The American Naturalist.

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ee D. S. JORDAN, LL.D., Seabed Univer

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THE

AMERICAN NATURALIST.

Vor. XXXVIII. January, 1904. No. 445.

ADAPTATIONS TO AQUATIC, ARBOREAL, FOS-

SORIAL AND CURSORIAL HABITS

IN MAMMALS.

IV. CURSORIAL ADAPTATIONS.

RICHARD SWANN LULL.

Or all portions of an animal's body to undergo specialization,

those which have to do with locomotion show perhaps the most

varied adaptations. Speed is so essential to a great number of

forms, either for escape from the enemy or for the chase of prey,

that its degree of development has much to do with the fitness

of the creature for survival. This adaptation is most manifest in

the modifications undergone by the feet and limbs, and to a less

extent in the lengthening of the head and neck in long limbed

forms as a necessary correlation. Speed adaptation is further

shown in the moulding of the contour of the body to lessen the

resistance of the air, an increase in the capacity of the heart and

lungs to meet the more rapid expenditure of energy, and finally

in saltatorial forms an increase in the length and weight of the

tail.

Most terrestrial mammals can run; but in comparatively few

orders is there any special adaptation for speed. Offensive flight

2 THE AMERICAN NATURALIST. [Vor. XXXVIII.

occurs among predaceous mammals such as the carnivorous mar-

supials and the true Carnivora, while defensive flight is found

among herbivorous forms both among the marsupials and in the

placental orders Rodentia, Perissodactyla, and Artiodactyla.

Feet and Limbs : — The main foot adaptations are shown in

the passage from a primitive plantigrade toa digitigrade or to

an unguligrade condition, and in the reduction of the number

of digits; the last being often accompanied by a close apposition

or even by a fusion of the remaining bones of the metacarpus or

metatarsus, and a reduction of the number of bones in the wrist

and ankle. Cursorial adaptation leads to the formation of true

ginglymoid joints in the carpus and tarsus, the motion being lim-

ited to flexion and extension though the angle of movement is

increased. This motion is confined to the proximal podials, while

the distal ones become flat and may either fuse with each other

as in the Pecora, or with the metapodials as in the Tragulide.

The development of tongue and groove joints mars the efficiency

of the limb for other purposes than running.

The laws which govern digital reduction among vertebrates

lead to an interesting grouping of the Mammalia with the

Amphibia in which the order of reduction is first digit I then

digit V as contrasted with the Sauropsida, the reptiles and birds

in which the fifth digit is invariably the first to disappear, fol-

lowed by digit I.

The axis of the mammalian foot may lie in digit III as in the

perissodactyls and in most rodents though not in the Lepo-

ridæ ; between digits III and IV as in the Artiodactyla and Car-

nivora, or in digit IV as in the diprotodont marsupials.

Another marked cursorial adaptation is the increase in length

of the lower leg and foot both absolutely and in relation to the

length of the femur ; the lengthening of the limb increases the

stride while the raising of its centre of gravity quickens the

motion. This modification reaches its highest expression in

creatures of moderate size such as the medium sized antelopes

for in larger animals the increase in weight demands greater

structural strength which limits the degree of such elongation. `

The Carnivora whose need of cursorial adaptation, outweighed

by a greater need of varied motion, is less than that of other

No. 445.] HABITS IN MAMMALS 3

orders under consideration show the most generalized condition

of feet and limbs ranging from the ancestral canid Cynodictis of

the Oligocene and lower Miocene, in which both manus and pes

are pentadactyl though with functionless pollex, to Lycaon in

which structural tetradactyly prevails. In most of the Canidze

the digital formula is manus 5, pes 4; the former being however

functionally tetradactyl. A curious reversional condition is seen

in many high bred domestic dogs in which a functionless hallux is

present without skeletal connection with the rest of the pes,

similar to the dew claws of cattle. The author has invariably

observed this hallux claw in Saint Bernard dogs since he first

noticed it, and it seems to occur in about fifty per cent. of fox

terriers. Occasionally it is observed in other breeds but not so

constantly and it probably never occurs in low bred individuals.

I have recently observed a Saint Bernard with zwo hallux claws

on each hind foot. Cats, with the exception of Cynzelurus, the

hunting leopard, are not addicted to running, as their run consists

merely of a series of bounds, the creature slowing down as soon

as possible, hence special cursorial modifications are hardly to be

looked for outside of the genus mentioned. In Cynzelurus the

compact feet, poorly retractile claws and dog-like proportions and

musculature show an interesting case of convergence toward the

Canide.

As in Lycaon the hyzenas have also reached a condition of

structural tetradactyly, the most extreme case of digital reduc-

tion to be met with in the order.

Among the polyprotodont marsupials the Tasmanian wolf,-

Thylacinus, which lacks only the hallux shows thus the same

digital reduction as in the majority of Canidz though the feet

are much less specialized. The diprotodont marsupials on the

other hand exhibit extreme modifications both for running and

jumping.

Of the diprotodont marsupials the Peramelidz, the bandi-

coots, exhibit decided cursorial modifications. Disparity of size -

exists between the fore and hind limbs, and there is aside from

this fact a further kangaroo-like reduction of the pes. The hal-

lux is very much reduced ; digit IV is on the contrary the domi-.

nant one, while digits II and III are syndactylly united to offset

4 THE AMERICAN NATURALIST. [Vor. XXXVIII.

digit V. In Peragale the lateral digits, except the hallux, are

subfunctional while in Choeropus the pes is functionally monodac-

tyl although not structurally so. The manus of Peragale has

five digits the three median ones being functional, with III as

the dominant finger, while the external digits are functionless.

The ungal phalanges on II, III, and IV are long and deeply

cleft. In Choeropus digits I and II are reversional, digit IV

being vestigial, while II and III are functional, III being as in

Peragale, the dominant one.

_ From the digital modification observed in the Peramelide to

that of the kangaroos is but a step, for while in the latter the

manus is more generalized the pes has reached a higher degree of

specialization in the total reversion of the hallux, the plan of modi-

fication being precisely as in the bandicoots. The more general-

ized pentadactyl manus, which shows no cursorial modification,

would seem to indicate that the bipedal gait was acquired before

speed requirements were met.

The Rodentia have five or six families in which true speed adap-

tation occurs ; one, the Leporida, which have a combination of

cursorial and saltatorial gait, the Dasyproctide and Caviidæ

which are purely cursorial, and the m and Pedetida which

are saltatorial.

In the Leporidz, the hares and rabbits, the gait is a curious

mixture of leaping and running. The wood rabbit, Lepus sylva-

ticus, when in a full gallop progresses in a manner similar to that

of most quadrupeds, while the jack rabbit, Lepus campestris,

which is a true hare, moves by a series of bounds, irregular in

length, with all of the limbs moving synchronously, though the

weight borne upon the fore limbs must be very slight, the power-

ful hind limbs giving the impetus to the body. The Leporidz

show no especial digital modification other than the simple reduc-

tion of the hallux. There is an elongation and greater compact-

ness of the metapodium as in the Canidz among the Carnivora,

and the axis of the foot lies between digits III and IV.

Of the purely cursorial types the Dasyproctidz are the more

generalized though they exhibit two distinct stages in the reduc-

tion of the digits. The first is that of Coelogenys, the paca, in

which the manus and pes are both structurally pentadactyl

No. 445.] HABITS IN MAMMALS. 5

though the pollex is reduced, while in the foot but three digits

are functional, the lateral digits being vestigial.

In Dasyprocta, the agouti, however, the hand remains dis-

tinctly pentadactyl though digits I and V are subfunctional, while

in the foot the digits are reduced to three. Thus the foot is

both structurally and functionally tridactyl, the metatarsals being

closely pressed together though not fused. The compact pes

thus formed is tending toward that possessed by the Dipodidz

(vide infra).

The Caviidze, represented by the Patagonian cavy, Dolichotus,

show a further reduction over Dasyprocta in that in the former

the manus is tetradactyl while the pes is in the same stage of

reduction in each.

Among the truly saltatorial rodents a range of specialization

is shown, starting from Perodipus, in which the manus and pes

are each pentadactyl though there is considerable disparity of

size between the fore and hind limbs, the progression being

by leaping with the hind feet. Dipodomys, the kangaroo rat,

belonging, together with Perodipus, to the American family

Heteromyidz, has much the same proportions, but the pes is

tetradactyl. In Pedetes, the African jumping hare, the pes is

tridactyl, the median digit being much the longest ; while in the

true jerboas, as Dipus, the disparity between the limbs reaches

its greatest development and the elongate metatarsals are fused

into one very bird-like bone. The digital formula is manus 5,

pes 3; but the clawless pollex is evidently undergoing reduction.

Among the ungulates I know of none in which cursorial

adaptation is manifest, which have not already functionally lost

the pollex and hallux, that is with the exception of Phenacodus

among the condylarths which was pentadactyl and undoubtedly

could run though exhibiting no very marked cursorial adaptation.

In general, while the pes is often more specialized than the

manus there is far more uniformity in the plan of modification

of fore and hind feet than was observed in the rodents and

diprotodont marsupials. The artiodactyl and perissodactyl

stems have modified the feet in such different ways that it

becomes necessary to give each group separate treatment.

Among the Perissodactyla, the rinocerotine group, mostly of

6 THE AMERICAN NATURALIST. [Vor. XXXVIII.

unwieldy build, contains but one family, the Hyracodontide,

ranging in North America from the Bridger to the White River,

in which a running type developed. Here the manus is tetra-

dactyl, the pes tridactyl, and as Osborn ' says: * Tridactylism is

rapidly acquired with a tendency to monodactylism in the lower

Oligocene." They strongly suggest the primitive horses in

general contour.

The Equidz are too well known to require more than a brief

review. The pentadactyl ancestral form is as yet undiscovered

and must be looked for in the Cretacic, for in the lower Eocene

there appears Eohippus with a tetradactyl manus, the pollex

being represented by a splint, and a tridactyl, elongated pes

which bears a splint of digit V. The other Eocene horses

exhibit the same stage of digital reduction as in Eohippus ; but

Mesohippus of the middle and upper Oligocene is tridactyl in

the manus as well as in the pes, the fifth digit of the former

showing a splintlike metatarsal, digit I being vestigial. Here

all of the digits are functional the laterals finally losing their

contact with the ground in Merychippus of the middle” Miocene

and in Neohipparion of the upper Miocene, a beautiful specimen

of which has lately been added to the American Museum col-

lection. Hypohippus of the middle Miocene with subfunctional

lateral digits and, in the manus, the vestige of metacarpal V is

an instance of arrested evolution owing probably to marsh

dwelling habits which necessitated a spreading foot.

Finally the monodactyl type of the Pleistocene and Recent is

represented by the genus Equus in which digits I and V are

reversional and digits II and IV vestigial, being represented by

the metapodial splints alone.

The Equidz are curiously paralleled in foot reduction by the

South American Litopterna in. which the tridactyl condition

with functionless lateral digits is shown in Proterotherium from

the Santa Cruz formation, Lower Tertiary of Patagonia. This

creature seems to parallel Merychippus, the main distinctions

being that the former has rather more slender phalanges in the

middle digit while those of the lateral digits are proportionately

lOsborn H. F. The Extinct Rhinoceroses of North America. Jem. Amer.

Mus. Nat. History, Vol. I, Part 3: P- 93.

No. 445.] HABITS IN MAMMALS 7

more robust. The metapodials are shorter and stouter than in

the horse, those of digits II and IV particularly being much

more prominent.

Thoatherium from the same beds is monodactyl, the lateral

metapodials being even more vestigial than in Equus which it

parallels, and as in Proterotherium, the phalanges, especially the

proximal and ungal of the remaining digit are much more

slender than in the horse, the ungal being cleft. A curious

admixture of perissodactyl and artiodactyl characters is seen in

the feet of the Litopterna for they have the odd toed feet of the

Perissodactyls together with. the characteristic double tarsal

joint, though not to so great an extent, of the artiodactyls.

The Artiodactyla early lose the hallux and pollex, for except in

Oreodon and Agriocheerus we have no instance of their survival

and while digits III and IV are equally well developed, II and

V. suffer all degrees of reduction from that seen in the swine to

the total disappearance in the camel and Antilocapra.

The swine are four toed, the lateral digits being sub-functional.

Dicotyles, the peccary shows an advance over most Suidz in

that digit V of the pes is entirely wanting giving an asym-

metrical foot, of uncommon occurrence in the order though

found in the Anoplotheres as well. In Dicotyles the metacar-

pals are slightly fused at their proximal end while in the

metatarsals the fusion extends over half the length of the bones.

The Pleistocene genus Platygonus shows a still greater special-

ization as it is structurally didactyl, but a splint of the fifth

metatarsal remaining. The metapodial bones show a greater

degree of fusion than in Dicotyles.

The Tragulide or chevrotains are in a sense transitional

between the swine and the true deer for, while four toed, the

lateral toes are functionless although in the existing genera

Tragulus and Dorcatherium (Hyomoschus) the lateral meta-

podials are entire. Fusion of the median metatarsals to form a

canon bone is found in Tragulus, but not in Dorcatherium which

together with its somewhat better developed lateral digits pre-

sents a more generalized condition than does Tragulus. Gelo-

cus, an extinct form ranging from the Eocene to the Oligocene,

is more specialized than either of the existing genera in that the

8 THE AMERICAN NATURALIST, [Vor. XXXVIII.

lateral metapodials are incompletely ossified. The metacarpals

are not fused, while the presence or absence of fusion of the

metatarsals is a specific variation.

The extinct Oligocene genus Protoceras gives us an inter-

esting example of the acceleration of the specialization of the

hind limbs over the fore, for while the latter have four well

developed functional digits those of the former are reduced to

two only, with closely applied metatarsals which do not fuse,

though strongly tending so to do. The lateral metatarsals

are represented by proximal vestiges only.

In the Pecora or true deer the lateral digits are reduced,

being functionless in most genera though sub-functional in

Moschus and in Rangifer probably due, in the latter genus at

any rate, to the necessity of a broad plantar surface for support

on the mossy tundras or on the snow, a condition analogous to

that of Hypohippus among the horses. In the deer the lateral

metapodials are incomplete, their distal ends always occurring

while only in certain genera as Cervus and Cariacus are the

proximal extremities also retained.

The Bovida exhibit an almost complete reduction of the

lateral digits, the dew claws being dermal appendages only, the

proximal phalanges being invariably absent, while the final stage

of total reduction of the lateral toes is found in the camels, the

giraffe, and in Antilocapra. In the Bovida as well as in the

later Camelida and the other forms mentioned the fusion of

the metapodials to form a canon bone is complete.

In the later camels there is a retrograde descent from the

unguligrade to the digitigrade condition, wherein the phalanges

lie prone upon the ground, giving the characteristic broad, pad-

like foot of the modern camel.

There are no instances of saltatorial adaptation among the

ungulates though some antelope and deer are wonderful

jumpers.

The lemurs among the primates present several instances of

saltatorial adaptation, notably in the sub-family Galagininz and

in Tarsius, family Tarsiidae; but here instead of an elongated

metatarsus, which has been the rule heretofore, it is the tar-

sus which is modified, for the calcaneum and navicular become

No. 445.] HABITS IN MAMMALS 9

lengthened and cylindrical as do the calcaneum and astragalus

in the frog. The hallux is large and opposable while digits II

and III are somewhat reduced, digit IV being the longest. In

Tarsius digits II and IV are clawed while the others bear flat-

tened nails.

The Skull— Cursorial adaptation has its effect upon the

skull only in the correlation that exists between long limbs and

dolichocephaly, brought about by the necessity of reaching the

ground on the part of a grazing animal. This is strikingly

illustrated in the horse series where the increase in the length

of the skull parallels the lengthening of the limbs.

Saltatorial forms which, like Dipus, have lengthened the hind

limbs only, do not exhibit marked dolichocephaly, as the feeding

habits of the creature do not require it. The grazing kangaroos

however have a moderately elongate skull.

The Vertebral Column.— Cursorial adaptation among mam-

mals is shown in the lengthening of the cervical vertebrz, espe-

cially in dolichocephalic forms, strikingly illustrated by the giraffe

and by Alticamelus of the Loup Fork of Colorado! a camel

showing the most remarkable convergence toward the giraffe,

although the latter is derived from a totally different stock.

Among the saltatorial forms, especially those with brachy-

cephalic skulls, the tendency is toward the shortening of the

neck accompanied by a greater or less degree of immobility. In

Pedetes cervicals 2 and 3 are so closely articulated as to elimi-

nate motion, in Perodipus the axis and the next two vertebra

are fused, while in Dipus all of the cervicals except the atlas

are coóssified as in whales. There is no increase or diminution

in the number of cervicals as a result of speed adaptation. The

dorso-lumbar series seem to suffer little alteration in cursorial

forms, though the lumbars increase in size in saltatorial types.

The high number of vertebrae found in the horses is also found

in other perissodactyls and in the Proboscidea, and so is not to

be considered a modification coming within the scope of the

present discussion. In some saltatorial forms, as the jerboas,

an exceedingly short back is found; but saltatorial adaptation

can exist without this feature.

! Matthew, W. D. Mem. Amer. Mus. Nat. History, Vol. 1, Part 7, p. 429, pl.

XXXIX.

IO THE AMERICAN NATURALIST. (Vor. XXXVIII.

The tail is generally reduced as a result of cursorial adaptation,

though in coursing dogs, as the grayhound and pointer, it aids in

keeping the balance when the creature changes its direction and

this may be a secondary cause for its retention. In saltatorial

forms on the contrary, the tail becomes an important organ for

use as a counterpoise ; for in truly saltatorial mammals the tail

increases in length and in weight directly with the increase in

proportion of the hind over the fore limbs. It is in such bipedal

forms as the kangaroos and the jerboas that the caudal counter-

poise reaches its highest expression, for in the former the tail is

long and heavy while in the latter the somewhat less proportion-

ate weight is compensated for by the extreme length of the

organ and the tuft of hair at its tip. The tail of the African

jumping hare, Pedetes, is long and feather-like, like that of a

squirrel. The development of a caudal counterpoise in bipedal

mammals is paralleled among dinosaurs of the Mesozoic though,

as the author will show in a forthcoming memoir,! dinosaurs are

never saltatorial, but always progress by alternating strides.

This is what one would be led to infer, for whatever the increase

in speed may be, I know of no reptile which runs at a gallop

that is with each pair of limbs moving nearly in unison, while

among the mammalia this is the common method of rapid pro-

gression. The jerboas wa/k on the hind limbs with alternate

strides, hopping only when speeding while the kangaroos have

lost the more primitive alternate footed gait and use the hop for

all degrees of rapidity. The hop may thus be considered as a

sort of bipedal gallop. a

Among the struthious birds, the cursorial habit evidently

having been acquired after the loss of the reptilian tail, the

counterpoising function is subserved by the wings which bear

up the anterior part of the body and at the same time lighten

the creature's weight.

COLUMBIA UNIVERSITY, DEPARTMENT OF ZOGLOGY,

November 30, 1903.

1Lull, R. S., Fossil Footprints of the Jura-T'rias of North America. Mem.

Boston Soc. Nat. Hist. Vol. s, no. 10.

No. 445.] HABITS IN MAMMALS II

BIBLIOGRAPH Y.

BEDDARD, F.

*02. eie e Cambridge Natural History. Vol. 10. London.

FLowER, W. H.

'85. An Introduction to the Osteology of the Mammalia. London.

FLowER, W. H. AND LYDEKKER, R

'91. Introduction to the study of Mammals Living and Extinct. London.

LYDEKKER, R.

:00 e New Natural History. Vols. 2-3. New York.

MATTHEW, W. D

:01. Fossil Mammals of the Tertiary of Northeastern Colorado. Mem.

Amer. Mus. Nat. Hist. Vol. 1, Pt

VERMES FOE

The Extinct Rhinoceroses of North America. Mem. Amer. Mus.

Nat. Hist. Vol. 1, Part 3.

WOODWARD, A. S.

'98. Outlines of Vertebrate Paleontology. Cambridge.

ON THE OSTEOLOGY AND SYSTEMATIC

POSITION OF THE PYGOPODES.

R. W. SHUFELDT.

IN a series of papers contributed to Zhe Journal of Anatomy

and Physiology of London (1889-1890) appeared a number of

my drawings of the bones of pygopodine birds. Among these

the osteology of certain grebes and loons was illustrated, but the

material at hand then was meagre. Moreover, through an

oversight the plate illustrating the bones of the lower extremity

in the loons was omitted. The osteology of the pelvic limb in

a loon is in a way more deserving of our consideration than per-

haps other parts of the skeleton in those typical divers, and as I

had fortunately preserved the aforesaid plate, I take occasion to

publish it here.

In an article entitled ** Concerning some of the Forms assumed

by the Patella in Birds” (Proc. U. S. Nat. Mus. Vol. vii, 1884,

pp. 324-331) I published two or three figures of the leg-bones

in loons and grebes, but the descriptive text-matter had reference

only to the morphology of the patella.

In the present memoir all of my previous work has been

amplified and practically rewritten, while I have added my inves-

tigations upon the osteology of the Pygopodes.

In my classification of birds the Pygopodes appear in the

scheme as a suborder, thus : —

Suborder: — PYGOPODES.

SUPERFAMILIES : —

Podicipoidea.

Urinatoroidea.

FAMILIES : —

Podicipide.

Urinatoride.

14 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Newton in his classic ‘Introduction’ to his Dictionary of

Birds (p. 111) makes the statement that the “ group known as

Pygopodes has been often asserted to be closely akin to the

Impennes, and we have seen that Brandt combined the two

under the name of Urinatores, but of their essential difference

there can now be no doubt, and indeed it is hard to look upon

Pygopodes as a natural group, so many are the differences between

the Podicipedide or Grebes and Colymbide' or Divers, though

recent morphologists agree to unite them, while the affinity of

the Divers to the Auks seems to be still more uncertain, and

there appears to be ground for considering the Ade to be

much modified relatives of the Laride.”

The discovery of the toothed Hesperornithida of the middle

cretaceous of America has doubtless modified the opinions of

systematists regarding the affinities of the Pygopodes.

I agree then essentially with Fiirbringer in confining the

families Colymbidz (loons) and Podicipidze (grebes) to a group

* Colymbo-Podicipites,” and closely associating the latter with

the families Enaliornithidz and Hesperornithidz in a Suborder

Podicipitiformes. Therefore I can proceed to the consideration

of the osteology of the grebes.

THE SKELETON OF THE GREBES.

Grebes may have the superior osseous mandible longer than

the cranium, or they may have it shorter than that part of the

skull. Of the first-mentioned, Colymbus holbelli is a good

example, while Podilymbus podiceps exhibits the latter character-

istic. In C. holballi the long, straight and acutely-tapering

superior osseous mandible is fully one fourth longer than the

cranium, and either narial aperture is suboval in outline, being

equal in length to the end of the bill which extends beyond its

anterior termination, This narial aperture is rather acutely

holorhinal posteriorly, and the dentary margins are cultrate for

1“ American ornithologists have lately used this term for the Grebes, to the great

disturbance of nomenclature. It is apparently from the ancestors of the Colym-

bide, before they lost their teeth, that Hesperornis branched off as a degenerate,

bulky and flightless form." — A. N.

No. 445.] OSTEOLOGY OF PYGOPODES. 15

their entire extent. No part of the nasal septum ossifies in the

skull (a feature common to all grebes that we have examined).

Each nasal bone has a form much as we find it in the loons, its

.processes being flat and rather broad. The region over the

cranio-facial hinge is moderately concaved, where the nasal

processes of the premaxillaries are seen to be persistent through-

out life, and their sutures plainly visible in the adult.

Posterior to this space the frontals between the superior

orbital margins are much narrowed, and the supraorbital glandu-

lar depressions barely discernible, being distinguishable in the

dried cranium only along their posterior moieties. The external

superficies of the cranial vault in the parietal region are smooth

and rounded ; the crotaphyte fossæ are extensive and practically

meet, mesially, over the large rounded supra-occipital promi-

nence, though no median crest or line stands between them.

More laterally, and upon either side, the occipital crest is raised

and prominent. In the grebe now under consideration it is quite

as thin, and lamellar-form as it is in the loons.

Upon lateral aspect of the skull, the post frontal and squa-

mosal processes are much absorbed, and the valley between them

wide. The aural entrance is extensive, very open and exposed,

being overarched by its somewhat thickened postero-superior

border.

Either zygomatic bar is straight, transversely flattened, and

tapers slightly as it proceeds forwards where it assists in making

a schizognathous articulation with the other bones.

The interorbital septum is markedly deficient in bone, and the

anterior cranial walls about the exit of the nerves hardly less so.

This deficiency is even greater than it is seen to be in the

loons. Pars plana is weak and feebly developed, and exter-

nally, it passes upwards and forwards to fuse with the nether

aspect of the frontal. A transverse perforation may normally

exist just posterior to the true mesethmoid. This latter ossifica-

tion terminates rather abruptly in front by a broadish face with

a small median crest extending down it as far as the sharpened

anterior apex of the rostrum, over which it is carried forwards.

A lacrymal is a fair-sized bone with a very narrow superior

limb, closely articulating with its entire mesial border with the

16 THE AMERICAN NATURALIST. (Vor. XXXVIII.

frontal and nasal, while the rather larger descending portion of

the bone is plate-like, being transversely compressed, and does

not reach the maxillary below. At its apex it supports a spicu-

liform os uncinatum, as we find in the Urinatoride.

The foramen magnum is large, looks almost directly back-

wards, and is of an acute cordate outline, with the apex above.

The occipital condyle is well developed, completely sessile, and

barely notched superiorly. Passing to the basitemporal area we

find it somewhat contracted, nearly level and smooth, while its

anterior apex underlapping the double entrance to the Eusta-

chian tubes. There are no evidences whatever of basipterygoid

processes, and the long, straight pterygoids stand well away

from the sphenoid. One of these bones has cultrate inner and

outer edges or borders, and is peculiar in the way it articulates

with the quadrate. The latter bone throws out a well devel-

oped apophysis, mesially, the summit of which is rounded to be

received into the articular cup existing on the posterior end of

the pterygoid. In most birds the pterygoids articulate upon the

inferomesial border of the os quadratum. These bones in the

grebe hardly touch each other anteriorly, where their palatine

heads are to some extent expanded. The sphenoidal rostrum is

comparatively slender and is carried to a sharp apex in front.

The palatines have their postero-external angles completely

rounded off, while their lower inner and outer margins are

moderately bent downwards,— the inner one rather abruptly so.

When articulated zz sizu these bones are in contact with each

other all along beneath the rostrum. The antero-mesial portion

of the post-palatine part of the bone, curls upwards and inwards

towards the mesethmoid, and in front its mesial process runs

forward as a long slender spine for the accommodation of the

vomer. The prepalatine portion of a palatine is long, narrow

and vertically compres: `d. Extending a long ways to the front,

rapidly tapering to a point as it does so, the prepalatine under-

laps the maxillary and maxillo-palatine, and passes along close

to the inner aspect of the dentary part of the premaxillary,

being thoroughly fused there in the adult. The suture, how-

ever, remains visible throughout life, For the size of the bird,

Holbeell’s grebe has one of the longest vomers at present

No. 445.] , OSTEOLOGY OF PYGOPODES. 7

known to me. It is lamelliform, thin and narrow, its surface

being in the middle plane, while behind it is moderately bifur-

cated, to be carried to.a sharp apex anteriorly. Either maxillo-

palatine is a subconcavo-convex oval plate of bone, of some

little size. Its mesial surface looks inwards and upwards,

the anterior fourth being fused almost indistinguishably with

the nasal, maxillary, palatine and premaxillary. Palatines and

maxillo-palatines are well separated from each other in the mid-

dle line, and from the vomer.

An os quaratum is rather a large bone in the grebes, with a

long, gently-inturned orbital process. Its mastoidal head sup-

ports two facets of articulation, being separated from each other

by a shallow sublongitudinal valley. Transversely, the quadrate

is much compressed, and I have already described above the

process at its infero-internal angle to accommodate the hinder

end of the corresponding pterygoid. The mandibular portion is

much excavated centrally on its nether aspect, with a small

articular facette on either side of the concavity. There is also

an articular line bounding this depression posteriorly. The

bone appears to be pneumatic.

Passing to the consideration of the mandible we find it to be

of the very acutely V-shaped pattern, with the ramal vacuity

completely closed in. The articular ends are enlarged and

abruptly truncated behind, where they show each a flat triangu-

lar surface. The ramal sides posteriorly are thin, lofty and flat,

to become narrower and thicker as they pass tapering forwards

to the acute apex. The symphysis is short, slightly excavated

above, and rounded below. It is only the posterior extremities

of the mandible that are at all pneumatic, the usual pneumatic

orifice being at the end of the inturned, stumpy articular termi-

nation of the bone. Aside from, the brevity of the superior

osseous mandible in the short-billed grebes, the skull characters

as given above for Colymbus holbwli are substantially repeated

in them. In Podilymbus podiceps, however, I observe that the

anterior extremity of the vomer terminates in a small, rounded,

disc-like nib, and its quadrates are rather more delicately fash-

ioned. It also has the mesial notch on the upper side of the

occipital condyle, and a mid-longitudinal raised line on the supra-

18 THE AMERICAN NATURALIST. [Vor. XXXVIII.

occipital prominence, which, as we shall see, is so much better

marked in the loons. Finally, the supra-orbital glandular depres-

sions are hardly perceptible in these dabchicks.

Grebes possess a hyoidean apparatus in some respects peculiar.

It is well exemplified in Podilymbus, where we find the glosso-

hyal performed entirely in cartilage, and the first basibranchial

represented by an expanded suboval disc of bone. At the

hinder margin of this the short second basibranchial, as a deli-

cate osseous rod, articulates in the middle line, while the long,

slender cerato-branchials, one on either hand, articulate close to

it. The epi-branchials are short and spiculiform. We find a

somewhat similarly fashioned first basibranchial in the tongue

of the kingfishers, but such a form of it is rare among birds.

The sclerotal plates in the eye-balls of the Podicipoidea have

their usual ornithic characters, being of moderate size only,

Squarish in form, and overlapping each other in the ordinary

manner.

THE TRUNK SKELETON IN THE GREBE.

Birds of this superfamily vary, even for the genera, with

respect to the number of vertebrz in the spinal column, and the

corresponding vertebrze themselves vary much in form and

character. Species such as Colymbus holbelli and Podilymbus

podiceps have 19 vertebrz in the cervical region of the spine,

the 19th bearing a pair of ribs that do not articulate by costal

ribs with the sternum. ;

But Æchmophorus occidentalis has 21 vertebrz in the cervical

region, with the free ribs on the 21st as they occur on the rgth

in Podilymbus. This last mentioned species has the first four

dorsal vertebrae fused into one piece, but the fifth one, standing

between this piece and the pelvis is free, and its ribs articulate

with the sternum by costal ribs. There is also a pair of pel-

vic ribs, the haemapophyses of which do not usually meet the

sternum. All have large epipleural appendages, save the last-

named; they being even found on the cervical pair. They do

not fuse with rib borders.

In /Echmophorus the dorsal vertebrz do not fuse, although

No. 445.] OSTEOLOGY OF PYGOPODES. I9

the interarticulations are very close. This grebe. has swe

pairs.of pelvic ribs, the haemapophyses of the first pair reaching

the costal borders of the sternum. All grebes have large

hypapophyses on the last two or three cervical vertebra, and on

all the centra of the dorsal vertebrae; they are very large in

AEchmophorus, the first two being represented by flattened

and out-spreading discs of bone of an irregular form. This

species is also peculiar in having the neural spines of the 19th,

20th and 21st vertebrae much modified for muscular attachment.

They resemble the ploughshare in form, being greatly increased

in size, and the excavation occurring behind. The first (19th)

has this modification most pronounced, while it is least marked

in the ultimate one (21st). Parapophysial spines are quite

aborted, or are represented by mere nibs of bone. A‘chmo-

phorus has the hypapophysial carotid canal extending through

twelve vertebrae ; it being generally closed in completely on the

gth and roth one of the series. These vertebra are the 4th to

the rsth inclusive.

In Podilymbus podiceps I found 49 vertebrae in the spinal

column. Nine free vertebrae and a pygostyle compose the tail

of this bird, and when they are articulated zw sz/z, they form a

peculiar sigmoid curve, dipping downwards, then upwards, as the

letter S. The pygostyle is very small and its characters much

aborted.

Grebes have their caudal vertebrae considerably compressed

jn the transverse direction. In the dorsal region the tendons

of the spinal muscles ossify and fuse with the summits of the

neural spines of the vertebra, and metapophysial spiculae may

also occur upon the transverse processes, as we find them in

other water birds. Another thing is worthy of attention here,

and that is the general form and outline of the skeletal parietes.

Further along it will be seen that in the auks and puffins this is

elongated,— the sternum being long, and the ribs sweeping far

backwards beneath the pelvis. In the grebes this is not usually

the case, for in Podilymbus the form of the thoracic skeleton is

much as we find it in the gulls; in /Echmophorus, however, it

is again more as in the Alcz ; it is quite so among the loons.

The form assumed by the pelvis among the Podicipoidea is

20 THE AMERICAN NATURALIST. (Vor. XXXVIII.

noteworthy ; though in its general pattern it closely approaches

what we find in the Urinatoride. The pelvis in 4:chmophorus

occidentalis well exhibits all the characters of this compound

bone among the podicipidine types. In that species is much

elongated and compressed laterally, especially its pos-acetabular

portion. In front of the acetabulze the fused sacral crista rises

far above the fore part of the ilium on either hand,— which

latter, each have the form of an oar-blade with a squarely

truncated anterior extremity. An extensive antitrochanter

surmounts either cotyloid cavity, while posteriorly the post-

acetabular surface faces almost directly outwards. Along the

dorsal middle line, for the posterior third, of the pelvis the iliac

borders are closely pressed together, marking the uro-sacral

vertebrae. Behind, a deep cleft indicates the division which

originally marked the terminal point of union between the ilium

and the very long and narrow ischium. The obturator foramen

completely merges with the obturator space, and the much-

extended, flat, ribbon-like post-pubic rod is widely separated

from the lower ischiac border, being carried far back almost

opposite the pygostyle. The ends of these bones of the pelvis,

however, are not dilated as are the postpubic bones in the

loons. The ischiac foramen is large, and of an elliptical out-

line. Podilymbus presents almost the same pelvic characters

as those just described for /Echmophorus, and in this species

the os innominata fuse completely with the “ sacrum,"— though

posteriorly the superior iliac margins do not quite meet over the.

uro-sacral vertebrae.

The sternum is very characteristic. In Figures 1, a and 1, 6

I have drawn it for Colymbus nigricollis californicus where its

podicipidine features may be seen. Generally speaking it is

broad and short, with a large subelliptical notch cutting out its

xiphoidal portion on either side of the keel. This gives rise to

a pair of flaring external xiphoidal processes, which curve out-

wards, then inwards, extending rather beyond the mid-xiphoidal

prolongation. They are long and narrow in the short-billed

grebes, and broader and relatively shorter in Aichmophorus and

Colymbus. The mid-xiphoidal process always shows a triangular

notch, which is much deeper in Podilymbus than in other species,

No. 445.] OSTEOLOGY OF PYGOPODES. 2I

and it may also show certain foramina in the hinder part of the

sternal body. The keel is triangular with a somewhat acute

carinal angle produced in front, and closely approached by the

os furcula when the bones of the shoulder-girdle are articulated

wn situ, Usually sir articular facettes are found upon either

costal border, and a costal process is large and subtriangular.

No manubrium exists, and the bone is depressed where it

occurs in the sternum when it is present. The costal grooves

are noteworthy, for they are very deep, and their superior and

inferior borders are produced well forwards.

As in the rest of the skeleton, save the hinder part of the

skull and lower jaw, the

sternum in the grebes is

completely non-pneumatic.

Upon comparing the

sterna of our various spe-

cies of grebes, I find but

few characters of marked

difference beyond the mat-

ter of size. Colymóus

auritus possesses a ster-

num most like that of

Podilymbus podiceps, and

next to it, in that respect,

comes Colymbus hollell.

In the several bones of

the shoulder-girdle, — the

salient characters are the

same for the various spe-

cies of this group. The

os furcula is always found

to be of the broad U-pat-

tern, much bowed to the

front, without hypoclidium dare fe bishi vw tiga dae Win Maiani di

A n. californicus), and

with narrow, laterally compressed limbs. — Superiorly, the clavic-

ular limbs taper out to acute points, and when the elements of

the girdle are articulated zx sztu, one of these rests by its outer

22 THE AMERICAN NATURALIST. (Vor. XXXVIII.

aspect against the head of the corresponding coracoid, while the

apex passes far over the anterior end of the scapula. Thusa

piece of the clavicular end, including the apex, is above the

scapula but not being in contact with it;— the actual point of

contact between these two bones being several millimeters

beyond or anterior to the apex.

The scapulz are quite long, and narrow, being gently curved

throughout their length in the vertical plane, the convexity

being along the dorsal aspect. For the most part the bone is

of uniform width, the head alone being somewhat thickened.

It offers only a moderate articular surface for the coracoid,

and the os furcula rests upon its upper side.

The scapule are especiall slender in CoZymóus auritus.

Comparatively speaking, the coracoids are usually long and not

very stout; they may, however, be only of moderate length.

The summit of one of these bones offers us the usual ornithic

characters, being peculiar only in having such a small scapular

process, and being rather compressed laterally. The distal end

of the bone is dilated and much flattened in the antero-posterior

direction. This expansion is carried some distance up the outer

side of the shaft, and, owing to the fact that the sternal end of

a coracoid sets so deep in its articular groove in the sternum,

the corresponding articular surface on the bone is carried up

some little distance both in front and behind, — most so upon

the latter aspect.

When the bones of the shoulder-girdle in a grebe are articu-

lated as in life, there is quite an interval between their sternal

ends, mesiad. As I have said above, this interspace on the

sternum is concave and its convexity is coextensive with the

inner border of the coracoid upon either side. A wide interval

in the same location exists in ZZesperormis regalis, the great

diver of the Cretaceous epoch in America.

THE APPENDICULAR SKELETON OF THE GREBES.

As an example of the skeleton of the pectoral limb of an adult

specimen of Colymbus holbelli we have chosen No. 17815 Coll.

U. S. Nat. Mus. In this specimen the humerus is 10.6 cms.

long, with nearly a straight, subcylindrical shaft, the extremities

No. 445.] OSTEOLOGY OF PYGOPODES. 23

of which are but moderately enlarged. At the proximal end

the radial crest is seen to be much reduced, and the excavation

overshadowed by the low ulnar crest which is unusually shallow,

with no evidences of pneumatic orifices. The articular surface

of the humeral head has the ordinary avian character. Distally,

the oblique and ulnar tubercles are prominent and offer consid-

erable articular surface for the antibrachial bones. The ulna is

10.2 cm. in length, and considerably compressed subtrans-

versely ; the long, slender radius when articulated, zz sítu, with

it, is in contact with its shaft for its distal moiety, thus much

reducing the *interosseous space," which, in reality, only exists

proximally. Manus has a total length of 8.3 cms., and the two

usual free carpals are present in the wrist. Carpo-metacarpus is

peculiar in being so comparatively long and slender, and for

having the index and medius metacarpals so close together, and

so nearly parallel. The phalangeal digits are long and slim, and

I fail to find any * claws" upon the distal extremities of any of

them. The proximal phalanx of the index digit is also elongated

and remarkably narrow ; the expanded portion and digital shaft

being indistinguishably merged with each other.

The skeleton of the wing in Podilymbus podiceps has the same

essential characters as in the wings of the long-billed grebes.

In all, the bones are well-proportioned and harmonize in their

lengths and calibres with the bones of the pelvic limb, in any

given species.

Altogether one of the most beautiful adaptive structures is the

pelvic limb of a grebe. When properly articulated, the short

femur has itslong axis directed from the acetabular center,

downwards, outwards, and slightly backwards. By the structure

of the knee-joint this brings the long axis of the tibio-tarsus

almost parallel to the long, mesial axis of the pelvis. Now the

tibio-tarsal articulation permits the exact play of the foot, by a

fore and aft motion, at right angles to this long axis of the tibio-

tarsus. It is an avian oar. The tarsus is compressed to the

last degree consistent with strength, — so that when it and its

blade-like toes make the forward stroke, the minimum amount of

podal surface is offered to the water in resistance. But in the

backward stroke of the foot, the articulation permits of the

24 THE AMERICAN NATURALIST. (Vor. XXXVIII.

reversal of this act, and the toes being turned, and to some

extent the tarsus, the maximum amount of surface thus afforded

is brought into play as in an oar. Femur and tarso-metatarsus

have about equal lengths, and they each equal /a/f the length of

the leg-bones, measuring from the apex of the patella to the mid-

lower point of the arc

of the distal tibial con-

dyles (Podilymbus podi-

ceps). The mid-anterior

toe in the adult meas-

ures. 5.5 cm. and the

tarso-metatarsus only

3.8 cm.; these are about

the usual proportions.

