDA AND CEPHALASPIDA. 865

WHITEAVES, J. F

':8 On some remarkable Fossil Fishes from the Devonian Rocks of

Scaumenac Bay, P. Q. with Descriptions of a new Genus and

three new Species. Can. Nat., Vol. 10, No. 1.

WHITEAVES, J. F.

':86. Illustrations of the Fossil Fishes of the Devonian Rocks of Canada.

Trans. Roy. Soc. Can. Section 4.

WOODWARD, A. S.

'91. Catalogue of the Fossil Fishes in the British Museum.

AN OUT-DOOR EQUIPMENT FOR COLLEGE

WORK IN BIOLOGY.

JAMES G. NEEDHAM. .

THE study of living nature is chiefly manifest at the two

extremes of our educational system in the establishment of

public school gardens and university summer laboratories. Of

what is being done in this line between the grades and the

graduates less is heard. Perhaps it is because the high school

and the college are less prone to advertise themselves by novel-

ties in their educational programs: perhaps, because they are

seeking to develop new methods instead of creating them full

fashioned outright.

Thus far, the grades that have acquired good school gardens

seem to have the better of it. For, besides having established

an inexhaustible and ever accessible base of supplies for nature

study work, they have at the same time set the pupils enthusi-

astically to educating themselves, and by the historic method —

by doing over again in the garden such work as was done when

the mental fibre of the race was first toughening.

The situation in high schools seems less fortunate. While

many of them have books outlining ecological phenomena, very

few of them have proper opportunity for the study of such

phenomena. The gtounds of the average high school are the

most drear and barren waste within the city limits. The life

that belongs to the soil has been exterminated. Only trees are

cultivated, and these are not often native trees. The birds are

English sparrows: the animals are stray cats, mice and roaches.

If there be parks in the city no natural assemblage of native

plants and animals is to be found therein: for the taste of

the average town has not evolved beyond the painting-the-lily

stage. By means of long rides on street cars at infrequent and

uncertain intervals classes are taken out to see some rem-

nants of nature and are shown things. The best thing found

7 867

868 THE AMERICAN NATURALIST. [Vor. XXXVII.

is often fresh air. How far this falls short of the individual

work that may be done in the school garden or at the labora-

tory table! The too common result of giving suggestions for

seeing things that there is no opportunity for seeing is a relapse

to book-and-recitation methods.

The field laboratories of the universities offer for the most

part excellent opportunities both in natural environment and in

facilities for its study ; but these are far from home and available

| k es

er. Ne

asd ios

«ew i

oae

e a NND y

Fic. 1.

only in the summer vacation, and the rank and file of university

students miss altogether the sort of training they afford.

Very little is heard of the movement toward the utilization of

living nature in college work, yet it is exerting a powerful influ-

ence over present methods. The anatomical work which was

the beginning, the continuation and the end of the old time

courses laid out by morphologists — and morphologists must be

given credit for having laid out the first practical laboratory

courses — no longer monopolizes all the time of the general

student. The elements of biology that make for culture are

far from being confined to the dissecting table. And more and

more college students are being encouraged to study nature in

the field and by those methods that in the last century yielded

our most important generalizations.

No. 444.] OUT-DOOR EQUIPMENT. 869

This article is intended to set forth some of the ways in which

living nature is utilized at Lake Forest College, where the work

is not that of a summer season spent far from home, but that of

daily college life, done about the doors of the college halls; not

the research work of graduates, but the general-culture work of

undergraduates ; not for the few who can afford it, but for the

many who need it.

The plan here has been to use things near at hand. Ina

large measure, therefore, the situation accounts for the things

that have been done. The accompanying map (Fig. 1) of the

campus and its immediate environs shows some of the deter-

mining physiographic conditions. The campus is situated on

the scarcely perceptible eastward slope of a terminal moraine,

which parallels the shore of Lake Michigan, and is covered with

fine oak woods. It is intersected by sharp ravines that have

been cut by puny postglacial streams. The ravine shown in the

map is scarcely more than a mile in length, and attains a depth

of about 70 ft. where it reaches the lake level at its mouth. On

the ridge at the head of these streams is a series of shallow

ponds, many of them temporary, and some of them doubtless

occupying old * buffalo wallows." Half a mile farther westward

the Skokie winds its leisurely course through the marshes at the

foot of the more abrupt westward slope of the moraine. The

Skokie and its marshes, the ponds, the upland woods, the

ravines, the crumbling outer face of the “bluff ” and the Lake

Michigan beach, each furnishes its own peculiar fauna and flora,

and all are within easy walking distance of the campus. But

the woods, the ravines, and an artificial pond are upon the

campus, and are as easily entered for study as are the labora-

tories: and, naturally, these are most used. In the map c is

college hall, the building in which the biological laboratories are

located, and > is the pond — perhaps the most important singie

feature of biological equipment —a veritable aquarium, perma-

nently stocked and self sustaining — teeming with a multitude of

forms of animal and plant life. Its proximity may be judged

from the view shown in the first figure of the plate, which view

was taken from the window of the general laboratory. The

heavily shaded portions of the campus (a and 4) comprising sev-

870 THE AMERICAN NATURALIST. (VoL. XXXVII.

The winter house.

No. 444.] OUT-DOOR EQUIPMENT. 871

eral acres extent, have been set apart by the trustees for the

purposes of a biological garden. Both plots are wooded and

traversed by deep ravines. Plot æ being at the front of the

campus is to be used chiefly for ornamental planting of native

shrubs and trees. Plot 4 is more secluded, and is more freely

used for the ends of biological instruction. It contains the

pond, and the winter house shown in the third figure of the

plate. Its ravines exhibit, especially on the shady side, a luxu-

riant tangle of shrubbery and vines, of flowers and ferns, so

little disturbed by civilizing influences that the native Cypripe-

dium regine and Adiantum pedatum — usually the first victims

of their loving friends — still flourish there abundantly. The

first thing done in this plot was taking measures to preserve

the native species still present, and to restore to it a number

that had been already exterminated. The next thing, was the

assembling of those biological and ecological types especially

useful for illustration in general course work. Plants are raised

here not for themselves alone but for the sustenance they afford

to the forms of animal life desired to be retained with them.

Care has been taken to provide a constant succession of wild

fruits for birds, the proper plants to sustain aphid colonies with

change of hosts, and for the food of particular animal species —

even for a solitary aboriginal family of woodchucks. Thus,

where nature had done much, and where the material. needed

was all near at hand, attention has been given to making things

as readily available for study in the field as they are in the lab-

oratory, to the end that field studies that are really worth while

might be undertaken.

But two things that are obviously artificial have as yet been

introduced into the garden: a plankton apparatus and a winter

house. Probably no college teacher has witnessed a good plank-

ton catch at a summer laboratory or field station without wish-

ing that such quantities and variety of the simpler organisms

might be available for his class-room work. Many of them may

be obtained, to be sure, with any simple sort of towing net ; but.

an apparatus that will get all the life at a definite depth and

serve for quantitative measurement of it has generally been

accounted too complicated and too expensive for the equipment

$72 THE AMERICAN NATURALIST. (Vor. XXXVII.

of an average college. But an apparatus that will do all these

things and that will gather in a few minutes use more plankton

than can be even qualitatively examined in a laboratory period,

was constructed by several student assistants a Lake Forest

College at a cost of about ten dollars.

The exterior of the apparatus is shown in Fig. 2 and the

net itself is seen in the second figure of the plate. A net of the

usual form was made of no. 20 silk bolting cloth. A large milk

can with loose breast and heavy steel cap was used to sustain the

net and to hold the slack water in which it should rest while

Le, +

Fic. 2.—A simple pl pp

receiving the inflow stream. The net itself is suspended from

two perforated circular brass strips soldered within the upper

and lower edges respectively of the breast. To a short tube

inserted into one side of the breast is attached the waste pipe g.

The intake pipe 7 is attached to the cap; but a three-way cock

is inserted in the couplings at vc, and a short pipe > is attached

to its lateral aperture, with a spray nozzle on the end. The

cock may be turned so as to stop the flow altogether or to direct

the stream laterally through the spray pipe g (as shown in action

in the second figure of the plate) or to direct it through the cap

No. 444.] OUT-DOOR EQUIPMENT. 873

into the net and out through the overflow pipe g. The rate of

flow is readily measured at the end of the overflow pipe g. The

net is placed for use on a platform at the foot of the dam that

confines the campus pond and the water is brought to it by

siphoning over the dam with a long garden hose. The upper

end of the hose is fitted with a funnel-shaped, screen covered

intake, and this can be held in any position and at any depth as

long as desired. The flow is perfectly uniform, even such

active and light loving forms as Corethra and Corixa being

readily taken down the pipe. This simple apparatus has made

itself an indispensable adjunct to the work of several courses.

Its use at once revealed the presence of vast numbers of Din-

obryon, Notholca, Polyarthra, etc. not previously known to

exist in the pond.

Near to the plankton platform in the ravine is the winter

house, in which are kept the plankton and other field apparatus

and garden tools. Here are proper quarters also, for live ani-

mals desired for study during the winter: tanks in the floor for

salamanders, frogs, crayfishes, earthworms; shorts-bins on the

wall for meal worms, sand beds for seeds and for plant cuttings,

etc. This most useful little house, shown in the third figure

of the plate, was built to fit the landscape and not to disfigure it.

Individual apparatus for field work is much more simple.

Air and water nets, cyanide bottles, jars and vials, garden

trowels, etc., are supplied every student ; for field work at Lake

Forest is just as individual as is laboratory work, and just as

definite results are expected from it. How much less expensive

is this out door equipment than the indoor equipment that all

the colleges have !

But the first requisite of satisfactory field work on the part of

college students is neither books nor apparatus, but living

nature near at hand. Fortunately, the native plants that are

needed will grow near at hand, if given room, and their natural

animal associates — at least those most useful for study — will

come and dwell again with them. Fortunately, also, they may

be made to add both beauty and interest to the grounds devoted

to them. Though the environment at Lake Forest is exception-

ally favorable for all this sort of work, it is to be borne in mind

874 THE AMERICAN NATURALIST. [Vor. XXXVII.

that nine tenths of what is undertaken here could, with proper

provision, be done anywhere. Ecological types are as widely

distributed and as available for study in different places as

are morphological types: and biological phenomena are as tan-

gible and as real as are systematic, while certainly not of less

interest or educational value.

BIOLOGICAL DEPARTMENT,

Lake Forest College.

PRELIMINARY REPORT ON THE “PALOLO”

WORM OF SAMOA, EUNICE

VIRIDIS (GRAY).

W. McM. WOODWORTH.

SINCE a Monograph of Samoa would not be complete without

some account of the “ Palolo,” at Dr. Kramer’s request I have

prepared the following summary, reserving for a subsequent

publication a detailed account of the “Palolo” and other

annelids of the coral reefs of the Pacific. In this preliminary

paper I can only touch upon historical matters and the often

written story of the “rising” and “fishing” of the “ Palolo,”

referring the reader to the publications of Collin; Friedlaen-

der, Kramer,* and Ehlers.’

The “ Palolo " has been known to naturalists for more than half

a century and much has been written about it in a fragmentary

way. It was, however, during the period of Kramer’s investi-

gations in Samoa that its true history was brought to light, and

''This provisional account prepared for Krümer's monographic work on the

Samoan Islands (Krämer, Augustin, Die Samoa Inseln etc. Stuttgart, E. Nügele,

1903, Bd. 2, pp. 399-403) and translated js him into German, is repri po here

with some changes and corrections. The author has in preparation an extended

study of the life history, morphology and disicibution of the ** Palolo,” ii ‘allied

Eunicide.

? Collin, A. Bemerkungen über den essbaren Palolowurm, — viridis

beds Appendix to Krümer's Bau der Korallenriffe pp. 164-17

Friedlaender, B. Uber den sogenannten Palolowurm. Blot Centralblatt.

Bd. 28, ‘i gue 1898. — /dem, Notes on the Palolo. Jour. Polynesian Soc.