The head of the fe-

mur is large, and much

excavated for the round

ligament. At the sum-

mit the articular surface

is rounded, and the tro-

chanter does not rise

above it. Its shaft is

very slightly bowed to

the front, and its con-

dyles markedly promi-

: nent behind, with an

Fic. 2.— Leg-bones and patella of Colyméus auritus. unusually distinct and

inl tite fa Oe Ponce GNI D Lee Se O Eo oe

and patella are thrown backwards out of position. head of the fibula.

a rotular crest of tibia. P, patella; F, femur; Fé

fibula; T, tibio-tarsus. Natural size. One of the most im-

portant characters of

the tibio-tarsus is the upward extension of its cnemial crest,

which is carried up to an apical process considerably above the

summit of the shaft, but in direct line with the forepart of it.

A large patella backs this at its supero-external aspect. It has

something of the same form as the cnemial crest, above which

it is slightly extended when articulated zz s zzz. The procnemial

ridge of the tibio-tarsus is extended as a sharp border down the

No. 445.] OSTEOLOGY OF PYGOPODES. 25

shaft of the bone; the latter being straight, flat anteriorly and

rounded behind. Having the usual ornithic form, the tibial

condyles are set rather obliquely on the distal end of the shaft.

They are about parallel to each other. The fibula is very long

with its lower end fused with the side of the shaft of the tibia.

Superiorly, it is broad and flattened in the antero posterior direc-

tion, and stands well away from the tibial shaft (Figs. 2, /.).

Among the short-billed grebes the transverse compression of

the shaft of the tarso-metatarsus is not nearly so evident as it is in

such species as Æchmophorus occidentalis and Colymbus holbælli.

Moreover, in the dabchick there is a very slight twisting of the

shaft upon itself, and this I have not observed in other grebes,

either fossil or recent. At the summit of the bone, the articu-

lar excavations for the tibio-tarsal condyles are deeply sculpt,

and the inner lateral border of the internal one may be conspic-

uously raised (Podilymbus). The subcubical hypotarsus of the

tarso-metatarsus in most all grebes is deeply once-grooved in

the mid-longitudinal line behind; and in front of this groove it

presents one complete cylindrical perforation (also for the pas-

sage of tendons) with a similar, though smaller, perforation upon

either side of it, situated more posteriorly.

This is quite different from what we find in the loons (see

Fig. 18, of the Plate). Of the distal trochlez the middle one is

the lowest on the shaft; the outer one next; while the inner

one is very distinctly elevated. They are all more or less

drawn towards the rear aspect of the bone. The hallux digit is

feeble and flake-like, as is the small, free metatarsal to which it

is attached. It is considerably elevated upon the shaft.

Generally, the podal joints show more or less compression,

while the terminal or ungual ones are positively scale-like, and,

as it were, greatly flattened. Usually, the basal phalanges are

the longest, and the others in any toe gradually diminish in this

particular proceeding towards the extremities. Their arrange-

ment as to the number on each toe is upon the more common

ornithic plan of 2, 3, 4, 5 to the 1-4 toes respectively.

26 THE AMERICAN NATURALIST. [VoL. XXXVIII.

THE SKELETON IN THE LOONS.

In all of its essential characters, the skull of the loons agrees

with that of the grebes. In the present description I have

chosen the skeletons of Urinator imber and Urinator lumme,—

the first being the bird known all over the world as że loon, the

second, also a largely cosmopolitan type, is the red-throated loon

or diver. In the latter the superior osseous mandible is fash-

ioned upon the same plan as we found it in the long-billed

grebes ; it differs principally in curving very gently upwards, a

feature not present in the beak of U. imber. The culmen in

both species is convex and evenly rounded. Loons have the

cranio-facial region depressed, best seen in U. imber, where the

sutures between the nasal processes of the premaxillary and the

nasals remain more or less open throughout life.

All the Urinatoridze are holorhinal, and the dentary process of

the nasal dips downwards and forwards in a gentle curve, thus

including the’ large and somewhat elliptical osseous nares.

They are devoid of any osseous nasal septum. Each lacrymal

articulates to a very limited extent with the corresponding

frontal, and almost entirely with the outer margin of the nasal.

The os uncinatum at the inferior apex of its descending limit,

fuses with that bone, but the lacrymal does not reach down to

the maxillary in U. Zuzime, although it often does so in U. zmóer.

In many of the gulls this process is much shorter, and is bent

backwards and often anchyloses with the pars plana; in the

Urinatoride the pars plana is not ossified, and the meseth-

moidal plate is thin, showing a raised welt at the usual site of

the base of this wing where it occurs in other groups.

A large vacuity is seen in the middle of the interorbital

septum, but the optic and other foramina usually retain their

integrity. The track of the olfactory nerve is commonly roofed

over for its posterior third or more by an extension of the bony

wall which covers the anterior aspect of the rhinencephalon.

This arrangement is not seen in a specimen of the skull of

Larus glaucus, and its interorbital septum is thick and entire.

The quadrate has a long, plate-like, and upturned orbital proc-

No. 445.] OSTEOLOGY OF PYGOPUDES. 27

" ess, and two prominent facets upon its mandibular foot, at about

the same level.

Regarding the skull of any of the Urinatoridz from above,

the most striking features are the deep, sharply-defined, supra-

orbital glandular depressions. These are extensively perforated

by minute foramina over their posterior halves, while a large

irregular foramen occurs at each anterior end. Over the fron-

tals they are separated by a thin crest of bone in the median

line, while their posterior halves curve regularly outwards, to

extend upon each roof-like post-frontal projection. In U. imber

we find them giving rise to a raised superior orbital margin,

connecting the lacrymal and the aforesaid post-frontal projec-

tions, as in Alca and Uria. In Urinator lumme this rim is

sometimes incomplete. Fully as marked as these supraorbital

depressions are the extraordinary crotaphyte fossae. These are

very broad antero-posteriorly, and strongly-marked throughout.

On the top of the skull they are separated by a raised median

line of bone, being the simple backward extension of that

smooth central area of the vault, which has remained unmuti-

lated by depressions. From this these fossa sweep on either

side in increasing depth downwards and forwards beneath the

overhanging post-frontal roof and over the top of the external

auditory wing.

In a specimen of U. imber belonging to the U. S. National

Museum (No. 18256) I find the superficies of the superior orbi-

tal margins very much roughened. In the middle line there is

also a “parietal foramen " at the hinder termination of a longi-

tudinal gutter that extends as far forwards as the thin crest

dividing the supraorbital glandular depressions.

Viewing the skull of the red-throated diver from behind we

find that the large dome-like supra-occipital prominence is over-

lapped by these broad crotaphyte fosse, and the median line

separating them is extended directly backwards as far as the

superior margin of the foramen magnum. This prominence is

also transversely crossed about its middle by the raised crest

that separates the crotaphyte fosse from the occipital area.

The plane of the foramen magnum is nearly vertical, and the

reniform condyle projects directly backward from a thick-set

28 THE AMERICAN NATURALIST. [Vor. XXXVIII.

pedicle, its convex surface being inferior. We shall see further

on that this posterior aspect of a diver's skull resembles much

less the same view of the skull of any of the Laridze than Alca

does; indeed, the posterior view of the skull of the razor-bill

very closely resembles a like view of the skull in several of the

gulls.

On the under side of the skull of Urinator lumme we find

the arrangement of the palate and other elements agreeing in

all essential particulars with the gulls, auks, or guillemots ; i

other words, its structure is that of a typical cecomorph of

Huxley's classification, so far as these parts are concerned.

In a well-cleaned skull the palatines can easily be traced t

their anterior endings, and this is equally true of U. imber.

Behind they are long and narrow, showing a double carination

with a concavity dividing them. This is again divided by a

transverse ridge near the middle of the body of each bone on

its under side. The ascending processes of the palatines are

embraced by the hinder ends of the vomer, and between them

both rides the thin inferior edge of the rostrum. Anteriorly

the vomer among the Urinatoridz is more or less pointed, while

above it is longitudinally grooved for its entire length, and the

edges of this groove are well curled outwards.

The maxillo-palatines are thin concavo-convex plates raised

above the horizontal portions of the maxillaries, and otherwise

arranged as in the Alcidæ and Laridz.

The posterior heads of the palatines are in contact, and form

a groove between them above for the rostral bar of the presphe-

noid. They are embraced in a peculiar manner by the anterior

ends of the pterygoids, which are fashioned like little two-toed

feet to hold them, the larger claw being above and the smaller

one below, the seizure being of such a nature as to limit the

motion to a fore and aft one. Coues noticed this arrangement

in the loon, and alludes to it in his memoir, The posterior end

of each pterygoid is much enlarged and makes an extensive artic-

ulation with the quadrate of the corresponding side.

The foramen ovale opens laterally in the red-throated diver,

and still more so in the loon, and the posterior wall of each orbit

is marked by an outwardly concave, nearly vertical ridge, which

No. 445.] OSTEOLOGY OF PYGOPODES. 29

seems to limit the depression of the crotaphyte fossa upon that

aspect on either side. Among these divers the mandible is very

much alike.

It is shaped so as to be in harmony with the form of the supe-

rior one, being carried to a sharp point anteriorly. Opposite

tothe posterior ends of the dentary each ramal side is deep from

above, downwards, and the vacuity found in other birds upon

this surface is completely closed in by the mandibular elements

of the vicinity — the splenial and dentary — principally the latter.

As among the Alcide, however, we find a large elliptical for-

amen in the surangular in most divers, but rather a small one

in the loon in the same situation. Both the upper and lower

ramal borders are rounded and the coranoid processes fairly well

developed.

Viewed from above, we find the mandible to be V-shaped,

with rather a short symphysis. The articulate facets for the

quadrate are large and included in a squarish area in each man-

dibular end. Each angle is truncate from above, downwards

and backwards, its emarginated lateral borders behind, enclosing

a rather deep concavity, seen upon direct posterior aspect. Now

the outer of these two borders on either ramal angle is produced

upwards, forwards, and outwards as quite a prominent peg-like

process. Thus I consider the angles of this mandible as being

both truncate and recurved, and it is easy to conceive how by

gradual steps this condition in the Urinatoridz could be so mod-

ified as to have the truncation subordinated or even disappear,

while the process became the feature of the mandibular angle, as

in such a form as Lunda cirrata, wherein but little further

change is required to produce the process as found in the Gal-

line. The mandible of Larus glaucus before me. has no such

process, and the parts that give rise to it are not present, the

mandibular ends being reduced to their simple requirements for

articulation with the quadrates.

The skull and mandible in the Usiistorides are non-pneu-

matic, though apparently not always so in U. imber.

I regret very much to find that the hyoid arches belonging to

the skeletons of these divers in my hands have been unfortu-

nately lost, and I am unable to say anything about their struc-

30 THE AMERICAN NATURALIST. (VoL. XXXVIII.

ture in these birds from personal investigation. But in U. zmber

it essentially agrees with what we found in the grebes.

A complete skeleton of Urinator lumme (No. 13,646 Smithso-

nian Collection) before me, has 43 vertebrz in its spinal column.

Of these the fourteenth is the first to bear a pair of free ribs ;

the succeeding six movable vertebrz connect with the sternum

by costal ribs; the next seventeen unite as a **sacrum" with the

pelvic bones; then follow six free caudals and a pygostyle

containing several more.

The dorsal ribs are broad, and bear large, freely articulated

epipleural appendages. Two pairs of ribs also come off from the

sacrum, and meet long, sweeping haemapophyses, that reach the

costal borders of the sternum. This specimen has also a “ float-

ing costal rib," which is very small and delicate. It is seen on

both sides. The form of the skeleton of the thoracic parietes

agrees to some extent with the shape it assumes in the Alcidz,

with its hinder ribs sweeping beneath the pelvis. This latter

bone is of extraordinary form and dimensions in all of the

Urinatoridze, even excelling the grebes in some of its peculiar-

ities. The anterior portion of an ilium is short and depressed

in comparison with its extensive backward reach. The neural

‘crest of the sacrum appears above the pelvic bones for its entire

length, and posterior to the large elliptical ischiac foramen the

ilium looks directly outwards, then outwards and upwards. A

small prepubis is present, while the post-pubic element is long

and slender, its posterior extremity, curving beneath the pelvis

behind, is dilated and paddle-shaped. It nearly meets the fellow

of the opposite side, where both are completed by an emargina-

tion of cartilage. It differs from the grebes in that it articulates

with the postero-inferior angle of the ischium upon either side.

Five of the last caudal vertebrae, together with the pygostyle,

are shown in side view in my above mentioned paper (Jour. Anat.

and Physiol). The three first chevron bones there exhibited are

freely articulated over the joints of the centra when they are

present ; the ultimate ones, however, become anchylosed to the

ander side of the rear vertebra in each case, the last one really

forming the antero-inferior process of the pygostyle.

Coues in his examination of the skeleton in Urinator imber

No. 445.] OSTEOLOGY OF PYGOPODES. 31

found 13 vertebra in the cervical portion of the spinal column,

with nothing especially peculiar in their articulations,! and he

says that although they “ possess characters which most readily

separate them from those of any other portion of the column,

they yet differ greatly from each other, in different portions of

the neck... Beginning with the third vertebra, and pro-

ceeding backwards, we find that the length of the bodies

increases successively to about the 8th or 9th, when it again

decreases rapidly, so that the last one is not as long as the

third. The body of the third is thin, being exceedingly com-

pressed vertically ; and coincidently with the lengthening of

each one successively to the 8th or 9th, they grow wider, and

comparatively not so deep vertically ; those that follow, however,

do not again grow more compressed as they shorten ; but on the

contrary become broader and broader, so that the last one is as

- wide as deep, and very stout and strong. With this widening,

there is also, towards the posterior extremity of this portion of

the spine, a very high development of the transverse processes

of the anterior extremities of each vertebra. This is so consid-

erable, that the width across these transverse processes much

exceeds the length of the whole vertebra. These processes are

also exceedingly stout, with several roughened eminences for

muscular attachments ; and the foramen for the vertebral artery,

which their two roots form, is as large as the spinal canal itself.

Now as we proceed up the neck to the head, these transverse

processes project less and less from the bodies of the vertebrze,

and become less robust and angular, at the same time that they

are antero-posteriorly elongated; and possess regular lamelloid

walls, so as to form rather canals than simple foramina for the

artery.

“The ‘styliform processes’ or Viii ribs' appear to

arise from the posterior aspects of the summits of each of the

transverse processes, beginning with the third vertebra. They

are directed backwards, exactly parallel with the axis of the

!Coues, E, The Osteology of the Colymbus torquatus, with notes on its

Myology.— Mem. Bost. Soc. Nat. Hist., i, pt. ii, Nov., 1866, pp. 131-172, fig. 2,

pl. 5. This time-honored and excellent paper was also afterwards separately

issued with a slight change in its title, but apparently without revision.

32 THE AMERICAN NATURALIST, (Vor. XXXVIII.

column, and, according to their length, form a more or less com-

plete osseous covering and protection to the vertebral artery

during its passage between any two contiguous foramina."

Then after describing the neural and haemal spines of the

“cervical vertebrae"; the axis and the atlas, Coues proceeds by

saying that, “If we consider the dorsal as corresponding in

number with the ribs, we should assign ten to this portion of

the column. The last three ribs, however, correspond to verte-

bree which are completely anchylosed to the sacrum as well as

to the iliac bones, and at the same time they differ in several

respects from the dorsal ribs proper. .... e

“The transverse processes of these [dorsal] vertebrae are as

usual very broad, long, and thin; their posterior border concave,

their anterior convex, and their postero-external angles pro-

longed backwards into a short ‘ styliform ' process, more or less

intimately connected with the next succeeding vertebra. The

horizontal lamella of the transverse processes of the last four

vertebrae are pierced by a quite large foramen."

* The superior spinous processes of the vertebrz are so long

that they nearly touch each other by their anterior and posterior

borders; only a slight space being left between them. They

are quite regularly rectangular in shape, having straight, flat

superior borders at right angles with the anterior and posterior

borders. They are connected with each other by dense and

strong ligaments, and probably become more or less completely

anchylosed with age.” He then carefully describes the exor-

mous development of the hzemal spines of the dorsal vertebrae,—

and although of much the same shape, the one for U. lumme

(Jour. Anat. and Physiol.) gives but a feeble idea of their remark-

able development in Urzgator imber, the subject of the memoir

from which I have been quoting. Among themselves the

motion of the vertebrae during life in this dorsal region of the

column is wonderfully restricted, and as I have shown above, in

some grebes the dorsal vertebra all fuse into one common piece.

Coues made out 15 vertebra as being fused together in the

pelvic sacrum ; but in a specimen before me, after careful count,

there appear to be sixteen. Marsh found but 14 in the sacrum

of Hesperornis regalis.

No. 445.] OSTEOLOGY OF PYGOPODES. 33

From specimens at hand it would appear that in Urinator

lumme, the ilia anchylose with the sacral vertebrae for the entire

length of the sacrum, whereas in a specimen of U. imber before

me (No. 18256, U. S. Nat. Mus.) fusion only exists opposite the

acetabulae and thence on anteriorly to the fore end, inclusive.

In Hesperornis it was only opposite the acetabul that fusion

took place. |

As much alike as the pelves of U. /wmme and imber are, there

is still another interesting difference between them, for in the

former the anterior ends of the ilia are seen to be quite obliquely

truncated, — they are more or less rounded in the latter species.

Marsh says of those bones in ZZesperormis regalis, that “the

anterior extremity of the ilium is thin, and rounded in outline "

(Odontornithes, p. 69), but fundamentally the pelves of all these

divers are much the same.

In describing the ribs in Urinator imber, Coues remarks (p.

144) that they “are ten in number. Of these nine articulate

with the spine, and eight with the sternum. Seven only are

dorsal ribs proper; the eighth and ninth being articulated with

the sacral vertebrae posterior to the tip of the crista ilii, and the

tenth being connected neither with the spine nor sternum. .. ..

« As usual, the ribs consist of vertebral and sternal portions,

movably articulated with each other. Both of these portions

grow successively longer from before backwards ; but the ster-

nal portions much more rapidly than the vertebral. Thus while

the sternal portion of the second rib is barely three fourths of

an inch long, that of the seventh is fully three inches. The

angle at the junction of these two portions, of course, varies

with every stage of an inspiration and expiration ; but at any

given moment the angles become successively more acute from

before backwards, — from the increasing length of the vertebral

as well as the sternal portions. ... . " The last rib differs from all

the others in being unattached at either vertebral or sternal

extremity. It consists merely of two extremely slender elastic

bones, tapering to a fine point, somewhat larger and broader at

their bases, where they are joined to each other. The sternal

portion is longer than the vertebral. Close by the junction of

the two, this sternal portion sends off from its posterior border

34 THE AMERICAN NATURALIST (Vor. XXXVIII.

a small, slight process, which curves directly outwards and for-

wards, lying parallel with the posterior border of the rib, which

it joins again about an inch from its origin, — leaving a space

filled up only by membrane. This may very possibly be

regarded as the rudiment of an eleventh rib, of which the

vertebral portion is wholly wanting. It is sometimes entirely

obsolete."

“The latter ribs project so far backwards, that the thoracic -

parietes are prolonged some distance behind the acetabula, and

consequently the femur in its normal position lies directly over

thelast three or four ribs, and moves backwards and forwards

upon them. The angle of the last rib reaches within less than

two inches of the posterior extremity of the elongated obturator

foramen ” (oc. cit. pp. 145, 146).

Marsh says that “the ribs of Hesperornis present no marked

features to distinguish them from those of modern birds. They

are composed of dense bone, but some of them contain irregular

cavities. The articulated vertebral ribs of Hesperornis regalis

are nine in number, on each side. The first three of these

were attached to the last three cervical vertebrae, and had their

distal ends free. The remaining six are all well developed ribs,

which were connected by means of the sternal ribs with the

sternum ” (loc. cit. p. 63).

In comparing Marsh's figures of the sterna of Hesperornis

regalis and Hesperornis crassipes, I find upon either costal bor-

der of the sternum of the first-named species but four facets for

articulation with the costal ribs, whereas in the latter form there

are five represented; and from this I am led to believe that

there was quite as much, if not more, variation in this matter of

ribs among those now long extinct types as there is among their

existing affines. (Compare Marsh’s Plates vi and vii, Figs. 1, 2,

and I, 2 respectively.)

Grebes, loons, and the great toothed divers of the Cretaceous

period all vary in this particular. And sometimes, too, as we

know, they are apt not to agree even in the number of facettes

on the costal border of the same sternum. Frequently the

number varies for the genus Urinator now under consideration,

of the family Urinatoridz, — sometimes in the same | Bpeciely

but more often among different species.

No. 445.] OSTEOLOGY OF PYGOPODES. Xs

_A clavicle of the pectoral arch has a broad head, but is as thin

as a knife-blade, the outer aspect of which, when articulated,

simply rests against the summit of the coracoid, while its poste-

rior end rides over the head of the scapula. This expanded part

of the clavicle rapidly contracts in width as it descends, until it

becomes quite rod-like, square on section, to curve abruptly

towards the sternum, where it unites with the fellow of the

opposite side to support in the median line rather a long, peg-

like hypocleidium. Viewed from in front, the furcula is a wide

U-shaped bone, with its lower arc curved more than usually

upwards. In common with other bones of the arch, it is non-

pneumatic.

The lower part of a coracoid is much expanded laterally, with

smooth and evenly concave articular lower margin for the sternal

groove. As in some of the auks, the lower lateral margin of the

bone develops a prominent upturned laminated process. The

shaft of the coracoid is transversely elliptical on section, and its

scapular process may, or may not descend, upon its inner side,

sufficiently far as to be pierced by the foramen, which likewise

occurs in the Alcidæ. The head rears to a considerable extent

above the glenoid cavity, and its tuberous summit curls over

towards the median plane.

Larus glaucus possesses a coracoid that has the foramen in

the scapular process, as well as the laminated externo-lateral

apophysis as in Urinator, but its furcula resembles that bone as

we find it in the auks and guillemots.

The scapula in the red-throated diver is short, and doubly

truncate behind. Its curvature may be quite abrupt just beyond

the head in some specimens. This latter is transversely narrow,

and thickened from above, downwards; it occupies the entire

upper margin of the scapular process of the coracoid.

In the Yellow-billed loon (U. adamsii), when the pectoral

arch is articulated zz szfu, if the line of the long axis of the cora-

coid were produced downwards, it would cut the lower margin of

the keel of the sternum at the junction of its middle and ante-

rior thirds ; the scapulae are much tilted upwards, and the aborted

hypocleidium of the furcula is over the tip of the carinal angle

and separated from it by about a centimetre.

36 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In Urinator imber the U is by no means as broad as it is in

U. lumme, the upper expanded parts are even still thinner, and

relatively somewhat larger, and finally, it totally lacks the hypo-

cleidium. All this agrees with the os furcula of the loon, the

skeleton of which Coues described (/oc. cit. p. 148).

The coracoid of U. zmber very closely agrees with that bone

as we find it in U. Zume, but the foramen that pierces the scap-

ular process in zz/er, is a constant feature in that species. The

scapule of these two divers are essentially quite alike, except in

point of size. A point to be noticed in this latter bone, is the

fact that the head and neck is bent at a rounded angle, with

greater or less abruptness with the continuity of the blade of the

bone. This flexure (U. Zumme, Spec. No. 13646 coll. U. S.

Nat. Museum) is greater in one scapula than it is in the other ;

the angle being more acute upon the left side. It is very open

in other specimens.

In Hesperornis regalis the clavicles in the adult did not fuse

with each other at the middle point below, simply articulating at

the point of contact. This is the condition of those bones in the

very young of Urinator imber, an embryonic condition, as it were,

that persists throughout life in the great extinct ancestor of our

loon.

Comparatively speaking, in all the loons, we find the sternum

to bea very large bone. It is twice as long as it is broad, and

it has great triangular costal processes. An extensive oval

notch on either side of the keel behind gives rise to lateral

xiphoidal processes, while the mid-portion, shaped like a shield

or an escutcheon, extends considerably more posteriorly, and

does not entirely ossify around its hinder border until late in

life. It may be pierced by a few foramina, where ossification

has not been quite thorough. This part of the sternum is

unkeeled, the keel at the best being very low, but with prom-

inent and projecting carinal angle in front.

As I have said above, the usual number of facets upon either

costal border is eight ; there may, however, be but seven. The

manubrium is broadly wedge-shaped and nearly aborted; its

triangular, anterior face is slightly concaved. Costal grooves

are long and deep, but relatively, not as deep as they are in the

No. 445.] OSTEOLOGY OF PYGOPODES. 37

grebes. They almost meet each other in the middle line, the

interval between them being less than the width of manubrial

base. On its thoracic aspect the bone is concaved, being most

so anteriorly, and gradually shallowing as we approach its hinder

part. Both this surface and the ventral one are very smooth.

Upon the latter the ‘pectoral muscular line’ extends from the

mid point of the lower lip of the outer third of the costal groove

obliquely to the carina meeting it at the juncture of its anterior

and middle thirds. In U. zmerthe bone averages 20 centime-

tres for its greatest length, and 8.5 cms. for its greatest breadth ;

— the last measurement being taken across the lateral xiphoidal

processes. Though very light and elegantly proportioned the

base is absolutely non-pneumatic throughout the superfamily.

Its form is well shown in the sternum of Urznator lumme (Spec.

No. 16628 of the U. S. Nat. Mus. Coll. ?) but its pattern may

vary considerably, being remarkably narrow and long in some

individuals. "When thus fashioned it reminds one very much of

the sternum in certain of the auks. But among some water birds

skeletal characters crop out very strangely sometimes, and even

to the casual observer the sternum of an albatross, a cormorant,

Plotus, a fulmar, and a grebe all more or less closely resemble

each other upon a direct pectoral view, and to a less extent in

several of those forms, when viewed from the side.

In concluding his account of the sternum of the loon, Coues

says: * Viewing, now, the sternum as a whole, we have to

notice how great an extent of surface is secured with a trifling

increase of weight. Posteriorly, this is attained by means of the

great lateral projection of the apophyses, as well as by their

length, and by the breadth and projection backwards of the thin,

almost cartilaginous xiphoid. Anteriorly, where the sternum is

not so wide, the deficiency is atoned for by the great depth of

the keel, and its projection forward; at the same time the out-

line of the crest of the keel is such that when the inequalities

of the bone is all filled up with muscular tissue the resulting

surface becomes flat, and broad as well as long, affording the

best possible outline for contact with the water." (doc. cit. pp.

I47, 148.)

According to Marsh * The sternum in Hesperornis somewhat

38 THE AMERICAN NATURALIST. (Vor. XXXVIII.

resembles in general form the corresponding bone in the genus

Urza, but in other respects is more like that in the Aazz/z. It

is thin and weak, and entirely without a keel. It is expanded in

front, especially between the costal processes, and has two deep

grooves for the reception of the coracoids. These grooves are

placed obliquely, converging anteriorly, and are widely separated

from each other. The sternum has a rounded mesial projection

in front, which is somewhat thickened, but there is no true

manubrium." ....* The sides of the sternum in Hesperornis are

concave in outline, and in Zesperormis regalis, there are four

articular projections on each side for the attachment of sternal

ribs. These processes are all on the anterior half of the

sternum. Behind these the lateral margins are nearly parallel.

The posterior end of the sternum is quite thin, and had two

shallow emarginations. In Hesperornis crassipes the sternum

had five articular faces on each side for the sternal ribs. The

posterior margin in the same species is less excavated than in

Hesperornis regalis" (Ordontornithes. p. 60.)

If you will refer to Plate VII of Marsh's work, from which I

have just been quoting, and examine Fig. 3 of the sternum of

Hesperornis crassipes, it will not be difficult to believe that per-

haps the sternum of that species had lateral xiphoidal processes

something like those found in Colymbus cristatus. In the.

specimen they look very much as though they had been broken

off, an accident very likely to occur in the sternum of a fossil

bird, and frequently seen even in the sterna of our existing

birds in the collections in the museums. The xiphoidal part of

the sternum was cleft by a shallow triangular notch, in Hesper-

Ornts crassipes, precisely

as we find it now in many of our:

existing grebes.

THe PECTORAL LIMB IN THE URINATORID.

All the bones of the upper extremity are non-pneumatic in

this family.. They are heavy, and when simply cleaned in the

rough, they soon become dark and the oily substances contained

in their cavities ooze out upon their outer surface in no incon-

siderable amount.

No. 445.] OSTEOLOGY OF PYGOPODES. 39

The palmar aspect of the proximal end of the humerus has a

large subcircular elevation upon it that is quite characteristic.

This projects in such a manner that upon the reverse side it is

seen extending beyond the border of the bone, near the shallow,

pseudo-pneumatic fossa.

The radial crest from its size and length is more than usually

conspicuous ; its free border is a long convexity, and this plate-

like process is carried well down the shaft, occupying fully one-

third of its length. Below it, the shaft for its middle third

becomes subcylindrical, showing a large nutrient foramen upon

its ulnar aspect.

The distal end of the bone is not spread much in a transverse

direction, but otherwise rather bulky. Two wide and shallow

furrows mark it on the anconal side for the passage of ten-

dons, and a large oblique and ulnar trochlea stand out upon the

other. The ectocondyloid process is barely noticeable:

. The radius is straight, and the major part of its shaft nearly

cylindrical; its articular ends present the characters of the bone

as seen in the majority of the class. "When articulated, these

alone meet the ulna, giving rise to a long, narrow, interosseous

space. Towards this the larger bone of the antibrachium pre-

sents a concave border of a moderate degree of curvature. Its

. Shaft, too, is quite cylindrical, and faintly shows the row of

papillae for the quills of the secondaries. It develops in a trans-

verse direction not an inconsiderable ledge at its distal end, upon

which the expanded end of the radius rests in articulation.

The carpus is composed of the two elements found in most

birds; they are here simply somewhat modified in form for the

family, and to accommodate themselves to the shape of the

other bones with which they come in contact. They in turn

having their own specific cast.

One of the first things that forces itself upon our attention in

examining the skeleton of the hand of one of these divers is the

unusual length (comparatively speaking) of the metacarpals.

Of these, the one for the pollex digit is of an extraordinary

length ; much more than a third the length of the index one,

and co-ossified with it in the usual manner.

I do not recall an instance among birds where the comparative

40 THE AMERICAN NATURALIST. (Vor. XXXVIII.

lengths of these two metacarpals is anything like it. The prox-

imal phalanx of the pollex is also long and compressed. It

bears a claw upon its extremity. Both of the other metacarpals

are long and very straight, allowing but a narrow interval to

exist between them.

The blade of the proximal phalanx of the index is meagre,

being flat anconad and faintly pitted upon the opposite side.

Its distal joint also bears a claw.

The phalanx of the middle finger is fully half as long as the

expanded one of index alongside of which it lies.

These observations upon the pectoral limit of the Urinatoridze

have been jotted down during my examination of this part of

the skeleton in a specimen of U. /wmme, and in it I find the

skeleton of the manus, just described, equalling in length the

bones of the antibrachium.

The humerus in this diver has a length of about 14.5 cm.;

the radius 11.4; the ulna 11.65; manus 11.8, of which latter |

the carpo-metacarpus claims 7.5 cm. From this it is seen

that when the skeleton of the limb is in a position of rest

and closed alongside the chest, the humerus projects beyond the

bones of the anti-brachium for some little distance. This is not

the case among the Laridz, whereas it agrees with A/a torda,

Uria, and, I expect, the Alcidae generally.

THE PELVIC LIMB IN THE URINATORID.

The skeleton of the pelvic limb in the Urinatoridz is a very

interesting structure, and highly characteristic of the family.

Coues has described its mechanism and structure in U. imber

in his memoir before alluded to, and I will here record a few

observations that I have made upon this limb as found in

Urinator lumme.

The femur is short and thick, being about as long as the

cnemial process of the tibio-tarsus above the articulation. Its

short shaft is somewhat cylindrical near the middle, bowed to

the front, and scarred in many places by tuberous projections

for muscular insertion. The head is large and globular, sessile,

and deeply marked by the pit for the ligamentum teres. Dis-

No. 445.] OSTEOLOGY OF PYGOPODES. 4T

tally, it is much expanded in a transverse direction, the inner

condyle being small and elevated, the outer one being very

large, strongly cleft behind for the fibular head, much the lower

of the two, and separated from its companion in front by a deep

rotular fossa.

The patella of the red-throated diver and other loons is gener-

ally considered to be the flake-like bone articulating at the

posterior base of the cnemial process of the tibia. Its form and

exact position I have given in my paper on the patella in birds

referred to in the second paragraph of the present paper. This

illustration also presents the outer aspect of the femur, tibio-

tarsus, and fibula; the latter two for their proximal two-thirds

only.

Nothing could be more interesting than the form assumed by

the tibio-tarsus of this diver. Its cnemial process is enormously

produced, having a deep, longitudinal concavity between its pro-

and ecto-cnemial ridges in front, and the two sides meeting in a

median ridge behind. The pro-cnemial ridge is carried down as

a wing for some distance on the side of the shaft. This latter

is somewhat flattened from before backwards for its entire

length, but better marked in this particular at its distal extrem-

ity, just before we arrive at the condyles, where also it is

marked by the broad, shallow tendinal groove. This is bridged

over by the usual bony span for the deep extensors.

The condyles are very prominent in front, but approach each

other as low, sharp ridges behind. Coues found the fibula in

the loon, *for an inch or so, quite separate from the tibia; is

then united with it for some distance, becomes again distinct for

about an inch, and then finally merges as a slender spiculum

into the side of the tibia, rather more than an inch above the

joint. A slight crest, however, gives an indication of it, which.

can be traced quite to the external malleolus of the tibia." This

description agrees with one of the specimens of U. Zumme, but

in another it is carried down distinct and prominent to terminate

in a well-formed malleolus upon the lower antero-lateral aspect

of the shaft. The lower portion is anchylosed with the tibio-

tarsus, but could, with but little difficulty, be separated from it

with a good sharp knife. In other words we find specimens of

Urinator lumme wherein the fibula is complete.

42 THE AMERICAN NATURALIST. (VoL. XXXVIII.

Four figures of the Plate illustrating the present paper are

devoted to the extraordinary tarso-metatarsus of the Urinatoridce

as seen in U. /umme. These give various aspects of the bone,

and distinctly show all the characters it possesses. Chief

among these is the great amount of lateral compression of the

shaft and trochlear end. The former is grooved both in front

and behind for its entire length, forming a guide as well asa

harbor for the passage of tendons.

The hypotarsus is very large, it being composed of a posterior

arcade of bone with three distinct foramina piercing its substance

in front of it. Occupying a position above the base of the mid-

trochlea, the inner one of these three compressed protuberances

projects the most posteriorly. The remaining two are separated

by a cleft, which is continued above by a groove on the anterior

surface, to be pierced obliquely from above, downwards, by the

usual arterial foramen. The mid-trochlea is the lowest of all

three, and rather the most anterior. They are all strongly

marked by median grooves intended for the corresponding sur-

face on each proximal phalanx of the digits.

A scalelike first metatarsal is suspended by a ligament

attached to its entire anterior free border, to the inner inferior

posterior margin of the shaft of the bone. It supports a feebly

developed phalanx and claw representing the hallux digit. As

for the three anterior toes, they are composed upon the usual

formula for the number of joints as found in this member in the

majority of the class. All of the ungual phalanges are in this

diver flat and scale-like. .

The proportionate lengths of these joints in the skeleton of the

pes are shown in the following measurements : — Hallux joint

has a length of but 1.1 centimeters, its claw but 0.5; the proxi-

mal phalanx of the inside toe measures 4.0 centimeters, the next

joint 2.1, and its claw 0.95. The proximal joint of the middle

toe 3.6, next joint 2.0, next 1.8, the claw 0.9; finally the proxi-

mal joint of the outside toe 2.8, next 1.6, next 1.4, next 1.8, and

the claw 0.85. . |

Aside from the osteology and other interesting points of

structure in the pelvic limb of red-throated diver, a notable

feature is to be noticed in the great number of fibrous loops

No. 445.] OSTEOLOGY OF PYGOPODES. 43

attached to the long bones at a number of points, which serve

to surround and guide the various tendons on their passage to

the toes and prevent them from slipping from their places as

they pass these narrow bones, when the limb. is brought into

. vigorous action.

D'Arcy Thompson in his very excellent memoir * On the

Systematic Position of Hesperornis”’ in contrasting the charac-

ters presented in the pelvic limb of Æ. regalis with the corre-

sponding ones as found in the hind limb of V. imber says of the

former that “ Firstly, the extreme shortness of the femur is a

very Colymbine feature; that bone is in. CoZymbus | [ Urznator|

and Hesperornis about one-quarter the length of the ilium ;

whereas in the Ratites, except in exceptional cases, such as

Dinornis clephantopus, the two bones are nearly of equal length.

Secondly, and of greater importance, the patella, which, small

and double in the Ostrich, is rudimentary or absent altogether in

the other Ratites, is of immense size and peculiar shape in

Hesperornis. In this bird it is a long trihedral pyramid, pointed

at its superior extremity, concave on its outer surface, bearing

at its lower extremity special and separate articular surfaces

for the tibia and femur, and lying in a line with the long axis

of the femur. Except that it is perforated for the tendon of

the ambiens muscle (as in the Gannet), it is extremely like

the patella of the Grebe, and practically identical with that of

Colymbus [Urinator|, except that in this latter it is fused with

the upper extremity of the tibia. The existence of a small

additional sesamoid in the knee-joint of Co/yméus [Urinator]

(Owen, Comp. Anat. II., p. 83) does not invalidate the homology

here adopted of the long ‘rotular process of the tibia’ with the

patella. (pp. 15 12)

If Thompson means by this that in the loons (Urinator) the

patella originally was separate as it now is in the grebes and

held the same relative position toan elongated rotular process of

the tibio-tarsus, as in the latter birds, and that since, in the

loons, such a patella has come to be fused with the aforesaid

process of the tibio-tarsus, the present writer is inclined to agree

with him, although he formerly held the opinion that the small

flake-like bone described by Owen was the only patella possessed

44 THE AMERICAN NATURALIST. [Vor. XXXVIII.

by the Urinatoride. Further along I shall refer to this matter

again.

The study of the patella in birds is a very interesting, not to

say, an important one, and, as has been noted above, as long ago

as 1884 the writer published an article in the Proceedings of the

United States National Museum on the subject (Vol. VII, pp.

324-331) in which was figured the patellz of certain penguins,

mergansers, gannets, grebes, divers, fulmars, Hesperornis, crows

and cormorants ; and to that article the reader is referred for

information touching what has just been said above.

There can be no question about the existence of the patella in .

the grebes, nor in Hesperornis, nor in the cormorants, but as I

have already shown, morphologists are not thoroughly agreed

upon the nature of the flake-like sesamoid found at the knee in

a loon, nor homologically speaking, its significance. Granted that

a large patella in the Urinatoridz has fused with the long cnemial

process of the tibio-tarsus, then it would hardly appear that the

small flake-like bone in the tendon of the extensor femoris muscle

should be considered a patella at all, although in the matter of

position it agrees with that sesamoid as it is found in all birds

that possess it. It would hardly seem reasonable that Urinator

had zwo patella at either knee-joint, and such very dissimilar

ones. In my opinion the last word upon this subject remains yet

to be said. The embryology of the Urinatoridz, as well as the

morphology of the structures involved in specimens of nestlings

and subadults of the species in all stages of their growth, requires

investigating.

In the Journal of Anatomy (London) (Vol. XXIV, January,

1890, and other volumes) I published a “ Brief Summary of the

Principal Osteological Characters of the Urinatoridze " to which

I refer the reader for further details regarding the osteology of

the grebes and loons.

It now remains for me to compare the principal osteological

characters of the loons and the grebes. "These may be conven-

iently arranged for reference in the following manner : —

No. 445.] OSTEOLOGY OF PYGOPODES. 45

A FEW OF THE OSTEOLOGICAL CHARACTERS WHICH DISTIN-

GUISHED THE PODICIPOIDEA AND THE URINATOROIDEA.

Podicipoidea.— Pars plana ossifies.

Urinatoroidea.— Pars plana does not ossify.

Podicipotdea.— Supra-orbital glandular fosse but faintly mark

the skull.

Urinatoroidea.— Supra-orbital glandular fossae deeply mark the

skull, being within the superior border of the

orbit and separated from each other mesially by

a thin, longitudinal crest of bone.

Podicipoidea— Twenty-four (24) or more dorso-cervical verte-

bree.

Urinatoroidea.— Twenty (20) dorso-cervical vertebrae. Not more.

Podicipoidea.— Sternum skort and broad, with the lateral

xiphoidal processes extending more posteriorly

than the mid-xiphoidal piece, which latter is tri-

angularly notched in the middle line.

Urinatoroidea — Sternum nearly twice as long as it is broad,

with the lateral xiphoidal processes not extend-

ing more posteriorly than the mid-xiphoidal

piece, which latter is unnotched and rounded

off posteriorly.

Podicipoidea.— Posterior free extremities of os furcula very nar-

row and pointed.

Urinatorotdea.— Posterior free extremities of os furcula very

broad, laterally compressed, and apices bluntly

rounded off.

Podicipoidea.—— Posteriorly the ischium does not articulate with

the superior margin of the very long post-pubic

style, anterior to its free end; and the latter is

not perceptibly dilated.

Urinatoroidea. — Posteriorly the ischium does articulate with the

superior margin of the very long post-pubic

style, anterior to its free end; and the latter

is considerably dilated and paddle-shaped.

Podicipoidea. — Pollex metacarpal short.