Vol 4-46; Wellington, N. Z. 198. — 72e», Nochmals der Palolo etc.

Bil. nra Bd. 19, pp. 242-269.

Jher den Bau der Ki i bt ind die Plankion- erteiing an der

en. Kiel

Mise Küste nebst mi ia tps Bemerkungen. und Leipzig 1897.—

Idem, barones d Biolog. Ce: iir cioe Bd. 19, pp. 15-30. 1899.—

Jdem, Paloloun mim: im October und November 1898 in Samoa. /did.

Pp- 237-239.

5 Ehlers, E. s Palolo (Eunice viridis Gray). Nachr. A. Ges. Wiss. Göt-

tingen. Math.-naturw. KT. 1898. pp. 400-415.

875

876 THE AMERICAN NATURALIST. [Vor. XXXVII.

much of our knowledge of this interesting worm is due directly

to him and to the stimulus of his work. The first extended

account was written by Collin! as an appendix to Kràmer's

earlier work on Samoa. In this account Collin, with previous

writers, considers the * Palolo” to be the posterior part of

Lysidice viridis (Gray), a few detached heads of which had

from time to time been taken with the “ Palolo " at the ‘fishing’

season, and as no other annelid heads were taken with the

“ Palolo”’ and all “ Palolo " were headless, it was natural, for want

of better evidence, to ascribe the “Palolo” to the genus Lysi-

dice. The discovery of the origin of the “ Palolo” was made

independently by Kramer and Friedlaender, although the latter

was the first to publish an account of his investigations.? Fried-

laender succeeded in obtaining from the reef rock at Samatau

several specimens of **Palolo" together with the head ends of

an annelid of different appearance and much larger size belonging

to the genus Eunice. His material was afterwards studied by

Ehlers? who recognized an extreme case of sexual dimorphism

and showed the “ Palolo” to be the epitokal posterior portion of

Eunice viridis (Gray). Ehlers says, “ Ich ergiinze das im Voraus

damit, dass ich die Eunice, die nun den Namen Eunice viridis

(Gray) erhält, in den Kreis der Eunice siciliensis Gr. bringe und

an ihr die Ausbildung des “ Palolo " als eine Form der Epitokie

auffasse, wie sie zum ersten Male aus der Familie der Euniciden,

und in ihrer Besonderheit abweichend von allen Erscheinungen

der Epitokie, die von Borstenwürmern bekannt sind, sich darstellt.

Demnach ist in der Art eine atoke und epitoke Form, in der

letzteren eine atoke und epitoke Kórperstrecke zu unterscheiden."

It was my good fortune to be at Levuka in the Fiji Islands

during the * rising " of the ** Palolo " * in November, 1897, where

I gathered much material and information, and in the following

year went to Samoa to learn more about the history of this

! Op. cit.

2See Thilenius, G.' Bemerkungen zu den Aufsätzen der Herrn Krämer und

Friedlaender uber den sogenannten Palolo. Biol. Centralblatt, Bd. 20, pp. 241-

242, 1900

3 Op. cit.

4 Bololo, pronounced Mbololo in Fijian.

No. 444] PALOLO WORM OF SAMOA. 877

mysterious worm. I arrived at Apia on October 20 and was

fortunate in meeting Dr. Kramer who placed at my disposal the

notes he had collected during three years in the islands. I made

my headquarters in the village of Falelatai on the South side of

Upolu a little to the eastward of Samatau where Friedlaender

obtained his material. After several days of fruitless search on

the reef between Samatau and Falelatai my native friends took

me to a shallow bay called Fagaiofu about two miles east of our

village. The bay lies between two small promontories and is

about one quarter of a mile wide, the distance from the shore to

the edge of the fringing reef, which fills the bay, is not more

than 150 meters. The place is so shallow that at low tide one

can wade from the shore to the edge of the reef. The reef plat-

form, which is composed chiefly of dead coral and honeycombed

reef rock, is interrupted by two narrow deep channels or passages.

The reef at Fagaiofu proved to be literally alive with “ Palolo.”

They were discovered by prising off, with a crowbar, masses of

the rock at the edges of the channels. They could be seen

dangling from the freshly exposed surfaces and wriggling free

into the deeper water of the channel to be carried seaward by

the retreating tide, to the astonishment of my natives who had

never seen the “ Palolo ” before the time appointed for its appear-

ance — this was three days before. Owing to the great length

of the entire worm, its fragile structure and intricate association

with the cavities of the honeycomb rock, the operation of freeing

unbroken specimens is a delicate one. With the aid of chisels

and forceps I succeeded with great difficulty in obtaining, in

addition to other material, three worms complete from head to

tail.

My experiences confirm the discoveries of Kramer and Fried-

laender as to the origin of the “Palolo.” The accompanying

figure, which is drawn to scale, shows the complete animal, the

broad anterior atokal portion being sharply marked off from the

more attenuated and much longer posterior epitokal part which,

when free-swimming, is known as the “Palolo.” The total

length averages 400 mm., about one fourth of this length being

in the anterior atokal part. 429, 359, and 250 atokal segments

were counted; the first two in male specimens the latter in a

878 THE AMERICAN NATURALIST. (VoL. XXXVII.

female. These figures are not accurate as a dense gelatinous

secretion in the posterior part of the atokal region makes it diffi-

cult to count the segments. The greatest diameter of the atokal

region is 4 mm. and that of the epitokal region 1-14 mm. dimin-

ishing gradually at the anal end, and more abruptly at the junc-

tion of the atokal and epitokal parts. The color of the male is

myers

ere

Ty SOF

SP e v g

XC fe.

aaepe m

ACT oO x

OTET AP UNO,

POEM pea?

eh EP

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A ETL L

Sapa’, go: $39.

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Wen Pe "AXI [SN

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M. ATE à

oon m

Eunice viridis (Gray). 'The narrow posterior epitokal part when

1 had RE H H $a L h “ Palolo.”

reddish brown, that of the female bluish green. These colors,

which are very marked in the epitokal portions are due to the

colors of the sperm and ova; after the discharge of these ele-

ments the collapsed integument is translucent and colorless.

These distinctive sexual colors are also present in the atokal

parts but are not so marked, the female being more greenish in

No. 444] PALOLO WORM OF SAMOA. 879

hue than the male; the colors here are integumentary. Each of

the epitokal segments bears on its ventral surface a prominent

pigmented spot, the * Bauchauge " of Ehlers. These eye spots

can be traced into the atokal part through about 20 segments,

diminishing in size toward the anterior end; they are lacking on

the anal segment and are usually absent in 2—6 of the preanal

segments.

A similar swarming of marine annelids, and at corresponding

seasons, is known for other islands of the Pacific, though the

worms have not everywhere been identified. ^ Powell' speaks

of them in the Gilbert Islands where they are known to the

natives as ze nmatamata and Codrington ? gives a detailed account

for Mota in the Banks Islands where they are known as un.3

Brown * mentions an annual appearance of a “Palolo” on the

East coast of New Ireland, and the wawo of Rumphius which

occurs at Amboina in the Moluccas is doubtless the same, as has

been pointed out by Collin. Seeman ê mentions the occurrence

in the New Hebrides, and it is known in Fiji and Tonga. It is

reasonable to suppose that a systematic search would show the

** Palolo ” or some allied form to have a wider distribution in the

coral reefs of the Pacific than has been as yet recorded. That

the annelid is best known from Samoa and Fiji is accounted for

by these two groups of islands having been most visited and

longest inhabited by whites. It is significant also that such

records as we possess from other places, though meager, have

come to us through the missionaries, the pioneers of intelligent

whites in the islands of the Pacific.’

! Powell, T. Remarks on the Structure and Habits of the uc Reef Annelid

Palolo viridis. Journ. Linn. Soc. London. Vol. 16, pp. 393 3-

? Codrington, R. H. Zhe Melanesians. Studies in their unas and Folke

Lore. Oxfor

3,Doubtless the a’007z of McIntosh.

4Brown,G. Notes on the Duke of York Group, New Britain and New Ireland.

Journ. Roy. Geog. Soc. Vol. 47, pp. 137-150. 1877.

5 Op. cit.

9$ Seeman, B. Viti. An Account of a ee Mission to the Vitian or

Fijian Islands in the Years 1860-1867. Cambridge 1

7Cases of swarming associated with extreme irai dimorphism have been

desccibed fora ence from Florida (Mayer, A. G., Bull. Mus. Comp. Zoil Vol.

37, 1900) and for one of the Lycoridz from Japan ( idm A., Journ. Coll. Sci.

imp. Univ. Tokyo, 1903).

880 THE AMERICAN NATURALIST. [Vor. XXXVII.

The “Palolo” makes its appearance in Samoa in the months

of October and November during the last quarter of the moon.

This is the time of the lowest or spring tides when the reef flats

in shallow places are uncovered or only awash, and at this sea-

son the sun is nearest to the zenith. I must reserve for my

final paper a discussion of the causes of the swarming of the

* Palolo " and will only say here that I am inclined to believe in

some thermotropic or heliotropic reaction of the eyes borne on the

ventral segments of the epitokal part of the worm. These eyes

have recently been studied histologically by Hesse! on material

collected by Kramer. Hesse states that from their structure

the eyes probably do not form images, but function rather in

reacting to light of different intensities, the direction of light

and possibly to different colors. It is significant that these eyes

are found only on a few of the posterior segments of the atokal

sedentary part and are not well developed; while on the other

hand they are highly developed on all but the anal segments of

the epitokal active part which leads such an ephemeral free

existence.

This spring season is recognized as the period of ripeness and

sexual activity throughout the Pacific Islands and where the

* Palolo " occurs the season and even the months are named for

it. All of the many other kinds of annelids inhabiting the reefs

are sexually mature as shown by the extensive collections made

by Kramer and myself in Samoa and Fiji and this is true also of

the reef fauna in general. The spawning time of the land crabs,

the occurrence of certain fish, etc., is reckoned by the natives as

so many days before or after the * Palolo," and so for the appear-

ance of blossoms the ripening of fruits and tubers. In Samoa

the “Palolo” season is called /aumafamua (i. e. the time of

much to eat), in the Banks Islands they say **/au matua the

season of maturity, yams can be eaten.’”

The ** Palolo-time " in Samoa embraces three successive day s.

When in the last quarter of the moon in October and November,

more especially the latter, the water on the **Palolo-grounds "

! Hesse, R. Untersuchungen uber die Organe der Lichtempfindung bei

niederen Thieren. V, Die Augen der polychaeten Anneliden. Zeitschr. wiss-

Zool., Bd. 65, pp. 459, 1899.

? Codrington, of, cit.

No. 444] PALOLO WORM OF SAMOA. S81

has a turbid or riled look, with floating patches of scum, the

natives know that two days later the “ Palolo " will ‘rise.’ This

first day is called sa/efu. The second day is marked by the

swarming of a small annelid, headless like the ** Palolo," and the

sexes distinguished by the same yellow and greenish tints. This

day is called motusaga. The third is the /a/e/ega when the

* Palolo" swarms and the natives come many miles to the

favored places to gather it. With “ Palolo” of the /atelega day

many of the small annelids of the motusaga occur, and a few

** Palolo" appear on motusaga day. A microscopical examina-

tion of the sa/efu scum shows it to consist of a gelatinous slime

in which are grains of sand, appendages, fragments and casts of

Entomostraca and a varied: detritus of the seething life inhabiting

the reefs, including many ova of various kinds in different stages

of segmentation. The sa/efu may be looked upon as a manifes-

tation of the awakening of the “ Palolo " previous to its swarm-

ing or marriage-swim ; an annual activity of countless numbers

of annelids resulting in a discharge into the water of the deposits

accumulated in the galleries and crevices of the reef-flats. The

small annelid of motusaga day is Lysidice falax Ehlers, the Z.

viridis (Gray) to which the “ Palolo " was so long ascribed.

CAMBRIDGE, Mass.

August, 1902.

FURTHER NOTES ON. THE HABITS OF

AUTODAX LUGUBRIS.