Urinatoroidea. — Pollex metacarpal remarkably long.

46 THE AMERICAN NATURALIST, (Vor. XXXVIII.

Podicipoidea. — Possessed of a large patella, co-existing with an

elongated cnemial process of the tibio-tarsus.

Urinatoroidea. — Possessed only of a very small, flake-like sesa-

moid, which occurs in the tendon of the extensor

femoris muscle at its insertion ; and probably the

true patella has coóssified in the adult with the

elongated cnemial process of the tibio-tarsus.

AFFINITIES OF THE PYGOPODES.

Taken in connection with many other good characters pre-

sented in the structure of grebes and loons, we must believe

that the differentiating osteological ones just given above, point

to the fact that the relationship now existing between these two

well-defined groups of birds can best be appreciated by creating

for them a superfamily in each case. To this end I consider the

grebes to compose the superfamily Podicipoidea, and the loons

the superfamily Urinatoroidea.

In 1884 (Proc. U. S. Nat. Mus., vol. vii, p. 331) I considered

the representatives of the extinct genus of cretaceous toothed

birds, Hesperornis to be “powerful divers" and the “ancient

ancestors" of our present existing grebes and loons. Essen-

tially, this still remains my opinion; and, at a later day, after

carefully comparing the osteological characters of the Podici-

poidea and Urinatoroidea with the corresponding ones in the

skeleton of Hesperornis regalis and H. crassipes as given by

Marsh, I again said that the result of those investigations ‘ con-

vince me of the fact that, however widely separated now, our

existing loons and grebes are derived from the same ancestral

stock to which Æ. regalis belonged ” (Jour. of Anat. London,

Jan., 1890, p. 169).

Our existing grebes and loons then are derived from, or are

the descendants of great toothed divers long since extinct. Pos-

sibly the Hesperornithidz were an offshoot family of a superfam-

ily, — the Hesperornithoidea, the latter the more typical of these

extinct divers, and from them our present Pygopodes were

derived, but we yet lack the necessary material to place such a

question beyond dispute. From a consideration of the osteolog-

No. 445.] OSTEOLOGY OF PYGOPODES. 47

ical characters I consider the Podicipoidea to be an earlier off-

shoot of the pygopodine stem than the Urinatoroidea, and more

nearly related to Hesperornis than are the latter birds. The

morphology of the pelvis and the pelvic limb, as well as certain

characters in the skull and trunk skeleton, point, I think, in

favor of this view.

No doubt but what the Hesperornithidz were in their turn

derived from still more ancient ancestors possessed of the power

of flight, and in the ages to come our present-day Pygopodes,

if it be their fate to have descendants, direct or indirect, those

descendants may in turn again become flightless forms through

a gradual loss of their pectoral limbs.

So far as the affinities of the Pygopodes are concerned with

other groups of existing birds, we shall see in other memoirs I

propose to publish on the subject that they present a number of

osteological characters exhibited in common with the Alcz and

the Longipennes.

48 THE AMERICAN NATURALIST. [Vor. XXXVIII.

EXPLANATION OF THE PLATE.

(Limb-bones of Water Birds: all drawn by the author, about five-sixths

natural size.)

Fic. r. Right femur of Urinator lumme; anterior surface.

Fic. 2. Right femur of Urinator lumme; inner surface.

Fic. 3. Right tarso-metatarsus of Larus delawarensis; anterior aspect.

Fic. Right tarso-metatarsus of Larus delawarensis; distal extremity viewed

from below. ;

Fic. 5. Right tarso-metatarsus of Zarus delawarensis; proximal extremity

viewed directly from above; nat. size. Figs. 3, 4, and 5, all from the same speci-

Fic. 6. Right tarso-metatarsus of Hematopus bachmani ; anterior aspect.

Fic. 7. Direct view from below, distal extremity, same bone as in Fig. 6.

Fic. 8. Direct view from above, proximal extremity of the right tarso-meta-

tarsus of Æ. bachmani, same bone as shown in Figs. 6 and 7.

à The same bone as in last figure seen upon the inner aspect of its dis-

tal extremity. (All from specimen 13636, coll. U. S. Nat. Mus.)

IG Right tarso-metatarsus of Chiornis minor, anterior aspect. (From

Dr. Kidder's type specimen.)

Fic. 11. Direct view from above, proximal extremity, same bone as in last

figure.

IG. 12. Direct view from below of the distal extremity of the right tarso-

metatarsus of Chiornis minor. »

Fic. 13. Right femur of Chiornis minor; anterior surface. ;

Fic. 14. Right femur of Hematopus backmani; anterior surface. (No. 13636,

coll. U. S. Nat. Mus.)

Fic. 15. Right femur of Zarus delawarensis; anterior surface.

Fic. 16. Right tarso-metatarsus of Urinator lumme; inner surface.

Fic. 17. Right tarso-metatarsus of Urinator lumme; anterior surface. Same

specimen. :

Fic. 18. Direct view from above, proximal extremity, of the same bone (U.

lumme).

Fic. 19. Right femur of Urinator lumme; posterior surface.

IG. 20. Direct view from below of the right tarso-metatarsus of Urinator

lumme, distal extremity.

Fic.21. Right tibio-tarsus and fibula of Urinator lumme; anterior aspect.

Fic. 22. Right tibio-tarsus of Chiornis minor.

Fic. 23. Right tibio-tarsus and fibula of Hematopus backmani ; anterior sur-

face. (No. 13636, Coll. U. S. Nat. Mus.)

FIG. 24. Right tibio-tarsus and fibula of

face. From the same skeleton that furnishe

above.

Larus delawarensis; anterior sur-

d bones for the other figures given

No. 445.] OSTEOLOGY OF PYGOPODES. 49

Ne. Bot. Garden

1905

THE ERUPTION OF MOUNT PELEE, 1851.1

TRANSLATED FROM THE FRENCH OF LEPRIEUR, PEYRAUD AND

Rurz BY

T. A. JAGGAR. JR.

TRANSLATOR’S NOTE.

THE following account of the eruption of 1851 is a valuable geo-

graphical record for students of the recent volcanic happenings in

Martinique, and the only printed copy of the original report known to

the writer is in the library at Fort de France. The region described

as the seat of activity in 1851, a gorge adjacent to that containing the

Etang Sec, is practically identical with the site of the crater of 1902,

or parts of it, and the very careful description of earlier conditions

there throws light on many of the phenomena of the later eruptions.

The investigating commission of 1851 report (1) abundant pumice

in the old rocks of the mountain, (2) no activity at the summit lake,

(3) unusual amount of water in the Etang Sec, (4) entire absence of

lava or incandescent material, (5) fragments of diorite ejected,

(6) hot waters, steam and dry rock-dust from the vents, (7) a local

tornado, (8) sulphuretted hydrogen, (9) a buzzing noise or intermit-

tent detonations. Ina very suggestive footnote they comment on the

superficial quality of the explosions, and although not themselves geol-

ogists, they conclude that Mount Pelée has never ejected molten lava

in any censiderable amount, but has always belonged rather to the

cinder-cone type of volcanoes. There are, in fact, some ancient lava

flows and intrusive sheets, but they are rare, the tuffs and agglomer-

ates everywhere predominating.

No attempt has been made to eliminate errors in the original

account. There are some geological and chemical phrases that are

! Official Report of 1851.— Tbe manuscript was apparently drafted by Dr. Rufz,

The translator obtained access to the printed copy of this report, kept in the lib-

rary at Fort de France. Eruption du Volcan de la Montagne-Pelée. Pub. in

1851. E. Ruella & Ch. Arnand. Imprimeurs du Government, Rue du Bord de

Mer, 94. République Frangaise.

52 THE AMERICAN NATURALIST. [Vor. XXXVIII.

antiquated and obviously inaccurate. ‘The same is true of the open-

ing sentence — there is certainly good historic evidence of the activ-

ity of Pelée before 1851. The notation for magnetic directions is

obscure, and in such cases the original symbols are reproduced in

the footnote. Making due allowance for trifling inaccuracies, the

report as a whole is the most complete and scientific record extant

of the topographic details of the southern slope of Pelée prior to the

eruptions of 1902—03.

October, 1903.

THE OrFiciaL REPORT, 1851.

A tradition without historical foundation records Mt. Pelée to

be a volcano. The conical form, the crater lake, the pumiceous

soil, all fostered this supposition. It was known also that in one

of the gorges there was sulphur, and the inhabitants living near

called this the Soufriere. The druggist Peyraud made a scienti-

fic excursion there in 1838! and brought back stalactites of

pure sulphur attached to the leaves of a tree.

! Letter to the * Directeur de l'intérieur," by Peyraud : —

Monsieur le Directeur général.

Jai l'honneur de vous addresser un petit flacon contenant de l'eau que j'ai rap-

portée de la source minérale qui se trouve sur les terres de M. Huc, à la naissance

de la Riviére Claire. Je dois vous observer que cette eau que l'on mavait dit

être à 60° Reaumur, nést dans ce moment qu'à 16°, et l'air ambiant à 18°. J'y

joins un échantillon de la matiére jaune qui tapisse les bords du morne d'ou cette

eau découle, puis un morceau de bois recouvert d'un mousse qui répand une forte

odeur de marée. Ce bois a été recueilli à

minérale, un peu à gauche en faisant face à

Plus tard je ferai en sorte de vous

endroit òu se trouve la source. Si vous désirez une plus

grande quantité d'eau je me ferai un vrai plaisir de vous en faire puiser, que je

renfermerai dans les flacons à l'émeri afin d'éviter la perte du gaz.

Je compte aller, dans quelques jours, parcourir la route de la Trinité afin de

visiter l'endroit qui répand une odeur sulfureuse et vous rendrai compte immédi-

atement du résultat de mes recherches. '

Je suis avec respect, Monsieur le Directeur général.

Votre tres humble serviteur,

Saint Pierre, le 20 Mai, 1838. P. PEYRAUD.

No. 445.] ERUPTION OF MOUNT PELEE. 53

Recently the inhabitants of the Précheur heights have com-

plained of a strong sulphurous odor. Some of them have visited

the Soufriére and report a fuming hole which has appeared, the

vapors mixing with the clouds and thus being imperceptible to

sight at a distance. On the 3rd of August the patrolman Car-

bonel brought in a partridge found dead near the new opening,

and he said there were many others, that all the birds dropped

dead when they flew over the fatal cavity. Like the Avernus

of the Ancients,

Quam super haud ullae potevant impune volantes

Tendere eter pennis: Talis sese halitus atris

Faucibus effundeus supera ad convexa ferebat ;

Unde locum Graii dixerunt nomine Avernon.

Virgi.

This year (1851) from the roth of May, there were no earth-

quakes in Martinique, but Guadeloupe had many of them and

was in continual fear. August 5, St. Pierre was peaceful ; the

weather had been fine. Towards 11 p. m. a dull, distant

sinister noise began, like thunder; it was mistaken for thunder,

or for the roar of the river in flood. The noise increased, waked

many people and caused alarm.

I was in my villa of Fonds Canonville, very near the source

of the sounds. I thought it was thunder, but was astonished at

its continuance; [ heard the workmen call me from without.

They shouted * Do you not hear that noise?" I answered,

“Ves, it is thunder!" “No, it is the Soufrière.” I rose and

looked towards Mt. Pelée, but could see nothing: the noise con-

tinued. The rest of the night was passed in great anxiety, and

torches were seen indicating the flight of many people. No

one knew anything definite and the answer to all questions was,

* C'est la Soufrière qui bout !”’

St. Pierre was equally frightened. In the morning roofs,

pavements, leaves of trees, all were covered by a thin layer of

grayish cinders, which made the town look like a European city

covered by the white frost of the early days of autumn.

These ashes covered the country between the city and Mt.

Pelée, covered Morne Rouge, and extended to Carbet. The

54 THE AMERICAN NATURALIST. [Vor. XXXVIII.

stream called the Riviére Blanche, because of the color of its

water (caused, like the * yellow baths " of Guadeloupe, probably,

by the presence of an iron hydrate) became black, charged with

ashes or mud, and this coloration of the water could be seen far

out to sea, as in time of flood.

The spirit of the city was an anxious curiosity, dissimulating,

according to the light-hearted spirit of the country, under many

jests. A few brave spirits made a reconnaissance of the moun-

tain, and from their description the first accounts were published

in the ** Courrier de la Martinique " and in * Les Antilles."

The government appointed an investigating commission ; Le

Prieur, chief Pharmacist of the colonial hospitals, who had

already made several explorations in Guiana: Dr. Rufz, and

pharmacist Peyraud. The present document is the report of

this commission.

The general aspect of La Montagne Pelée, seen from St.

Pierre, is that of a great cone, from whose summit descend

sharp spurs down to its base, these spurs being separated by an

equal number of gorges or valleys. It resembles the cone

formed by a pleated filter paper. In order to reach the summit

of the mountain it is necessary to follow one of the spurs, for

the valleys are often precipitous gorges. On August 28, we

took what is considered the most direct route, by way of the

sugar plantation “ Rivière Blanche," near the farm Paviot, and

came out at the habitation Ruffin. Ruffin is a farm 14 hours

horseback from St. Pierre, 551 meters above the sea. We slept

there: 7 a. m. the morning of Aug. 29, the thermometer marked

in air 23° Centigrade and 22.5? in the earth. At St. Pierre at

this hour, 27° or 28°. We descended by a zigzag path to the

bottom of the ravine of the Riviere Claire. This path is cut in

* punites," or local pumice stones which are white and friable,

in little fragments. The Rivière Claire is so-called by way of

contrast to the Rivière Blanche. The latter, whose waters are

always milky, comes from one of the creases in the mountain

separated from the Riviére Claire by a steep spur: both flow

separately above, but unite below, and continue to the sea under

the name of the Riviére Blanche. It appears that now the

Riviere Claire receives the volcanic mud and blackens the

No. 445.] ERUPTION OF MOUNT PELEE. 55

Rivière Blanche, no longer deserving its name of “clear.” At

the bottom of the ravine where we were, the Riviére Claire forms

a cascade 2 or 3 meters high: it is only a brook that one can

easily jump over. Its banks were covered with from I5 to 20

centimeters of mud. The water has ceased to be drinkable.

After having crossed the bed of the Riviére Claire, it became

necessary to climb the opposite slope which is a very abrupt

escarpment, where the first explorers had to construct a path, by

means of a ladder of ropes and lianas: for a half-hour one goes

upstairs in this fashion. The slope becomes easier and we are

in virgin forest. There is no path. In order to advance in the

direction of the hill one must push through palms, and tree-ferns,

the long spines of which make treacherous support. This is

the forest zone called the “little woods” in the colonies as

opposed to the “great woods" where there are great trees

remarkable for their height and the size of their trunks. About

ten o'clock we reached the point where the first traces of the

eruption were visible. The foliage of the ferns, bananas and

other plants which form the vegetation of these places was dried

and reddened as if it had been burned ; on the leaves and on the

soil also there was a thin layer of dried mud, the remains of

volcanic ejections. Here the barometer registered a height of

846 meters. From there on the volcanic mud became more and

more abundant. It covers the leaves in a dry adherent mass,

and on the ground it is viscous and sticky and appears like a

gray clay, in some places more than a foot thick : little scintil-

lating points may be distinguished on the surface which the

lens, and later a chemical analysis, showed to be globules of iron

sulphide. As we had noticed that the leaves, branches and

trunks of trees were plastered with mud only on the side toward

the volcano, we thought at first that the weight of the mud was

sufficient to explain the inclination of all these objects; they

seemed to be depressed violently, but the quantity of mud on

the leaves was only a few millimeters thick and not sufficient to

break the branches; we soon saw that something more than the

weight of mud must be called in to account for the disorder,

and a sort of chaos through which it soon became necessary to

thread our way. Here were great trees broken, overturned,

56 THE AMERICAN NATURALIST. (Vor. XXXVIII.

twisted, not only in one direction but in all directions, and often

in a direction the reverse of that affected by the weight of the

mud. This brought to us the conviction that something like an

explosion had taken place. The air, displaced first by hot

vapors blown out of the volcano, and rushing into the vacuous

places so formed, became involved in a whirl or local tornado :

we say local for the signs of disturbance were only in the

vicinity of the volcanic opening. The people who fled with

torches from the houses lower down the slope did not have their

flambeaux blown out by the wind.

Climbing higher, the mud layer became thicker making the

walking difficult and woe betide those with light or ill-made

shoes! At the same time the crest of the spur we were fol-

lowing became narrower ; a moment arrived when we had on the

east, on the right-hand side, the ravine of the Rivière Claire.

It was this last we were following, guided by the vapors that

were rising from this gorge. Beside it we walked on a high

crest rising above the bottom of the valley 50 to 60 meters ; at

our feet the valley of the Riviére Claire was plainly visible.

From this point we obtained a full view of the effects of the

eruption of the 5th of August, in all its extent and horror. The

verdure is absolutely gone from the place which was formerly

the scene of densest foliage. Trees, leaves, flowers, all are

buried as though under a gray shroud. It is the sadness of

winter with the trees denuded and smutted with a black snow.

Though we found no dead birds, we also heard none of them

singing. The mountain whistler, whose sweet pipe is associated

with the melancholy grandeur of our “grands bois,” the whistler

of our mountains has fled from these his former haunts; we did

not even meet the deadly trigonocephalus whose home is here.

A dread silence, a sky obscure by vapor, an atmosphere charged

with a strong odor of sulphuretted hydrogen, complete this

scene worthy of Tartarus. The slope of the bed of the ravine

in the midst. of all this desolation, is very steep; it extends

from the east toward the west from the summit peak of Mount

Pelée called * Morne LaCroix,” but the bottom is not continuous ;

it is interrupted by several cliffs; here and there, in the midst

of the general grayness rifts may be seen where the soil is rent

No. 445.] ERUPTION OF MOUNT PELEE. 57

bare; these are crevasses made perhaps by the steam of the

mountain or by earthquakes about the rent. Ancient pumices

may be seen of various colors, reddish or grayish, and among

these are some rare masses of gray dioritic rocks, or of por-

phyries that the ancient fires have hurled out of the entrails of

the earth along with the pumice.

In the upper part of the valley there are rising dense sheaves

of whitish smoke which mark two volcanic vents from which

came the mud that covers the valley and its environment. At

eleven o'clock we arrived at the level of these two craters, from

which we were separated only by the valley itself. A barometer

reading here gave an elevation of 966 meters above sea-level.

The temperature of the air was 23°.5 at twenty-five minutes after

eleven o'clock ; after removing the mud, the temperature given by

a thermometer buried in the soil was 217.5; this was 44, more than

the heat found at the lower station, where the persisting vegeta-

tion protected the soil from the direct action of the sun's rays.

But this was not the upper limit of the action of the volcano.

It was easy to see that mud had been thrown several hundred

meters above the orifices, for the line of green verdure did not

begin to appear except at the very summit of the mountain.

We wished to reach the summit in order to see better the actual

extent of the lands covered by erupted material, but our guides

assured us that the ascent here was not possible, that we should

be cut off. by cliffs and that there was no path. Looking over

the scene of desolation, we estimated that the action of the vol-

canic eruption took place within a perimeter of something more

than eight or nine hundred meters.

It was necessary to go down to the bottom of the valley in

order to explore and’ examine the orifices themselves; to do

this we had to slide down, with the aid of roots and lianas, an

abrupt cliff 15 or 20 meters high; the younger and more agile

members of the party decided to do this under the leadership of

Mr. Peyraud. As they approached the fuming vents ahead of

them, they found the mud deeper; the valley was divided in two

branches by a spur from the summit of the mountain. In the

right hand branch are the two craters; they lie in a N-S line,-

separated by a space of 25 to 30 meters, in the midst of which

58 THE AMERICAN NATURALIST. (VoL. XXXVIII.

the waters which descend from the mountain have worn a chan-

nel 12 to 15 meters deep. At this time the vents were almost

free from vapor — something hitherto unknown since the day of

the eruption. Therefore it was possible to examine them

thoroughly. The left hand vent opens in the right slope of the

spur which divides the upper part of the valley; it is of irregu-

lar, circular form and three or four meters in diameter. From

this opening the thick vapors are discharged with the greatest

force; these appear brilliantly white in the sunlight, spreading

to leeward in a billowy plume, but they are denser and blacker

in the immediate vicinity of the orifice; the emission of vapor

is not preceded by any subterranean noises. The detonating

sounds heard take place at the instant the vapor escapes from

the hole, and this leads us to the belief that this noise is the

result of the expansion of steam in air, a phenomenon entirely

comparable to the detonation of a cannon.

This opening is reached by a sort of open gallery about a

meter and a.half wide excavated in a reddish pumiceous rock

covered with dark gray mud, which was unstable and so hot that

it was impossible to hold it in the hands more than a few seconds,

especially that on the immediate rim of the crater. Here the

attempt was made to sound the depths of the cavern by means

of a zinc pail attached to a cord. But when the cord was with-

drawn the pail had disappeared without evidence as to whether

the loss was due to insecure knots or the melting of solder.

The cord was impregnated with a strong smell of sulphuretted

hydrogen. Stones dropped into the hole were heard to fall

quickly with a noise as of striking a liquid. An alcohol ther-

mometer buried in the soil rose almost to its highest limit.

The barometer gave an elevation of 88 3 meters, indicating that

these rents were 83 meters below the summit of the spur, where

the rest of us had remained.

This crater had already been visited by Mr. de Maynard but

that was in the first days of its formation when steam action was

so violent that details could not be seen.

Across the gulch, but separated as we have said by twenty-

five or thirty meters, is the other higher orifice, and this is also

the greatest vent, — it is harder to reach than the first, lying at

No. 445.] ERUPTION OF MOUNT PELEE. 59

the bottom of a great four-sided funnel behind which is a very

high cliff; this funnel opens against the wall of the cliff like a

great chimney. One of the guides at that point venturing a

little too near slipped on the steep slope leading to the orifice

and was almost thrown in, but happily he checked himself just

in time and we were saved from a horrible tragedy.

It was noon, and the openings, which up to that time had been

giving off verv little vapor — so that no steam could be seen

from St. Pierre — began to puff and give off an odor of sulphu-

retted hydrogen more strongly than heretofore, and at the same

time detonations could be heard. We thought of returning, but

we wished to visit a third crater seen fuming some hundreds of

meters lower down the same ravine, which was said to be the

seat of the ancient Soufrière.

Two routes appeared practicable: the one following down the

steep crest that we had come up by, in order to find, a little

lower down, a less steep slope; the other going directly from the

upper orifices to the one below following the same ravine longi-

tudinally; the first of these routes was, according to the guides,

the only practicable one; the other passing escarpments and

cliffs led to an inaccessible point above the vent. Mr. LePrieur

and I followed the guides, Mr. Peyraud and some others who

were with us wished to try the more difficult route, but after

much trouble, they: were obliged to retrace their steps and

rejoin us. They had been cut off by those precipices which on

the mountain are invisible a very short distance away, but which

prove when one reaches them that the straight line is not always

the shortest road between two points.

After going down a half hour we arrived at the level of the

third crater, but it then became necessary to drop down into the

ravine for a depth of fifty feet by sliding on a slope inclined

about thirty degrees. Finaly we reached the bottom and

found the bed of the ravine four or five meters wide, in the

middle of which a swift brook runs through a bed of grayish

- mud. This mud has a thickness of half a meter. On the two

banks rise wall cliffs eight to ten meters high. The bottom of this

gorge is thus in a fashion shut in. To the east, above, in the ,

direction of the higher vents that we had just left, the ravine is

60 THE AMERICAN NATURALIST. (VoL. XXXVII.

shut off by a rock wall at the summit of which gushes out the

steam of the third crater; along with the steam, a cascade of

water escapes, which falls into the ravine and forms the brook

mentioned. Unfortunately it was not possible to climb to the

edge of this hole and get a good view of its dimensions.

The cliff which rises above it at the back is hollowed out in a

sort of cavern, and before it the crater forms a v shaped breach

whence comes the water of the cascade; it is probable that the

arrangement of these places changes from time to time under

the different forces at work, for Mr. Peyraud who had been here

in 1838, found the place unrecognizable in 1851. Besides the

crater of to-day which was quite new to him, there was lacking

a cold spring which our companion had promised us and which

for several hours we had counted on for quenching our thirst.

We found only springs of hot water, the coolest of them at 37°.

On the heights above the ravine, the Abbé Lespinasse, during

the first days of the eruption, had planted a cross, in order to

reassure the frightened populace. This is the only barrier that

man dares oppose to threatening nature at such a time. The

presence of hot water here appeared to us an important basis of

more extensive study. On our right looking to the east, about

a meter and a half from the bottom of the ravine and three

meters from the rock wall which cuts it off, there is a hot spring

having a temperature of 70°; its taste and its odor indicate the

presence of sulphuretted hydrogen. The principal jet is about

an inch in diameter and all about it are tiny rills at the same

temperature: two meters above this first spring and a little

further along there is a second spring of the same quality with

a temperature of 46°, which falls, by several little cascades, on

the rock, from which it rebounds below. The action of the air

liberates a great part of the sulphur contained and this is depos-

ited in a light powder on the lower rock, which is coated yellow-

ish white, but in falling lower down on another ferruginous rock

It 1s again decomposed and the sulphur this time combines with

the iron to form a black sulphide of iron.

On the same side going higher another spring is found with a

temperature of only 22°; this has a still more sulphurous taste.

On the opposite bank two meters from the bottom and one

No. 445.] ERUPTION OF MOUNT PELEE. 61

meter from the end cliff there is another sulphurous spring with

a temperature of 90°. The muddy water coming down the

brook has very variable temperature ; during our exploration,

Aug. 29, it was 37°, but on the 30th it was 65° (for the study of

the springs about this third crater was found so interesting that

Messrs. LePrieur and Peyraud returned twice to take the tem-

perature and to get some bottles of water for analysis). It is to

these gentlemen that we are indebted for detailed information

concerning the springs; they determined that the temperature

of the different springs coming out of the rock was always the

same, while that of the main brook varied; this difference

according to them is due to the ‘fact that the materials thrown

out of the crater mix with the water of the brook, the source of

which is rainfall. This is properly the head-waters of the

Riviére Claire. According as the ejectamenta are more or less

abundant the water is more or less hot. This we had oppor-

tunity to prove Aug. 29th, for while we were in the ravine

making observations, suddenly several detonations were heard

and at the same time blacker and more abundant fumes came

from the crater; the water of the brook changed quickly in

temperature rising from 27° to 47°. There were twenty of us

crowded together in the ravine. Nearly every one was afraid

and each sought safety or the adjacent cliff a few meters high.

But as this could be reached only by one very narrow path, in

single file, the crowding added to our panic. To the credit of

Messrs. LePrieur and Peyraud be it said they did not share our

fright and remained behind facing the eruption and continuing

their observations, and they did not rejoin us until a long time

afterwards. At the moment of the noise and the ejection of

vapor a cloud of grayish mud was thrown into the air which fell

in a very fine powder on our hats and clothing. The brook

increased in volume to a blackish gray boiling mass. It did not

appear to ever reach a depth of more than one meter, for the

mud-stains on the wall were not visible at any point above this

height. After three successive explosions the crater became

quiet and the vapor became whiter. It was at this time again

determined that the noise took place entirely at the orifice of

the vent by the expansion of steam in air, and was in no sense

62 THE AMERICAN NATURALIST. (VoL. XXXVIII-

subterranean, for the puff of vapor and the noise were exactly

simultaneous. We saw no ejections of fire, stones, or even sand ;

the only accident that was somewhat remarkable was the detach-

ment of a block of ferruginous rock from the right side of the

ravine, which rolled into the depths and broke into pieces. This

rock measured twelve to fifteen cubic meters. On the sixth

of September, Messrs. LePrieur and Peyraud on their second

return hither determined that in spite of the thick masses of

vapor thrown out since the 29th of August, and in spite of

rain which had fallen twice on the mountain, the level of the

brook had not risen. They said that our foot-prints of the last

excursion were still visible in the mud of the bottom of the

ravine, some of them containing a little water, evidently from

rainfall: in the interval between the two visits there was no

change, and the different springs showed the same temperature ;

in the spring at 70? one could boil eggs, and some of the native

crawfish were found cooked.

A copper bucket was thrown into the crater securely attached

to a cord; three times the vessel was thrown against the rock

wall and was recovered dented, but it was not possible to obtain

the slightest particle of liquid; the material of the interior was

found to be reddish pumice which accounted for a reddish color

that had been at one time observed in the vapor coming from

this orifice. While we were there, however, this steam and that

from the upper crater always appeared to us white or blackish,

the last condition being due to particles of sulphurous mud.

In order to complete the exploration of the scene of eruption

it remained to examine the valley where the first openings were

made. For the openings that we have just described are not

those which opened first. Those which are in action today

opened on Saturday, the oth of August, aftera series of detona-

tions more violent than those of the sth. The first openings

ae situated in a lateral valley which joins the one where the

active craters are, but much lower down. They are today

entirely quiescent. From the calm which reigns in their vicinity

one would never suppose that a month ago they had served to

give vent to the first manifestations of a volcano. These

openings are ten in number, along a single line, and trend in the

No. 445.) ERUPTION OF MOUNT PELEE. 63

same direction as the upper vents; behind them is a high cliff

which forms the right side of the ravine they occupy. The

first is at the same time the lowest and smallest ; its diameter is

0.60 meter to 0.70, and its depth about 0.30 m. The bottom is

covered with a reddish ferruginous sand and the rocks there-

abouts are brownish and tumbled in disorder, doubtless the

product of the eruption. The ravine has been deeply excavated

by water which has removed the greater part of the soil and left

only denuded rocks. The different openings are on a rather

steep slope. The tenth which is the highest of all is also the

widest and deepest; it is four meters long and a meter and a

half wide. Above there is a tree supported by its denuded

roots; this vent is quite deep and appears to contain water still,

but a line forty feet long failed to reach the bottom ; the third

opening, counting from below, is also of elongate shape; it is.

not remarkable except that it is half covered by a boulder which

has not been displaced. There are also some plants which have

not been totally destroyed and their roots hang denuded in the

opening. The bottom of this cavity is, like that of the other,

covered by sediment colored with iron oxides. An elevation

taken at half past three in the afternoon in the middle of these

small openings gave 816 meters above the sea. The thermom-

eter gave 20.5? in the air and 23.5? in the soil ; much rain fell on

that day.

In returning to the habitation Ruffin and passing the Morne

Plumet, — the most elevated point of this part of the mountain

and hence named Gros Morne,— one does not encounter any

trees, the only vegetation being shrubs, grasses and sedges;

at this height (812 meters above the sea) there is a fine view ;

directly opposite is the ravine of the Claire within which are the

three active openings, to the left extends all the district of

Précheur with its picturesque farm houses situated each on its

little hill, and to the right in the distance may be seen the wide

landscape of St. Pierre, the city itself with its reddish roofs, and

the vessels anchored in the roadstead.

The general direction of trend of the ravine where the active

steaming vents are situated is ENE.’ From the summit of the

! The French notation used is N. E. 1/4 SE.— W. 1/ 4 W. (ne 1/4s, e-o 1/4 0).

64 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Morne Plumet the orifice nearest to St. Pierre (that from which

the thickest fumes come) is to the east 30^ north ; the second is

32^ north and the third which opens in the ravine is to the east

36° north. St. Pierre lies to the south 10° east! at a distance

of about 10 kilometers as a bird flies; the town of Précheur is

about 7 kilometers directly west. The Ruffin house, which is

the nearest dwelling, is about 3 kilometers from the upper

opening, and 2 kilometers from that situated above La Sou-

friére; the sugar factory Canonville is 5 kilometers away.

From the Grande Rue du Mouillage the upper opening is north

4° east ;? it is this which is best seen from below and from this

escaped the densest vapors; these entirely mask the second

opening which in this direction is behind the first; as to the

vapors thrown out by the opening situated in the ancient

Soufrière, they cannot be seen from St. Pierre; the high crest

which separates Riviére Blanche from Riviére Claire hides this

opening entirely.

The 29th of August we returned at two o'clock to the Ruffin

House. Our excursion had occupied about seven hours ; at the

cost of some fatigue we bore with us the memory of one of the

most imposing spectacles man had ever seen. But our task was

not finished. We had to assure ourselves that there were no

other points in the mountain where changes had taken place in

consequence of the eruption of the fifth of August. Of course

it was important to determine what had happened to the hot

waters which exist in the part of Mt. Pelée called Montagne

d'Irlande where Mr. A. Desnoux de Messirny has built a bath

establishment. Sept. 2, Messrs. Le Prieur and Peyraud betook

themselves to that locality and found the water of the spring

itself at 35°.8, and at the first faucet of the first bath of the

establishment (which is ten minutes walk distant) we found the -

temperature to be 33°. The weather was clear ; — after heavy

rains have fallen on the pipes, the loss of heat between the

spring and the baths is still greater. Dr. Dutroileau who was

at the establishment on account of his health on the night of the

fifth of August assured us that no change happened to the water

! ^ Du compas " — presumably magnetic.

26N 1/44 E."

No. 445.] ERUPTION OF MOUNT PELEE. 65

either in temperature, volume or limpidity. The air here as in

all the Précheur district is strongly impregnated with the odor

of sulphuretted hydrogen and all silver pieces turned brown as

well as those paintings which contained compounds of lead. In

the night of 5th to 6th of August and the oth of the same month

strong but short shocks of earthquake were felt ; since that time

the earth has remained at rest, but from time to time strong

detonations may be heard, similar to cannon shots in the dis-

tance. At the same time it is stated that a movement is felt

such as might be produced by a powerful blow struck beneath

the soil. This sensation was also perceived at the Ruffin House

but at my residence Fond Canonville, which is not on the mas-

sive rock of the hills but on the seacoast, I perceived nothing

of the sort.

All along the road leading to the baths and at the spring

Messrs. Le Prieur and Peyraud perceived no landslips, even

though the walls of the ravine by this road are formed of tufa

20 to 25 meters in height, frequently deeply trenched. This

soil is composed of pumice in masses or fragments on which

the water has deposited in certain places incrustations. Above

the bathing establishment toward the Carbonal House many

rocks are found high up the slope which are friable and fine

grained, horizontally bedded, and resting on the fragmentary

pumice ; these rocks are variously colored and serve as support

to the pumice; this suggests that often eruptions had taken

place, throwing out pumiceous rocks and that in the intervals

the flowing water in its turn had deposited the particles held in

suspension during the calm periods. Along the lower part of

the road all these beds are very well shown, especially in those

portions artificially trenched ; there are there several good sized

heaps of ancient volcanic cinders of a faint violet tint and rather

more sandy in quality than the muds thrown out today ; there

are no metallic particles ; some are reddish like the sands found

near the little extinct vents, or on the steep slope back of the

vent called La Soufriére.

There remained another important point to determine:

namely, what had happened to the actual summit of the Mon-

tagne Pelée, where there is a lake supposed to be the seat of an

66 THE AMERICAN NATURALIST. (VoL. XXXVIII.

ancient crater. Seen from St. Pierre the mountain did not

appear to have changed at all in height, but had nothing hap-

pened to the waters of the lake? Noone knew, for no one had

been there since the eruption of August sth, and on this account

Messrs. Le Prieur and Peyraud resolved to explore the summit

on 4th Sept. The trail to the summit of Mt. Pelée is more

travelled than that which leads to the craters. The inhabitants

of St. Pierre sometimes make picnic excursions to the summit.

Leaving the Eynard House near the base of the mountain, the

spur is followed which leads beside and overhangs the ravine of

the Riviére Séche. For three quarters of an hour one passes

cultivated lands, the earth there being loose and formed of frag-

mentary pumice covered by a thin bed of vegetable mold which

is very permeable ; vegetables are raised on these slopes. Then

the traveller comes to the great woods, fig trees covered by

vegetable parasites, and long lianas which climb to the very

summits of these forest giants and then swing back to earth

where they throw out roots and form a dense mass of vegetation

entirely distinctive of the tropics. On leaving these woods at

the end of an hour, the trees are seen to grow smaller and are

gradually replaced by low shrubby and herbaceous vegetation ;

but none the less the botanist finds here too a constant source

of delight. I saw, wrote Mr: Le Prieur, superb flowers worthy

of hot-houses, especially two superb cromelias, the one with a

long spike of flowers, the other with yellow and red flowers ;

there are the Brazilian huckleberries, with violet-red flowers ;

three beautiful Species of lobelia, with great flowers which recall

some of the fields of certain portions of France. This last plant

is found about the lake, and on the humid slopes leading to

Morne LaCroix, the culminating point of the mountain — so

named because a cross has been placed there. In general, of all

the floras of the Antilles, that of Martinique is least known

because of the trigonocephalus, whose terrible reputation fright-

ens away the hardiest botanist. Our explorers started at seven

o'clock and reached the lake at half past ten. According to the

guides who are accustomed to the place no change has taken

place in the lake. It is some three hundred paces in circumfer-

ence ; the thermometer gave a temperature in air of 19° and in

No. 445.] ERUPTION OF MOUNT PELEE. 67

the water 207.5. Nevertheless, the temperature seemed much

colder on account of a north wind blowing strongly at the time,

and the dense fog over the mountain. This prevented them

from viewing the magnificent spectacle ordinarily seen when the

weather was fair—a view inclusive from the Grenadines to

Antigua. The water of the lake was as abundant and as clear

as usual. A maceration within it of certain vegetable matter

gives it a grassy taste. Before arriving at the lake it was nec-

essary to cross a crevasse 40 m. wide, which crosses the whole

width of the spur that they followed, and is well known to those

who have taken this excursion. This crevasse has not been

changed at all nor widened. Neither on the road nor from any

point of view was any trace of disturbance seen.

The bottom of the lake is carpeted with a layer of thin mud

and this rests upon a heap of fragments of pumice of yellowish

gray color partly decomposed and recemented by a little ferrugi-

nous clay. On the southwest border of the lake a small beach

has formed, composed of very fine grains from the debris of

these pumiceous rocks which the movement of the waves, raised

by the north wind, bring there continually, for there is not on

that side any elevation to protect the rocks from the action of

the wind. At the summit of Morne LaCroix the barometer

gave an elevation of 1277 meters above the level of the sea.

This is the highest point of the island. The thermometer in

air gave 18°.5 and in earth 19^.2. From this point steam could

be seen toward the west, coming out of the upper craters 400

meters lower down. A little more to the left not quite so far

down there was seen from time to time a water surface showing

bluish reflection, filling the basin called formerly by the guides

the “dry pond” (étang sec), because ordinarily this engen is

empty.

Even on the plants of the summit of Morne La Croix, traces

of volcanic cinders were found which had been carried to this

point. Messrs. Le Prieur and Peyraud, not being willing to

leave the mountain without a visit to the dry pond — which is

commonly believed to be another more ancient crater of the

volcano — visited it during their descent by means of a trail

rarely used, very difficult and seldom visited before they went

68 THE AMERICAN NATURALIST. (VoL. XXXVIII.

there. They found this supposed * dry " pond filled by a consid-

erable mass of water and according to their estimate five times

greater than the upper lake: the guides attributed the presence

of this water to the abundant rains which had truly been

extraordinary during the “hivernage” of 1851; they asserted

that during the previous lenten season this pond had been dry.’

The barometer here gave an altitude of g21 meters above the

level of the sea. Thus this dry pond is at almost the same

elevation as the upper vents of the volcano which are in a ravine

beyond. Nothing else was found changed in these localities

according to the guides, and nothing in the way of fissures nor

disturbances. After this last expedition our official work was

finished ; we had learned that the action of the volcano did not

extend beyond the limits we had explored, and that it was con-

fined to the ravine where the Riviére Claire takes its rise and

that immediate vicinity; It was useless to examine the north-

ern slope of mountain toward Macouba, for the inhabitants of

that quarter observed nothing extraordinary except an odor of

sulphuretted hydrogen, which, it is said, was perceived even as

far as St. Marie. At Macouba the leaves of the trees were

coated with only the barest trace of those cinders which caused

such a fright in St. Pierre. É

1Some naturalists think that the heat of certain volcanoes comes from no

great depth and the water which they throw out is merely rain water which

penetrates by means of the fissures in the earth and accumulates in subterranean

cavities ; several of the observations made during our eruption of the fifth of

August lend support to this opinion: (1) the years 1850-1851 had been very

water and this has become for us a sort of pluviometer ; (2) the shocks of earth-

ensa r the soil were felt only on the slopes of

the mountain, in those little estates situated probably above the level of the

as at my house in Fond Canonville) although they are

volcano, nothing of the sort was felt. Moreover at .

Fond Canonville the Sprin i

like the Fontaine Chaude, have not been in any way changed. It seems to me

furnace is in the body of the moun-

No. 445.] ERUPTION OF MOUNT PELEE. 69

Conclusions.

Doubtless no dissertation will be expected from us on the

cause and nature of the Mt. Pelée volcano. This study would

require a professional knowledge other than we possess; we

have merely tried to fulfill the request of the government and

report upon the extent of the accidents occasioned by the

eruption of the 5th of August and certain exact details. We

will vouch for nothing beyond the facts which we ourselves

have determined. If these facts, compared with those pos-

sessed by science already à propos of the other volcanoes of

the earth (more than two hundred in number) can throw some

light on the nature of this great phenomenon we shall be well

satisfied. But it is not our task to enter upon such matters.