WM. E. RITTER.

Wiru the information about the breeding habits of this sala-

mander obtained by Mr. Miller and myself during the summer

of 1899! we supposed that in succeeding seasons we should

have little difficulty in securing sufficient eggs to enable us to

make a fuller study of its development. The next summer, con-

sequently we, and particularly Mr. Miller, searched for the eggs

constantly and carefully under logs and rocks and in half

decayed stumps all about the San Francisco Bay region, but

not a single egg rewarded our efforts. I have kept up the

quest each year since, but not until the present summer has

anything but failure come of it. Now, wholly by accident, the

usual breeding place of the species, for this locality at least,

appears to have been found. This turns out to be holes in

trees. The one positive, and one doubtful, instance reported by

us of egg-laying in the ground seems to have been exceptional.

In caring for the oaks (Quercus agrzfolia) on the campus of

the University of California this summer the trees have been

subjected to a treatment they have never before received.

This has consisted in the careful cleaning out of the decayed

wood and foreign accumulations from all accessible corners and

cavities, and of painting the walls of these with coal tar, and

then filing the smaller cavities with Portland cement. The

men engaged in the work have taken from these holes about 100

specimens of the salamander itself, and twelve bunches of its

eggs.

The form-habit of this oak commonly spoken of as the “live

oak," is well known to all who are acquainted with the environs

! Ritter, Wm. E., and Miller, L. H. A Contribution to the Life History of

Autodax lugubris Hallow, a California Salamander, Amer. Wat., Vol. 33, 1899,

p- 69r.

883

884 THE AMERICAN NATURALIST. (VoL. XXXVII.

of San Francisco Bay. The tree is short trunked, and diffusely*

branched, the contorted branches being wide spreading rather

than high reaching, so that a height of fifty feet is exceptional.

The salamanders are found as high in the trees as there are

holes suitable for their dwelling places. Some have been taken

from holes at the height of thirty feet at least. In some of the

largest cavities as many as twelve individuals were found; more

commonly, however, a hole contained two, or occasionally but a

single one.

Several facts indicate pretty clearly that in some cases all the

inhabitants of a single chamber were close of kin, constituted in

fact, a family. Where a considerable number of individuals were

together it invariably happened that the majority were small, and

the particularly significant thing is that the small ones were all of

about the same size, their length being about 50 mm. Besides

these individuals of minimum size there occurred in nearly all

the inhabited holes whether containing the small ones or not, a

few, usually two, individuals of maximum size. Then in addition

to those of maximum and minimum size, there were frequently

found, in the same hole, several others of intermediate size.

Those of minimum size constituted in all probability a single

litter, and were at this season of the year, viz., early autumn,

yearlings. Furthermore, I strongly suspect they were fre-

quently, if not always, the young of the individuals of maximum

size occurring in the holes with them. If this interpretation of :

the meaning of the presence of the small individuals together is

correct, it would follow that they had probably never yet in the

year of their existence left the tree in which they were hatched.

It is hardly to be supposed that they could make nightly excur-

sions to the ground and return to the same hole to spend the

day. As was pointed out in the paper above cited the species

is distinctly nocturnal. Asa rule the cavities occupied by the

animals while of ample dimensions for creatures of their size,

had but very small orifices by which entrance could be effected

from the outside world. Frequently the opening was not more

— than three or four centimeters across, barely larger than neces-

sary to admit the body of the full grown animal. But for the

directions the workmen had received to carefully clean to the

No. 444] AUTODAX LUGUBRIS. 885

"bottom all the cavities they could find, even though the

entrances had to be cut larger in order that the remedial

treatment might be administered, the dwelling places of the

salamanders would surely not have been found. Wide

mouthed cavities were rarely found occupied.

The egg clusters, each containing from twelve to eighteen

eggs, and each egg with its own pedicle about two centimeters

in length were usually suspended from an overhanging surface

where the parent was able to bring its body into contact with

them. This it did by winding itself around the egg clusters

in much the same way that several other species of Urodela are

known to do. More than one bunch of eggs rarely occurred in

the same hole.

Several observations make the question of the extent to which

parental care of the eggs and young may go in this species, one

of much interest. In the first place it appears that the male

and female may both together participate in the office. Prob-

ably a majority of the cavities in which eggs were found con-

tained two animals of maximum size. In some instances at least

these were certainly male and female. Whether this was always

or usually the case or not I am not yet able to say, since, this

point not having been raised until most of the animals taken

had been put together in a common terrarium, it was impossible

. to tell which were inmates of the same hole. It may be noted

in this connection that there are no secondary sexual distinctions

in this species, so far as I have been able to determine.

Again the animal seems to exercise more or less of an active

defence either of itself or of its eggs or both. Its unusually

large teeth has been a subject of comment by nearly all zoólo-

gists who have written about it,' and in the paper by Miller and

myself attention was called to Cope's statement that **this Auto-

dax is probably more capable of inflicting a bite than any other

of the American salamanders." At that time we were obliged

to say, however, that ** we had not been able to get any positive

! Cope for example, (The Batrachia of North America, p. 182), remarks that

*this curious genus is furnished with by far the most powerful dentition of any

existing salamanders, and resembles in this respect the genera of the Coal Meas-

ures, Brachydectes, Hylerpeton, and Hylonomus.”

886 THE AMERICAN NATURALIST. (VoL. XXXVII.

evidence on the point." The workman who found most of

the specimens in the holes tells me, without having been ques-

tioned with reference to the matter, that the old ones with the

eggs usually “showed fight " when he first came upon them.

He says they seized a stick or his finger when held toward them

with decided energy. I have, furthermore, the direct testimony

of another person, in this instance a student well experienced in

out of door natural history, that an individual once found by

him under a rock or log in the field, seized his finger in a dis-

tinctly vicious manner.

Thus is brought to light more of the peculiar traits that have

attracted the attention of several observers of this unsalamander-

like salamander. So far as I am aware this is the only urodele

that could properly be called arboreal.

Thése peculiarities are the more interesting in that there can

be no question as to the real affinities of the species. Its close

kinship to the other plethodons can not be doubted for an

instant. All that is unique about it, it has acquired so far as

can be seen all by itself. Experimental study on its behavior

should, consequently, yield unusually interesting results.

UNIVERSITY OF CALIFORNIA,

Sept. 15, 1903.

A IKIF TO THE TRUCHAS PEAKS, NEW

MEXICO.

WILMATTE PORTER COCKERELL.

Tue high mountains of Colorado are continued southward into

New Mexico without a break as far as the region of Santa

Fe and Las Vegas. Some distance north of this, however,

the range forks, presenting roughly the form of a reversed Y.

One of the arms of the Y is known as the Las Vegas

Range, the other as the Sante Fe range. The Truchas

Peaks are in the Sante Fe Range and represent almost the

southernmost extension of the Arctic-alpine zone.

The first week in August, 1902, I visited these peaks and

collected both flowers and insects. There is no road into the

region; and from Blake's ranch, which is just inside the Pecos

Forest Reservation we secured a guide, saddle horses and pack

animals, The first day we passed over the Las Vegas Range,

travelling through beautiful alpine meadows and great stretches

of spruce forests. Travelling through this country was very

delightful and the long vistas of mountain slopes, with here and

there the sparkle of an alpine lake, more than compensated for

hard parts of the trail where the horses must be guided carefully

as they jumped over the fallen trees. In places where the fire

had destroyed the trees the hillsides looked much like a board

covered with jack-straws and even our skilful guide sometimes

led us into boxes out of which our horses could not climb.and

we were obliged to retrace our steps and try a new path.

Our first camp was on the Mora fork of the Pecos River: this

is a small but very rapid stream that heads in the Truchas, and

it is famous through all northern New Mexico for the size and

number of trout that it contains. We saw trout (Sa/mo spilu-

rus) in great numbers but the stream was low and very clear

so that only a few were hooked.

Some dark purple bells (Campanula uniflora) grew here

887

888 THE AMERICAN NATURALIST. [Vor. XXXVII.

among the short grass which covered the open spaces of the

valley. The next day's journey took us over grassy slopes

dotted with bunches of mountain daisies; growing solitary or

rarely in groups of two or three the yellow Mariposa lily (Ca/o-

chortus gunnisoni perpulcher, Ckll) opened wide its petals

showing the purple and yellow center. Small dark bees indus-

triously gathered nectar and pollen from these flowers. It was

interesting to see the quick motions with which they pushed

the hairs from the glands and sucked the nectar or climbed the

filaments and loaded themselves with white pollen from the

bursting anthers. The store house was so easily accessible and

the stores so abundant that often the small bee overloaded itself

and sank into the grass; then there was great buzzing and

scrambling and throwing overboard of a part of the load before

it could safely take to wing again. Here and there we saw the

purple columbine (Aquilegia cerulea) with great splendid flowers

of purple and white, though the plants were dwarfed like the

plants of all high mountains.

Our second camp was made on the Pecos River, which in this

region is about six feet across and very rapid. The river course

is bordered with spruce with here and there an open grassy

glade covered with a sort of bunch grass which proved excellent

food for our horses. At the roots of this grass we found several

small mice which moved with a curious quick creeping motion.

This grass was full of'grass-hoppers, many of the species

having no wings. Several large Megachile bees (M. sape//onzs)

were busy on the flowers here and Argynnis eurynome and A.

electa flew from flower to flower. It was curious to see these

butterflies going from glade to glade, for they would often rest

in the boughs of the spruce trees, usually on the bright silver

tips and here the wind would gently rock them back and forward

and the sun would glisten on the silver spotted wings !

Along the river bank I collected a gigantic Pedicularis pro-

cera; it was almost six feet tall and had flowers an inch and a

half long. Here we found great patches of Polemonium growing

so closely bunched that no other plant could grow among them.

There was one such bunch covering a space about ten feet

square. A beautiful species of Actæa grew here but though

No. 444] TRIP TO -THE TRUCHAS PEAKS. 889

the plants were very large there were very few in numbers.

Two species of Ribes, A. wolftt and R. lentum, occurred in the

same vicinity.

August the second we climbed the peak to the northeast, and

two days later the one farthest southwest. The slope was

gradual so that by following the hogback we could ride quite

out of timber. Here we found good grass for our horses and

after securing them with lariats we left them to feed and walked

to the mountain top. Just at the edge of the forest a great

patch of gentians were growing along the trail; we had seen

the fringed gentian and the bell gentian but this was our first

sight of this high alpine flower. The leaves are short and form

a thick mat on the ground and from this dark green mass the

light lavender bells dotted with dark purple are lifted on a

slender flower stalk two to four inches high. The species is

Gentiana frigida.

A few hundred feet below the gentian field we passed over

hillsides covered with splendid dark purple larkspurs (Delphinium

subalpinum) there were acres and acres of this flower which

varied from a deep purple marked with black to a very light

purple ridged with white, and like the Polemonium the larkspur

grows in thick clusters crowding out even the grass at its roots.

The southwest Truchas we found very difficult to climb and

in places we pulled ourselves up by the bunch grass and held

our places by digging our heels into the soft earth. All the

plants were now reduced to mossy mats hardly more than an

inch through.

A tiny primrose grew among the rocks; beautiful dark blue

forget-me-nots (Eritrichium argenteum) with white woolly leaves

and stems were abundant ; a Sedum covered large acres and a

little phlox (Phlox condensata) grew in the shade of the rock.

A bright reddish humming-bird rested on my shoulder. The

butterflies flew with swift steady strokes, and were impossible

to secure when on the wing. A large black Papilio flew past

fluttering for a moment over the rock monument which marked

the highest point of the mountain. The Parnassius smintheus

flew about in numbers; some very worn and broken from long

struggles with the high winds which usually prevail in these

890 THE AMERICAN NATURALIST. (VoL. XXXVII.

mountain heights. Argynnis eurynome, very light green with

yellow and silver markings underneath with dark ferruginous

above trimmed abundantly with black, flew over the hillsides ;

the yellow and pink coloration of Colas scudderi added to the

beauty of the scene. Bumble-bees and flies buzzed about our

heads, and great gaudily colored saw flies tangled themselves in

our hair. The ants were just swarming and under the rocks

we found several species of beetles and spiders.