The following we can vouch for : —

The eruption of the fifth of August was entirely a local event

bounded by the ravine of the Riviére Claire, devastating an area

800 to 9oo meters broad at the outside.

The effects of the eruption were at first a continuous buzzing

sound, then a series of intermittent detonations, and simultane-

ously there was thrown out a jet of white or black vapors which

made a deposit wherever they spread in the shape of a grayish

mud or cinder; and these vapors produce in their vicinity a

strong odor of sulphuretted hydrogen.

It is not possible to assert whether this material is always

thrown out in the form of cinders or powder, or not sometimes

in the form of a rain of mud. It is more probably in a powdery

condition and when it falls on the trees it is moistened by steam,

or when it falls on the ground it becomes mixed with rain water

and forms a sort of clay. ; ;

The eruption of the fifth of August was not accompanied by

any noticeable earthquake in Martinique, even in the Prêcheur

district. Since that time no shocks have been felt. In this

respect the opening of these vents seems to be for our island a

happy event, a kind of safety valve giving vent to subterranean

gases and vapors and so protecting us from those earth commo-

tions which formerly produced such desolation here. “It hap-

pens," says Buffon, “that in the lands subject to earthquake,

79 THE AMERICAN NATURALIST. [Vor. XXXVII.

when a new volcano breaks out earthquakes cease and are only

felt during the violent eruptions of the volcano; this has been

observed on the island of St. Christopher; and the great

encyclopædia begins its article on “volcanoes” as follows :

« Volcanoes are a beneficent device of nature, etc. etc."

The buzzing sounds of detonations are not produced by sub-

terranean ebullition, but they take place simultaneously with

the ejection of vapor and are produced at the orifice of the vent.

The cinders or muds are the only materials thrown out by

the volcano. We have found neither lavas, nor even stones of

the smallest possible dimensions which could be identified as

eruption products. - |

The geological structure of Mt. Pelée, as far as our incomplete

observations go, shows no lava flows: “for we must not include

under the name of lavas,” says a geologist, “all the materials

ejected from the throat of a volcano such as cinders, pumice

stones, gravels, sand; but only those which, reduced by the

action of heat to a liquid condition, form on cooling solid masses

the hardness of which is greater than marble." These lavas

exist principally in the vicinity of volcanoes which eject fire.

Now we find about our volcano only pumice, generally fragmen-

tary, and some deposits of cindery substance, in the middle of

which appear diorite fragments, torn out of the interior of the

earth in preceding eruptions.

This geological structure of Mt. Pelée leads to the belief that

the earlier eruptions (which show at least two craters, the dry

pond and the lake above) have been of the same nature as that

of the fifth of August. Everything goes to show that this

volcano should be ranked with the cinder, or mud volcanoes,

and not with fire volcanoes (volcans de feu).

Compared with the common notion of the Soufrière of Guad-

eloupe, what we have learned concerning the new vents of Mt.

Pelée is closely similar. It is probable that the conditions are

about the same in the case of the Soufriéres of St. Lucia,

Dominica and Montserrat.

As to Guadeloupe, there have been eruptions several times,

notably in February 1837, and December 1846.

_ These eruptions have always opened new fumaroles and

ejected cinders and thick mud.

No. 445.] ERUPTION OF MOUNT PELEE. 71

Mt. Pelée when closely examined shows no fissures, landslips,

nor displacement of waters, and hence the action of the eruption

of the fifth of August was very local.

The city of St. Pierre situated more than ten kilometers away,

and the town of Précheur distant seven kilometers, appear to be

out of danger from eruptions even considerably greater than the

one which has taken place. Nothing in the land where they

are situated indicates great catastrophes. Even to reach the

Ruffin House or the Eynard House, which are nearest to the

vents, a disturbance would be necessary very different from the

present one. The matter thrown out even in the immediate

vicinity of the vents did not reach a depth of more than a meter.

This material finds a natural path of flow in consequence of the

steep slope and the gorge of the Riviére Blanche whose waters

naturally carry it off to the sea. Further the planters who fled

at first have since returned to their dwellings, and their work,

and have no further fear of the noises heard from time to time

nor of the odor of sulphuretted hydrogen that is continually

perceptible.

This odor impregnates the atmosphere all about Mt. Pelée

and extends even beyond. It augments or diminishes in certain

places according to the direction of the wind; it is sometimes

comparable to the odor of burnt gunpowder, at other times to

that of stirred up swamp mud ; up to this time this odor appears

to be merely disagreeable and has had no unsanitary effect either

upon men or animals. However, for some time I have been

Struck with the large number of persons of the Précheur district,

especially on the habitation Beligny and in my own plantation,

who have consulted me about skin troubles and insomnia which

they attribute to the sulphurous emanations. I have seen cows

drinking the muddy water of the Riviére Blanche and the pro-

prietor assures me that they suffered no ill effects. I have not

observed any flight of the birds away from the district though

one would suppose their respiratory systems must be very sen-

sitive. Silver pieces in all the estates of the Précheur quarter

turn brown, and so do all paintings which contain compounds of

copper or lead. It is worthy of note that at Guadeloupe the

exhalations of sulphuretted hydrogen are not perceived in the

72 THE AMERICAN NATURALIST. (VoL. XXXVIII.

vicinity of the Soufriére — hence we may hope that here also

they may cease in due time. No great deposits of pure sulphur

have been found either here or in Guadeloupe. Everywhere the

sulphur appears to be in a state of combination.

Moreover it should be observed that the presence of sulphu-

retted hydrogen in the atmosphere in all those localities where

there are thermal sulphur waters is not regarded as unhealthy ;

the waters of Vernet and of Cauterets give out these exhala-

tions and they are believed salutary for people with pulmonary

trouble, therefore we may hope that the Précheur district, already

renowned for its therapeutic qualities in the treatment of these

diseases, may acquire a new title to fame in the eyes of such

invalids.

But the most remarkable result of our excursions and the

most interesting is the information about the place formerly

called La Soufriére. There in a very small area four springs of

different temperature occur close together, of which three are

sulphurous at temperatures of 9o", 70°, and 46°. The other is

cold with a temperature of 22.5? and while not being very good

to drink is not the less potable. The hot springs contain free

sulphuric acid (acide sulphydrique) which volatilizes in conse-

quence of the heat and leaves a residue on evaporation weighing

two drachms for each meter of water; this residue of soluble

salts contains sulphuric acid and hydochloric acid combined with

soda, potash, magnesia, lime and iron as well as a small quantity

of silica.

It is to be hoped that at some time when calm shall have been

re-established at the seat of eruption, and in the minds of the

inhabitants, that some bold speculator will turn to our advantage

that which has given us such a fright and will build an establish-

ment of thermal waters in the midst of these places now so des-

olate. These waters could be conducted to a reservoir so dis-

posed, that their high temperature might be reduced without the

loss of their beneficial properties.

Summing up the volcano La Montagne Pelée, it appears to be

merely one more interesting curiosity added to the natural his-

tory of Martinique — a curiosity that foreigners will wish to visit

and which with fitting industry on the part of the natives may

No. 445.] ERUPTION OF MOUNT PELEE. 73

be made a source of health and wealth — in calm weather the

ships coming from France will see from a distance the long bil-

low of white vapor rising straight toward the heavens, and will

find this a picturesque addition to the landscape — the last touch

needed to complete the majesty of our ancient Montagne Pelée.

NOTES AND LITERATURE.

ZOOLOGY.

Birds in their Relation to Man. — Mrs. Florence Merriam Bailey

in her “ Birds of Village and Field " included much matter relative to

the food of birds, based on the reports of the Dept. of Agricul-

ture; nothing has since been done to put before the public a system-

atic compilation of the results of the work which this Department

has done. Prof. Weed attempted to give a senior class in the New

Hampshire Agricultural college a course in economic ornithology.

His book * Birds in their Relation to Man”? has developed from the

series of lectures which he wrote for this object. The work should

be an effective agent in informing the general public of the part birds

play in the garden, field and forest. Man in his relations to birds

comes in for treatment, and it is only thanks to a few honored names

such as Wilson, Forbes, Beal and Palmer, that the prevailing record

of short-sighted selfishness and ingratitude is lightened.

The book begins with a chapter on methods of studying the food

of birds, the only really satisfactory ones being that of the Dept. of

Agriculture, the examination of stomachs ; and that of Prof. Herrick,

the study of nestlings from a teut; the development of the study of

economic ornithology is then briefly treated. The next three chap-

ters treat in general the vegetable and animal food of birds ; then

follow chapters on the amount of food of birds ; birds as regulators of

outbreaks, and the relations of birds to predaceous and parasitic

insects. Nearly half the book is devoted to the food of the separate

families of birds as illustrated by typical examples. The bulletins of

the Dept. of Agriculture naturally form the basis of this part of the

work, with the result that while the passerine orders are well-covered,

the food of the water-birds, among which little systematic work has

yet been done, is very inadequately treated. Interesting chapters

follow on the “conservation " of birds including an account of legisla-

tion lately enacted, and helpful suggestions for resisting the attacks

! Weed, C. M. and Dearborn, N. Birds in their Relation to Man. London.

Lippincott. 8vo, pp. 380. With numerous full-page illustrations, cuts, diagrams,

etc.

76 THE AMERICAN NATURALIST. (VoL. XXXVIII.

of injurious birds without indiscriminate onslaughts on the whole

species. There are four valuable appendices, the first three dealing

with protective legislation, the fourth a bibliography of economic

ornithology.

There is naturally considerable unevenness of treatment in

the account of the food of birds due to the gaps in our present

knowledge of the subject, but the matter at Prof. Weed's and Mr.

Dearborn's command, is presented clearly and in a scientific spirit.

The doubtful birds, the crow, the crow blackbird, the bobolink, etc.,

are treated in an unprejudiced spirit : — they receive justice tempered

with mercy. The book is intended not so much as a storehouse of

facts, as a powerful argument, which cannot fail to have a very bene-

ficial influence with the reading public.

The illustrations will give the book a little more favor with the

people, and thus area help. The cuts on pages 137 and on 139,

however, are of doubtful value and the full page illustration on page

59 called American Long-eared Owl looks very much like a Shori-

eared Owl.

RoR.

BOTANY.

The Morphology of Angiosperms.!— For several years past there

has issued from the Botanical Department of the University of

Chicago a series of studies upon the embryo-sac and related topics,

some of which have been important contributions to the subject.

These papers form the basis of the present volume, which has been

prepared by the head of the department, with the assistance of Dr.

C. J. Chamberlain. While the book contains little material that has

not appeared before, nevertheless it can claim to be based, to a con-

siderable extent, upon work done under the supervision of the authors.

Evidently an enormous amount of literature has been gone over, and

on the whole, the summarizing of the results has been well done,

and the book will be very useful to the student who wishes to know

the present status of the subject. One would feel more confidence in

some of the conclusions reached by the authors, if these were based

to a greater degree upon first-hand observations ; but it is quite pos-

! Coulter, J. M. and Chamberlain, C. J. Morphology of Angiosperms. N.Y.

Appleton & Co. 1903. pp. vii4- 348.

No. 445.] NOTES AND LITERATURE. 77

sible that there is less personal bias than would be the case in a work

based mainly upon the personal investigations of the writers.

The title of the book is, perhaps, somewhat misleading, as it deals

only briefly with general morphology; but we think the authors have

done well to restrict it mainly to the sporangium, gametophyte and

embryo, since a general morphology of the vast group of Angiosperms

could hardly be compressed within the limits of a single volume.

The book comprises seventeen chapters, of which the first nine will

be found of the greatest value to the student for reference. In these

the general morphology of the flower, the microsporangium, macrospo-

rangium, male and female gametophyte and embryo are treated in

detail, and on the whole extremely well.

The chapter on the flower is in our opinion one of the very best in

the book. The author (we assume the senior author) shows here a

sureness of treatment which comes only from an intimate first-hand

knowledge of his subject, this being by no means so evident in some

of the succeeding chapters, especially the one on the microsporangium,

which immediately follows this chapter on the flower.

The chapter on the microsporangium opens with the remarkable

statement that the zerosporangium is derived from the periblem. To

the reviewer is credited the statement that in Naias the sporangium

arises from the plerome — a statement which it may be remarked, he

did not make. There seems to be a curious confusion in the authors’

minds between the terms “sporangium " and “sporogenous tissue."

We cannot accept the view here set forth, that the microsporangium

in the Angiosperms is an endogenous structure. The author seems

to have in mind the obsolete theory of the imbedding of an originally

superficial structure, a view which is directly contrary to the conclu-

sions of the most recent studies on the development of the sporan-

gium. It is now pretty generally admitted that the eusporangiate

type, such as that of the angiosperms is the more primitive form of

sporangium, and the authors themselves assume the origin of the

angiosperms from some form of eusporangiate pteridophyte. The

close resemblances in the development of the sporangia between the

latter and the angiosperms are familiar to every one who has made a

diréct study of the subject. We do not believe that the assumed

difference in the origin of the archesporium is so fundamental as the

authors claim. :

It is strange that the most important work of recent years, bearing

on the comparative development of the sporangium should be quite

ignored. It seems hardly possible that the authors are not

78 THE AMERICAN NATURALIST. (VoL: XXXVIII.

acquainted with Bower's magnificent series of monographs on this

subject, but we can find no reference to them in the book.

The idea of an imbedded sporangium seems to have been taken

from the older German texts; but a careful study of the context, in

either Goebel or Strasburger, will show that both of these authors

consider the whole of the superficial tissue of the loculus, as forming

the wall of the sporangium, and the whole pollen-sac as the direct

homologue of the microsporangium of the pteridophytes.

It seems to be also assumed, although we can see no warrant for

this, that the nucellus represents something more than a macrospo-

rangium.

As might be expected, the development of the embryo-sac is given

very complete treatment, the chapter dealing with this important topic

comprising fifty pages, of which four are devoted to the bibliography.

Much of the matter in this chapter is taken from the numerous papers

which have been issued from the botanical laboratory of the Univer-

sity of Chicago, and many of the copious illustrations are drawn from

the same sources. This chapter will probably be found the most

useful in the book: The extensive literature of the subject has been

carefully reviewed, and on the whole, little exception can be taken to

the references selected to form the bibliography appended to the

chapter. A great many facts are presented, and although the very

number may be rather confusing to one unfamiliar with the subject,

the chapter will nevertheless, give the student an excellent idea of the

present status of our knowledge of the development of the embryo-

sac.

The male gametophyte, naturally, has less space devoted to it, but

is sufficiently complete. We should like to call the authors’ attention

to a mis-statement. The male prothallium of Sparganium is not

referred to at all in the preliminary paper quoted, but was first

described in the more complete monograph published subsequently,

and which seems to be unknown to the authors.

Chapters seven and nine are concerned respectively with Fertiliza-

tion, The Endosperm, and The Embryo.

The chapter on Fertilization is well up to date, and gives a clear

account of the latest studies upon this important topic. The chapter

on the Endosperm is not so satisfactory, a number of more or less

Important omissions being noted. Thus no mention is made of the

peculiar behavior of the lower endosperm nucleus in Naias.

‘Campbell, D. H. Studies on the Flower and Embryo of Sparganium. roc.

Cal. Acad. Sci. Botany, Vol. 1, No. 9, 1899.

No. 445] NOTES AND LITERATURE. 79

The treatment of the embryo is very satisfactory and leaves little

to be desired. The discussion of parthenogenesis and polyembryony

is especially good.

Except for the chapter on the phylogeny of the angiosperms, the lat-

ter chapters might have been entirely omitted without the value of the

book being seriously impaired. The chapters on classification are

entirely too brief to be of much value to the beginner, and the special-

ist will prefer to consult Engler & Prantl's Watiirliche Phlanzenfamil-

ie, from which the substance of these chapters is borrowed.

The chapter on geographical distribution is very fragmentary, and

leaves something to be desired, also, in the matter of accuracy. For

instance, we doubt whether the statements as to the relative numbers

of Archichlamydez Sympetale and Monocotyledons will bear close

examination. Thus the statement that the Archichlamydez and

monocotyledons are relatively more numerous in the tropics than in

temperate regions may be questioned. In round numbers the species

of monocotyledons. Archichlamydez and Sympetale are 20,000,

60,000 and 40,000. A tabulation of the number of species in the

Northeastern states is given in Britton & Brown — the numbers are

respectively 1058, 1601 and 1361. It is thus seen that while the

Sympetale are relatively slightly in excess, this is very much more

marked in the monocotyledons, which our authors assert are rela-

tively more numerous in the tropics. Two tropical floras were

examined, Hawaii and the West Indies. In the former the figures

are taken from Wallace's /s/and Life. The numbers are mono-

cotyledons, 137; Archichlamydez, 271; Sympetale, 318. There is

thus a marked predominance of Sympetale, and a deficiency of

Monocotyledons and Archichlamydez, directly the reverse of the

Statement given by the authors. In the West Indies (Griesebach,

Flora of the British West Indies) the numbers are approximately,

Monocotyledons, 713; Archichlamydex, 1456; Sympetalae, 91:3.

The monocotyledons in both cases are relatively less abundant than

in the strictly temperate flora of the Northeast United States.

The statement that the Archichlamydex have developed no charac-

teristically boreal group, while the Ericales are essentially boreal, is

not in accordance with the facts. The authors themselves have

called attention to the peculiarly austral family of Ericales, the Epa-

cridez, and scattered ericaceous genera occur in the tropics, both of

the old and new worlds. We should certainly consider the Salicales

as quite as distinctively a boreal group as the Ericales.

The chapter on the phylogeny of the angiosperms contains much

8o THE AMERICAN NATURALIST. [Vor. XXXVIII.

of interest, and is clearly written. The recent speculations upon the

relation of angiosperms and gymnosperms; the connection between

monocotyledons and dicotyledons, are given due attention. In the

discussion of the question of the possible monocotyledonous affinities

of Podophyllum, it may be said that the suggestion that the two appar-

ent cotyledons are possibly one, morphologically, was not suggested

by Holm. Much stress is laid upon the somewhat dubious “ Pro-

angiosperms” of the lower Cretaceous. These are supposed to have

arisen from some eusporangiate filicineous stock and to have given

rise, independently, to the monocotyledons and dicotyledons. If we

are to assume that the angiosperms are monophyletic, it is considered

that the monocotyledons are probably derivations of the dicotyledons.

The recent mutation theory of De Vries is also given due attention.

Prof. Jeffrey's contribution of two final chapters on the vascular

system has some value in itself, but comparatively little bearing upon

the morphology of angiosperms. Of twenty pages, less than six deal

with angiosperms, and of thirty-three figures only five represent this

group. ‘These chapters seem to us unnecessarily loaded with tech-

nical terms, and are by no means easy reading.

D.C.

Notes. — No. 25 of the new series of “Contributions from the

‘Grey Herbarium of Harvard University,” issued as No. 5 of the

current volume of Proceedings of the American Academy of Arts and

Sciences, on Sept. 25, is an important paper by Greenman on Mexi-

can and Central American Angiosperms, mostly of recent collection.

An account of an ecological study of Big Spring Prairie, Wyandot

County, Ohio, by Bonser, is published as no. 7 of the Special Papers

of the Ohio State Academy of Science.

Vol. 3, fascicle 1, of Urban’s Symbole Antillane, dated September

16, contains the first part of a “Flora Portoricensis,” by Urban.

Fascicle 126 of the Fra Brasiliensis, issued in December, 1902,

concludes Vol. 5, part s, of the work, dealing with orchids.

The long-delayed number needed to complete the 1901 volume of

the Bulletin de la Société Botanique de France, dealing with the I9OI

session in Corsica, contains important data on the flora of that island,

including an especially full account of the fungi.

An account of the vegetation of Corsica, with photograms, by

Rikli, is published in the ViertelJahrsschrift der Naturforschenden

Gesellschaft in Zürich, of Apr. 11, 1903.

No. 445.] NOTES AND LITERATURE. 81

Vol. 2, fascicle 5, of Coste’s Flore descriptive et illustrée de la France,

de la Corse et des Contrées limitrophes, carries the work into Solanacez.

With volume 8, issued in April, 1903, the Rouy, Foucaud and

Camus Z7ore de France is brought to a conclusion, and correction

sheets for the earlier parts are added to this volume.

An account of the flora of the Sundribuns, by Crain, is published

as Vol. 2, no. 4, of the Records of the Botanical Survey of India; and

from the Government Printing Office at Calcutta is being issued a

Flora of the Upper Gangetic Plain, and of the adjacent Siwalik and

Sub-Himalayan Tracts, by Duthie, —the first part covering Ranun-

culacez to Cornacee.

The first fascicle of an illustrated quarto treatise on the botany of

the middle and lower Congo, by de Wildeman, has been issued as a

part of the Annales du Musée du Congo, of Brussels.

Volume 35 of the Zransactions and Proceedings of the New Zealand

Jnstitute, as is usual with that publication, contains a number of

important botanical articles.

Recently issued parts of Engler's Das Pflanzeureich are the follow-

ing : — 12, Pfitzer, Orchidacez-Pleonandrze ; 13, Ruhland, Eriocaula-

cez ; r4, Grosser, Cistacez ; 15, Mez, Theophrastacez.

A paper on root, stem and leaf structure of Æschscholtzia californica,

by Denniston and Werner, is contained in Vol. 6, no. 8, of Phar-

maceutical archives.

Under the name Wittia Amazonica, Schumann describes and fig-

ures in the Monatsschrift fiir Kakteenkunde, of August, a new generic

type of cactus from Peru, of the aspect of Phyllocactus but with

small red flowers.

The Revista do Centro de Sciencias, Letras e Artes de Campinas, of

July 31, contains a description and figure of AAipsalts pilocarpa, by

Lófgren, and diagnoses and illustrations of a number of other new

Brazilian plants by Edwall.

The Monatsschrift für Kakteenkunde, of September 15, contains

descriptions of several new cacti.

Cereus gummosus, as it. grows in Lower California, is figured by

Schumann in the Monatsschrift für Kakteenkunde, for July 15.

The Gardeners’ Chronicle of August 8 may be called an Opuntia

number, with numerous illustrations of this genus as cultivated at

La Mortola.

82 THE AMERICAN NATURALIST. (Vor. XXXVIII.

A cristate tree of Cereus giganfeus, in situ, is figured by Mrs.

Drennan, in oral Life for September.

A good figure of Yucca glauca, as it grows in Colorado, is printed

in Floral Life for August.

Excellent practical instructions for the collection of herbarium

material of palms are given by Dammer in the WVofizblatt des K.

botanischen Gartens und Museums of Berlin, of July ro.

A paper on the phylogeny of Angiosperms, resuming publications

in the Botanical Gazette, is reprinted by Coulter from Vol. 10 of the

Decennial Publications of the University of Chicago, under date of

April 1.

The endogenous adventive buds of several genera of phanerogams

are considered by De Candolle in a separate from the Archives des

Sciences Physiques et Naturelles of Geneva, for July.

The structure of a number of the woods of Borneo is described by

Bargagli-Petrucci, and illustrated with photograms, in fascicle 6-8

of Malpighia for 1903.

The first part of a study of the comparative anatomy of the ine

of the Salicacez, by Perrédés, is published as No. 39 of the papers

from the Wellcome Chemical Research Laboratories, of London.

The root anatomy of Angiosperms is the subject of a paper by

Kroemer, forming Heft 59 of Bibliotheca Botanica.

Variation in the androecium of .Sze//aria media is discussed, with

curves, by Reinóhl, in the Botanische Zeitung, 1 Abteilung, of

September 16.

A paper on the development of the fruits in heterocarpic Com-

posites is published by Patané in Malpighia, Vol. 17, fascicle 9.

Dr. Chamberlain’s paper on Mitosis in Pellia, contained in the

Botanical Gazette of July, is concurrently issued from Vol. ro of the -

Decennial Publications of the University of Chicago.

The application of the kinematograph to the class presentation of

plant movements is continued by Miss Scott in a paper on the move-

ments of the flowers of Sparmannia africana in the Annals of Botany

of September,

The spurting of water from the leaf-tips of Colocasia nymphafelia

is considered by Molisch in the Berichte der deutschen Botanischen

Gesellschaft of September 21.

No. 445] NOTES AND LITERATURE. 83

The phenological relations of sun-spots form the subject of a short

note, with curves, by MacDowall, in JVa/ure of August 27.

An illustrated account of the lumber industry of the Northwest

coast, bv Lamb, is contained in Out West for October.

A nicely illustrated popular article on Elm and Tulip trees, by

McFarland, is contained in Zhe Outlook of October 3.

An economic account of Zizania, by Brown and Scofield, is pub-

lished as Bulletin No. 50 of the Bureau of Plant Industry of the

United States Department of Agriculture.

The differentials of varieties of Avena, and those of Triticum, are

respectively discussed by Dufour and Dassonville in the Revue

générale de Botanique of July 12, and Scofield in Bulletin No. 47 of

the Bureau of Plant Industry of the U. S. Department of Agriculture.

An account of rattans and the rattan industry is contained in the

Agricultural Bulletin of the Straits and Federated Malay States, for

April and May last.

An illustrated paper on origin and distribution of camphor in the

camphor tree, by Homi Shirasawa, is. published in Vol. 5, no. 3,

of the Bulletin of the College of Agriculture of the Tokyo Imperial

University.

| In the Pharmaceutical Review, of September, True shows that

under the name of Pink-root Ruellia ciliosa occurs in the trade, as

well as Spigelia marilandica.

A well illustrated account of Castilla elastica and the Central Amer-

ican production of rubber from it, by Cook, forms Bulletin no. 49 of

the Bureau of Plant Industry of the U. S. Department of Agriculture.

The first of a series of notes on latex-yielding Apocynacez of the

Congo, by De Wildeman, has been issued at Brussels asa publication

of the Independent State of the Congo.

An account of the root-parasitism of the sandal tree, by Rama Rao,

is contained in Zhe Zndian Forester of September.

An account of the Isoetes of southern California is compiled for

the West American Scientist of September.

Helpful notes on the collection and cultivation of cryptogams for

use in the biological laboratory are contained in current numbers of

the Journal of Applied Microscopy and Laboratory Methods.

34 THE AMERICAN NATURALIST. [Vor. XXXVIII.

An annotated edition of the Codex of Clusius, the original of which

appeared three centuries ago, has recently been issued by Istvanfh,

of Budapest, and is illustrated by colored reproductions of the origi-

nal water color sketches of fungi, by Clusius.

A key to the species of Rhizopus, with description of a new

parhogenic species, Æ. eguinus, is published by Costantin and Lucet

in the July number of the Bulletin trimestriel de la Société myco-

logique de France.

A note on Costa Rican edible fungi is contained in No. 24 of the

Boletin el Instituto Fisico geografico de Costa Rica.

Several edible fungi are illustrated in the Gardener’s Chronicle of

September 19. |

Inoculation experiments with graminicolous species of Claviceps

are described by Stáger in the Botanische Zeitung, Abteilung 1, of

July 30.

Some decay of stored apples is shown by Eustace, in Bulletin No.

235, and the popular edition of the same, of the New York Agricul-

tural Experimental Station, to be caused in part by a species of

Hypochnus following Fusicladium, and in part to be of an obscure

origin, without any parasitic cause so far as determined.

Oudemans and Koning have distributed two recent papers on a

new Sclerotinia of tobacco, from the Amsterdam Academy.

An economic account of the wilt disease of tobacco, caused by a

species of Fusarium, forms the subject of part 1 of Bulletin No. 51, of

the Bureau of Plant Industry of the United States Department of

Agriculture, by McKenney.

An important paper on the bacteria of tilled soil, by Hiltner and

Stormer, forms Vol. 3, Heft. s, of Arbeiten aus der Biologischen

Abtheilung fiir Land und Forstwirthschaft am K. Gesundheitsamte,

of Berlin :

Bacteria and the Nitrogen Problem is the title of an article by

Moore, separately printed from the 1902 Yearbook of the Depart-

ment of Agriculture.

An extended consideration of the significance of bacteriological

methods in sanitary water analysis, by Winslow and Nibecker, is

contained in the Zechnological Quarterly for September.

No. 445.] NOTES AND LITERATURE. 85

An account of the lichens of the northern boundary of Minnesota,

by Fink, is separately printed from Minnesota Botanical Studies, under

date of July 3.

Vol. 4 of De Toni's SvZoge Algarum, recently issued, continues.the

Floridez.

An account of the algz which contaminate public water supplies,

by Moore, is separately printed from the Yearbook of the Depart-

ment of Agriculture, for 1902.

A most interesting portrait gallery of distinguished botanists is

contained in Professor Wittrock's * Catalogus Illustratus Iconothecæ

Botanice Horti Bergiani Stockholmiensis," issued as Vol. x NU 2.

of the Acta Horti Bergiani.

A portrait of Celakovsky forms the frontispiece to the Sitzung-

sberichte der k. bohm. Gesellschaft der Wissenschaften — Mathematisch-

naturwissenschaftliche Classe, for 1902.

A short account of the San Salvador botanical garden is contained

in the Anales del Museo Nacional of that Republic, of August 1.

A sketch of the botanical gardens at Frankfurt am Main, by

Mobius, is separately printed from the 1903 Bericht der senckenber-

gischen naturforschenden Gesellschaft.

The Journals.— The American Botanist, for July, contains the

following popular articles: — Bradshaw, “The Castor-oil Plant a

Heselbarth, “The Walking Fern”; and Clute, “ Pollination of the

Sunflower.”

The Botanical Gazette, for August, contains the following: — Law-

son, "Studies in Spindle Formation”; Frye, “The Embryo-sac of

Casuarina stricta”; Coker, * The Gametophyte and Embryo of Tax-

odium —concluded”; Bliss, “The Occurrence of two Venters in

the Archegonium of Polytrichum juniperinum” ; and Cook, * Polyem-

bryony in Ginkgo."

The Botanical Gazette, of September, contains the following arti-

cles : — Ganong, “The Vegetation of the Bay of Fundy salt and

diked Marshes ” ; Shull, “Geographic Distribution of /soetes sac-

charata ? ; Parish, * A Sketch of the Flora of Southern California " ;

Thom, * A Gall upon a Mushroom”; and Coker, “Selected Notes,

IL — Liverworts.”

86 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The Bryologist, for September, contains the following articles : —

Gozzaldi, “ Thomas Potts James,” with portrait; Holzinger, “ Fabro-

Jeskea austini in Europe"; Chamberlain, “ Mounting Moss Spec-

imens,” and “ Buxbaumia aphylla” ; Harris, * Lichens — Nephroma-

Solorina”; Hill, “ Branched Paraphyses of Bryum roseum”; and

Lindberg, “ Stereodon plicatulus.”

The Bulletin of the Torrey Botanical Club, for August, contains

the following : — Murrill, “ The Polyporacez of North America — V,

The Genera Cyrtoporus, Piptoporus, Sentiger and Porodiscus ";

* Mez, Bromeliacez Nicaraguenses Nove”; Berry, “The American

Species referred to Thinnfeldia”; Wiegand, “Some Notes on Jun-

cus” ; and Nash, " Revision of the Family Fouquieriacez.”

The Bulletin of the Torrey Botanical Club, for September, contains

the following : — Best, “ Revision of the North American Species of

Leskea”; Eastwood, “New Species of Western Plants”; Mac-

Dougal, * Some Correlations of Leaves”; and Cushman, * Desmids

from Bronx Park, New York.” :

The Fern Bulletin, for July, contains the following articles : —

Fitzpatrick, “ The Fern Flora of Iowa”; Clute, * Fernwort Notes,

III”; Eaton, “The genus Equisetum in North America, XIV,

E. hiemale” ; Gilbert, “Campbell Easter Waters " (with portrait) ;

Gilbert, “ Asplenium muticum” ; Flett, “The Fern Flora of Washing-

ton”; Christ, “Can Scolopendrium lindeni, Hook. “be separated .

from S. vulgare Sm?" and Gilbert, * Two New Varieties of the

ternate Botrychium.” |

The Plant World, for August, contains the following articles: —

Safford, “ Extracts from the Note-book of a Naturalist on the Island of

Guam, IX”; Straw, “Ferns of Smugglers’ and Nebraska Notches " ;

Spaulding, “The Relations of Insects to Fungi”; Barrett, “The

Birthplace of Agriculture”; Lindahl, “A fasciated Tulip”; and

Waters, “Some Summer Observations.”

The Plant World, of September, contains the following articles : —

Baum, “The Breadfruit ”; Bailey, “Lianes”; Safford, “ Extracts

from the Note-book of a Naturalist in Guam, X"; Goetting, “ The

Leucocrinum " ; and Barrett, “Three Ecological Problems."

Rhodora, for August, contains the following articles : — Fernald,

« : :

American Representatives of Zuzula vernalis ”; Phelps, “An Hour

in a Connecticut Swamp”; Webster, “A Beautiful Pluteolus ”;

No. 445.] NOTES AND LITERATURE. 87

Collins, “Some Notes on Mosses”; Eames, “The Dwarf Mistletoe

in Ct.”; and Collins, * Notes on Algæ, V."

Rhodora, for September, contains the following articles : — Eames,

“The Dentarias of Connecticut”; Collins, ** Isaac Holden"; Cush-

man, “Notes on New England Desmids, I”; Fernald, * Arabis

drummondii and its relatives”; Collins, “Notes on Alge, VI”;

Robinson, * On the Twelfth List of New England Plants"; and a

short notice of the late Charles James Sprague.

Torreya, for September, contains the following articles : — Berry,

« Liriodendron Notes"; Robinson, “The Distribution of Fucus

serratus in America”; Earle, * Key to the North American Species

of Galera"; Richards, * An improvised horizontal Microscope uf

Griggs, “ A remarkable Physalis” ; Cockerell, “ Two Orchids from

New Mexico”; Small, * The Habitats of Polypodium folypodioides "l

and Barnhart, * Duplicate Binomials."

CORRESPONDENCE.

To the Editor of the American Naturalist :

Sir:—In The American Naturalist June, 1903, p. 385, Mr. J. H.

Powers says that the metamorphosis of Amdlystoma tigrinum is due

to nutritive causes. Be pleased to rectify that the same conclusion

was published by me August, 1899, in Za Waturaleza, Ser. ii, F. iii,

p. 369. The title of my notes is as follows:

“El Ajolote sufre la metamorfosis general en la clase de los

batracios, par aumento de nutricion y no por cambia de medio.”

Experiments and observations in the latter where Amblystoma was

first observed, show that metamorphosis is very slow in artificial

or natural conditions, if the animals are feeble, and very rapid in the

best conditions of nutrition as in deep waters of Xochimilco, a lake

that never becomes dry.

The dogmatical views on adaptation of Amblystoma are wrong

and the paper of Mr. Powers has only attested my own published

conclusions,

I am very respectfully yours,

A. L. HERRERA.

Mexico, Oct. 12, 1903.

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Co.,1903. x-- 190 pp., maps and illustrations.— BAKER, F.C. Shells of Land

and Water. A Familiar Introduction to the Study i nA Mollusks. Chicago,

A. W. Mumford, 1903. 8vo, xvii J- 17 opi pls. and many text figs.—

BuRKETT, C. W. STEVENS, F.L 1: dd “Hit: . H. Agriculture for Beginners.

Boston, Ginn & Co., 1903. 8vo, xii + 262 pp., 215 figs. — COMSTOCK, ANNA B.

Ways of the Six-Footed. Boston, Ginn & Co., 1903. Svo, xii4- 152 pp., 47

figs.— DopGE, C. W. General Zoólogy. Practical, Systematic and Compara-

tive. Being a Revision and Rearrangement of Orton's Comparative erie

New York, American Book Co., 1903. 8vo, 512 pp., 379 figs. $1.80.— GAR-

DINER, J. S. The Fauna and Geography of the Maldive and Laccadive ven

Being the Account of the Work Carried on age of the Collections made of an

Expedition during the years 1899 and 1900. Vol. ii, Part 1, pp. 473-588, pis.

26-34.— HiGGINSON, E. Map of the oe af Per London, 1903.—

Hunter, S, W. and VALENTINE, M. C. Laboratory Mid of Biology. New

York, Henry Holt & Co., 1903. 8vo, xii +4 215 pp.— Jones, L. H. The

Jones Readers 1-5. Boston, Ginn & Co., 1903. 8vo, 1, 160 pp.; 2, 208 pp.; 3,

286 pp.; 3, 416 pp. ; 5, 496 pp. Many illustrations, partly abr aa. OGG,

V. L. First Lessons in Zoolo ogy. .New York, Henry Holt & Co., 190 8vo,

x + 363 pp., 257 figs.— MILLER, D. R. The Criminal Classes. Cnoi and

Cures. Dayton, United Brethren Publishing House, 1903. Svo, ix -+ 227 pp.,

portraits. $1.50 — MORELEY, MARGARET W. The Insect Folk. Boston,

Ginn'& Co., 1903. 8vo, vi-204 pp., many figs. Noé, J. Recherches sur la vie

osscillante. Essai de biodynamique. Paris, Alcan, 1903. 8vo, 372 pp., 40 figs.

7 francs.— WATERS, C. E. Fems. A manual for the Northeastern States with

analytical keys based on the Stalks and on the Fructification. New York,

Henry Holt & Co., 1903. 8vo, xi + 362 pp., over 200 figs. $3.00.— WELCKER,

A. A Dream of Sedes Beyond Us. San Francisco, Cubery & Co., 1903.

8vo, 38 pp.— WuNDT, W. Sese und Psychologie. Leipzig,

Engelmann, 1903. 8vo, 126 pp., 3 ma

ALVARD, H. E. and PEARSON, R. A The Milk Supply of Two Hundred

Cities and "ons. U. S. Dept. Agr., Bureau Animal Industry Bull. No. 46.

210 pp. — AMEs, O. A New Species of Habenaria from Cuba. Proc. Biol. Soc.

Wash. Vol. xvi pp. 117-118.— ARNOLD, R. The Paleontology je Strati-

graphy of the Marine Pliocene and Pleistocene of San Pedro, California. Mem.

Cal. Acad. Sci. Vol. iii, 420 pp., 37 pls.— BEAN, B. A. Notice of a Small Col-

lection of Fishes, including a Rare Eel, recently received from H. Maxwell

Lefuoy, Bridgetown, Ret West Indies. Proc. U. .S. Natl. Mus. Vol.

xxvi, pp. 903-964, 1 fig. — BAKER, C. F. On the Gnathodus Species » the

go THE AMERICAN NATURALIST. [Vor. XXXVIII.

Abdominalis Group. /uvertebrata Pacifica. Vol.i, pp. 1-12. — BARROWS, F. W.

Economic Zoólogy. dm Buffalo Soc. Nat. Sci. Vol. viii. pp. 16. — BOHR, C.

and HASSELBALCH, K. Om Fosterets Varmeproduktionog Stofskifte. Over-

sigt. Kgl. Danske ru Forhandl., 1903, No. 3, pp. 314-348, 3 figs.— CLARKE,

S. F. An Alaskan Corymorpha-Like Hydroid. Proc, U. S. Natl. Mus. Vol.

xxvi, pp. 953-958, 5 figs. — Cops, J. N. The Sponge Fishery of Florida in ba

Rept. U. S. Fish Com: for 1902, pp. 161-175, pls. 6-9. — Comstock, F. M.

small Esker in Western New York. Amer. Geol. Vol. xxxii, pp. 12-14, b.

2-3. — DALL, W. H. Synopsis of the Family of Astartidae with a Review of

the American Species. Proc. U. S. Natl. Mus. Vol. xxvi, pp. 933- 256 ps

62-63.— DAVENPORT, C. B. Cold Spring Harbor Monographs. II. Th Col-

lembola of Cold Spring Beach, with Special Keierence to the Movements of ros

Poduride. Bull. Brooklyn Inst. Arts & Sci. 1903. 32 pP- pl.

MANN, C. H. The Fresh-Water Fishes of Western Cuba. Bull. U. S. Fish

Com. for 1902, pp. 211-236, pls. 19-21 and 15 text figs.— ELLIS, Mary. Index to

Publications of the New York State Natural History Survey and New York

State Museum 1837-1902. Bull. N. Y. State Mus. Miscellaneous No. 2

239-653. — EVERMANN, B. W. Statistics of the Fisheries of the Middle Atlantic

States. Rept. U. S. Fish Com. for 1902. pp. 433-540.— FISHER, W. K

of the Laysan and the Leeward Islands, Hawaian Group. Bull. U. S. Fish Com.

for 1902, HE 1-39, pls. 1-10. — GaRciA F.— Shade Trees and other Orna-

mentals. Bull. New Mex. Agr. Exp. Sta. No. 47, 55 pp» 29 figs. — GILL, T

On some neglected Genera of Fishes. Proc. U. S. Natl. Mus, Vol. xxvi, pP-

959, 962.— GILL, T. On some Fish Genera of the First Edition of Cuvier's

Règne Animal and Oken's Names. Proc. U. S. Natl. Mus. Vol. xxvi, pp. 965-

967. — Goss, A. Ash Analyses of some New Mexico Plants. Bull. N. Mex.

T Exp. Sta., No. 44, 14 pp.— GRANT, M. Moose. Seventh Ann. Rept. Forest

: he

omm. State N. Y., pp. 225-238, 6 — GUTHRIE, J. E

Collembola of men ete Geol. Nat. Hist. Surv. Minn., Zool. Ser. iv, 110

pp. 16 pls. — HELLER, E. and Snopcrass, R. E. Papers from the Hopkins

Stanford Galapagos rins. 1898-1899. XV. New Fishes. Proc. Wash.

Acad. Sci. Vol. v, pp. 189-229, pls. 2-20. — HERRICK, C. J. The Degree and

Sense of Taste in Fishes. Ju. U. S. Fish. Com., for 1902, pp. 237-272. 3 figs-—

SCHIMURA, T. On the Formation of Anthocyan in the Petaloid Calyx of the

Red Japanese Hortense. Journ. Coll. Sci. Imp. Univ. Tokyo. Vol. xviii, 18 pp»

1 pl.— JENNINGS, H. S. Rotatoria of the United States. II. A Monograph of

the Rattulidae. Bull. U. S. Fish Com. for 1902. pp. 273-352, pls. 1-15. —

Saati. O. A. Aquatic Nematocerous Diptera. V. Y. State Mus. Bull.

8. pp. 327-448, pls. 32, 5o. — JoxEs, J. W. L. Sociality ues Sympathy.

Puli Contributions to Psychology. Val iii, Nos. 3-4. — JORD ^

Review of the Fishes of Japan belonging to the Family of UR Nese te Proc.

U.S. Natl. Mus. Vol. xxvi, pp. 1003-1013, 3 figs. — JorDAN, D. S. and EVER-

MANN, B. W. Descriptions of a New Genus and two New Species of Fishes from

the Hawaian Islands. Bull. U. S. Fish Com. for 1902. pp. 209-210.— KELLOGG,

.Ll. Feeding Habits and Growth of Venus Mercernaria. Bull. N. V. State

Mus., Zool. No. 10, 28 pp., 8 figs. — KENDALL, W. C. Notes on Some Fresh-

Water Fishes from Maine. Zu. U. S. Fish Com. for 1902, pp. 353-368, 5 figs—

Prien to C. Habits of some of the Commercial Catüihes, Bull. U. S.

or 1902. pp. 399-400. — LAMBERT, J. Description de Échinides

No. 445] PUBLICATIONS RECEIVED. 91

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Bruxelles. Mem. Mus. Roy. Hist. ks fu Tom. II, 150 pp., 6 pls. — Macoun,

J. Catalogue of Canadian Birds. Birds of Prey, Woodpeckers, i

catchers, Crows, Jays and ee URS the following Orders: Raptore

Coccyges, Pici, Macrochires, and part of the Passeres. Geol. Surv., pp. MIT

MansH, M. C. A More Complete Description of PA trutte. Bull. U. S.

Fish Com. for 1902, pp. 411-415, pls. 1-2. — MILLER, G. S., Jr. A Nataline Bat

from the Bahamas. Proc. Biol. Soc. Wash. Vol. xvi, pp. 119-120. — MOENK-

HAUS, W. J. Description of a New Species js Darter from Tippecanoe Lake.

Bull. U. S. Fish Com. for 1902. pp. 397-398, 1 — Morse, C. M. South Buf-

falo Floods and d ie Api Bull. bee Soc, Nat. Sci. Vol. viii, pp.

16, maps. — NEEDHAM, J. G., MacGiLLivRAY, A. D., JOHANNSEN, O. A. and

DAVIS, KR. C, gom Insects in New York State. Bull. N. V. State Mus.,

Entomology, No. 18. pp. 200-517, pls. 1-52.— NELsoN, E. W. New PUMA

from Central America. Proc. Biol. Soc. Wash. Vol. xvi, pp. 121-122. — NORD

QUIST, O. Some Biological Reasons for the Present Distribution of Erea a

Fish in Finland. Fennia, Vol. xx, pp. 29, — OBERHOLSER, The

North American Forms of Aag opu (Say). Proc. Biol. Soc. Wash.

Vol. xvi, pp. 113-116. — PERKINS, O. Report on the Collections of Fishes

made in the Hawaian Islands a Descripi ona of New Species. Bull. U. S. Fish

Com. for 1902. pp. 417-511, pls. 1-4.— PERKINS, R. C. L. The Leaf-Hopper of

the Sugar Cane. Board of Commissioners Agr. & Forestry Terr. Hawaii, Div.

Ent. Bull., No. 1. 33 pp. — RICHARDSON, HARRIET. Isopods collected at the

d d of the United States Fish Commission ag Albatross.

Bull

"sh Com. for 1902. pp 47-54, 8 figs. — SCHALLER, W.T. Spodu-

mene ka ye Diego Co., California. Univ. Cal. use go gov Geol. Vol.

iii, pp. 265-275, pls. 25-27. — SCHWARZ, G. E.— The Diminished Flow of the

Rock River in Wisconsin and Illinois and its Seat to the Babe Forests.

U. S. Dept. Agr. Bureau of Forestry Bull. No. 44, 27 pp. 6 pls. — SHAR

Report on the Fresh-Water Ostracoda of the United States National aderat

including a Revision of the Subfamilies and Genera of the Family TTR

Proc. U. S. Natl. Mus. Vol. xxvi, pp. 969-1000, pls. 64-69. — SHERRA

A Working Plan for Forest Lands in Hampton and Beaufort b de; South

Carolina. U.S. Dept. Agr., Bureau of Forestry Bull. No. 43. 54 pp»; 13 pls. —

SMALLWoop, M. E. Cold Spring Harbor Monographs. I. The Beach Flea;

Talorchestia uc Brooklyn. Bull. Brookl. Inst. Arts & Sci., 1903. 27

pp., 3 pls. — SurrH, H. M. The Common Names of the Basses and Sun Fishes.

Rept. U. S. Fish pi for 1902, pp. 353-366. — SNow, JULIA. The Plankton

Algz of Lake Erie, with Special Reference to the Chlorophyceze. Bul. U. S.

Fish Com. for 1902, pp. 369-394, pls. 1—4.— STEJNEGER, L. A New Hognose

Snake from Florida. Proc. Biol. Soc. Wash. Vol. xvi, pp. 123-124. — STEVEN-

SON, C. H. Aquatic Products in Arts and Industries. Fish Oils, Fats, and

Waxes. Pudim from Aquatic Products. Rept. U. S. Fish Com. for 1902, pp.

177-279, pls. 10-25. — STEVENSON, C. H. Utilization of the Skins of Aquatic

Animals. Rept. U. S. Fish Com. for 1902, pp. 281-352, pls. 2 26-38. — STEWART,

J. H. and Arwoop, H. Experiments with Buckwheat and Oats. Bwll. W. Va.

Agr. Exp. Sta. No. 84. pp. 467-480, lem — Stewart, J. H. and Hire, B. H.

Commercial Fertilizers. Bull. W. Va. Agr. Exp. Sta. No. 85, pp. 110.— STONE,

Injuries to Shade Trees from Electricity. Bull. Mass. Agr. Exp. Sta. No.

92 THE AMERICAN NATURALIST. (VoL. XXXVHI.

91. 21 pp. 12 figs.— TRuE, F.W. Notes on a Porpoise of the Genus Prodel-

phinus from the Hawaian Islands. Bul. U. S. Fish Com. for 1902, pp. 41-45,

pls. 1-2. — VERNON, J. J. and LEsrkER, F. E. Pumping for Irrigation from

Wells. Bull. New Mex. Agr. Exp. Sta. No. 45, 67 pp., 35 figs. — VERNON, J.

J. and TiNsLEY, J. D. Soi Moisture Investigations for the Seasons of 1901 and

1902. Bull. New. Mex. Agr. Exp. Sta., No 46. 46 pp. — WHITEEAVES, J. F.

Notes on Some Canadian EH IR of *Lituites Undatus." Ottawa Nat. Vol.

xvii, pp. 117-122. — WiLCOX, W. A. The Fisheries and Fish Trade of Porto

Rico in 1902. Rept. U. S. Fish Com. for 1902. pp. 367-395. — Y ^sUDA, A. On

the Comparative Werne of the Cucurbitacee, Wild and Cultivated in Japan.

Journ. Coll. Sci. Imp. Univ. Tokyo. Vol. xviii, No. 4, pp. 56, 5 pls.

Aéronautical World, s Vol. I, No. 11. — American Inventor, The. Vol.

xi, No. 2. — Annales de la Société Royale Malacologique de Belge. Tom. xxxvi. —

Boletin de la Academia Nacional de Ciencias en Cordoba. Tom. xvii, No. 3. —

Boletin de la Comision de Parasitologia Agricola. Yom. ii, No. 2. — Bulletin of

the Children’s Museum. rooklyn Institute of Arts and Sciences, N a. 12 pt.

Bulletin Johns Hopkins Hospital. Vol. xiv, Nos. 149-151. — De: sicher Arbeit.

Jahrg. i, Heft 12, Jahrg. ii, Heft 10.— Journal of Applied Microscopy and Labora-

tory Methods. Vol. vi, No. 7, July.— Journal of Geography, The. Vol. ii, No. 6,

June.— Missouri Botanical Garden. Fourteenth Annual Report.— Naturaleza,

La. ‘Yom. iii, Nos. 5-10.— Nuova Notarisia, La. Ser. xiv, July.— Papoose,

The, Aug.— Plagas de la Agricultura Las. Nos. 9-10, 2 pls., 3 text figs.— Pro-

ceedings of the Rhodesia Scientific Association. Vol. iv, Pt. 1, July.— Revista

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Oct.

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THE

AMERICAN NATURALIST.

Vor. XXXVIII. February, 1904. No. 446.

RECLASSIFICATION OF THE REPTILIA.

HENRY FAIRFIELD OSBORN.

History OF CLASSIFICATION.!

Philosophy of Classification.— The history of the classification

of the Reptilia resembles that of the classification of other forms

of vertebrates in its gradual approximation to the truth.

The general progress has been from superficial to profound

characters, from purely adaptive characters to those which are

phylogenetic and indicate real affinity. For a century and a

half superficial resemblances and analogous adaptations have

been the pitfalls out of which the final classification is slowly

emerging.

Every classification, moreover, has had its underlying phi-

losophy. The “special creation” philosophy underlays the

Linnzan system, but in so far as Linnzus, Cuvier, De Blain-

ville, Owen perceived a real order in certain profound charac-

ters, their systems will stand.

The philosophy underlying modern classification is the

! Presented at the first meeting of the Society of Vertebrate Palzontologists,

Philadelphia, December 29th, 1903. Abstract, in part, of a memoir entitled The

Reptilian Subclasses Synapsida and Diapsida and the Early History of the

Diaptosauria. Mem. Amer. Mus. Nat. Hist. Vol. I, Pt. VITI, Nov. 1903.

94 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Lamarck-Darwin theory of the law of descent which involves

a branching or phylogenetic scheme of relationships. Toward

this we are slowly progressing. The final classification will be

a formal or tabular statement of the tree of descent, in which

only so much pre-Darwinian classification will survive as was

based upon the perception of real phylogenetic characters.

The evolution philosophy held out a strong temptation to

rapid generalization in phylogeny. It is a striking fact that the

evolutionists, Huxley, Cope, Heckel, perhaps because they

attempted to generalize too rapidly, have proved less fortunate

in their arrangement of the Reptilia than Owen, whose pre-

Darwinian systems of 1839 and 1859 have best stood the

test of time and of discovery.

Both Cope's and Huxley's systems are largely wrecks today ;

Huxley's because while entirely logical in method it outran the

state of knowledge and discovery. Cope was less logical; his

fatal error was over reliance on single characters without dis-

criminating whether they were primitive or adaptive. Marsh

was gifted with unerring taxonomic judgment as to real phylo-

genetic relationships; the chief defect of his system was that he

partly or wholly ignored the rules of priority, renaming and

redefining groups which had previously been defined with suf-

ficient clearness to be recognized. It is with real regret that

I feel compelled, as a matter of historic justice, to revive some

of the older names for certain groups of which our knowledge is

almost entirely due to the fundamental contributions of Marsh.

Priority — Palzxontological discovery is constantly sweiling and

expanding the groups of fossils discovered long ago; it is con-

sequently necessary either to abandon these groups or to raise

or lower their grade. For example. Owen’s “family” Cyno-

of clearness or as a matter of historic justice.

No.446] RECLASSIFICATION OF THE REPTILIA. 95

The history of classification is usually presented by giving

the complete schemes published successively by various anato-

BOK eet eae ae BN E an

G 1.— A primitive Plesiosauroid Synapsidan, Lariosaurus balsami. After

Boulenger. x à

mists. The student will perhaps gain as valuable a lesson by

considering the anatomical philosophy, true or false, which has

prompted different systems of classification.

96 THE AMERICAN NATURALIST. [Vor. XXXVIII.

FALSE PRINCIPLES OF CLASSIFICATION.

Superficial Resemblances.— Yn Brogniart's system of 1799 the

Lacertilia and Crocodilia are wrongly united on limb structure

as (II) Saurii, as distinguished from (III) Ophidii, and (1)

Chelonii.

The similar scaly covering led Latreille (1820) to rightly

unite the Ophidia and Lacertilia as Squamosa ; previously Oppel

(1811) grouped the Lacertilia and Ophidia as Squamata ; this,

however, is the single instance in which epidermal resemblance

happens to coincide with underlying fundamental characters.

As instances of errors based upon epidermal characters, we

may cite the union of the Testudinata and Crocodilia by Klein

as Cataphracta ; or the union of the same animals by Merrem

as Loricata.

Resemblances in Single External Organs.— So able a palæ-

ontologist as von Meyer (18 30) attempted to classify the

reptiles by resemblances in foot structure. He thus divided the

Saurii, or limbed reptiles into (1) Dactylepodes, including Lacer-

tilia and Crocodilia ; (2) Nexipodes, including the Ichthyosauria

and Sauropterygia (Plesiosaurus, Nothosaurus) ; (3) Pachypodes,

including the Iguanodontia and Megalosauria; (4) Pterodactylii,

including the Pterosaurs — a false system.

Classification of Analogous Adaptations — De Blainville (1835)

united the Ichthyosauria and Sauropterygia as Enaliosaurii, or

sea lizards. Owen (1839) adopted the order Enaliosauria as

embracing the Ichthyopterygia (1859) and Sauropterygia (1859),

but remarked that these animals « do not form a strictly natural

group."

Classification by single Internal Characters —This method was

especially characteristic of Cope. In 1869 he defined the Arch-

osauria as differing from the Monimostylica of Müller only by

the exclusion of the order Testudinata; he observed that close

sutural attachment of the quadrate bone “

was the important

feature which characterizes the order "

; by this feature he

united the Sauropterygia (Nothosaurus, which was selected as a

type) the Crocodilia, the Thecodontia (suborder of Dinosauria),

the Dinosauria, the Anomodontia, and the Rhynchocephalia ; a

totally unnatural and transitory grouping, because based upon

No.446] RECLA.SSIFICATION OF THE REPTILIA. 97

the possession of a single primitive character, namely, the

fired quadrate.

We find that almost every attempt to classify the reptiles by

superficial characters, by external organs, by general external

adaptations, by single internal organs, has proved unnatural.

TRUE PRINCIPLES OF CLASSIFICATION..

The conclusion is that there are three ruling principles in

classification.

First, as to priority, we owe it to our palzeontological forebears

not to abandon the lower or higher groups they have proposed

except in cases of absolute necessity. In some instances we

must give a group a higher or lower rank than the author

originally assigned to it, or a different position in the system.

Priority has no force where a group is based on a false con-

ception or on a false grouping of types, as in the definitions

of Theromorpha and Archosauria by Cope..

Second, as to phylogeny, Classification is a formal expression

of our knowledge of phylogenetic relationships ; it must, there-

fore, constantly shift and change as new relationships are

discovered. The final classification will be the phylogenetic

tree. This being the case, it is desired to include within a

group its ancestral forms as soon as they have definitely

branched off toward it. For example, Hyracotherium should

not be placed in the Lophiodontide, as was done by Cope, but

in the Equide. Again, if it should be demonstrated that the

Protorosauria are ancestral to the dinosaurs and to no other

reptiles, they should be placed in the superorder Dinosauria. —

Third, as to definition, classification, like phylogeny, should be

based on a number of characters of different parts of the body

having different functions, in order to diminish the danger of

being misled by analogous evolution, otherwise known as par-

allelism, convergence and homoplasy.

The neglect of one or other, and in some cases of all these

three principles and the loss of the clarifying mind of George

Baur have led to great and rapidly increasing confusion in the

arrangement of the Reptilia in recent years. Smith Woodward,

Broom and von Nopsca, have been working in the right direction.

98 THE AMERICAN NATURALIST. [Vor. XXXVIII,

p-cor. F

Fic. 2—

: Sain ma mn pees of shoulder-girdle. 4-D, after Seeley; E-F, after Howes. 4,

az; ^

views of Ornithorhynchus ^d juae wees ©, P D, Rhopalodon; E, ventral, Æ’, lateral

No. 446.] RECLASSIFICATION OF THE REPTILIA. 99

PROPOSED RECLASSIFICATION.

At the Washington meeting, 1902, of the American Associa-

tion, I presented a joint paper with Dr. J. H. McGregor on the

diphyletic arrangement of the reptiles, based on comparison of a

large number of characters. I have since made a more searching

study of the same problem, designating these two groups as the

subclasses SYNAPSIDA, or primitively single-arched reptiles, and

DiAPsIDA, or primitively two-arched reptiles, and have grouped

all the most primitive forms of Diapsida in the superorder

DIAPTOSAURIA, a group equivalent in taxonomic rank to the

Squamata or the Dinosauria. I now propose to briefly describe

the reptiles which fall within these two groups.

SUBCLASS SYNAPSIDA Osborn.

The chief distinction of the single arched reptiles is that there

is either no opening at all in the temporal region (Cotylosauria),

or a single large supratemporal opening (Anomodontia, Plesio-

sauria, Testudinata) as in the upper view of the skull of a

plesiosaur, a turtle, or a mammal. This supratemporal fossa is

large, because the cranium or brain case is long while the facial

portion of the skull is relatively short, these proportions being

directly reversed in the Diapsida. The temporal arch consists

primitively of two arches combined. The squamosal is always a

large element. The quadrate is correspondingly more or less

reduced; it is never movable, and is functionally supported by

the squamosal. In the shoulder girdle the coracoid and procora-

coid are separate, or united by suture. The phalangeal formula

is primitively 2. 3. 3. 3. 3, like that of mammals.

I. Order CorvLosAvRiA Cope

Pareiasauria Seeley.

These are the most primitive of reptiles, retaining many

Stegocephalian (amphibian) characters, and a solid cranial roof

with temporal openings rudimentary Or not developed at all.

IOO

Subclass SYNAPSIDA Osborn 1903.

Order CoryLosauRIA Cope 1880

(Pareiasauria Seeley 1889)

Superordr ANOMODONTIA Owen

(Theromorpha Cope 1878, in

part.)

Order THERIODON'TIA Owen 1876.

| geni Therocephalia Broom

roni Cynodontia Owen

an DICYNODONTIA Owen 1860.

Order PLACODONTIA auct. ex H.

von Meyer? 1863 Incerte Sedis.

Order SAUROPTERYGIA Owen 1860.

Suborder Simosauria auct. ex

Gervais ! 18

(Nothosauria kid 1882.)

Suborder Plesiosauria auct. ex

Quenstedt? 18<2

Order TEsSTUDINATA auct. ex

Shaw‘ 1802

Suborder Pleurodira auct. ex

Duméri and Bibron € 1835.

Suborder Trionychia? auct. ex

Pictet 1853.

‘ “ Simosauriens."

"TN and authors assigned for

the eurtiest | recognition of the super-

generic rank of several of these

may be altered by future in-

vestigations.

THE AMERICAN NATURALIST. [Vor. XXXVIII.

Subclass DIAPSIDA Osborn 1903.

Superorder DIAPTOSAURIA Osborn

1903.

[Order PRocoLoPHoNiA Seeley

1867.

Order PROTOROSAURIA Seeley

1887

Order PROGANOSAURIA Baur 1887.

4 Order GNATHODONTIA Owen 1680.

(Rhynchosauria Osborn 19053).

Order PELYCOSAURIA Cope 1878.

" — CHORISTODERA Cope 1877.

* RHYNCHOCEPHALIA Gün-

ther 1868.

Order PARASUCHTIA Huxley 1875.

Suborder Aëtosauria Nicholson

and Lydekker 1889.

Suborder Phytosauria Baur 1894

ex Jaeger 182

Order ICHTHYOSAURIA Blainville

1835 ex Jaeger 1824.

(Ichthyopterygia Owen 1860.)

rder CRocopiLiA? Wagler (?)

1830.

peni

Mesosuchia Huxley

75.

Suborder Eusuchia Huxley ies

Thalattosuchia Fra

I9OI.

Superorder DINOSAURIA Owen

1840

( Order THEROPODA Marsh 1881.

Suborder Megalosauria ex Fitz-

inger 1843.

(Thecodontia Owen 1860.)

Suborder dcs sium Cope

< 1867.

(Compsognatha Huxley 1870.)

Order OPISTHOCŒLIA Owen I

(Sauropoda nae: 1881.)

Order ORTHOPODA Cope

(Predentata ec pe ).

Superorder SQUAMATA Oppel 1811.

f Order LACERTILIA? Owen 1839.

| Order MOSASAURIA auct. ex Ger-

3 vais? 1845.

Lone: OPHiDrA? Brogniart "ox:

Order PTEROSAURIA auct. ex Kaup

1834.

No.446] RECLASSIFICA TION OF THE REPTILIA. IOI

The large South African Pareiasauridae are more specialized

than the Texan Pariotichide and Diadectide, the latter being

the type of the order.

Theoretically some unspecialized members of this order gave

rise to all other reptiles including both Synapsida and Diapsida.

II. Superorder ANOMODONTIA Owen.

This was originally defined by Owen (1860) as an ‘order’

embracing the “families” Dicynodontia, Cryptodontia and

Gnathodontia (Rhynchosaurus). The reference of the latter

‘family ’ proves to have been an error. In 1861, in the second

edition of his Paleontology, Owen included in the Anomodontia

the ‘family’ Cynodontia, based on the types Galesaurus and

Cynochampsa ; thus raising the Anomodontia to the rank of a

superorder which is equivalent in part to the superorder Thero-

morpha Cope.

This ‘superorder’ includes two orders, which represent an

adaptive radiation from more primitive truly reptilian types

(Therocephalia Broom) into the more mammal-like Cynodontia,

both with full sets of teeth, and finally into the highly special-

ized Dicynodontia, in which the teeth are greatly reduced. All

these animals retain, however, some primitive or cotylosaurian

and stegocephalian characters.

I. Order THERIODONTIA Owen.

1. Suborder Therocephalia Broom.

Broom has recently published an admirable paper on the

* Classification of the Theriodonts and their Allies," ! in which

the Therocephalia? are defined from the types Scylacosaurus,

JElurosaurus, Ictidosuchus, Deuterosaurus, Titanosuchus, Gor-

gonops, as representatives of six families. These are medium

sized reptiles, and apparently the most primitive of the Anomo-

1 Rep. So. Afr. Assoc. Adv. Sci. 1903; PP- 362-369. es

2 This is defined as an order by Broom and may prove to be of full ordina

rank; it is here provisionally grouped with the Anomodontia as a suborder of

Theriodontia.

102 THE AMERICAN NATURALIST. [Vor. XXXVIII.

dontia. The teeth are differentiated as in mammals into

incisors, canines (sometimes double), and molars; the molars,

however, are simple, and the palate is simple, like that of the

Cotylosauria, that is, there is no secondary palate.

2. Suborder Cynodontia Owen.

This was originally defined as the * family " Cynodontia by

Owen in 1861, and may be embraced within the order * Therio-

gre-frontat.

Fic. $c Synapsid Type. Palatal and superior views of the skull of

Dicynodon, showing the elements as interpreted by Seeley.

Note : especiaily the exposure of the prevomer, the large

extension of the squamosal, the pre- and postfrontals, the single

~— illary After Seeley.

dontia " Owen of 1876, which was based on the same types,

viz.: Galesaurus and Cynochampsa.

These are intermediate anomodonts of medium size. In

contrast with the Therocephalia the squamosal is more expanded

and the quadrate is greatly reduced. Approaching the mammals

also, there is the secondary palate, formed of the maxillaries and

palatines, also the double condition of the occipital condyles

No.446] RECLASSIFICA TION OF THE REPTILIA. 103

which are single in Therocephalia. These cynodonts have lost

several of the other more primitive characters of the theroceph-

alians, such as teeth in the pal-

ate; and their specialization is in

the direction of the mammalia.

The phalangeal formula is 2. 3.

3. They include three

families, typified respectively by

the genera Lycosaurus, Cynog-

nathus, and Gomphognathus.

2. Order DICYNODONTIA Owen.

This term was originally used À 1

as a “family " term, under Ano- j |

modontia, by Owen in 1859; pes

Huxley raised it to the rank of

an order from the types Dicyno-

don, Oudenodon. The latter,

Owen (1859) had placed in the

“family” Cryptodontia, in ref-

erence to the absence of teeth.

Despite the high specialization Frc. 4—Synapsid Type Top view of

of the dentition, these animals w LE ES xb rie

retain the more primitive fea-

tures of the single condyle, of the cleithrum or epiclavicle, of

the large quadrates. On the other hand, like the cynodonts,

they show a rudimentary secondary palate. They approach the

mammals also in the loss of the prevomers and development of

the vomer (parasphenoid).

In the most primitive family of Endothiodontidz one or more

series of molar teeth are present on the maxillaries and dentaries ;

the interclavicle is a rounded plate as in the Stegocephalia. In

the more specialized Dicynodontide, maxillary teeth are absent,

or present as a pair of tusks, and there are no teeth in the lower

jaw; the interclavicle is elongated, and a cleithrum is present.

The third family, Lystrosauridze, exhibits a small interclavicle,

and no cleithrum. A fourth family, Cistecephalidz is doubtfully

ranked here by Broom.

— -———-

104 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Broom has contributed a large number of papers on the

structure and arrangement of the

South African reptiles, which for the

first time throw a perfectly clear

light on their relationships to each

other.

3. Order PLACODONTIA.

(Zucertg sedis.)

The position of this order as typi-

fied by the genus Placodus, is still

very doubtful. It is characterized by

very large teeth in the palate, and by

the absence of teeth on the maxil-

laries and premaxillaries. It includes

littoral, shell-eating forms, which may

be an independent offshoot of the

Anomodontia, or may be more nearly

related to the Sauropterygia.

Fic. 5.—Synapsid Type. Dorsal as-

pect of skull of Mothosaurus

ndriani? A primitive plesio-

sauroid reptile. After Cope.

III. Order SAUROPTERYGIA Owen.

The theory of the relationship of

the plesiosaurs to the Synapsida

and especially to the Anomodontia

and Testudinata still requires con-

firmation.

The skull structure is

typically synapsidan. The shoul-

der girdle structure, so far as

known, in the Triassic plesiosaurs

is certainly more synapsidan

than diapsidan. Numerous resem-

blances to the Testudinata have

been pointed out. On the other

hand, certain of the oldest Triassic

plesiosaurs,

such as Lariosaurus,

Fic. 6.— Synapsid Type Dal de (Fig. 1) closely approach the Diap-

of skull of Triony

sida in the phalangeal formula.

No.446] RECLASSIFICATION OF THE REPTILIA. 105

IV. Order Testupinata (ex Shaw.)

The kinship of the Testudinata to the Synapsida is indicated

both by the skull structure and by the phalangeal formula. The

shoulder girdle structure, like that of the plesiosaurs, is still in

dispute; the main question being as to the homologies of the

cos. st.

Fic. 7.— Diapsidan types of shoulder-girdle. A, Paleohatteria. After Credner. The cartilaginous are

are entirely restored. Restoration by J. H. McGregor. X 4. B, Sphenodon juv. (15 cm.); modified

from Howes. By J. H. McGregor. X Î C Pleurosaurus. Cartilagi

Dames. x 2. D, Mesosaurus tenuidens. Modified from Gervais. X1

Cl, clavicle; z. cZ, interclavicle ; sc, scapula; s. sc, suprascapula ; cor, coracoid; f. cor, procora-

coid; f. cor, coracoid foramen ; 4, humerus ; FE entepicondylar (ulnocondylar) foramen.

(

ginous elements omitted. After

D L4 od ”

anterior ventral processes variously known as the “ procoracoid

or * proscapula."

SUBCLASS DIAPSIDA Osborn.

All these animals are readily distinguished by their general like-

ness to the existing Hatteria. In contrast to the Synapsida the

cranium is short ; the temporal region is primitively fenestrated

by two distinct openings, the supra- and latero-temporal fenestre,

106 THE AMERICAN NATURALIST. (VoL: XXXVIII.

bounded by the supra and latero-temporal arches, one or both of

which may secondarily disappear. In turther contrast with the

Synapsida, the squamosal is a relatively small element, fre-

quently separate from the prosquamosal, and never entering into

articulation with the lower jaw; the quadrate, on the other hand,

is a relatively large element, uncovered, and sometimes second-

arily movable. In the shouider girdle we find a most distinctive

character in the early coalescence of the coracoid and procoracoid

into a single bone, or in the degeneration of the procoracoid.

Another highly distinctive character is the phalangeal formula,

2. 3. 4. 5. 3-4, which is secondarily modified in the aquatic

forms. ;

There appear to have been two great adaptive radiations

among the Diapsida. The first is that which occurred during

the upper Carboniferous and Permian, branches of which are

Fic. 8.—The most primitive known Diapsida

trigoniceps Owen (Order PRocoLopHonta). After Broom

n. A,manus, B, pubis and ischium, and C, pes, of Procolophon

eoe

already well separated in the Permian and have been collectively

grouped in the superorder Diaptosauria by Osborn. The

second, or later radiation of the Diapsida, partly sprung from

terminal branches of the first, is known in the Triassic, and

includes the great orders Parasuchia, Ichthyopterygia, Croco-

dilia, the superorder Dinosauria, the superorder Squamata, and

finally the Pterosauria.

No.446] RECLASSIFICATION OF THE REPTILIA.

MAI A W

MAMM

107

Superorder DiAPTOSAURIA Osborn.

This superorder embraces the Rhyn-

chocephalia of Günther, the Pelycosauria

of Cope, which certainly represent dis-

tinct orders of reptiles, and the more or

: x less distinct orders or suborders Protoro-

as sauria, Procolophonia, Proganosauria,

EN Choristodera, Gnathodontia, Rhynchoce-

oS BR | | phalia. They have appropriately been

Doer

lif

yon

eon

opp

termed ‘rhynchocephaloid’ reptiles by

Broom.

The common characters of these prim-

itive or stem Diapsida, are as follows :

cranium with two complete temporal

arches ; vertebrae typically amphiccelous

and often perforated by a notochordal

canal; hypocentra frequently present

throughout vertebral column ; ribs on

< all vertebrae from first cervical to eighth

at V caudal inclusive, generally single-headed

aN or incipiently two-headed ; large abdomi-

GER nal ribs or plastron always present ;

EX coracoid and procoracoid early uniting

AN into a single bone; pubis and ischium

DN ventrally in continuous contact or Sec-

pis ondarily fenestrated.

The adaptive radiation of these mostly

= >

small sized animals into ambulatory, lit-

toral, amphibious, and fully aquatic types,

together with specializations of the skull `

and dentition for a great variety of feed-

ing habits has resulted in a divergence

sufficiently profound and ancient to

form seven groups which have been

variously assigned the rank of orders

or suborders as follows :

CLL FE SCS S FC EEO 4

ia longicau-

5 g—A primitive diapsidan. P:

chet Aa x b Order

estoration by J. H. McGregor.

PROTOCOSAURIA.

108 THE AMERICAN NATURALIST. (VoL. XXXVIII.

1. Order PROCOLOPHONIA Seeley.

This includes the most primitive of the Diaptosauria ; those

closest to the Cotylosauria, and distinguished by the entire

absence of lateroemporal fen-

estre,' by the persistence of

the epiotics and auditory

notch of the cotylosaurs,

and other very primitive char-

acters. These animals are

thus far recognized in the

‘Permian of South Africa

only.

2. Order PROTOROSAURIA

Seeley.

This land group includes

Protorosaurus, Palzeohatteria,

Kadaliosaurus, distinctively

ambulatory and in part leap-

ing reptiles, certainly carniv-

orous ; distinguished by the

straight limbs, strong develop-

ment of the hind limbs, corre- -

lated with a dorsally expanded

ilium and from two to three

sacral vertebrae.

These animals show all the

characters which we should

expect to find in the ancestors

of carnivorous Dinosauria ;

Fr rrr dd IEEE as

RSS yD

)N el EEE

ps we

AA. ete

ssas

C CE

E t Ca SA

Sr N7 AN A

CN T

IN SE m WA

jt \\ N US H d NA

vu Ñ SM ü A

AR v

R 24

NY» WG

Wh AEF”

WRK?

PROGANO-

the three genera known are Fic. xo. Stereosternum tumidum. Restoration by

: x $. Order

too far specialized in the

direction of ambulatory and

H. McGregor.

SAURIA.

predatory types to have given rise to any of the other known

Diaptosauria.

! Dr. R. Broom has just made this important observation.

No, 446.] RECLA.SSIFICATION OF THE REPTILIA. IO9

3. Order PROGANOSAURIA Baur.

This aquatic or amphibious group, which has been confused

with the Protorosauria, is at present represented only by the

genera Mesosaurus Seeley and Stereosternum Cope, closely

allied forms from the Permian of South Africa and South

Fic. roa.—Mesosaurus tenuidens. After Gervais. X j. Order PROGANOSAURIA-

r amphibious types,

e teeth, very

the

America. These are typical swimming o |

with greatly elongated rostrum, delicate, prehensil

heavy ribs with highly modified capitular attachments ;

vertebrae are also highly distinctive and unique in structure.

IIO THE AMERICAN NATURALIST. [Vor. XXXVIII.

These forms also are too specialized to give rise to any of the

higher Diapsida; they represent an isolated and dying out

group.

4. Order GNATHODONTIA Owen.

Rhynchosauria Osborn.

Owen first proposed the “family” Gnathodontia in 1859,

typified by Rhyncho-

saurus, at the same time

that he proposed the

“ family” . Dicynodontia.

It seems proper that this

term should be given pri-

ority over the order Rhyn-

chosauria proposed by

Osborn in 1903.

Here again we have a

highly specialized division

resembling the Procolo-

phonia in general body

and cranial form, but dif-

fering from these animals

widely in the concentration

of pavement-like teeth on

the pterygopalatines and

the development of a large

edentulous bony beak.

They were probably lit-

ea shell eyeing ae de: at oe Aver Burckhardt

e Aeolus ie the true ne i ceo

ynchocephalia.

5. Order PELycosaurRiA Cope.

This land group, developed in the Permian of Texas and

Bohemia, is distinctively ambulatory and carnivorous. It is

characterized by the abbreviation of the tail, the enormous

development of the spines of the dorsal vertebrae, also by the

No. 446.].. RECLASS/FICATION OF -THE REPTILIA. III

the persistence in certain forms of a suture between the coracoid

and procoracoid.

This line also became very highly specialized, and died out in

the Trias.

_ Fic. r2.— Restoration of Ranbolophoras (Order Petycosaurta). About fẹ nat. size. After Case.

6. Order CHORISTODERA Cope.

These amphibious animals, found thus far only in the Creta-

ceous and in the Lower Eocene, represent a sharply defined

division with a greatly elongated gavialoid rostrum, teeth acro-

dont and internally folded, dorsal ribs two-headed. They include

the American Champsosaurus and the European Simoedosaurus ;

the latter being distinguished by more distinct adaptation to

aquatic life were made the type of the order Simoedosauria by

Dollo. The analogies are with the Proganosauria and the aquatic

true Rhynchocephalia, but there are no phylogenetic relation-

ships with these forms.

II2 THE AMERICAN NATURALIST. [Vor. XXXVIII.

7. Order RuvNcHocEPHALIA Günther.

These are the * Rhynchocephalia Vera ” of Boulenger. They

represent by far the most conservative of all the Diaptosauria

because even the recent Sphenodon is in certain respects more

primitive than most of its Permian relatives.

The order includes the Jurassic radiation of terrestrial, semi-

aquatic, and fully aquatic types, with modifications paralleling

ycosaurian (Embolophorus) B, Profile

view of conjoined scapula and procoracoid of same. This is one of the few

Diapsida in which the pr id is still separated by suture from the cora-

coid. C. Pelvis of same. After Case. x }.

those in the Choristodera and Proganosauria, also the existing

terrestrial genus Sphenodon. The Jurassic forms are in many

respects less primitive than the existing genus.

II. Order PARASUCHIA Huxley.

This amphibious fresh water group, typified by Phytosaurus and

Aétosaurus, placed in the suborders Aétosauria and Phytosauria

respectively, has long been treated in connection with the Croc-

odilia, owing to Huxley's influence and authority; McGregor

has Shown that it has practically no affinity to the Crocodilia,

its relationships being closer to the Ichthyosauria, although it

constitutes an independent order, probably of freshwater, littoral,

carnivorous, short snouted (Aétosaurus) or long snouted (Phyto-

No. 446] RECLA.SSIFICA TION OF THE REPTILIA. $14

saurus, Mystriosuchus) forms, analogous in their habits to the

modern Crocodilia.

The Parasuchia are found by McGregor to represent an

undoubted modification of the rhynchocephaloid or diaptosaurian

type.

III. Order Icutuyosauria Blainville.

The ichthyosaurs are also distinctively Diapsida, both in

shoulder girdle and in basicranial structure. The latero-tem-

poral fenestra, however, is closed, perhaps secondarily. The

posterior position of the nares and the elongation of the snout

in front, is analogous to that in the Parasuchia and may be

indicative of divergence from a common stem.

The most primitive form, Mixosaurus affords a transition to

the ambulatory limb type of the Diaptosauria. None of the

known orders of Diaptosauria, however, can as yet be considered

ancestral to the ichthyosaurs.

IV. Order CROcopiLiA Wagler.

We must exclude from the Crocodilia the Parasuchia of

Huxley and embrace only the Mesosuchia and Eusuchia of

Huxley with the addition of the typical marine forms, the

Thalattosuchia, recently monographed by Fraas.

V. Superorder DivosAvRiA Owen.

It is a mistake to raise this group to the rank of a DES

as has been done by some authors, because its three great sub-

divisions certainly lead back to a common stem form in the

Permian which was not dissimilar to the type represented by

the Protorosauria.

The ordinal nomenclature still requires further study. At

present I am disposed to place the carnivorous forms in the

order Theropoda Marsh, including two suborders, (1) Mega-

losauria for the large types with solid, hour-glass shaped

II4 THE AMERICAN NATURALIST. (Vot. XXXVIII.

vertebra, (2) Symphypoda Cope for the smaller types with

hollow, cylindrical vertebrae.

The Opisthoccelia of Owen although proposed as a ‘ suborder ’

of Crocodilia appears to enjoy priority of definition over the

Cetiosauria of Seeley or the Sauropoda of Marsh.!

Similarly the Orthopoda of Cope is distinctively prior to the

admirable term Predentata of Marsh. In the selection of these

terms we cannot be governed by our preferences ; we are bound

to stand by the law of priority.

VI. Superorder SovAMATA Oppel.

This superorder ranks in value with the Diaptosauria and

Dinosauria inasmuch as it includes the very wide adaptive radia-

tion of three groups of animals which were undoubtedly closely

related in origin, namely : (1) Lacertilia, (2) Mosasauria, (3)

Ophidia. |

The radiation of the Mosasauria from the Lacertilia is analo-

gous to that which we have observed occurring independently in

three orders of the Diaptosauria, namely, the Proganosauria, the

Choristodera, and the aquatic Rhynchocephalia of the Jurassic.

VII. Order PTEROSAURIA Kaup.

There is no question as to the Diapsidan relationships of the

Pterosauria and as to their original derivation from Rhyncho-

cephaloid types, although their specialization has carried them to a

very great extreme of separation from any known Diaptosauria.

CONCLUSIONS.

I trust that the reclassification of the Reptilia here outlined,

and the order of arrangement here adopted will be found to

simplify their study, Memoirs now in preparation by Broom on

the Procolophonia, by Case on the Pelycosauria, by Brown on

SUNY M AUNT

Riggs, E. S. Structure and Relationships of Opisthoccelian Dinosaurs. Part

» Apatosaurus Marsh. Field Columbian Museum Publ. No. 82, Aug. I, 1903.

No. 446.] ARECLASS/FICATION OF THE REPTILIA. II5

the Choristodera, by McGregor on the Parasuchia, will further

elucidate the still numerous and perplexing questions of phy-

logeny.

Origin of Aves.— The birds probably originated from a

group of Diaptosauria identical with or closely related to that

which gave rise to the Dinosauria. It is not true that birds

have descended from dinosaurs, but there is very strong evi-

dence that birds and dinosaurs are descended from à common

stock.

Origin of Mammals.— There is no question that the mam-

mals are affiliated with the subclass Synapsida rather than with

the Diapsida ; both in skull and shoulder girdle structure and in

the phalangeal formula they are Synapsidan.

As to their nearer relationships they appear to be rather with

the superorder Anomodontia and with the order Cynodontia or

Theriodontia. The divergence of the mammal stem from these

typical reptiles will probably be found to have occurred in the

Permian or Lower Trias of South Africa. In fact Broom has

recently described what he believes to be a mammal jaw,

Karoomys, from the Karoo Beds of South Africa.