The view from the top was magnificent. Toward the west

the great Rio Grande Valley could be seen bordered by hills

which looked to be made of blue mist, so far away they lay.

To the north, the south and the west were rugged peaks and

wooded ranges everywhere marking the horizon off by curious

zigzag lines. Two bald eagles flew through the space to our

right —down and then up, darting and sailing and then across

until they disappeared from sight.

NOTES.

List of Plants collected above timber line. Determined by T. D. A.

Cockerell and Aven Nelson.

Eritrichium argenteum Wight, Mertensia celestina Nelson & Ckll.

(a new species found on this trip 1), Saxifraga chrysantha Gray, Delphinium

nov. Sp., Trifolium sp., Epilobium angustifolium L., Senecio holmii

Greene, determined by Mr. Greenman, Gentiana frigida Haenke, Sedum

stenopetulum, Veronica wormskjoldii R. & S., Phlox condensata (Gray) E-

Nelson, Oxyria digyna (L.) Camptdera, Draba s., Castilleia haydeni (Gray)

Ckll

List of insects collected above timber line.

1. Coleoptera. Determined by H. C. Fall. Amara brunneipennis, A.

femoralis, Tachys nanus, Aleochara sp., Pachybrachys hepaticus, Hyper-

aspis fimbriolata, Galeruca externa, Aphodius aleutus, Balaninus sp.

2. Diptera. Determined by D: W. Coquillett. Lasiophthicus pyrastri

Linné, Peleteria ænea Stæger, Trypeta occidentalis Snow

. Hemiptera. Determined by O. Heidemann. Corisekzia nitiduloides

Wolff, Mysius angustellus Blanch, Geocoris limbatus Stål., Lygeus turcicus

var. kalmii Stål., Irbisia sf. near drachycerus, Uhler.

4. Lepidoptera. Determined by T. D. A. Cockerell. Argvnnis eury-

nome, Brenthis helena, Parnassius smintheus, Pyrameis cardui, Sciaphilz

1 Described in Proc. Biol. Soc. Washington. March, 1903, p. 46.

No. 444] TRIP TO THE TRUCHAS PEAKS. 891

argentana (det. by Dr. Dyar), Colias scudderi flavotincta, Lycena rustica,

Plusia hochenwarthi, Colias eurytheme.

5. Orthoptera. Gomphocerus clavatus var. clepsydra, Melanoplus excel-

sus (det. by Mr. Rehn).

6. Hymenoptera. Determined by T. D. A. Cockerell. Bombus frigidus,

Bombus ternarius, Bombus flavifrons, Clisodon terminalis, (on thistle).

7. Ants. Determined by Prof. W. M. eeler : — Formica fusca sub-

sericea, Formica n. sp. (near subpolita), Leptothorax canadensis, n. var.,

Myrmica rubra, subsp.

Other species of various orders were collected, but have not yet been

studied. The above is the first list of the fauna and flora of the Arctic-

Alpine zone in New Mexico, excepting the records of insects taken on Taos

Peak about thirty years ago by Lieutenant Carpenter.

Las VEGAS, NEw MEXICO,

April 4, 1903.

! Bombus flavifrons is new to New Mexico. On the same trip, my wife took it

also on the top of the Las Vegas Range (about 11,000 ft.). A male from the lat-

ter place represents a new variet

. flavifrons var. veganus. Anterior part of thorax with yellow hair only; scu-

tellum with a good deal of black; first two abdominal segments with yellow hair;

third with aie slightly mixed with red; fourth with red; the rest with black.

bdomen with yellow hair; some black hair on front, and on

iio of face: some pen niae at at oe of Maa. Malar me long;

first and third joints of flagellum about equal, secon Tibiz

with yellow, tarsi with orange hairs. The yellow hair throughout i is bright rather

pale canary color. Mr. Viereck writes me about this: “There isn't a d in the

lot of B. flavifrons in the Cresson collection which tallies with your description of

var. veganus, though the structure is the same. There are a few females which

must look more like what you have, so far as pubescence goes. One small

worker (9 mm. long) taken at Beulah by Dr. Skinner agrees even better than the

specimens just cited, but has all the pale hair whitish. — T. D. A. COCKERELL.

QUARTERLY RECORD OF GIFTS, APPOINTMENTS,

RETIREMENTS AND DEATHS.

EDUCATIONAL GIFTS.

Amherst College, $100,000 from various sources for the astronomical obser-

vatory ; an annual income of $1500 from Miss Pratt of Brooklyn.

Chicago University, $300,000 from various sources for researches in

Egypt and Babylonia.

Columbia University, $40,500 toward the fund for the purchase of South

Field.

Cornell University, $100,000 and the residuary estate by the will of Fred-

rick W. Guiteau.

Harvard University, $154,000 from various sources for Emerson Hall;

$25,000 and half the residuary estate, for the Medical school, by the

will of Dr. George Haven; $25,000 by the will of Richard W. Foster.

New Haven, a conditional gift of $300,000 from Andrew Carnegie for a

public library.

University of Michigan, $4000 from J. B. Whittier for a fellowship in

botany.

University of the Pacific, $100,000 raised by the California M. E.

Conference.

University of North Carolina, $25,000 from Judge Wm. P. Bynum, $4000

from J. S. Hill.

Vassar College, a conditional gift of $200,000 from J. D. Rockefeller ;

$50,000 from other sources.

Washington & Lee University, $10,000 from Mrs. Cyrus H. McCormick.

Wesleyan University, $25,000 from Cephas B. Rogers.

Western Reserve University, $50,000 from various donors for the Women's

College.

Wooster University, $100,000 from L. H. Severance.

Yale University, $100,000 by the will of Sarah B. Harrison; a metallurgical

laboratory from John Hayes Hammond ; $300,000 for a dormitory from

F. W. Vanderbilt.

APPOINTMENTS.

Dr. Ralph Arnold, assistant on the U. S. Geological Survey.— C. F.