AMERICAN el SEUM OF NATURAL HISTORY,

Jecember 28th, 1903.

THE EARLY DEVELOPMENT OF DESMOGNATHUS

FUSCA!

HARRIS HAWTHORNE WILDER.

In a former number of the American Naturalist (March, 1899)

I presented what seem to have been the only published observa-

tions on the development of one of our commonest and most

generally distributed salamanders, Desmognathus fusca, but as

I was then unable to describe the early stages, a most essential

gap in this history remained unfilled.

The eggs which furnished the object of my former sketch

were laid in the laboratory terrarium on or about June 1, 1898,

but as the first observations were made on them@June 11, at

which time they were in the form of well-formed embryos coiled

about enormous yolk-masses, the first eleven days of the devel-

opment remained unknown, a period which includes the

extremely important cleavage stages, the formation of the

blastopore and the beginning of the head and tail folds.

Since that time a number of specimens of Desmognathus

have been kept in our terrarium each spring, and the favorite

hiding-places investigated daily during the egg-laying season, but

with no success until the present year (1903) when on June 22

at 1.00 P. M., there was found a batch of twenty freshly laid eggs

associated with a small but evidently mature female. Atthistime

the eggs were in the early cleavage stages, and varied from the

two-celled stage with the second cleavage forming to that of 16

cells, as represented by the first five rows of Fig. 1. Nine of

these were preserved at once in 5% formaline, and the remain-

der were killed, one or two at a time at intervals representing

the most important stages. The eggs were, however, rather

few in number, and in spite of considerable conservatism in the

daily sacrifice, there were but two left when cleavage was com-

1 From the Zoological Laboratory of Smith College.

117

118 THE AMERICAN NATURALIST, (Vor. XXXVIII.

pleted and the blastopore was about to be formed. These two,

compelled on June 27 to make a journey with me to Maine, in

spite of the utmost care, did not survive the hardships of travel

and the experiment came to an end. I am thus able to record

the development only during the first three days of existence,

leaving the period from the fourth to the eleventh still unknown

save through analogy with allied forms, and waiting to be written

when occasion may offer.

This early history includes only surface study, mainly of

preserved material, but seems to be of especial importance as

showing a genuine holoblastic type of cleavage when from the

later form of the embryo a meroblastic form was to have been

expected.

The various cleavage stages are represented in Fig. 1, in

which the first perpendicular column represents them as seen

from the upper pole, the second from the side, and the third

from below; the figures in each horizontal row representing

several views of the same egg. Of these stages the first five

occurred simultaneously at the time the eggs were found, 1.00

p. M. June 22, and as no eggs were found the day before, they

probably represent stages of 6-15 hours, resulting from eggs

laid at intervals during the preceding night.

The successive changes and the descent of the various cells

seem evident, and may be traced with an approximate certainty

as far as the 23-cell stage, as indicated by the lettering. In the

first stage shown, the first cleavage is complete, separating the

egg into its two halves, A; B+C+D and E+F+G+H, while

the second cleavage, beginning at the upper pole, has proceeded

nearly to the equator and is separating the halves into the com-

ponents A+B, C+D, E+F and G+H, a stage which is seen

completed in the second row.

The third cleavage, at least in the eggs examined, is not the

typical horizontal one which might be expected, but consists of

a set of four meridional ones, which start near the upper pole as

so many separate fissures from one of the two former cleavage

planes, probably the first. Similar fissures were observed in

one instance near the lower pole and it is evident that those

from above and those from below meet one another, and result

No. 446.]

1st cleavage completed,

2nd cleavage begun,

1.00 P. M

f |

[Th gg

at this date, and then

that of 16 cells.]

2nd cleavage com-

pleted, 4-celled stage.

June 22, 1.00 P. M.

Reo}

1l clea

H 2

kad v

age. June 22, 1.00

P. M.

3rd cleavage completed,

8-celled stage. June

22, 1.00 P. M.

4th cleavage completed

16-celled stage. June

22, 1.00 P. M.

28-celled stage, with 16

romeres, 12 mac-

Tomeres. June 22,

4:30 P. M.

42-celled stage, with 26

ui m 2 mac-

dei - June 22,

11-00 P. M.

DESMOGNATHUS FUSCA.

Fic. 1.— Eggs of Desmognathus fusca [1903 material]. ! ce es

stages, observed June 22, between 1.00 and 11.00 pM. T :

and column shows upper poles ; the next side views ; and the right-

H x 6. ! i:

120 THE AMERICAN NATURALIST. [VoL. XXXVIII.

in the formation of an eight-cell stage, which is bilateral along

the plane of the first cleavage, and consists of the four cells A.

B. C. and D. upon one side, and of E. F. G. and H. on the

other (fourth row of Fig. 1). These eight meridionally arranged

blastomeres do not seem to have equal value in respect to size,

for through a slight obliquity in these four new lines of cleavage

there result four smaller blastomeres which alternate with four

larger ones, and of these the latter alone usually attain the lower

pole while the former ones do not reach it. This is seen by a

study of the lower pole views in the fifth and sixth rows of Fig.

1, where the smaller blastomeres A. C. E. and G. intrude like

wedges along the lines of the first two cleavages, but do not

reach the pole save in the single instance of A. in the fifth row,

an unusual condition.

The next cleavage is an horizontal one, forming an approxi-

mate circle about the upper pole, and cutting off small segments

from each of the eight blastomeres of the preceding stage (fifth

row). This results in a 16-cell stage, consisting of eight micro-

meres, a, b, c, etc., clustered about the upper pole, and eight

macromeres, the remaining parts of the original cells. This last

cleavage takes place so far above the equator of the egg that it

does not change the aspect of the lower half, and thus the

drawing of this egg (the third of the fifth row) would answer

equally well for the preceding one.

By a comparison with Eycleshymer's studies of Amblystoma,

(Journal of Morphology, Vol. X, 1895) it becomes evident that

this latter cleavage is the one which is described as typically the

third in Amphibian eggs, and that the four meridional cleavage

lines which result in the formation of the 8 cell stage, together

form the usual fourth. Indeed, this transposition of the two

cleavages occasionally occurs in Amblystoma, and as my obser-

vation rests upon the study of but two eggs, it cannot be

asserted that the order described is the typical one in Desmog-

nathus. It is, however, identical with the method shown and

figured by Kerr in Lepidosiren, and his figures of the third

cleavage (by means of the four short lines) as copied by Ziegler

(Figs. 213-314 of his Entwicklungsges. der. niederen Tiere.

1902) would serve in every respect as better pictures of Des-

No. 446.] DESMOGNATHUS FUSCA. I21

mognathus eggs in the corresponding stages than I have been

able to draw. The occurrence of the third meridional before

the horizontal cleavage seems also to be the rule in Acipenser

and in Amia.

The next stage, that of 28 cells, is shown in the sixth row, and

appears to consist, first, of a division of each micromere,

increasing their number to 16, and, secondly, of a longitudinal

division of the smaller macromeres A, C, E, and G, into A’. A."

C'. C". etc., while the other four B. D. F. and H. remain

undivided. The subdivisions of the smaller macromeres may or

may not be visible from the lower pole, as is seen in the third

figure of the row.

Beyond this the subdivisions cannot be followed with certainty,

but the last figure shown (seventh row of Fig. 1) which consists

of 26 micromeres and 16 macromeres, 42 in all, appears in some

June 23, 10.30 A. M. June 23, 10-30 A. M. June 24, 11.30- M.

ide view. Pole.

Fic. 2.— Eggs of Desmognathus fusca [1903 material]. Later segmentation stages, represent-

ing a direct continuation of the series shown in Fig. 1. X 6

respects to show some regularity, although in others it is dis-

appointing. We would expect, indeed, to account for the

increase from 12 to 16 macromeres by a subdivision of each of

the remaining ones, B. D. F. and H, but while this can be seen

to be the fact in the case of F and H, thé two other new macro

meres cannot well be explained by a subdivision of B and D.

Instead of this, they appear as those marked x and y, and the

relation of those and of B and D to the smaller intruded macro-

meres is such as not to allow the desired interpretation. Less

difficulty appears in the interpretation of the 26 micromeres, =

while we would naturally expect a doubling of each of the 16 0

the former stage, and a consequent 32 in this, its smaller coe

ber may well be accounted for by the supposition that some

the original 16 have not yet divided. No definite interpretation —

122 THE AMERICAN NATURALIST. (Vor. XXXVIII.

can be made, however, for this 42-cell stage, since but a single

specimen was available for study, but it may be seen from the

above sketch of this and the preceding stages that the eggs of

Desmognathus will furnish a very interesting and convenient

object for the study of cleavage problems in Amphibia.

Sketches of two later cleavage stages are given in Fig. 2 and

may be seen to consist of a greater and greater subdivision of

both micro- and macromeres, apparently without special regu-

larity. The last figure (June 24) represents the lower pole of

the oldest stage I succeeded in obtaining.

The above observations, which establish the fact that the

FiG. 3.— Farly embryos of Desmognathus fusca [1898 material]. The figures in the upper

Tow represent different aspects of a single egg of the stage described in the previous

r as“ A.” Similarly, the two figures of the lower row represent different aspects

of the stage described primarily as “ B.” x 6.

eggs of Desmognathus are holoblastic, will serve to correct the

statement to the opposite effect given in my previous article on

the subject, and while it is always unpleasant to be proven ina

mis-statement, it is more satisfactory to the investigator to be

able to furnish his own proof than to leave it to others. At the

time of writing my previous article, I possessed no embryos

younger than those represented in the upper row of Fig. 3, and

it then appeared to me impossible that such a relation between

embryo and yolk as the one shown here could result from an ess

of the holoblastic type. A slightly older stage, that represented

in the second row of Fig. 3, seemed still more convincing in

this respect, since by this time a set of branching blood vessels

No. 446.) DESMOGNATHUS FUSCA. 123

had made their appearance on the surface of the yolk, reminding

one forcibly of similar embryos e. g., those of certain Teleosts,

that result from meroblastic eggs. Too sharp a distinction

between the holoblastic and meroblastic type cannot, however,

be drawn, since there are numerous transition forms, as that of

Salamandra maculosa, in which the egg is 4-5 mm. in diameter,

and the first two cleavage planes, although they slowly cut their

way around the egg on the outside, often fail to divide the yolk

internally, * so dass die Furchung in den ersten Stadien partiell

ist" (Ziegler Joc. cit, p. 249, on the authority of Grónroos,

1903). A similar condition has been produced experimentally

in the frog's egg by O. Hertwig (1897). The eggs of the

Gymnophiona, which are the largest of all Amphibian eggs,

(7-8 mm. in Hypogeophis rostratus) are at first typically mero-

blastic, with a germinal disk that spreads gradually over the

yolk; but during and after gastrulation the yolk becomes

divided up into large cells, so that the eggs are ultimately

holoblastic.

In these latter, also, as well as in the eggs of Salamandra and

Desmognathus, there is a noticeable array of blood vessels upon

the yolk, which at first sight closely resembles a true yolk circu-

lation, but which in reality consists of the vessels which normally

appear superficially in the region distended by the yolk mass,

and thus, although they may receive nutriment from the yolk,

they are in no sense true vitelline vessels. The veins of this

region, though not the arteries, could be closely followed in the

two series from which Figs. 4 and 5 were taken, and are shown

to be three in number, one median and two lateral. The median

vein lies along the mid-ventral line of the swollen abdomen,

scarcely noticeable posteriorly, but increasing anteriorly as it

picks up several lateral branches. It passes along the dorsal

(concave) aspect of the liver and enters the sinus venosus in

company with the hepatic vein. By its position and course It

undoubtedly corresponds to the abdominal vein of the —

and thus, although it may be also a potential portal, it cannot b

very definitely related to the true vitelline veins of gei

embryos. In the same way the two lateral veins sre xa T "

identical with the large cutaneous veins which lie along the sides

I24 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of the trunk. They empty into the Ductus cuvieri near the

entrance of the subclavian.

Thus, although it is seen that many Amphibian eggs are on

the border between the holoblastic and meroblastic types, and

although they form interesting transitions between the two,

especially useful in breaking down artificial distinctions, yet it

must be confessed that through the observations here recorded

the eggs of Desmognathus are at first almost typically holo-

blastic, although in the later relation of embryo to yolk they

greatly resemble meroblastic embryos. (cf. Fig. 3 of this paper

with the following in Ziegler's text-book, 1902, Fig. 304.)

Hypogeophis (Brauer); Fig. 188 herring (Kupffer).

In concluding this paper I wish to present a few sections

Sas =e £

LRE

Fic. 4.— Sagittal section,

a little to one side of the median line, of an advanced embryo of

Desmognathus fusca (1

X t5.

898 material] a little older than * Stage D” of the previous paper.

g P

taken through advanced embryos of Desmognathus (the 1898

material) which will show the actual relation of the embryo to

the large yolk mass and will demonstrate its cellular character in

the later stages.

Fig. 4 is a sagittal section of an advanced embryo, and shows

the median relationships in the ventral half, but strikes the

brain and the dorsal portion somewhat laterally. By combining

several adjacent sections, the pharynx, cesophagus and anus

were completed. In this the peritoneum may be followed

dorsally as a continuous line investing the rectum, the yolk mass

and the cesophagus, and enclosing a portion of the coelom ; and

ventrally around the yolk mass enclosing a large coelomic space

No. 446.] DESMOGNATHUS FUSCA. 125

ventral to heart and liver and a smaller one ventral to the anus.

The entire yolk mass, which is wholly cellular, is enclosed within

the peritoneum and is morphologically the equivalent of the mid-

dle portion of the intestinal tract in which a lumen has not as yet

appeared. Fig. 5 shows four cross-sections of an embryo a little

older than that of Fig. 4 and taken from a single series, the

first through the liver, the second through the yolk at its great-

est circumference, the third through the hip-girdle and rectum

and the fourth through the cloaca, with the hind-limbs at the

side. The numbers attached to each are those of the sections

as numbered in the series. These show the same relationships

as seen in Fig. 4, the second especially, in which two lateral

e p ©

fi; ie

YN»

Fic. 5.— Four cross-sections from an advanced embryo of Desmognathus fusca [1898

material] slightly oec than Fig. 4. The sections are 20 # in thickness, and the com-

aay sn from the pt the nose © the end of ihe cloaca Coun of 335 sections.

X 15.

cclomic spaces may be seen dorsal to the yolk-mass, separated

by a mesentery. In the first the alimentary canal (cesophagus

or beginning of stomach) is suspended between dorsal and ven-

tral mesenteries, in the latter of which lies the already well-

developed liver, while in the former is seen the beginning of a

pancreas. In the third section are seen the two lateral cœ-

lomic cavities, dorsal and ventral mesenteries, the hip girdles and

the heads of the femora, and in the fourth are the last (most

posterior) remnants of the coeloms, the open cloaca, and the two

legs, the one cut through the femur, and the other through tibia

and fibula. :

SMITH COLLEGE,

Nov., 1903.

THE HISTOLOGY OF THE LIGHT ORGANS OF

PHOTINUS MARGINELLUS.

ANNE B. TOWNSEND.

ORGANIC light has always been a subject of interest, both to

the scientist and to the casual observer of nature, and no other

photogenic organisms are of such wide range and easy access as

the fireflies. In some of its phases the light of the firefly comes

within the scope of the physicist and chemist. To the former

belongs the consideration of its nature, of its spectrum and other

physical properties ; to the latter come the problems of chemical

analysis concerned with the phenomenon, but to the student of

entomology remains the study of the delicate living structure in

which this wonderful process of photogeny takes place.

In spite of the abundance of these insects, little work has

been done on the structure of the light-organs in our American

fireflies. During the past twenty years some careful investiga-

tions have been made upon European and Cuban forms, but

there is practically no literature upon those of our own country.

The purpose of my study has been to learn something of the

histology of the light-organs of Photinus marginellus, the most

common firefly about Ithaca during June and July. This work

was begun at the suggestion of Dr. Wm. H. Seaman of Wash-

ington, D. C., whose advice during its progress has been most

helpful. I am also indebted to Professor Comstock and to the

members of his staff at Cornell University for their kindly

supervision of my work. |

The material from which this work was done was collected

near Cornell University during the summers of 1901 à

1902. The insects begin to appear by the middle ot June, but

are not abundant until July. The height of the Hyg esent

during the first part of July. The large majority of the insects

! Contribution from the Entomological Laboratory of Cornell University.

127

128 THE AMERICAN NATURALIST. (Vor. XXXVIII.

caught were males. A number of females were found, but

never on the wing. They were always either in the grass, or on

some low plant where they could easily have climbed.

Three methods of fixation were tried the first summer. For

two of these the insects were killed by dropping into hot water.

The caudal part of the abdomen was then clipped off and put

into the fixing fluid without removing any part of the chitin.

Half of this material was fixed for twenty-four hours in Flem-

ming's solution (strong). This proved almost worthless, with

the exception of a single slide. The other half of this material

was fixed in Gilson's mercuro-nitric solution for seven hours.

This gave very good preparations of the general features of both

layers of the light-organs.

The third lot of material was killed and fixed in hot 70%

alcohol. This was satisfactory for the gross structure of the

organs, though not good for the finer details.

The second summer the killing of the insects in hot water was

abandoned. The cadual portion of the abdomen was clipped off,

the tip cut away, and the dorsal wall removed to insure better

penetration of the fixing fluid. The living tissue was then put

directly into the fixer. Flemming’s fluid, after this treatment,

gave much better results than before. Hermann’s fluid gave a

fairly good fixation. The cells of the cylinders are definitely

outlined in these preparations, but the tissues in general are

opaque.

By far the best results for the tracheal stfuctures were

obtained by the use of osmic acid. The strength of the acid

was varied from 1% to 1%, and the time of fixation, from two

to thirty hours. The best preparations were from material left

for thirty hours in 1 h osmic acid.

Material fixed in Flemming's or Hermann's fluid, or in osmic

acid, was washed for twenty-four hours in running water. That

fixed in Gilson's fluid was put directly into 70% alcohol. All

the material was dehydrated by carrying it through the grades

of alcohol, from 70% to absolute. Cedar oil was used for

clearing. These sections were all cut in paraffin.

The greater part of the sections were cut IO p in thickness,

but a few thinner sections, 3 u and 5 p, were made for the deter-

mination of some finer structural points. |

No. 446.] PHOTINUS MARGINELLUS. 129

The material was all stained after the sections were cut. The

Gilson and alcoholic sections were stained for about two minutes

in Gage’s chloral hematoxylin, and for a half minute in eosin.

This gave satisfactory general results. A double stain with

alum carmine and picric acid did not prove a good stain after

Gilson’s fluid. Following fixation with osmic acid, Hermann’s

or Flemming’s fluids, safranin proved most satisfactory. The

sections were stained for twenty-four hours in a mixture of equal

parts of saturated aqueous and alcoholic solutions of safranin,

then differentiated in absolute or even slightly acid alcohol.

Iron hematoxylin is also a good stain for osmic acid material.

Carbol-xylene was used as a Clearer, and the sections were

mounted in Canada balsam.

Teazed preparations were made by dissecting out the light-

organs entire, placing them for from fifteen to thirty minutes in

-5% osmic acid, then for a few minutes in weak caustic potash,

and teazing in normal salt solution.

Experiments with methylene blue injection were not successful.

Only one insect showed any coloration of the central nervous

system, and in that the finer nerves could not be traced. It is

difficult to get a good injection with such small insects. How-

ever, my attempts in this line were begun near the close of the

collecting season, and with further experimenting it is possible

the results might have been better. The most successful speci-

men was killed an hour and a half after injection. |

Since the old idea that the firefly's light was dependent upon

the presence of phosphorus or some similar substance has been

abandoned, other theories have been advanced in attempt to

explain the phenomenon. The view generally accepted is that

the light results from the oxidation of a substance produced by

the metabolism of the light-organ cells. The nature of this

substance has not been determined, but that its photogenic prop-

erty is independent of the life of the cell is proved by the fact :

that when the organs are dried and reduced to a powder the

light reappears under the influence of air and moisture. When

the fresh photogenic tissue is crushed, the light ipcrenses m

brilliancy, and it is some time before it wholly disappears.

Radziszewski ('80) through his study of the artificial luminosity

I30 THE AMERICAN NATURALIST. (Vor. XXXVIII.

of lophin, discovered a series of carbon compounds similar to

those found in living organisms and capable of becoming lumin-

ous under conditions compatible with life. The conditions nec-

essary for this production of light he found to be the presence of

oxygen, an alkaline reaction and slow chemical action. Watasé

(96) states that in the firefly the phenomenon is due to the oxi-

dation, in alkaline media, of a granular substance secreted by

the cells of the photogenic tissue. He offers no further sugges-

tion as to the character of the substance than that it is **a secre-

tion of fatty nature." He gives as proof of the oxidation theory

the fact that when the photogenic material is crushed on a slide

and lowered into a jar of carbon dioxide the light disappears

instantly, but reappears when the slide is placed in a jar of oxy-

gen, or simply in the air. This may be repeated several times

with the same material. Watasé recognizes the necessity of

moisture as well as of oxygen in the process of photogeny.

Dubois stands almost alone in opposing the theory of oxida-

tion. As a result of experiments with ozone, nascent oxygen

and oxygenated water he states (95), that the action of ener-

getic oxidizing reagents at once and finally extinguishes the

light, without first causing any increase in brilliancy. How-

ever, the absence of oxygen seems to destroy the light, as it is

suspended when the light-organs are placed in a vacuum. From

his earlier work he concluded that the light was the result of a

process of crystallization. His later work ('982) has led him to

abandon this theory. He still rejects the oxidation hypothesis

as crude and unscientific, and offers in its place one of a reaction

between two substances to which he has given the names luci-

ferase and luciferine. The.accessibility of the material led him

to use Pholas dactylus, a marine mollusk, as the basis of his

study. The inner wall of the siphon of a large Pholas was

scraped with a knife and the resulting pulp crushed with sand

i 95% alcohol. After twelve hours it was filtered and a

liquid. obtained which was not luminous, even after vigorous

agitation with air. The alcohol was drained off from the resi-

Wes bd eye added. After some hours a second non-

inde RUE filtered off. A mixture of one part of the

ree parts of the second gave a beautiful

No. 446.] PHOTINUS MARGINELLUS. I3I

luminescence at ordinary temperature. By adding to liquid

No. 2 five or six times its volume of 95% alcohol, or by boiling

it, a white floccose precipitate was formed and the mixture of

the remaining liquid with liquid No. 1 no longer produced light.

He therefore considers the white precipitate as constituting one

of the two photogenic substances, the luciferase. Luciferine was

obtained in an impure state by evaporation of alcoholic liquid No.

I. Another experiment was tried with the luminous mucous

secreted by Pholas. Two portions of the mucous mixed with

water were taken, and one extinguished by agitation, the other

by bringing to the boiling point. The mixture of the resulting

non-luminous liquids was photogenic. A similar result was

obtained with the prothoracic organs of Pyrophorus noctilucus.

One was extinguished by crushing, the other by dropping it into

boiling water. When the latter was crushed and mixed with

the former, the light reappeared.

Dubois therefore states that he has established experimentally

that the light of living organisms is produced, in the presence

of water and oxygen, by the reaction between luciferase, an

instable proteid substance possessing in large measure the

general properties of an enzyme, and luciferine, a chemical

Substance. While Dubois confidently asserts that biological

light is not a result of oxidation, his experiments would not

seem to prove this conclusively. He himself admits the neces-

sity of oxygen, and even if the process is not one of simple,

complete oxidation, it would yet seem probable that oxidation

is the essential factor in photogeny. Td

The photogenic tissue of Photinus responds definitely to the

action of oxygen. A series of experiments shows uniformly a

decided increase in the brilliancy of the light when the tissue

is placed in a jar of oxygen. Tissue in which the light hast Been

wholly extinguished by the action of carbon dioxide becomes

instantly photogenic when placed in oxygen. Until there has

been more extensive experimental study of a large number of

organisms, generalizations on the subject of organic light are

unsafe. gu :

The fact that Dubois's work was done upon a marine organism

is significant. If, as one must assume, the photogenic Hen

142 THE AMERICAN NATURALIST. [Vor. XXXVIII.

has arisen independently in different organisms, it would not

seem strange that the light of such widely separated forms as a

marine mollusk and a terrestrial insect, though in both cases a

process of oxidation, might be produced in a different manner.

The physical properties of the firefly'slight have been studied

by Dubois, Langley and Very, Young, and Watasé, with essen-

tially the same results. The spectrum given by the light of the

Lampyrida is perfectly continuous, without any trace of lines,

either bright or dark. It lies within that portion of the spectrum

which most powerfully affects the organs of vision, though

having small thermal or actinic effect. Dubois has demonstrated

by photography the presence of some actinic rays in the light of

Pyrophorus. A single insect was used, and five minutes was

required for printing from a plate which would have taken only

a fraction of a second with sunlight. Dubois attributed the

presence of actinic rays to a fluorescent substance which he

found in the blood.

Most careful and elaborate experiments have failed to show

more than an infinitesimal amount of heat connected with the

light. One authority even goes so far as to say that not more

than one-thousandth of the energy expended in the flash of the

firefly is converted into heat waves. When one considers that

in our ordinary oil or gas lamps more than ninety-nine per cent

of the energy is lost as regards illumination, and that even in the

arc light only about ten per cent. of the waves are visible, the

interest which this *cheapest form of light " arouses from the

economic point of view is very apparent. It is also an alluring

problem to the student of physics to determine by what process

the medium wave lengths are produced independent of the longer

and shorter waves. If this “secret process" could be wrested

from nature, its economic value would prove almost inestimable.

While the phenomena of biological light early attracted the

attention of observers of nature, as Aristotle, Democritus and the

naturalist Pliny, it is only within the last century that any serious

study has been given to the organs which produce it. The dis-

covery of their cellular nature may be credited to Peters. In

1841 he refuted the theory of Carrara ('36) that the light was

dependent upon an air-sac extending from the mouth to the light-

No. 446.] PHOTINUS MARGINELLUS. 133

organs and acting as a bellows, and stated that the photogenic

tissue was made up of little spheres, regularly arranged and pen-

etrated by the tracheæ. Leydig and Kölliker in 1857 definitely

recognized the cellular structure of the light-organs. Their work

has been followed by that of several other European investigators.

During the past twenty years Wielowiejski and Emery have

made important histological researches upon species of Lampyris

and Luciola.

All recent workers agree in stating that the ventral light-plates

of the male lampyrids are composed of two more or less clearly

defined layers ; the dorsal, chalky, opaque layer, and the ventral,

or truly photogenic layer. The former is composed of fairly

regular, polygonal cells, filled with a great quantity of crystals of

urate salts. The ventral layer is composed of two distinct ele-

ments ; the tracheal structures and intermediate areas of paren-

chyma. The parenchyma cells contain fine granules of non-urate

composition. The main trachez of the photogenic segments

send vertical branches down through the light-organs. Aside

from their profuse branching they show no unusual features

until they reach the ventral layer. The tracheal structures

within the ventral layer differ in different forms, and the two

species upon which Wielowiejski and Emery based the bulk of

their work, Lampyris splendidula and Luciola italica. respec-

tively, show a considerable difference in this respect. In

Luciola each vertical tracheal axis is surrounded by a cylindrical

mass of semi-transparent tissue, within which it branches in an

arborescent manner. The method of branching in Lampyris 1$

fasciculate, rather than aborescent, and the trachez are much less

regular in their distribution. Max Schultze, in 1864, found in

osmic acid preparations from the light-organs of Lampyris sp -—

didula certain blackened bodies at the periphery oe —

These he found to be penetrated by the finer tracheæ. dvd

to find further continuations of the trachez beyond these res,

he called them the “ tracheal end cells.” Wielowiejski À pem

his study of the same species, found that instead - — : i:

ultimate endings within the so-called «end cells, ae daten

branch, sending out fine “tracheal capillaries — w ith k d

beyond the cylinder and in most cases —— i DT

134 THE AMERICAN NATURALIST. [Vor. XXXVIII.

adjoining cylinders. Emery, from his work on Luciola, confirms

the views of Wielowiejski in all points except as regards the

anastomosing of the capillaries. He found them always ending

free, never uniting with those from the same or another

cylinder.

In this work the nomenclature previously used has been

retained except in a few cases where a change seemed especially

advisable. The more familiar terms phosphorescent and lumin-

escent, with their nouns, are abandoned and photogenic and

photogeny substituted. Phosphorescent is objectionable as it

suggests that the light is due to the presence of phosphorus.

Photogenic — light-generating — gives a more definite idea of

the actual phenomenon than luminescent. The name “end

cells" was used by Max Schultze because he believed the

trachee had their ultimate endings within these cells. Now

that it is proved that the trachez do not so end, but merely pass

over into the tracheolar network, the name * transition cells," as

used by Holmgren ('96), is far preferable. The term tracheoles,

which is used elsewhere in insect histology to designate fine

trachael branches not possessing spiral thickening, is prefer-

able to * tracheal capillaries."

The light-organs of the male of Photinus marginellus are in

the form of two plates lying above the ventral body wall of the

fifth and sixth abdominal seg-

ments. (Figs. 1 and 2.) The

underlying cuticle is trans-

parent, allowing free emission

of the light. The plates lie

just beneath the central nerv-

ous system and directly upon

; the very thin hypodermis.

vibus ien. section through entire abdo- They are yellowish white in

color. In the female of the

Same species dics is only a single, somewhat spherical organ

in the centre of the fifth segment. In this work all descriptions

of structure refer to the light-organs of the adult male. The

main trachez of the photogenic segments send branches ventrad

through the light-organs. Thus the dorsal surface in fresh

No. 446.] PHOTINUS MARGINELLUS. 135

material is shown to be pentetrated by numerous trachee. The

vertical or oblique trachez continue to branch profusely in an

arborescent manner. This repeated branching is characteristic

of the trachez of the photogenic tissues. The tracheal epithel-

ium is composed of thin, flattened cells, with large flattened

nuclei. Prominent hair-like projections of the intima are abun-

dant in the lumen of the large tubes. These internal chitinous

hairs have been noted in Lampyris by Gersteecker, and in

Luciola by Emery. The light-organs are innervated by nerves

from the last two abdominal ganglia. These ganglia are both

situated in the fifth segment, over the more cephalic light-plate.

I failed to trace more than these primary nerve branches, as my

attempts at methylene blue injection were unsuccessful.

Each light-plate is composed of two distinct layers, in this

Fic. 2.— Longitudinal section through abdomen from fifth segment to

e o.

caudal end. x2

respect agreeing in structure with the European species which

have been studied. In none of my preparations have I been

able to detect any trace of a membrane, either surrounding the

light-organs or separating the two layers. | Wielowiejski (82)

States that in Lampyris splendidula each light-plate is surrounded

by a delicate film of connective tissue, in which small rounded

nuclei may be faintly seen in well stained material. He gives

no figures of this, however. Emery (84) says there is no indi-

cation of a membrane in Luciola. The two layers cannot be

distinguished in fresh material, but a difference in the two

surfaces of the light-organ is apparent. The dorsal surface 1s

a bright chalky white, while the ventral surface appears yellow-

ish and luminous. Examined with a low power of the micro- -

136 THE AMERICAN NATURALIST, (VoL. XXXVIII.

scope the dorsal surface is seen to be divided into polygonal

areas.

The appearance of the cells of the dorsal layer varies much

with the treatment of the tissue. Material fixed in alcohol and

brought in contact with water for but a short time in staining

with hematoxylin shows the cells filled with a dense content of

coarse granules. With reflected light these granules still show

their characteristic chalky whiteness, while with transmitted

light they are brown. Granules identical in appearance are also

found in the fat cells of the same region of the body. Material

fixed in any fluid requiring subsequent washing in water shows

a considerable decrease in the

granules of the dorsal layer.

DX En XI cds 2 CIAR ~

- Dra Tm T

eh ===. This verifies the statement of

Cg E m

Wielowiejski that these gran-

ules are insoluble in alcohol,

but soluble in water. Kolli-

ker,in 1857, proved them to

be crystals of urate salts, and

his results have been accepted

by Wielowiejski and Emery.

When the crystals have been

dissolved out the form of the

€ oe (nhi x both layers of light — Cells is easily determined.

: " uson’s fluid, hematoxylin and eosin. They are polygonal, fairly

regular in outline and simi-

lar in size. They average about 28 u by 25 m. Those upon the

upper surface are somewhat more spherical than those beneath.

Large nuclei are always present, but the cytoplasm seems to

have been almost entirely replaced by the granular secretion.

(Fig. 3.)

The dorsal layer not only forms a plate resting upon the

ventral layer, but it projects beyond the latter and extends along

Its caudal surface to the body wall (Fig. 2). There are two

groups of muscle fibres in each light plate, extending from the

dorsal to the ventral body wallin the lateral portions of the

plate. These muscles are surrounded by a layer of cells dis-

tinctly separated from the cells of the ventral layer and contin-

No. 446.] PHOTINUS MARGINELLUS. 137

uous with those of the dorsal layer. In material in which the

dark granules of the dorsal layer cells have not been dissolved

out, they are found equally in the cells surrounding these groups

of muscle. (Fig. 1.)

The ventral layer is composed of two distinct elements, the so-

called parenchyma cells, and the cylinders. The cells of the par-

enchyma differ from those of the dorsal layer in being very

irregular in shape and size. Occasional cells extend from the

dorsal surface of the layer to the body wall.

In some cases the depth is several times the

thickness, while some cells are almost spheri-

cal. In some places the ventral layer is found

to be several cells deep. (Fig.4.) The size

of the cells varies considerably. The granu-

lar secretion in these cells is much finer than

that in the dorsal layer. Max Schultze has

stated that these granules are of non-urate

composition. He examined the granules of 4 ay

both layers with polarized light, finding that Fic. 4—Cells of paren-

those from the dorsal layer were bi-refractive, m i? 2g

but that those from the ventral layer were

not. Those of the dorsal layer having been proved by Kólliker

to be of urate composition, he conclüded the granules of the

ventral layer were non-urate. : Wielowiejski verified the results

of these experiments, but did not agree with Max Schultze's

conclusions. He states that these experiments merely prove

the granules of the ventral layer to be in general amorphous.

He thought, however, that the difference in composition could

be readily demonstrated by reagents. He found the granules of

the dorsal layer to be soluble in water but insoluble in alco-

hol, while the opposite was true of those of the ventral layer.

Emery says that the granules of the ventral layer are not crystal-

line, and that they disappear altogether in balsam preparations.

The cells of the ventral layer appear to have more or less of

granular content in all of my preparations, including those from

material fixed in alcohol. It is only in osmic acid material that

cells are found comparatively free from such secretion. As all

my sections are mounted in balsam, this medium would not seem

138 THE AMERICAN NATURALIST. (VoL. XXXVIII.

to have any decided effect. This question, however, can be

satisfactorily settled only by the study of fresh tissue.

Between the areas of parenchyma are sharply defined, more or

less cylindrical masses of tissue surrounding the vertical tracheal

stems and their branches. It is about these cylinders and their

trachez that the greatest interest is centered. The cylinders are

from 23 x to 68 m apart, and average about 30 4 in diameter.

Their appearance differs greatly with the fixation. In material

fixed in alcohol or Gilson's fluid, and stained with hematoxylin

and eosin they appear as areas less granular, and consequently

less deeply stained, than the intervening parenchyma. They

contain a large number of small nuclei, especially abundant near

the trachez. After fixation in Hermann's fluid and staining

with safranin the cell outlines appear very distinct. (Figs. 10 and

11.) Preparations fixed in Flemming's solution and stained

with safranin also show cellular structure, though not so defi-

nitely as the preceding. Less indication of the structure of the

cylinders is shown in the osmic acid material.

If fresh material, placed for fifteen minutes in .5 % osmic

acid and then treated for a few minutes with weak. caustic

potash, is viewed from the ventral surface, the cylinders appear

as very distinct rings. In all preparations, both temporary and

permanent, the boundaries of the cylinders are sharply defined.

Within each cylinder is a main:tracheal stem which gives rise

to numerous branches in the characteristic aborescent manner.

There is no change in the structure of the trachez until near

the periphery of the cylinder, where each fine tracheal twig

breaks up into tracheoles. The number of tracheoles arising

from one tracheal twig seems to vary somewhat. Ordinarily

there appear to be only two, but three or four are not uncom-

mon. Emery gives the number of tracheoles in Luciola as

being uniformly two. In Lampyris, Wielowiejski found the

number variable, as many as six sometimes occurring.

The tracheoles are fine tracheal branches and are characterized

by having no spiral thickening of the intima. Their chitinous

structure is plainly shown by the fact that they persist in

material treated for some time with caustic potash.

Max Schultze, Targioni-Tozzetti (70) and Emery were all of

No. 446.] PHOTINUS MARGINELLUS 139

the opinion that the tracheoles do not contain air, but a colorless

fluid. Wielowiejski also found them filled. with a fluid, but

recognizing the extreme improbability of such condition existing

in life, he looked for some explanation of it. In dried, air-filled

material mounted in weak glycerine the tracheoles as well as the

larger tracheal tubes were filled with air. After about five

minutes the silvery, glistening lines of air became broken up,

———

Fic. ;5.— T 1 preparation, showing anastomosing of tracheoles.

and gradually, from the tracheoles in, the tracheæ became filled

With a fluid. This would seem to prove, what one must believe

a priori, that the entire tracheal system is filled with air. My

Observations agree with Wielowiejski's, for although ncn

preparations the tracheoles were always alr eady filled with liquid,

the penetration of the liquid into the larger branches was unmis-

takably from the tracheoles in. It may be noted. in Abit come

140 THE AMERICAN NATURALIST. [Vor. XXXVII.

nection that in the tracheoles of developing wings, structures

entirely similar to those of the light-organs, the presence of air

may be readily seen.

In sections parallel to the axis of the cylinder the tracheoles

are generally cut, so that their entire length cannot be followed.

The fact that they appear to end free cannot, therefore, be taken

as any proof. In rare cases they are seen to anastomose with

those of adjacent cylinders, and in sections transverse to the

cylinders, beautiful demonstrations of anastomosing may be

seen. Ifa light-organ from a freshly killed insect is placed for

a half hour in .5 osmic acid, then for a few minutes in caustic

potash, and then placed under the microscope, ventral side up,

the tracheoles can be easily seen. The cylinders stand out as

definite circular or. oval rings, and from the inclosed trachea

radiate the tracheoles. As the distribution of the cylinders is

fairly regular, the network of tracheoles has a notably uniform

pattern. Tracheoles from three cylinders unite at a point about

equidistant from their respective cylinders. (Fig. 5.) In prep-

arations from material fixed in Flemming’s, or Gilson’s fluid, the

same tracheolar network may be clearly seen. (Fig. 6.)

When the ventral surface of a fresh light-organ is studied

under the microscope in a dark room the light is found univer-

sally distributed through-

out the parenchymatous

area. The cylinders stand

out as non-photogenic spots

on the background of light.

This shows that the photog-

eny occurs in that portion

of the tissue where the tra-

cheolar network is found,

and where there is conse-

F1G. 6.— Anastomosing of tracheoles. Fleming’s fluid, t

Ma era g quently the most abundan

supply of oxygen.

Wielowiejski found anastomosing of the tracheoles generally

true in Lampyris, although he admits of some exceptions to the

rule. Wistinghausen and Holmgren found anastomosing of the

tracheoles in the silk-glands of. the caterpillar. Emery states

No. 446.] PHOTINUS MARGINELLUS. I4I

that in Luciola the tracheoles in all cases end free. This is

shown in his figures 4 and 7, Plate XIX.

In Lampyris Wielowiejski figures the tracheoles as winding

irregularly and twisted and looped about each other. In Pho-

tinus they are generally almost straight. They may be slightly

wound about each other, but for the most part they pass directly

from one cylinder to another.

Both Wielowiejski and Emery agree in considering that the

tracheoles pass between the cells of the parenchyma, although

positive proof is difficult. In no case has any portion of a

tracheole been found within the parenchyma cells, although they

have been seen closely applied to the exterior of the cells. In

surface sections of Photinus the areas between the tracheoles

would appear to correspond to cells, each possessing a nucleus.

It thus seems altogether probable that in their course outside

the cylinders the tracheoles are intercellular.

In 1864 Max Schultze studied the light organs of Lampyris

splendidula, using osmic acid as a fixer. He found the finer

tracheal branches losing their spiral thickening and passing into

star-shaped, finely granular bodies which he believed to be true

cells possessing distinct nuclei and cell membranes. He failed

to find any continuation of the trachez beyond these stellate

cells, and so assumed that these cells enclosed the ultimate

endings of the trachez. The name “tracheal end-cells," as

given by Max Schultze has since been generally used, even by

those who knew it to be a misnomer.

In his studies of the same species, Wielowiejski found the

tracheze passing into the stellate “ end-cells " of Max Schultze,

but instead of ending there, branching to form tracheoles which

penetrate the inter-cylindrical parenchyma. He believed these

* end-cells " to be true cells, much flattened and similar in form

to the endothelial cells of vertebrates. They extend about the

bases of the tracheoles in a web-like manner and are more or less

stellate in shape. These cells show a characteristic reaction

with osmic acid, causing a precipitate to be formed, especially

about the point of origin of the tracheoles. The tracheal twig

appears constricted at its apex, and is intensely blackened by

the osmic acid. The effect of the acid varies, in some cases the

entire end cell being uniformly blackened.

142 THE AMERICAN NATURALIST. [Vor. XXXVIII

In the female of Lampyris noctiluca and the female and larva

of Lampyris splendidula Wielowiejski found a somewhat dif-

ferent condition than in the adult male of Z. splendidula. The

tracheoles arising from the ends of the finer trachez are gen-

erally only two in number. They may also occur along the

course of the smaller trachez, instead of only at the ends, and

they may even arise from some of the larger tracheal branches.

It is obvious that under such conditions *'end-cells " like those

figured for L. splendidula would not be present. He states,

however, that there is a membrane spread out between the

tracheoles, although it fails to give the characteristic “ end- .

cell" reduction with osmic acid. From the larva of Z. noc-

tiluca he figures one of the larger trachez with its branches,

with a strongly developed, nucleated membrane surrounding it

much as a cylinder surrounds the tracheal axis in the imago of

Photinus. The epithelium of the large primary trunks of the

tracheze in the larva shows the power of precipitating osmic acid. .

Wielowiejski also studied tracheal endings in other parts ot

the adult Z. splendidula. In the fat body and reproductive

organs he found ** end-cells " in abundance, similar to the typical

ones of the light-organs in their reaction with osmic acid, but

differing considerably in shape and in the number of tracheoles

contained. In all these instances Wielowiejski interprets the

“ end-cells ” as being a special development of the epithelium of

the trachea. In his figures the trachez, before entering the

photogenic tissue, show well developed epithelium, this layer

being sometimes almost as thick as the diameter of the tube.

Wielowiejski neither figures nor describes cylinders in the

light-organs of Lampyris. After his study of Luciola italica

('86) he states that he found no such regular arrangement of the

tracheze here as occurs in the two species of Lampyris formerly

studied. He did not consider the mass of the cylinders in

Luciola as homologous with the ** end-cells ” of Lampyris.

In Luciola italica Emery found the ventral layer composed of

cylinders and intermediate areas of parenchyma much as has

been already described for Photinus. In osmic acid preparations

he saw, just within the periphery of the cylinder, small, irregular,

three-cornered masses, in which the distal ends of the trachea

No. 446.] PHOTINUS MARGINELLUS 143

and bases of the tracheoles appeared to be imbedded. When aa

tracheal axis was isolated these small bodies looked like ** grapes

on a stem," while from each * grape " two tracheoles proceeded.

As these browned bodies were found only in the osmic acid

preparations he believed them to be artifacts, and not the

“tracheal end-cells " of Max Schultze. He concluded that the

clear cell elements of the cylinder are the real *'end-cells."

Within these the tracheze undergo their final division, each

giving rise to two tracheoles. In Luciola only that part of the

cell which is in direct contact with the bases of the tracheoles is

blackened by osmic acid.

Emery agrees with Wielowiejski in considering that the “ end-

cells " are formed from the tracheal epithelium.

Two of the latest investigators of tracheal endings, Wisting-

hausen and Holmgren, have both worked on the silk-glands of

lepidopterous larvae. Both found the finer trachez passing

over into what they term the “tracheal capillary end-network, "

a network formed by the anastomosing of the tracheoles and

their branches. They agree in stating that the epithelium of

the tracheoles is extended in a webdike manner to form the

*end-cells." | Holmgren discards the term * end-cells," sub-

stituting for it the E

more correct name

of *transition cells,"

as these structures

form the transition:

between the tracheal

tubes proper and the

tracheolar net-work.

In the light-organs

of Photinus, fixed for

thirty hours in .1 46

“smic acid and

Stained with safra-

nin, the transition

cells may be seen

most plainly. They

show with varying fus Transiti isi ypi 4 i id paration. xaso-

144 THE AMERICAN NATURALIST. [Vor. XXXVIII.

clearness in all the osmic acid material, and in one insect fixed

in Flemming's fluid. In the typical osmic acid preparation they

appear as irregularly spherical bodies, blackened throughout but

most intensely so at the point of origin of the tracheoles. They

show no appearance of nuclei, but as the nuclei of the adjacent

cells show only faintly with this treatment, this is not significant.

The transition cells of Photinus as shown in osmic acid

preparations are more similar to the blackened, grape-like bodies

described by Emery, than to the stellate, endothelioid cells

figured by Wielowiejski. They occur at the apices of the finer

d. X 300.

tracheal twigs, and near the periphery of the cylinder. The

space between them and the tracheal axis, and the spaces

between the transition cells themselves appear clear. The edges

of the spheroid masses are generally irregular, and their whole

appearance suggests an artificial condition. (Fig. 7.) In sec-

tions where the effect of the osmic acid has not been as exten-

sive, the same blackening at the points of tracheolar origin may

be seen, but instead of finding spherical bodies surrounding these

points of furcation, the granular mass of the cylinder appears in

different condition. It extends along the periphery of the

No. 446.] PHOTINUS MARGINELLUSs. I45

cylinder, and follows the course of each tracheal branch, in

some cases almost to the main stem, so that instead of a struc-

ture resembling a cluster of grapes, one finds along the wall of

the cylinder a series of fan-shaped masses, one for each tracheal

twig, their apices toward the axis of the cylinder. (Fig. 8.)

As there is great irregularity in the form of these dark bodies

within the cylinder, and also in the shape of the intervening

clear spaces, it would seem that Emery is correct in considering

them an artifact.

Definite cellular structure can be seen only in the material

fixed in Hermann's and Flemming's fluids. In these prepara-

tions, as in those from osmic acid, the tissue seems to be

shrunken and distorted. To a large extent the nuclei appear

to have been separated from the cytoplasm, and to lie in the

spaces left by the shrinkage of the cells. The cells show a

tendency to shrink away from each other, and away from the

main axis of the trachea, thus becoming smaller, denser bodies

surrounding the distal ends of the tracheal branches, and in

contact with the periphery of the cylinder. (Figs. 9 & 10.)

In material fixed in alcohol or Gilson’s fluid there is no appear-

ance of cells within the cylinder, although an abundance of small

nuclei may be clearly seen.

Emery suggests that the cylinder, in Luciola, may be a syn-

cytium, but in both longitudinal and

transverse sections of the cylinders

in material of Photinus fixed in

Hermann’s fluid, the cells are

clearly demonstrated.

The “end-cells” of Lampyris

and the cylinders of Luciola are

stated to be a special development

of the tracheal epithelium. This

is not true of the cylinders in E

Photinus, as the avila can be yay sone rud i

definitely seen, not only in the tra- :

cheal axis, but even in the small branches. (Fig. 11.) The x d

thelium of the tracheze of the photogenic tissue is altogether dif-

ferent from that figured by Wielowiejski for Lampyris. Instead

146 THE. AMERICAN NATURALIST. [Vor. XXXVIII.

of being thick, it is very thin, with large flattened nuclei. In

both longitudinal and transverse sections, where a nuclear stain

has been used, the epithelium may be readily traced within the

cylinders, even to the finer branches. It is exactly similar in

appearance to that in the dorsal layer, and is wholly distinct

from the cellular elements of the cylinder. Small flat nuclei

are seen closely applied to the trachez, while the irregular

nuclei of the cylinder cells lie at a little distance away.

In his study of Luciola in 1886, Wielowiejski did. not find the

cylinder a syncytium, as

stated by Emery, but

believed it to be composed

of two elements, a nucleated

epithelium immediately sur-

rounding the trachea, and an

outer layer belonging to the

parenchyma. The latter he

found generally separated

' from the trachea by the

Fic. 1.— Section transverse to cylinder. Her- action of reagents, but still

arabes: X 235. 5

connected with the paren-

chyma cells. These observations would seem to be in agree-

ment with the conditions found in Photinus.

The bulk of the cylinder is thus shown not to be of tracheal

origin. The trachez pass into the cylinder cells, there pipe

to form “the tracheoles, so that the name “transition cells”

equally befitting here, although the structures to which it is

applied cannot. be considered homologous with those of the

"capillary end-network” of the caterpillar. If the cylinders

belong to the parenchyma, they are at least clearly distingue

from it by definite boundary lines. In some

cases there might seem to be a transition

between the cells of the dorsal layer and those

of the cylinder, but the cylinder cells are much

smaller than those of either the dorsal layer

or the parenchyma. They retain their spherical ugh cylinder

shape much more than either of the others showing _ epithel-

mentioned. It would seem probable that all m

Fic. rr.— Section

EURO

No. 446.] PHOTINUS MARGINELLUS. 147

three forms of cells are of the same origin and that their struc-

tural differences are due to difference in function.

Wielowiejski, from his work on Lampyris splendidula believed

in the possible transformation of the parenchyma cells of the

ventral layer into the cells of the dorsal layer through the physi-

ological effects of photogeny. Emery did not accept this theory,

and after his study of Luciola and two American species of

Lampyridze, Wielowiejski himself (89) stated that, for those

forms at least, it was untenable.

The conditions in Photinus are such as to apparently preclude

such an hypothesis. The two layers are distinctly separate in

all preparations and the relative thickness of the layers is fairly

constant. There is no indication of a transition between the two

layers, nor is there any apparent difference in the thickness of

the layers in material put up in early summer and in that taken

at the close of the flying season. Still more important are the

inherent differences in the two layers. In the dorsal layer there

is a solid mass of polygonal cells, similar in form and size, and

irregularly penetrated by tracheæ. In the ventral layer there is

a distinct division into two elements, the cylinders enclosing the

tracheal trees, and the parenchyma cells. The arrangement and

distribution of the trachez of the ventral layer is strikingly reg-

ular. The parenchyma cells are extremely irregular, both in

form and size. It would, therefore, seem difficult to suppose

that the dorsal layer could grow at the expense of the ventral

layer.

Several theories have been offered as to the origin of the

photogenic tissue. Kolliker (57) regarded the light organs as

“nervous apparatus.” | Owsjannikow (68) thought them of

epithelial origin. Wielowiejski (86) suggests their derivation

from the “kleine CEnocyten" which he finds absent in the

photogenic species of Coleoptera. The most general view,

however, is that the photogenic tissue is differentiated fat body.

This is upheld by a general similarity in structure, position and

cell content. It has been already noted that granules exactly

similar in appearance to the urate crystals of the dorsal layer

have been found in the fat body near the light organs.

A question of this character could be settled only by onto-

148 THE AMERICAN NATURALIST. [Vor. XXXVIII.

genetic study of the photogenic tissue. This has not been

attempted, so far as I can learn, by anyone besides Dubois ('98).

He has studied the development of both Lampyris noctiluca and

Pyrophorus noctilucus from the earliest stages. The eggs were

found to be luminous even before they were laid, so that the

light was transmitted in unbroken continuity from one gener-

ation to the next. Dubois followed the development of the

light organs through all the different changes occurring from

the beginning of segmentation to the emerging of the adult

insect, and his observations led him to state definitely that the

photogenic tissue is derived directly from the underlying hypo-

dermis, by a proliferation of these cells. He also states that in

the development of the organs a transformation takes place in

the protoplasm of the cells, the older cells toward the upper

surface of the light organs becoming filled with opaque, chalky

granules. The younger cells, in which this transformation has

not taken place, constitute the parenchyma.

While these results are not in line with previously accepted

ideas in regard to the derivation of the light organs, they are

based upon the only kind of study which can determine the

problem. Apparently Dubois’s work has not been generally

accepted, and it needs verification by other workers. Nothing

could be more profitable in our present state of knowledge than

extensive and thorough study of the photogenic tissue through-

out all its transformations. !

BIBLIOGRAPH Y.1

CARRARA, M.

'36. Sulla phosphorenza della Luciola comune (Lampyris italica L.)

Biblioth. italiana, Tom. 82, pp. 357-370.

Dupsols, R.

'86. Contribution à l'étude de la production de la lumière par les êtres

vivants. Les Elaterideslumineux. Bull. Soc. Zool. France. Vol.

II, pp. 1—275, pl. 1-9.

! As an exhaustive bibliography of this subject has already been published by

Gadeau de Kerville (87), I give here only such references as have direct bearing

upon my work. s

No. 446.] PHOTINUS MARGINELLUS 149

DuBois, R.

'87. De la fonction photogénique dans les oeufs du lampyre. Bull.

Soc. Zool. France, Tom. 12, pp. 137-144.

Dusols, R.

'93. Sur la mechanisme de la production de la lumiére chez l'Orya

barbarica d'Algérie. Compt. rend. Acad. Sc. France, Tom. 117,

pp. 184-186. Translation in Ann. and Mag. Nat. Hist. Ser. 6,

Vol. 12, pp. 415—416.

Dusols, R

'98. Revue Générale des Sciences pure et applicquées. Tom. 5, pp.

415-422 ; 529-534.

Dusois, R.

'96. Physiological light. Smithson. Rept. for 1894-95, PpP. 413-43! ;

pls. 23-26.

DuBois, R.

'96. Sur le luciférase ou zymase photogéne des animaux et des végétaux.

Compt. Rend. Acad. Sc. Paris, Tom. 123; pp. 653-654. Abstr.

by W. A. Nagel. Zool. Centralbl. Jahrg. 5 ; p. 5.

DunBois, R.

'98. Leçons de Physiologie générale et comparée. Paris.

Dusois, R..

'85. Recherches sur l'émission des radiations chimiques par les ani-

maux et les végétaux. Compt. rend. Ass. franc. avanc. Sci., Sess.

22; p. 298.

Dusols, R.

:00. Sur le méchanisme de la biophotogénése. Compt. rend. Soc. Biol.

Paris. T. 52; pp. 569-570-

EMERY, CARLO.

Untersuchungen über Luciola italica L. Zeitschr. wissensch.

Zool. Bd. 4o, pp. 338-355 Taf. 19.

EMERY, CARLO. à

La luce della Luciola italica osservata col microscopio.

Ent. Ital. Ann. 17; pp. 351-355 Tavis;

GADEAU DE KERVILLE, H.

'87. Les Insects phosphorescents. Notes complémentaires e

graphie générale. Rouen.

GADEAU DE KERVILLE, H.

'90. Les végétaux et les animaux lumineux. Rouen.

HOLMGREN, E. :

'95. Die trachealen Endverzweigungen bei den Spinndrüsen der Lepid-

opterenlarven. Anat. Anzeiger. Bd. 11, pp. 340-346.

KóLLIKER, A. :

'58. Die Leuchtorgane von Lampyris, eine vorläufige Mittheilung. Ver-

handl. d. phys. medizin. Gesell: Würzburg. Bd. 8, pp. 217-224-

Bull. Soc.

t Biblio-

150 THE AMERICAN NATURALIST. [Vor. XXXVIII.

LANGLEY, S. P., and VERY, F. W.

'90. On the cheapest form of light. Am. Jour. Sci. Ser. 3, Vol. 40, pp.

97-113.

LEYDIG, FR.

'57. Lehrbuch der vergleichenden Histologie. pp. 343 ff.

MATTEUCI, CARLO.

'483. Sur la phosphorescence du Lampyre d'Italie (L. Italica). Compt.

rend. Acad. Sci. France, Tom. 17, pp. 309-312.

MURAOKA, H.

'96. Das Johanniskáferlicht. Annalen der Physik und Chemic. Bd.

295, PP. 773-781.

OwsJANNIKOW, P.

'68. Ein Beitrag zur Kenntniss der Leuchtorgane von Lampyris nocti-

luca. Mém. Acad. Sci. St. Pétersb. Tome II, no. 17. P-

I pi.

PETERS, W

"1. Ueber das Leuchten der Lampyris italica. Archiv. f. Anatomie.

Jahrg. 1841, pp. 229-233.

RADZISZEWSKI, BR.

'77. Uber das Leuchten des Lophins. Berichte deut. chem. Gesell.

Jahrg. 10, pp. 70-75.

meee ey: Br.

Uber die Phosphorescenz der organischen und organisirten Korper.

Annalen der Chemie. Bd. 203, pp. 305-336.

ee Max.

Ueber den Bau der Leuchtorgane der Mannchen von Lampyris

splendidula. Sitzber. niederrhein Gesell. Natur. u. Heilkunde zu

Bonn. Abstract in Arch. Naturg. Bd. 30, pp. 61-67.

SCHULTZE, MAX.

‘65. Zur Kenntniss der Leuchtorgane von Lampyris splendidula.

Archiv. mikr. Anat. Bd. 1, pp. 124-137, Taf. 5-6.

SEAMAN, WM. H .

'91. On the Luminous Organs of Insects. Proc. Amer. Soc. Micro-

scopists, Vol. 13, pp. 133-157. Pl 1-5.

TARGIONI-TOZZETTI.

'70. Sullorgane che fa lume nelle lucciole volanti d'Italia. Boll. Soc-

Entom. Ital. Anno 2, pp. 177-189. Tav. 1-2.

WATASE, S.

'95. On the physical basis of animal phosphorescence. Biological Lec-

tures delivered at Wood's Holl. 1895, pp. 101- 118.

WIELOWIEJSKI, H. von.

'82. Studien über die Lampyriden. Zeitschr. wissensch. Zool. Bd. 37:

pp. 354-428, Taf. 23-24. :

No. 446.] PHOTINUS MARGINELLUS I51

WIELOWIEJSKI, H,

'86. Über das paure der Insekten. Zeitschr. wissensch. Zool.

43, PP- 512-536.

Wome, H., VON.

'89. Beiträge zur Kenntniss der Leuchtorgane der Insekten. Zool.

Anzeiger, Jahrg. 12, pp. 594-600.

WISTINGHAUSEN, C. VON.

'90. Über Tracheendigungen in der Sericterien der Raupen. Zeitschr.

wissensch. Zool. Bd. 49, pp. 565-583, Taf. 27.

YounG, C. A.

'"J0. Spectrum of the Firefly. Amer. Nat., Vol. 3, p. 615.

ORAL BREATHING VALVES OF TELEOSTS,

THEIR MODIFICATIONS AND RELATION

TO THE SHAPE OF THE MOUTH.

EVELYN GROESBEECK MITCHELL.

Ar THOUGH the oral breathing valves of teleost fishes are very

prominent and their function important, they have, strangely

enough, been almost wholly neglected. Owen (C66), suggests

that they “seem intended to prevent the reflux of the respiratory

stream." Macullum (84) mentions them in his article on Ami-

urus but ascribes to them no function. Stannius (39) has

described them, as has also Cuvier (36) who suggests that they

not only prevent the reflux of water but the escape of food.

Galton ('71), describes the valves and their working in detail,

much as Dahlgren ('98) did later. Howes ('83), refers to them

in the trout, explaining their function.

In 1898 Dahlgren, who knew nothing of Galton's article,'

took up the subject in detail and describes the valves in the sun-

fish, Eupomotus gibbosus. The valves are sheets of membrane

composed of elastic connective tissue covered with a mucous

membrane continuous with that lining the mouth. They are sit-

uated in the oral cavity just caudad of the maxillary and mandib-

ular teeth. In this species of sunfish there is a median muscular

thickening. The valves “are placed with their edges pointing

downward and backward at an angle of less than forty-five

degrees to the axis of the body. This angle is increased to

about eighty degrees when the valves are struck by the regurgr-

tating stream of water." :

The function of the oral valves is to aid in the act of breathing,

which has hitherto been described as “a kind of swallowing."

Dahlgren says, “The respiratory stream enters the oral cavity

by the mouth and leaves. by the two gill openings." - - - -

! His attention has since been called to it by the present writer.

153

I54 THE AMERICAN NATURALIST. [Vor. XXXVIII.

“ [n inspiration the stream enters at the mouth in response to

a dilation of the oral cavity produced by the outward lateral

movement of the opercular frames.

* At the same time water is prevented from entering at the

gill openings by the brancheostegal valves which although they

are attached to the opercular frames, move independently of and

contrary to them ; so that, while this outward movement of the

frames extends the gill openings, the brancheostegal valves close

them automatically by the action of the water which tries to

enter.

“In inspiration the water is forced out of the gill openings by

a corresponding contraction of the oral cavity. At the same

time the water is prevented from regurgitating through the

mouth, not by the contraction of the latter, but by the automatic

operation of the maxillary and mandibular breathing valves which

move as accurately and efficiently as any of the heart’s valves.

Caught on their posterior edges by the first movement of regur-

gitation, they snap together and completely prevent the water

from leaving the oral cavity by the mouth which, meanwhile, is

left partly open, almost as much open as during inspiration.

“That these valves are of value as breathing organs is evident

upon casual observation ; that they are of much importance is

shown by the compensatory action, brought about by injury ; that

they are not of immediate vital importance is proved by the

fishes' ability to get along without their services until they are

repaired."

In examining about 70 species of fishes, the writer finds two

types of valves. These she has called the crescentic and the U-

shaped types. (Fig. 1.) There is a

f 2 : third form which may, however, be

y considered as a modification of either

type and will be described later.

Fic. 1.—Types of valves. C, crescentic; The crescentic type of valve 1s

nl tae oe fo typically shown in. the catfishes,

Silurida, particularly in Schilbeodes

punctatus. This type of valve, as its name indicates, is in the

form of a crescent with tapering ends, and may be broad or

narrow. That of Eupomotus gibbosus is also of this type. The

No. 446] ORAL BREATHING VALVES OF TELEOSTS. I55

free edge of the crescent is not necessarily a perfect curve

(Fig. 2, a), but may be

interrupted by a projection

or a notch. (Fig. 2, 2, c, d.) b

In the U-shaped type, E

the ends do not taper but

are bluntly truncate. (Fig.

1, 4.) The free margin in e

these also may be perfect

or interrupted.

A number of interesting P d

modifications occur. As fea 2

before mentioned, the free How e btched ; of betas pose 7 hag

edges are often interrupted NETSA

and in some cases, as in Pomolobus pseudoharengus and Pomo-

lobus estivalis, are attached near or at the meson, in one or two

lines, to the roof of the oral cavity. (Fig. 3.)

The central thickening which Dahlgren mentions may be a

tooth-like projection, may extend only halfway to the free margin

or back between the tooth pads, may be straight, wedge-shaped,

semi-circular (in this case the flat side was against the attach-

ment), or may be round, this form being either at the meson on

the attachment or in the

center of the valve. (Fig.

4.) In one case, that of

Ameiurus natalis, a hori-

zontal thickening passed

through the round cen-

trally situated, mesal thick-

ening. (Fig. 4, 7.) The

thickenings may be found

in either or both valves.

They appear to be muscu-

weal can ote w AA” dessin lar. Their function may

attachment; ô, esfivalis (parallel attachment); c, be to strengthen, possibly

imei (madian ir Hee ee (po. tighten, she valves, as

they are found on the

larger valve when the valves are of unequal size, and this valve

156 THE AMERICAN NATURALIST. [Vor. XXXVIII.

is the more subject to strain and injury from its size and posi-

tion. What is meant by “position” will be explained in the

miim

RUN Bae

c TEN

Fic. 4.— F f l thickenings. 4, tooth-pads.

paragraph on the relationship of the valves to the shape of the

mouth. :

In the majority of species examined, papillze are found on the

valves. These papillæ vary widely in size and arrangement.

They may be in almost regular rows, scat-

tered, cover the whole valve or only a part

of it, even, as in Cynoscion nebulosus, run

back among the teeth. They may be

coarsely setiform, flattened, rounded, on a

stalk or, in one case, of a circumvallate

/ FSN appearance. (Figs. 5 and 6.) In some cases

P es the papilla are fine and closely arranged in

M ud sie rows, giving the appearance of short vertical

a, in lines; pase: rows or folds. (Fig.5c. These are easily

rows te that the ae" distinguished from the true folds found on

Me 4 einer ot diall DNE valves. These papillæ at first appeared

thickenings or folds. ^ to be one elongated papillus but proved to be

as described. In Ammbloplytes gruniens are found rugas sur-

mounted by small papillz.

A third kind of valve has been mentioned as being a modifica-

tion of either the crescentic or U-type. The writer was for a

long time greatly perplexed because many of the Cyprinide

No. 446.] ORAL BREATHING VALVES OF TELEOSTS. I57

apparently had no mandibular valve, yet, from the shape of the

mouth, there was every reason to expect a small one. Finally

in a large specimen of Semotilus atromaculatus it was seen that

the place of the valve was taken by several rows of tall papillæ.

(Fig. 6, d.) This was found :

to be the condition in all the Aff 88 3p

minnows where the mandibu- d itt zO

lar valve seemed absent. o

The function of the papillæ 80° JSt

may be to aid in the finding ios d js

of food. The best develop- (6 Fom of papilla. a, setiform; 6, verti

ment of papillae seems to be cal section and surface view of the flattened

papille; c, wounded; d, stalked, e, circum-

among the bottom feeders, CIE

where such sense organs i

would be most useful. Another modification of the valves is in

the pigment which is frequently found in one or both valves.

The pigment spots may be regularly arranged or scattered like

the papillæ the arrangement varying with the species.

Some valves are further distinguished by ruga or folds or by

Fic. 7.— Valves of A. grumiens. a, maxillary; C, mandibular.

very thin and transparent margins or both as in the case of

Ambloplytes gruniens (Fig. 7).

As before stated, the valves are not ever of quite equal size.

Their size and shape are dependent on the size and shape of the

mouth. For instance, in the case of the Siluridz, (Fig. 8, a)

the mouth is terminal, practically horizontal and very wide.

For this reason it does not need to be opened far to admit the

requisite amount of water, therefore the valves are long and

crescentic but only of moderate depth, (Z. e. the distance from

attachment to margin). The mouth being horizontal, the wot

gitated stream strikes the two valves with about equal force ;

therefore, both must be well developed.

158 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In the Catostomidz, or suckers, the case is very different.

Here the mouth is inferior, (Fig. 7, c), and the snout depressed.

In this family the maxillary valve is well developed and very

deep. There are two reasons for this. First, the mouth of the

sucker is not wide and must be opened farther; second, the

lower valve is entirely absent. The reason for this is obvious.

The depressed snout decurves the regurgitated stream, which

strikes the upper valve only, hence a lower valve is not neces-

sary, the edge of the upper valve catching against the lower jaw.

Ye 8. qw anitiel action a E Rd incurrent and excurrent streams and popirien 3 of valves

n horizontal mouth. 2, similar section in superior mouth; c, in

telerior. mouth.

The position of the maxillary valve in these fishes is nearly hori-

zontal, this being a better position to catch the returning stream.

In the case of the minnows whose mandibular valve is replaced

by papilla, and in the case of some other fishes with maxillary

valves much larger than the mandibular valves, the mouth is

apparently horizontal or even slightly oblique.. In all such cases,

however, either the snout is depressed or the premaxillary is

protractile and is held protracted when the fish is breathing

quietly, thus depressing the snout and deflecting the greater

part of the water against the maxillary valve.

As may be inferred from the foregoing, the fishes with true

oblique mouths possess large mandibular and small maxillary

valves. The latter are never completely absent but may be vey

small. The case of Astroscopus y-græcum, the “stargazer,

might be supposed to form an exception to this rule, as the

mouth is really superior. From analogy it might be supposed

that the maxillary valve would be lacking. Such, however, is

not the case. There isa well developed maxillary valve (Fig.

3), like two crescentic valves joined at the meson by their ends.

No. 46] ORAL BREATHING VALVES OF TELEOSTS. 189

The reason for this structure is easily seen when the floor of the

oral cavity is examined.

This floor is raised in the center and hollowed at each side in

such manner as to divide the outgoing stream into two, and the

corresponding part of the roof is so shaped and hollowed as to

deflect the stream somewhat, so that notwithstanding the supe-

rior position of the mouth, the upper valve is necessary.

In Echensts naucratis, where the mouth is truly superior, there

is a small maxillary valve, whose greatest width is on the parts

each side of the meson about half way to the ends. This also

can be accounted for by the hollowing out of the roof of the

oral cavity.

In Hemirhamphus unifasciatus the mouth appears superior,

but it is more truly horizontal, the appearance being produced

by the great prolongation of the mandible. This species has

the maxillary valve but little smaller than the mandibular.

Pomolobus pseudoharengus affords a very characteristic example

of the development of the mandibular valve. The mouth being

the valves are crescentic, the

mandibular valve being four

times the width of the maxil- Te ndn onary bec cesta he a,

lary valve. The valves are

covered with coarse papille. They entirely close the mouth

when the jaws are at full stretch, which has been noted in no

very oblique and narrow, the

mandibular valve is very deep

and *baggy." (Fig. 8.)

In Chilomycterus | sche pfi

other species. 7

A good example of deep valves is found in Oncorhynchus

nerka, a salmon. In this fish the jaws are hooked, long and

somewhat compressed. If the valves were narrow, there would

be a long, free margin. Consequently the valves would have to

be very heavy to resist the outward pressure from within against

the unsupported margins. As it is, the free edges lie far back in

the mouth, their length is much reduced and the surface of the

valve, which is far better able to stand the strain, increased and

hung in a better position for resistance between the bones of the

jaws. (Fig. 9.)

160 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The valves of predacious fishes seem to be the heavier and

more strengthened by muscular thickenings. This may be

because the feeding habits of such fish would be liable to injure

the more delicate membranes.

The valves vary widely in different genera and in different

species of the same genus. The generic variation depends on

general shape, size and appearance; the specific on papillze,

muscular thickenings and pigment. The general type of a fam-

-muscle

Fic. 9.— Valves of O. nerka. a, maxillary; C, mandibular.

ily, notwithstanding, is very characteristic, except, perhaps in the

minnows, in some of which the papillæ form valve occurs. By

general type, the general appearance is meant; not the crescen-

tic or U-types only. Individual variation seems comparatively

slight and generally resulting from injury. If one individual

shows a certain arrangement of pigment, papillæ, and central

thickening, it will, so far as it has been possible to ascertain, be

found in the rest of the species and nowhere else. For con-

venience of comparison and explanation the writer has, where

enough species and specimens were available, arranged results

in synoptical form.

The writer desires to thank Dr. Burt G. Wilder and Dr. H.

D. Reed for valuable hints and use of specimens and, more

especially, for their kind interest and encouragement.

CORNELL UNIVERSITY.

No. 446] ORAL BREATHING VALVES OF TELEOSTS. 161

CATOSTOMID.

A. Papillz present.

B. Small papille in fold-like rows over whole valve

Erimyzon sucetta

BB. Papillz not in fold-like rows.

C. Papille on proximal half' of valve Catostomus macrocheilus

CC. Papillae over whole valve.

D. Papillz crowded over whole valve; distinct, rounded, not

so large as those on lips . Catostomus catostomus

DD. Papille less crowded in distal 2 half or in a circular

mesal spot.

E. Circular mesal spot where papilla are flattened,

small and well separated. . Catostomus nigricans

EE. No such spot. - Proximal papillae about size of

those on lips and in several rows. Distal papille

less crowded, flatter, rather elongate

Catostomus commersonit

AA. ipa: absent.

Small thickenings, perpendicular in direction, resembling long, low

papilla in appearance . . . . . Moxostoma macrolepidotum

SILURIDZE.

A. Papillz on both valves.

B. Thickenings triangular and in Mand. V.

Amiurus vulgaris

CC. Thickening inverted-triangular, apex toward tooth pads.

D. Pigment on thickening and spreading in a crescent

through the valve on either side . Noturus gyrinus

DD. No pigment in either valve . . . Noturus miurus

BB. Thickening in Max. V. and not trian

E. Thickening a vertical rod. Pcia o on | Mand. V. larger and

closer set than those on Max. V. . . Amiurus nebulosis

EE. Thickening not a vertical rod.

Thickening shaped like a sphere with a rod running

through it parallel to the free edge

Amiurus natalis (Fig. 4, 7.)

FF. Thickening spherical, barely touching attachment and

covering $ the width of the valve. Papillae so small

that the valves at first appear smooth, on proximal

$ of the Max. V. sparsely scattered on M

Ictalurus punctatus

! Proximal, — near or toward the attachment.

- * Distal, — away from the attachment.

162 THE AMERICAN NATURALIST. [Vor. XXXVIII.

AA. Papillz on Mand. V. only.

Central muscular thickening passing between tooth pads

Noturus flavus

CLUPEID.

A. Max. V.notattached mesaly . . . . . . . . Clupea harengus

AA. Max. V. attached mesally.

B. Attached on one perpendicular median line.

C. Mand. V.a crescent, wide; the distal two thirds thin and deli-

cate, no pigment . . . . Dorosoma cepedianum

CC. Mand. V. a horseshoe with Solid ends and the free mar-

gin rather squared off at the apex of the curve not

perfectly curved like the attachment. Valve much

wider at apical part of curve than on sides, (Fig. 2

Opistheoma oglinum

BB. Max. V. attached mesally on two lines. (Fig. a. 2.)

D. Attachment V-shape. (Fig. 3,a.) Clupea pseudoharengus

DD. Attachment in parallel lines, margin between ends of these

lines free. (Fig. 3%) . . . . . Clufea estivalis

PERCADÆ.

A. Thickening in Max. V., a three quarters circle flattened against attach-

ment and covering most of width of valve . . Perca americanum

AA. No thickening.

B. Papille and pigment. . . . . . . . . Hadropterus aspro

BB. No pigment.

C. Leo saei iw Ts DUK etai OH ANE EEERM

CQ. vo. u bos ae ola ps as Seca tU MERE

CENTRARCHIDZE.

A. Max. V. extending between tooth pads.

B. Papillz in perpendicular rows on either or both valves.

C. Rows ridge-like (or fold-like).

D. Ridge-like rows on both valves . Ambloplytes rupestris

DD. Ridgedike rows on Mand. V. only

Pigment spots on the papille . . Micropterus salmoides

CC. Papille in rows on Mand. V. but not close enough to give

ridged appearance. Slight thickenings in V. under the

rows of papilla. Clear margin . Ewfomotus gibbosus

BB. Papillz not in perpendicular rows.

E. Papillz numerous on one or both.valves. :

F. Papille very large, flat and crowded on both valves, g1v-

ing tesselated appearance, and extending

tooth pads . . Apomotus punctatus

No. 446] ORAL BREATHING VALVES, OF TELEOSTS. 163

F.F. Papille on Max. V. rather flat, medium size, crowded

but not giving the appearance of those in F ; a few

pigment spots at meson; mesal fold extending but

slightly between tooth pads.

Mand. V. covered with fine rounded papillz little

crowded, pigment in proximal half. . Æ. auritus

EE. Papille few.

A few large papilla on Mand. V. along attachment;

central thickening extends through tooth pad and

half way to margin; well pigmented. PapillIze

in Max. V. fine and well separated

Chenobrytes gulosts

GG. No papilla in Mand. V. Mand. V. well pigmented

at base, and meson quite black. Max V. well pig-

mented, especially between ‘tooth pads. Papillæ

few and small . . . . . . Pomoxis sparoides

AA. Max. V. not extending between tooth-pads.

H. Papilla in vertical, foldlike rows . . . . Apomotus pallidus

HH. Papillæ not in vertical, fold-like rows.

I. Papille of Mand. V. very few and flat, hard to see. Those

of Max. V. blunt setiform. Mand. V. well pigmented on

proximal half . . . Micropterus dolomieu

II. Max. V. with fine, ande, avil distributed papillæ and

: slight mesal thickening. Mand. V. with fewer and some-

what larger papille; a few pigment spots n base

meson extending through tooth-pads ae eae

ARGENTINIDA.

Valves crescentic ; no papilla; mandibular valve somewhat the larger.

A. Mesal thickening on mandibular valve circular in shape and placed

against attachment of valve, covering about 4 the width of the valve.

Central third of roin and ieu half of mandibular valve

pigmented . . . Osmerus mordax

AA. No mesal hiai

Pigment spots near attachment of mandibular valve, few and

valable >= 5 9 2. . . s Hyfomesus pretiosus

r THE AMERICAN NATURALIST. [Vor. XXXVIII.

BIBLIOGRAPHY.

CUVIER, G.

'86. Leçons d'anatomie compareè. Vol 1, p. 497.

DAHLGREN, U.

'98. The Oral Valves of Teleost Fishes. Zool. Bull, Vol. 2, pp:

118—124.

GALTON, J. C.

"71. How Fishes Breathe. Pop. Sci. Rev., Vol. 10, p. 431.

Howes, G. B.

'83. Zoology and Food Fishes. London. International Fisheries Lit-

erature, Vol. 2, pl. 2.

MACALLUM, A. B.

: The Anatomie of Amiurus. Proc. Canad. Inst. Vol. 2, No. 3, pp.

387-457.

OWEN, R.

'66. On the Anatomie of Vertebrates. Vol. 1, p. 413.

STANNIUS, H.

'S9. Symbolæ ad anatomiam piscium Rostock.

(Ao. 445 was mailed March 21, 1904.)

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THE

AMERICAN NATURALIST.

VoL. XXXVIII. March, 1904. No. 447.

BREEDING HABITS OF CRAYFISH.

E. A. ANDREWS.

Or the breeding habits of the European crayfish much is

known, but this is not the case with the American species. The

€asy assumption that the habits here were essentially identical

with those there, has been, possibly, one reason why so little has

been put on record regarding our American forms. That there

are, however, some considerable differences will appear from the

following notes made upon that common species Cambarus affinis

in 1894 and 1900-1903. Though these observations were made

upon individuals kept in confinement in the Biological Labora-

tory, they may, for the most part, be taken as a guide to what

is to be expected from field observations, which still remain much

to be desired.

Sex ratio.— The specimens used were taken from the Poto-

mac River in Maryland and when attention was given to the

ratio of the sexes it was found that one lot in March 15, 1901, |

contained 26 females and 14 males while another lot, April 20,

1903, contained 39 females and 14 males. While this shows a

marked predominance of females over males at those periods and

at that locality it is not general, for a lot of eighty of the same

Species taken in October, 1903, from a pond in Baltimore, con-

165

166 THE AMERICAN NATURALIST. [Vor. XXXVIII.

tained 41 females and 39 males.. Even where the females are

more abundant there is no probability of the eggs going without

fertilization since it was found that one male unites with several

females. :

Time of sexual union— When specimens were taken in

February and put into tanks some unions occurred at once while

others were deferred till March. When the sexes were kept

separate till March 6 and then put into one tank, unions took place

March 6, 7, 8, 9, 10 and 11. After that there were occasional

unions, especially of the younger and smaller animals, up to

April 2. Besides these observed unions in February, March

and April it would seem that there is an Autumnal pairing, in

place of or in addition to the Spring pairing. Several small

specimens reared from eggs laid in the Spring were found to

pair early in October of the same year; while at that same time

pairs of larger size were taken in a pond in Baltimore. More-

over specimens from Washington, D. C., in 1894, in November,

were seen to pair as soon as they were put into a shallow dish.

Description of sexual union— The union of the sexes in

C. affinis was first observed in 1894, and briefly described in the

American Naturalist in 1895. The same phenomena have been

repeatedly witnessed in successive years and there is no doubt

that in this species of crayfish the sperm is transferred from the

male to an external seminal receptacle upon the female where

it is stored up till the time that the eggs are laid. There is no

copulation or use of intromittent organs such as takes place in

the crabs; nor on the other hand is there any such vague

attachment of spermatophores as has -been described in the

European crayfish, Astacus.

In captivity the union lasts from two to ten hours and either

the male or the female may repeat the process with other

individuals.

When a female is put into a dish in which a male has been

kept till he is accustomed to it he soon seems aware of the

presence of the female and does not act as he does when only

males are introduced. The preliminary steps toward union are

soon taken. The male advances with ready claws and seizes

the female, sometimes gently. The female retreats or, when

No. 447.] BREEDING HABITS OF CRAYFISH. 167

seized, often struggles to escape. Despite these struggles the

male holding the female with one of his claws fastened to her

antenna or to any projecting part of her head eventually succeeds

in turning the female over to lie upon her dorsal surface : if

there is no struggle the same result follows more directly and

methodically. When the female is turned over the male stands

over her ventral surface and later transfers the sperm to the

receptacle.

When the first seizure is not effective in leading to a ready

turning of the female the male exhibits considerable skill and

shows something like intelligence in commanding the new

circumstances. Two cases were seen in which the male mounted

upon the back of the female and seized her claws as is usual

after turning the female over, though in this case turning had

not been accomplished. In this unusual position the male

attempted to adjust the sperm-transferring appendages to the

female and then desisted: then the long antenne of the male

were bent down strongly against the dorsal side of the thorax of

.the female. In one case the exopodites of the third maxilliped

were also used in feeling the female. After getting some infor-

mation, apparently, by the use of these sense organs, the male

proceeded to turn the female over and finally continued a normal

sperm transfer. In this turning over the male had to deal with

the problem of revolving the female through 180° while under

his body and starting with his right claw holding her right claw

and his left holding her left so that finally his right held her left

and his left her right. In accomplishing this feat the male first

removed his left claw from the left claws of the female and with

it seized her rostrum and adjacent head region : he then turned

the female about 9o? so that she lay upon her left side while he

stood over her right side. Next the right claw released the

female's right claws and seized her left claws. He was now able

to turn the female through 90° on to her dorsal surface. Then

he transferred his left claw from her rostrum to hold all her

three right claws. After that the usual union could take place

and ten minutes later sperm was transferred and union con-

tinued for some hours.

NM as soon as the male has thrown the female upon her

168 THE AMERICAN NATURALIST. (Vow. XXXVIII.

back he seizes all her clawed legs with his two large claws, hold-

ing the three left ones of the female with his one right claw and

the three right ones of the female with his left. He then moves

forward over the supine female to the position indicated in Fig.