Baker, assistant professor of biology in: Pomona College, California.—

Franklin D. Barker, instructor in zoólogy in the University of Nebraska.—

Dr. Edward Bayer, custodian of botany in the Bohemian Museum at Prag.

893

894 THE AMERICAN NATURALIST. (VoL. XXXVII.

— Prof. V. von Borbás, director of the botanical gardens at Klausenburg.

— Dr. Max Borst, extraordinary professor of anatomy in the University at

Würzburg.— Charles J. Brand, assistant curator of botany in the Field

Columbian Museum.— Dr. F. Broili, docent for geology and paleontology in

the University at Munich.— Dr. Alfred Burgerstein, extraordinary professor

of botany in the University of Vienna.— Dr. Fridiano Cavara, professor of

botany in the University at Catania.— Dr. Otto Conheim, extraordinary pro-

fessor of physiology in the University at Heidelburg.— Dr. E. B. Copeland,

chief botanist of U. S. Philippine Commission.— Dr. Lucien Louis Daniel

to the chair of agricultural botany in the University of Rennes.— Dr. L.

Detre, docent for bacteriology in the University at Budapest.—Dr. Karl

Diener, extraordinary professor of paleontology in the University at Vienna.

— Dr. Francis A. Dixon, professor of anatomy in the University of Dublin.

— Dr. Hermann Diirck, extraordinary professor of anatomy in the Univer-

sity at Munich.— A. D. E. Elmer, botanical collector to the U. S. Philip-

pine Commission.— Dr. C. H. Gordon, acting professor of geology in the

University of Washington.— Dr. D. Hepburn, professor of anatomy in

University College, Cardiff.— Dr. Thomas Jehu, lecturer on geology at the

University of St. Andrews.— Dr. H. S. Jennings, assistant professor of

zoólogy in the University of Pennsylvania.— Dr. H. P. Johnson, associate

professor of bacteriology in the University of St. Louis.— Dr. J. N.

Langley, professor of psychology in the University of Cambridge.— Dr.

A. G. Leonard, professor of geology in the University of North Dakota.—

Osmond E. Leroy, of the Canadian Geological Survey, geologist to the

Chinese department of Mines.— Mary Isabel McCracken, instructor in

bionomics in Stanford University.— Dr. Burton D. Myers, instructor in

anatomy in Indiana State University.— Dr. Bohumil Nemec, extraordinary

professor of plant anatomy in the Bohemian University at Prag.— Wilmon

Newell, State entomologist of Georgia.— Edith M. Patch, entomologist in

the Maine Experiment Station.— E. C. Perisho, state geologist and profes-

sor of geology in the University of South Dakota.— Dr. Augustus Pohlman,

assistant professor of anatomy at the Johns Hopkins University. — Dr. Pom-

peckj, extraordinary professor of paleontology in the University at Munich.

` — Dr. Peter Potter, associate professor of anatomy in St. Louis University.

— R. C. Punnett, demonstrator of comparative anatomy in the University of

Cambridge.— Prof. W. M. Scott, pathologist to the Bureau of Plant Indus-

try, U. S. Dept. of Agriculture.— Dr. J. R. Slonaker, assistant professor of

physiology in the Stanford University.— Robert E. Snodgrass, instructor in

entomology in Stanford University.— A. I. leSouef, director of the Zoólog-

ical Garden at Sydney.— Dr. Percy G. Stiles, instructor in physiology in the

Massachusetts Institute of Technology.— Dr. Franz Stuhlmann, director of

the biological agricultural Institute at Amani, German East Africa.— Prof.

J. W. Toumey, director of the Yale botanical garden.— Dr. Armin Tscher-

mak, extraordinary professor of physiology at Halle.— Dr. Alfred bio :

docent for botany in the University at Freiburg, Switzerland.— Dr. A

No. 444.] GIFTS, APPOINTMENTS, RETIREMENTS. 895

Wilder, professor of geology in the University of lowa.— L. L. Woodruff,

assistant in biology in Williams College.

RETIREMENTS.

Dr. Howard Ayers, from the presidency of the University of Cincinnati

at the end of this college year.— Prof. Arnold Dodel, from the chair of

botany in the University at Zürich.— Homer H. Foster, from the chair of

botany in the University of Washington.— Dr. A. Hansgirg, from the chair

of botany at Prag, after forty years incumbency.— W. M. Scott, from the

position of entomologist to the state of Georgia.— J. E. Todd, from the

professorship of geology in the University of South Dakota.

DEATHS.

Dr. Eugen Askenasy, honorary professor of botany at Heidelburg, aged

58.— Dr. Franz Bauer, docent for geology in the Munich Technical School,

by a fall from the mountains near the Tegernsee, June 21.— John Allen

Brown, archaeologist, at London, Sept. 24, aged 72.— Cornelius Van

Brunt, botanist, at New York, Oct. 1, aged 76.— M. A. Certes, zoólogist, at

aris.— M. Meunier Chalmers, professor of geology in the Sorbonne.—

Mr. iiie Radcliffe Grote, the well-known entomologist, at Hildesheim,

Sept. 23.— Prof. Karl Haussknecht, botanist in Weimar, July 7.— Alex-

ander von Homeyer, ornithologist and entomologist, in Greiswald, July 14,

aged 70.— Dr. C. T. Hudson, the well-known student of rotifers.— Prof.

Emond Nocard, director of the veterinary school at Paris, and author of

works on parasitic diseases of animals, Aug. 3, aged 54.— Dr. A. G.

Ohlin, docent for zoólogy in the University at Lund, July 12, aged 36.—

Dr. Alphonse Renard, professor of natural history in the University at

Ghent, July 9, aged 60.— Professor Alexander Rollett, physiologist of the

University at Graz, Oct. 1, aged 69.— Dr. Frank Russell, anthropologist, an

associate editor of this journal, and formerly instructor in Harvard

University, in Arizona, Nov. 7.— Carl Wüstnei, an ornithologist of Meck-

linburg, Dec. 21, 1902.

(No. 443 was mailed Jan. 1, 1904.)

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