1. From this figure it is evident that the two crayfish are accu-

Fig. 1.— Male holding female and with fifth leg supporting abdominal appendages that are

bout to transfer sperm to tl lus of the female. 4 diameter.

x $d

rately and closely adjusted to one another; not only does the

male hold all the claws of the female, but his abdomen is tightly

bent around that of the female which is closely coiled up under

the male. While all the five right legs of the female may be

seen, there appear to be but four legs of the male’s set of five

left legs. The base only of the male's fifth leg is shown; poste-

rior to this are the peculiar long male pleopods, or appendages

of the first abdominal segment and of the second abdominal seg-

ment. These four appendages are elevated at an angle of about

45° and point toward the ventral surface of the thorax of the

female, forward and downward. A view of the right side of the

pair would, however, show the left fifth leg of the male project-

ing outward and backward between the fourth and fifth legs of

that right side. ;

The explanation of this peculiar arrangement is found in a

habit of the male which seems necessary for the accomplish-

ment of sperm transfer and is a very instructive example of

mechanical adjustment amongst. several rigid calcified organs.

After the male has come forward over the supine female there

is a period of ten to twenty minutes of apparent inaction before

No. 447.] BREEDING HABITS OF CRAYFISH. 169

the next move which is as follows: the male rises up away from

the female, but still holding all her claws, and deliberately passes

one of his fifth legs across under his body so that it projects

from the other side. "When the male again settles down against

the female it is found that the pleopods have the position shown

in the figure, whereas before this move they had the normal

position, being directed forward, horizontally under the thorax

of the male.

It is this forced and remarkable position of the fifth leg which

secures the necessary elevation of the male pleopods. These

pleopods might be compared to the blade of a pen knife half

open and tending to shut up into the handle when pressure is

exerted against the tip: and as such a blade might be held in

position by a pencil placed across between the blade and the

handle and held there, so the pleopods are held in position by

the crossed leg which lies anterior to them and between them

and the thorax of the male. All these parts are firm and rigid

and the pleopods articulate only where they hinge to the

abdomen. As the male draws himself down with force against

the female the pleopods are so held by the above device, that

their tips enter the annulus of the female and the pressure so

exerted would tend to shut them down into their resting posi-

tion, but this is opposed by the fifth leg which blocks the

pleopods firmly. That there is force exerted by the tips of

the pleopods against the annulus was shown in the case of a

female that had been reared from the egg in the Laboratory

and when put with a male in April of her second year was

covered by a dark blackish deposit after wintering. It was

soon found that in union with the male the edges of the opening

of the annulus had been scraped clean of the dark deposit and

stood out conspicuously against the rest of the dark exo-skeleton.

Though the male always uses one of his fifth legs as an

apparently necessary secondary sexual organ it is not always the

h right and others with

right or the left ; males were seen wit

left legs so used. Whether the use of right or left is fixed for

each male by circumstances OF whether it is a matter of chance

or whether inherent in the structure of each male was not

determined, but a student who kept many crayfish and sought to

170 THE AMERICAN NATURALIST. [Vor. XXXVIII.

determine this point reported to me that one male was seen to

use the same leg on one side, in several successive unions with

different females.

In one case the fifth leg was seen to project between the third

and fourth legs of the opposite side instead of between the

fourth and fifth as is the rule. Possibly this may have been in

connection with some difference in size between the female and

male. , Though the two pairing are about the same size there is

often considerable difference in length and to secure accurate

interadjustment of such rigid bodies with so many protuberances

and pairs of appendages is no small problem. To solve it the male

at times relaxes his abdomen and moves forward over the female

and when finally the tips of the pleopods have been introduced

into the annulus the male again envelopes the end of the

abdomen of the female and firmly holds it as in the hollow of a

hand. The persistent union of the two is made more complete

by the use of the hooks found on the legs of the males of this

and some other species of Cambarus ; apparently these fasten

the male so that the abdominal pressure exerted down the pleo-

pods against the annulus does not react and push the male

upward away from the female. In C. affinis there is one of

these hooks or spines on the third segment of each third

walking leg and the male fastens these two hooks into the base

of the fourth legs of the female. In the above figures the male

has raised the third left leg so that the hook is free and far from

its proper socket. When lowered into place the hook depresses

the soft membrane that forms the external aspect of the basal

segment and thus a temporary socket is formed. The outer

ventral edge of this socket is bounded by the stiff calcified ridges

that help form the hinge between the first and second segments

and it is against this rigid rim that the spine of the male's leg

catches. The use of these male hooks as secondary sexual

organs is thus established and we may expect to find that in

those species of cambarus in which more than one pair of legs

are provided with such hooks, that there will be a corresponding

increase'in complexity of the adjustments of sexual union.

The union of the male and female is now so firm that they

cannot be readily separated, and if thrown into actively boiling

No. 447] BREEDING HABITS OF CRAYFISH. 171

water the two may be fixed in almost normal positions and serve

to make most excellent museum preparations.’

Bound together in this way the transfer of sperm from male

to female takes place during several hours. Since the crayfish

may now be roughly handled or transferred from one dish to

another there is little difficulty in observing how the sperm-

transfer is effected and a lens may be used without causing the

union to cease. The terminal part of the vas deferens of the

male, on each side, is found in this period of union to protrude

horizontally into the water from the base of each fifth leg as a

short, soft, bent tube of translucent appearance. This organ fits

exactly into the^beginning of the long groove that passes down

the first male pleopod, right or left. These pleopods are in fact

massive, calcified and rigid tools, each with a deep sinuous groove

along it that is seen to lead the sperm from the above ending of

the vas deferens down to the tip of the pleopod. The special-

ized, sharp tips of these organs are inserted into the annulus.

The sperm seen to issue from the vas deferens tube glides down

the groove of the pleopod to the annulus in the form of long,

macaroni-like cords. Microscopic examination of these cords

reveals a central axis of real sperm made up almost entirely of

the sperm cells and an outer tubular case comparable to soft

macaroni. During this transfer of sperm the tip of each pleopod

of the second pair is closely applied to the pleopod anterior to it,

but no observation was made upon the mode of action of these

second pleopods. Each of them has terminal filaments that may

well be sensory and also a peculiar, soft, somewhat triangular

* spoon " or scoop that fits nicely against the groove of the first

pleopod. It would seem that the second pleopods may act to

convey sensory impressions and to protect and guide the sperm

masses which were found not to go astray but to be in some

way retained in the grooves of the first pleopods and forced on

into the annulus, probably with the guidance and direction of the

Second pleopods.

Probably both sides of the body in the male are active at the

| The same method may be used to advantage for museum specimens of females

etc., lessens the solubility of

“in berry" etc., since the coagulation of the liver,

substances that ordinarily discolor the alcohol. ;

172 THE AMERICAN NATURALIST. (VoL. XXXVIII.

same time in transferring. sperm, but this was not directly

observed. During the whole process of union the male is in a

state of excitement while the female is quite the reverse, as far

as could be judged. The action of the male in turning and

adjusting the female is greatly assisted by the state of passivity

simulating death that overtakes her soon after being seized by

the male. This inertia of the female extends even to the

respiratory movements, which seem absent in strong contrast to

the condition in the male. The female seems to be dead and

the only signs of the continuance of life that were seen were the

movements of the eye-stalks in cases where the efforts of the

male led to his claws coming against the eyes and, at times,

a slight convulsive tremor in the abdomen, possibly connected

with sperm transfer. The very small first pair of abdominal

appendages which lie often against the annulus and have no

probable use unless it be in connection with the phenomena of

union, may convey sensory stimuli and occasion the above

abdominal contractions. On the other hand the male is in a

strongly excited state during the entire period of union: when

the struggle and turning have ceased there are still quick vibra-

tions of the anterior maxillipeds and strong currents of water

thrown out from the gill chamber as well as the long continued

contractions of the limb and abdominal muscles and probably

those of the internal male organs. The process terminates

when the male moving backward and rising up crosses his fifth

leg back again under him into its own side. He then releases

the female.

The annulus.— 'Thus the result of union of male and female

is the storing up of sperm within the annulus or sperm receptacle

of the female. This is in strong contrast to what has been

described as taking place in the European crayfish, Astacus,

which has no annulus; for French observers state that the

male distributes sperm masses, or spermatophores, over large

areas of the under side of the female. On the other hand in

the American lobster a transfer like that in Cambarus doubtless

takes place since Bumpus discovered the sperm-containing

receptacle upon the female.

The structure which serves in the lobster to hold sperm is,

No. 447.] BREEDING HABITS OF CRAYFLSH. 173

however, apparently not homologous with the annulus of Cam-

barus but is different in position and construction. Whether

there may not be some kind of receptacle in Astacus, as. some

appearances there would suggest, something more like the

receptacle of the lobster, is a possibility that needs future con-

sideration. It is well known that the annulus differs in many

species of Cambarus, and now that we know its use as a sperm

receiver we may expect to find some of these specific differences

have their uses in the processes of union. The male pleopods

are also characteristically and often remarkably different in

various species of Cambarus, and these differences may find their

explanations in their uses with different styles of annuli.

In C. affinis the annulus may be described as a calcified region

on the ventral side of the thorax between the sterna of the

somites bearing the fourth and the fifth legs. The sternum of

the former somite is a wide plate, concave across the middle line

and rising up at its edges right and left as two high plates that

diverge posteriorly and stand close against the bases of the legs.

The sternum of the latter somite is a transverse band enlarging

at its ends against the right and left fifth legs and bearing at

its middle a transversely elongated rounded elevation. All these

parts are hard and calcified. The annulus fills all the space

between the above sternal plates and thus lies across the ventral

line at the interval between the fourth and the fifth legs. It is

close against the sternum anterior to it and may be moved

slightly as if hung to it by a stiff hinge. id

In shape the annulus, Fig. 2, is CUM

a transverse and elongated ellipse

with pointed ends right and left.

It is a calcified plate with two ele-

vations, or hills, of varying size ees

and shape near the middle—one "7 m pus gloat veg d

right, the other left. Between

these elevations is a longitudinal groove and at the bottom of the

posterior part of this groove is a narrow chink into which a fine

instrument may be forced. This chink opens posteriorly into a

deep groove or valley that runs from right to left across the

major part of the annulus. This big groove is just posterior to

174 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the long axis of the annulus and anteriorly it is overhung, some-

what, by the two hills, while posteriorly it is bounded by an

elevated lip or transverse ridge that forms the posterior edge of

the annulus and sometimes shows a faint crack in it as if made

of right and left hàlves welded together.

Though the hills are right and left the median slope of one

extends across the median line of the body of the crayfish so

that the chink between the hills is always asymmetrically placed,

sometimes on the right and sometimes on the left. In forty-one

females, all but two of which were young, probably but four

months old, examined in October, only three cases were found

in which the chink was on the left side; in the thirty-eight

other females it was upon the right side.

In four still younger females, 32-40 mm. long, the annulus

was found less well developed and with but slight transverse

groove.

Sections of this organ show that the chink opens into a small

rounded pouch or sac which as seen from a dorsal view projects

upward as a small, curved ridge. Its walls are stiffly calcified

cuticle and no opening could be found excepting the external

chink above mentioned. This cavity of the annulus serves as

the seminal receptacle, but it is only a small, specialized pit in

the external cuticle or exo-skeleton.

After sexual union the female shows a white plug of waxy

substance projecting from the chink and filling up the transverse

groove at that point as indicated in Fig. 2. This plug is

necessarily excentric, generally upon the right side, but in some

females upon the left side. It is the surplus sperm envelope or

macaroni-like case that came down the pleopod groove and was

forced into the chink of the annulus. This plug may remain for

weeks, but it disappears some time after the eggs are laid. It

may thus be used as evidence of foregoing union for a much

greater time than the vaginal plug of some rodents can be.

Examining the contents of the cavity of the annulus when the

plug is in evidence we find it full of a similar compact paste-like

material that may be quite hard and has the form of a tubular

sheath around a granular mass that proves to be the peculiar

sperm cells of the crayfish. The sperm in this sperm receptacle

No. 447] | BREEDING HABITS OF CRAYFISH. 175

is in very small amount as compared with the amount produced

by one male or with the great mass of eggs to be fertilized, but

the actual number of sperm cells is quite large and that they are

numerous enough to fertilize all the eggs seems certain though

not actually demonstrated. As a fact when females were iso-

lated as soon as union had taken place and kept apart from

males till eggs were laid these eggs developed as if fertilized.

No sperm was ever seen to be deposited by the male upon

other parts of the female than the annulus and if it were not

protected as in the annulus it is difficult to see how it could long

survive exposure to water which quickly destroys the sperm

cells. The laying of eggs, however, may not occur for some

weeks after union.

Again, in one case in which the annulus was removed from

a female just before laying, the eggs were deposited with normal

secretions and habits and though the female was left undisturbed

for five days the eggs then appeared shrunken and abnormal, to

the naked eye. Twenty-four days later there was only a mass

of mouldy dead eggs left on the female. Though there were

some cases in which normal females lost most of their eggs

by death and fungus yet the above eggs were thought to be all

bad from an early date and,it is probable they were not fertilized,

in the absence of the annulus.

On the whole the evidence seems strongly to favor the view

that the sperm received into the annulus in union is the sperm

which later fertilizes the eggs. In fact it is possible that spérm

taken in antumnal unions may be used for the eggs laid the fol-

lowing spring. Experiments to test this are not yet completed.

Mortality. — The crayfish kept in the laboratory were not

fed till after they had laid eggs; then they ate meat, raw and

cooked? raw hen's eggs, and pieces of earth worms, as well as

Chara and Hydrodictyon. After sexual union many died and

it was found that the males died in larger numbers than did the

females: thus in one lot seventeen died within two weeks of

union in March, and thirteen of those were males and only four

females. In several cases the males died within a few hours

after union. *

Date of laying. — After union there is quite a long petiod

I 76 THE AMERICAN NATURALIST. (Vor. XXXVIII.

before the eggs are laid : this period is often some weeks. When

about roo males and females were put together March 6-11

the first eggs were laid March 24th, while other females laid

from then on to April 15th, though those laid after the first

of April were probably unduly retarded by various artificial

influences. That March is the normal time of laying for this

species and at the Maryland station on the Potomac whence these

specimens came is shown by the fact that 39 females taken there

April 20th, 1903, all bore eggs in late stages of development that

were almost the same as the various stages then present in the

eggs laid in March by the previously captured specimens.

Preparation for laying. — 'The females tend to secrete them-

Fic. 3.— Female standing lil Sete and ci : td ith ll claws. x $.

selves in dark corners and four or five days before laying they

are noticeably excitable, an approaching object causes them to

raise their claws and to assume the defensive in a much more

active way than was previously the case.

These days are also taken up with great and peculiar activity

upon the part of the female resulting in a very thorough cleaning

of the ventral side of the abdomen. No matter how darkly dis-

colored the exo-skeleton may have become during the winter

it is now made white and clean over the under side which then :

comes to contrast strongly with the remaining dark exterior.

No. 447] | BREEDING HABITS OF CRAYFISH. 177

The process of cleaning was observed in a number of cases

and always ran the same course. Though the female is now

so alert that it is difficult to catch her in the act of cleaning

yet the attitude assumed is most noticeable when seen. As

indicated in Fig. 3 the body is raised high up from the usual

crouching, crawling position and stands like a tripod supported

above the bottom of the aquarium upon the two outspread large

claws in front and the oddly down-bent abdomen behind. The

other legs aid but little in support of the body and are concerned

with the cleaning of the abdomen. The fifth legs, and at times

the second and third also, were seen to be thrust back under

the abdomen and there carefully and patiently used to remove

ie. Z A

F ig a dl oth a ius f p

FiG. 4.— Comb and picks on end of fifth leg; used by female to clean abdomen before

ing. x .

all the dirt from the entire under surface including the pleopods.

Even the numerous long, plumose hairs on the pleopods lose their .

covering of dark * dirt," and the transformation wrought in the

appearance of the whole under side of the abdomen is so great

that one would suppose the crayfish had cast its shell and wore

an entirely new one.

The ends of the fifth legs are shoved against the pleopods

and other parts of the abdomen with considerable force but it is

only slowly that they accomplish perceptible cleaning. On

examining the tips of these legs we find they seem especially

well fit for such cleaning work. As indicated in Fig. 4 the

terminal segment is like a strong comb, as it has a series of spines

178 THE AMERICAN NATURALIST. [Vor. XXXVIII.

along one free edge. At one of the angles of the penultimate

segment there is a group of strong picks that look like the

horny tips at the ends of legs. In fact these two picks are born

upon a truncated process that suggests a homology with the

finger of the chelate legs, that process of the penultimate

segment which apposes the last segment to form the claw;

and we might regard this fifth leg as having lost, or not yet

acquired, a perfect claw. The penultimate segment also bears

beautifully serrated hairs,as indicated in the figure, recalling the

appearance of some setze of annelids. That these tufts of ser-

rated hairs may serve as scouring brushes seems not improbable.

We would then have a double pick to loosen dirt, a stiff comb

and a brush with saw-tooth hairs to accomplish the cleaning.

The fourth leg has the same structure and though it was not

seen in use it may well take part in cleaning. The second and

third legs bear well developed claws and these were seen slowly

plucking at the encrusted abdominal surface. As elsewhere

shown! these claws are well fitted for cleaning as they can pick

off objects and are provided with a long row of flat, serrated

plates on each limb of the forceps that should serve excellently

in cleaning the pleopods.

Time of laying. — In confinement the crayfish deposited their

eggs at night time with few exceptions; only one among thirty-

two laid eggs in the day time. Yet it was found possible to force

the laying of eggs in the day time by keeping the female every

night in barely enough running water to moisten the ventral

surface of the body and with no opportunity for the normal

submergence. When returned to deeper water in the day time

eggs were finally laid, at noon, upon the third day of such

treatment.

Laying. — The females were easily disturbed and never laid

when under observation so that the actual emission of eggs was

not seen but the following facts were observed just before

and just after the actual laying. A short time before laying

the female is sometimes found lying upon her back waving

the abdominal appendages back and forth in a rhythm of about

1 Biol. Bull. Jan. 1904.

No. 447.] BREEDING HABITS OF CRAYFISH. 179

one second. The endopodites of the third maxillipeds and

the anterior three pairs of legs are sometimes swung back

and forth also. Finally a peculiar secretion is passed forth

and the eggs are then laid. This secretion is furnished by the

* cement glands" of the under side of the female and needs

special notice. One of the things that make much for the

clean appearance of the under side of the abdomen when the

female is about to lay is the presence of large milk-white

areas on the basal parts of both endopodites and exopodites

of the sixth pair of abdominal appendages, forming very con-

spicuous white patches when the tail fan is expanded. The

other pairs of pleopods are pretty uniformly milk white but the

endopodites are more densely white and the glands in their

terminal parts are somewhat segmentally arranged. The

sternal plates between all six pairs of pleopods also stand out

as milk white areas. Anterior to the abdomen the only milky

gland areas are the sternal plate of the last somite, the annulus

and the edges of the two flaring sternal plates anterior to the

annulus. At other times of year these “cement gland " areas

seem inactive or at all events inconspicuous. When a portion

of one of the milk areas was removed from the tail fan

or from any of the smaller pleopods it furnished, under pres-

sure, a milky material which swelled up in water as a clear

jelly containing minute spherules as seen under Zeiss 2. D.

and as a somewhat milky glair as seen with the naked eye.

When first pressed out from a piece of the glandular area the

secretion also contains the minute spherules or droplets.

In one case a female was seen to stand with the body

raised high above the bottom of the tank and to wave the

pleopods back and forth while they gradually became covered

with a clear slime or glair. Forty-five minutes later, at 1.15 p.m.

the female was lying upon her back and all the eggs had

passed out of the oviduct. The general appearance of this

female that had just laid is indicated in Fig. 5. Lying upon

the back with the limbs stiffly extended and no motion visible

the creature seems dead unless the strongly bent abdomen

suggest muscular contraction. Passing forward from the

widely expanded tail fan is a faint film of slime or glair that

180 THE AMERICAN NATURALIST. [Vor. XXXVIII.

extends to the second legs and neighboring parts. Under this

veil a few eggs may be seen not far from the oviduct but

the great mass of eggs, several hundred, is concealed by the

bent abdomen which encloses them somewhat as a nearly

closed hand might a quantity of shot. The actual openings

of the oviduct are concealed since the abdomen is bent to its

utmost and the tailfan is carried very far forward over the

ventral surface of the thorax. Some slight tremors of globules

in the glair near the openings of the oviduct suggested rhythmic

contraction of the oviduct, about once a second, even after the

‘AI IOS oh

A TAN v | n

^S A E et INS

Fic. 5.— Female lying on back with legs held rigid and apron over eggs as they are being

aid. x 4.

eggs had been laid. Gradually the abdomen relaxed somewhat

and allowed more of the ventral surface to be seen ; while most

of the eggs lay in a mass enveloped by the abdomen, some

of them still extended forward on to the thorax as a flat layer.

After ten minutes there were some slight movements of the legs

and then of the second maxillipeds also. Six minutes later the

legs moved more actively and a minute later the crayfish turned

itself over on to its ventral side, though this was not easily done

in a smooth glass dish.

No. 447.) BREEDING HABITS OF CRAYFISH. 181

The eggs must have passed out of the oviducts in a short

space of time and have been received into the basket formed by

the bent up abdomen, a basket full of glair that would protect

the eggs from contact with the water. That this glairy sub-

stance is the secretion of milk-white cement gland areas seems

certain; when the material about the eggs is examined under

the microscope it contains droplets like those found in the

secretion of the cement gland.

The probable mode of laying may be inferred from the above

observations and from the following considerations. If we place

a female crayfish upon her back and bend the abdomen forward

over the thorax as far as possible we can see that if the eggs

were forced out by the contractions of the oviducts they

would issue in two streams from the mouths of the oviducts

which are on the bases of the antepenultimate legs : they would

then emerge into a median trianglelike depression formed by

the thoracic sterna and this would form an inclined plane down

which the eggs would flow into the basket formed by the

abdomen, which is on a lower level. When first seen the eggs

were very soft, apparently liquid and most easily deformed and

indented yet coming rapidly back to a spheroidal shape, owing,

partly at least, to the presence of a thin membrane. At first

the eggs were not spherical but pear-shaped or elongated ; but

when put into hardening liquids they took on a spherical form.

When the female is lifted out of the water the soft eggs crowd

together and have polyhedral shapes as if liquid, or plastic. In

the position assumed by the female it would seem that gravita-

tion acting upon the liquid eggs would bring them into the

abdominal basket.

The female that has laid continues to hold the abdomen flexed

and the eggs are contained in the basket of glair for some hours.

As seen from the side such a female, Fig. 6, seems to have

an apron of glair stretching from the second legs and that part

of the thorax back to the expanded tail fan and somewhat

bellied downward as time goes on. In this condition a remark-

able rhythmic habit was observed in several females and

regarded as a necessary element in the future success of the

eggs. This performance lasts several hours and may be spoken

B as : —

182 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Turning.— It consists in a long series of changes in position

from the right to the left : the female lies as if dead now on one

side and now on the other. After lying a few minutes upon the

right the female got back into the ventral position and in a min-

ute or so turned on to the left side, remained there a few minutes

and then went back to the ventral position for a minute or two,

then to the right sideand so on. The striking features in this

set of habits are the inert state assumed while lying upon the

sides and the great regularity and persistence of the alternating

rhythm. The following example will show the details of this

process as observed in female XX XII, April 15, 1903, from 1.35—

Fic. 6.— Female standing up after laying: apron connecting thorax and bent abdomen.

X 2.

5.50 p.m. As shown in the table this female, as soon as the

supine egg-laying position was given up, remained upon her

ventral side for three minutes, as in Fig. 6, then turned and

lay upon the right side for two minutes, then upon the left side

for six minutes, then upon the ventral side for one minute, upon

the right side for three minutes and so on.

V3, R2, Vo, L6, V, R4 Vr l4 V Reve h

VI, NL OVE Lag Va R3, V4, la Vi R3 Vi TS

, R$, Vt, L4 Vii, Rab Vt, L6, Vip R$

Vh RO, Vo? Ls, Vii a L4 Vo RLQ Vb DM

V$! R4j, V4, L4, Vi, “Rad, Vr, L6p Ved, Re; Vi, bol

Vig, R63, Vi, L63,V1, Rž, Vi, Ls, V2, Rz, Và L5i

V2, R8, Vi, Lob, Vr, Roe Vi La Và Ru. V

No. 447.) | BREEDING HABITS OF CRAYFISH. 183

Observations were discontinued with the animal upon the ven-

tralside at 5.50 p. m. In this long series there is only one break

in the rhythm at the point marked x when after 51 minutes of

turning in which there were ten alternations from right to left ;

the female then walked about the dish as if seeking some corner

more retired than the one she finally returned to and remained

in to make the following ten alternations. This walking away

from the corner otherwise so closely kept accounts for the long

interval of eleven minutes on the ventral side in place of the

usual one of only one or two minutes ; to it also may be due the

break in the close sequence, the turning upon the left side when

the right was the one to be expected ; unless indeed we assume

the time factor to be so dominant as to cause the female to go

over to the left side after eleven minutes because that would

have been time enough for, say, a ventral rest of 2 minutes, a

right rest of 74 and a left of 14 which would bring the period

around again to a left rest as next in order. At the point

marked + the ventral time is abnormally lengthened as it

includes the minute and a half spent by the female in struggling

to get her ventral position after accidentally rolling over upon

her back from the position on the right.

The result attained by this performance would seem to be the

proper fastening of the eggs to the pleopods, without which the

eggs would not develop. During this process the female stays

-in one corner of the dish and except when turning from one

position to another lies so still that it might be taken for dead ;

only rarely were any of the claws reached near to the abdomen

and there is no evidence of any manipulation to secure the

attachment of the eggs within the abdominal basket. It is well

known that ultimately each egg is firmly tied to the hairs of the

pleopods (a few also to some sternal hairs) by a strong string

continuous with a tough membrane that envelops the egg as a

sort of second shell. These strings and cases are gradually

formed, and from some of the cement gland secretions no doubt.

For twelve hours or more after the eggs are laid they may be

easily pulled away from the pleopods as they are stuck only by

a soft glair, but later this glair hardens and force is needed to

break the strings away from the pleopod hairs. in onc puse

184 THE AMERICAN NATURALIST. [Vor. XXXVIII.

when the glair, with many eggs, was taken away from the

abdomen soon after laying, new glair was formed within a few

minutes. When the egg is freshly laid it is covered by a soft

slime that may be squeezed off by rolling it under the cover

glass; this material looks like the glandular products above

described. The abdominal basket seems full of such material

and the eggs sinking down through it are seen in masses on

right or left sides according to the position of the female. The

pleopods hanging down into this mass are probably pouring out

over their hairs more and more of the glandular secretion which

will finally harden. By turning the female must let all the eggs

fall against now the right and now the left pleopods and again

when standing, ventral side down, allow the eggs to dangle down

in their coatings of slime till strings would, probably, be pulled

out above them attaching each to some part of the pleopod.

As there are four to five hundred eggs laid at one time,

in one case 631 eggs by count, it is no light problem to get

every one well fastened by its own stalk to the abdominal organs

that later serve for their protection and aération.

Any escape and loss of eggs during the turning action is pre-

vented not only by the abdominal basket of slime with its special

apron or surface in contact with the water but by the position

assumed by the female; when upon her side the legs under the

thorax raise it so that gravity would tend to hold the eggs in the

abdominal basket; when upon the ventral side the same is:

generally true since the abdomen is carried lower and the thorax

raised. :

After the turning has ceased the glair apron is still in evidence

and becoming more conspicuous from discoloration and accumu-

lation of dirt. But after some hours it becomes broken by

movements of the pleopods and by straightening of the abdomen

and is gradually removed. The claws often have masses of glair

upon them and perhaps they assist in removing the superfluous

mass. Shreds of glair remain attached to the edges of the tail-

fan as late as thirty-six hours after the eggs are laid.

Care of eggs. — Nery few eggs fail to become fastened to the

pleopods in the above process of turning and henceforth the

female bears them so fastened till they hatch. When the eggs -

No. 447] BREEDING HABITS OF CRAYFISH. 185

were removed and put into dishes of running water they died,

except when taken in very late stages with the embryos well

formed. The female takes a certain amount of care of the eggs

"which seems to greatly increase their chances of hatching ; as

a rule most of the eggs hatch, but in several cases the eggs upon

the abdomen became covered by a fungus which bound them all

together into one dead, discolored mass. Still the female bore

them till long after they should have hatched. That this fungus

attacks the eggs in the open was shown by the fact that one

of the 39 females taken ** in berry " was found to have moulded

eggs when received. The female after laying backs into the

darkest, most protected corner available and for a long time keeps

the abdomen more or less bent down under the eggs which are

then protected from dirt ; but at times the abdomen is straight-

ened out and the eggs hanging like bunches of grapes from the

pleopods are moved back and forth in a manner well calculated

to keep them clean and to insure better aération, Fig. 7..

Fic. 7.— Female aérating eggs by raising and straightening abdomen and waving pleopods

back and forth. x $.

Sometimes also the female may be seen reaching back among

the eggs with her smaller claws as if to examine or to clean

them. Some of the females died before the young hatched out

and this was more often the case amongst the females that had

laid in the open and not in the laboratory. If the eggs were

left upon the dead female they became overgrown by mould and

died; but if taken a week before hatching and put into

a McDonald fish-jar they hatched successfully in considerable

numbers.

Fertilization. — As most of the eggs hatch and as sections

of freshly laid eggs show sperm nuclei there seems no doubt

186 THE AMERICAN NATURALIST. [Vor. XXXVIII.

that the eggs are fertilized and probably the sperm in the

annulus is used for that purpose. But, unfortunately, the mode

of fertilization or even its actual occurrence has not been made

out. The sperm plug may remain visible for a few days after

laying but it then disappears. Its appearance also is changed,

its end broken, after laying. Examination of the contents

of the annulus after laying showed very few sperms. As the

eggs are laid they probably pass over the annulus and a relatively

small amount of sperm might fertilize all of them if it came out

of the sperm plug at the right time. Before laying the annulus

is covered with glair and possibly this may act to bring the

sperms out as well as to protect them from the water, which

produced marked and apparently destructive changes in the

sperms. Some osmotic factor may here be concerned in bring-

ing out the sperm. Other means of getting the sperm out from

the waxy tube that we have called the sperm plug and from the

interior of the annulus might be: the pressure that the sternal

plate between the fifth legs may exert upon the annulus when the

legs are forced forward, as sometimes seemed to. be the case

about the time of laying ; or some activity of the small pleopods

of the first abdominal somite. The former action, however,

seems only to force the annulus to face more vertically in place

of- horizontally while the necessity of the first pleopods as

instruments in fertilization was disproved by cutting them off

from a female about to lay and finding that the eggs under-

- went normal cleavage as if fertilized.

In support of the view that the sperm issue from the annulus

at the time of laying it was found that if the annulus was then

removed the female seemed not to be inconvenienced, going

through turning movements as usual, but the eggs did not

develop. This may, however, have been due to the fact that

the female was lifted out of the water, the glair disturbed and

thus both eggs, and presumably sperm, exposed to the water,

which may have prevented fertilization even if the sperm were

upon the eggs.

No sperm were found upon the eggs nor upon the annulus :

when sperm are taken from the male or from the annulus

they undergo changes of form which might make them- less

No. 447] BREEDING HABITS OF CRAYFISH. 187

recognizable upon the egg and moreover the egg is 250 times as

thick as the sperm and quite opaque.

The whole process of sperm and egg-meeting and union is

much in need of elucidation.

Sperm. — While the pairing habits of Cambarus are more

complex than those of Astacus the sperm cells are not as com-

plex in form as those described for Astacus by Herrmann.

In place of the many radiating arms he figures C. affinis has six,

as a rule, but sometimes 5, 7 or 8. This is also true

of C. bartoni. The remarkable bowl shaped vesicle of the

crayfish sperm does not have as complex a shape in either

of these American forms as it has in Astacus. The diameter

of the body is 8 » while the arms extend out four times that

distance on every side so that the entire spread of arms is

over 70 »; but only the refractive vesicle is conspicuous and

its longest diameter is less than that of the body.

Period of development, — Attached to the pleopods and

presumably fertilized the eggs go through the developmental

changes that have been described for Astacus by Reichenbach

and others. As these specimens of C. affinis were kept in con-

finement the times of various phenomena could be determined

with some precision and in general the eggs of any female

develop at about the same rate while the eggs of different

females progress differently. Eggs laid near the end of March,

1894, and in 1900, hatched late in May: eggs laid April rith,

1901, hatched the end of May. In 1903 eggs were laid from

March 23d to April 15th; when the eggs were kept in water

at 123? to 14? c some required eight full weeks to hatch ; others

in a warmer room where the water was not constantly running

hatched in six weeks, and some in just five weeks. Most were

hatching about May 18th and this was true also of eggs on the

females taken April 20th and then in late cleavage or even

embryonic stages; so that it seems probable that late in May

is a natural time for Music in the open where these crayfish

were taken.

Cleavage. — The nuclear multiplications and migrations

of “cleavage” take place but slowly in this heavily yolk laden

egg. From sections it seems that the sperm and egg nuclei may

188 THE AMERICAN NATURALIST. [Vor. XXXVIII.

be not far from the surface 12 hours after the eggs are laid,

while 24 hours after laying the cleavage nuclei are near the

centre of the egg and only about ten in number.

There are only a few cleavage nuclei half way out to the

surface 48 hours after laying, while 72 hours shows a large number

of nuclei near the surface.

In the live eggs some of the cleavage phenomena may be seen

with a pocket lens and are much as described by Herrick for

C. immunis. At about 72 hours there are rounded areas scat-

tered all over the egg, shimmering through the outer part of the

egg; each is about 15 y in diameter and some 30 of them may

be seen upon any face of the egg. When such eggs are crushed

under the microscope the dense, clear nuclear areas may be

found amongst the yolk spherules and also long bundles of fibres

connecting the chromatin bodies. i

Eggs taken at earlier times from different females differed

and also upon the same female. At about 48 hours a few

clear areas might be seen in a group deep within the sub-

stance of the egg; some eggs showed 5 or 6, some 30-40,

some none at all. Later there were similar areas nearer the

surface but upon one side of the egg only. Soon these areas

spread so that but part of the surface of the eggs failed to

show them and finally they are all over the egg.

After the end of the 3d day the increase of these areas by

division is plainly indicated by their shape. From 82 hours

on to 7 days there are some hundreds of areas each about

150 to 300 p, some elliptical, others dumb-bell shaped and others

in pairs of rounded areas 150 p in each diameter. There is

now a very striking appearance as of light beads floating above

the dark background of the egg yolk. As the areas increase

in number they become crowded till they finally touch one

another and now in profile stand up as hillocks bounded by val-

leys and projecting into a space of some 100 p that is formed :

between the egg and its case.

In late cleavage the color of the egg, which is olive or yellow

green at first but varies much in different females, is now

more dark sage green and this color with the clear space about

the egg gives the entire mass of eggs upon the female a more

No. 447] BREEDING HABITS OF CRAYFISH. 189

transparent look that enables one to pick out eggs of this period

with ease. Under the lens the eggs present a very attractive

appearance as the surface has come to be covered by polygonal

cell outlines that form a neat mosaic. These cells at first about

200 » in diameter and containing nuclear areas 100 y in diame-

ter become reduced to 50 y.

The eggs are often much flattened spheroids that readily

revolve in their cases so as to keep one face upward. The

cells flatten down and the eggs again seem smooth.

After this first week there is a period of apparently little

change, nearly a second week in which the eggs turn dark brown

and at first were thought to be dead. The surface cells no

longer show, while the yolk is arranged in large polygonal areas

about r50 m in diameter and, not quite in contact with one

another. Reagents, however, show the very many small nuclei

of the surface overlying these yolk pyramids.

Embryos. — But before the end of this second week the

embryo is outlined somewhat as in Reichenbach's Stages A and

B ; that is in gastrula stages.

In fhe third week Nauplius stages like Stages F and G

of Reichenbach were arrived at in from 18 to 28 days in different

batches of eggs. Apparently in connection with the very large

amount of yolk in these eggs the embryo is small, only 600 »

on an egg 2000 p.

About the 3oth day the embryo had grown to a length

of 1000 » and was about stage H of Reichenbach, that is with

some nine pairs of appendages. However one rapidly develop-

ing lot of eggs got to that stage in 21 days and hatched out

at the same time as a lot of eggs laid two weeks before this

rapid lot. Here a difference of temperature was present and

may have caused this unusually rapid development. In these

exceptional eggs about two days were spent in passing from

stages G to H and nine days from H to J. Reichenbach's final-

stage K was reached 15 days after J and hatching followed in

from one to two days. : ; ;

In C. affinis we thus found that cleavage took up the first

week, the beginning of an embryo the second week, to progress

as far as the Nauplius the third week and more, to enlarge the

190 THE AMERICAN NATURALIST. [Vor. XXXVIII.

embryo over one half of the egg a fourth week and more and

to perfect the embryo for hatching a fifth and sixth week or

more. The whole egg development required from five to eight

weeks in different sets of eggs under different temperature.

The heart-beat. — When the embryo has grown so large that

it occupies half of the shell and the yolk is correspondingly

reduced the beating of the heart is quite conspicuous, more than-

two weeks before the embryo hatches. Its beat is rapid and

may be more than one hundred to the minute and there is the

remarkable feature of periodic slowing down and even stoppage.

In one embryo the heart beat about 150 times in a minute

besides resting some ten seconds so that the rate was very

great. There were generally five rests in a minute each of one,

two or more seconds each and the intervals between rests were

taken | up with 26 to 36 beats.

Hatching. — The hatching of the eggs as seen in a watch

glass with 2-A.took place as follows. The stiff, transparent case

within which the embryo has developed splits open along the side

next to the embryo's back as a leather ball might if filled with

something that expanded. Before this there were seen some

muscular movements within the embryo and now the region

still containing yolk material was seen to jerk, the antennz

now and then contracted in jerks and the legs moved slightly.

The back of the creature became more and more exposed to

the water, I Fig. 8. The legs showed seeming spasms of con-

traction travelling along them and causing local shortenings.

The back of the head-thorax and of the abdomen protruded

more and more till only the ends of the body and the limbs

remained within the shell, II Fig. 8. The larva thus comes

into the world back first. In their development the legs and

the abdomen have been formed beneath the thorax and bent

forward parallel to it, II Fig. 8; but now the legs are

straightened out and raised up more nearly at right angles to

the thorax and the abdomen also is moved backward and this

aids in pushing the larva out to the position shown in I

Fig. 8.

These changes have taken some fifteen minutes and after about

five minutes more there is a sudden straightening out of the

No. 447.] BREEDING HABITS OF CRAYFISH. I9I

abdomen which throws the whole creature out into the water

as a long straight larva hanging with only. the tip of the

abdomen left inside the shell, III Fig. 8. The limbs now free

in the water kick about and at times the abdomen contracts

and since its tip is fast within the shell the result is thát the

body is brought up near the shell again. The fastening of the

abdomen within the shell is brought about by means of a larval

skin which is fast to the shell on the one hand and to the

Fic. 8.— Three stages in the hatching of young. In III the larva remains attached by abdo-

men to cast off skin within egg shell. X 12.

anal region of the larva on the other. This skin was seen

at the beginning of hatching as a delicate veil over the eyes

and doubtless some of the spasmodic contractions seen along

the legs were serving to free the limbs from this embryonic skin.

The larva thus moults and hatches at the same time and leaves

its cast skin within the egg shell to be made use of as a means

of keeping the larva from falling away from the mother, for

a time. Hanging thus from the egg the young animal soon

begins to spasmodically open and shut its large claws and when

the body is brought up to the egg case by the contraction

192 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of the abdomen, now and then, the claws finally clasp hold

of the stalk of the egg case. As these big claws have the

same larval character found in Astacus, that is recurved or

hooked tips, they become firmly fixed in the material of the egg-

string and apparently the larva could not get free again. The

larva is thus made fast fore and aft, its claws are fast to the

egg stalk and its anus is fast to the cast skin within the egg

Ld hua ra

Pur E AY \ NC

pan

KSN

Fic. 9.—Larva 24 hrs. after hatching: claws fastened to stalk of egg shell and abdomen

fastened to cast skin inside of egg shell. x 25.

case. As the egg cases are still firmly tied to the pleopods

by the strong egg stalks the larvæ live still pendent from and

for protection dependent upon the mother.

Such a larva 24 hours after hatching is shown in Fig. 9,

As in the European crayfish the young is hatched with a swollen

globular head-thorax with proportions strikingiy unlike those

of the adult, but in life the head thorax is not so dispropor-

No. 447] BREEDING HABITS OF CRAYFISH. 193

tionally swollen as it becomes in preserved specimens. In this

region there is still a large, colored, saddle-shaped dorsal area

where the yolk material is present.

All the appendages are present as in the adult except the

first and last abdominal appendages so that the abdomen ter-

minates with a very simple broad telson and has no wide tail

fan. The eyes are very large. The second antennz which

were packed within the egg shell along the edges of the carapace

external to the legs, I, II, III, Fig. 8, soon after hatching

come to be between the legs right and left, Fig. 9, and in

this dejected position add to the general helpless and incom-

plete appearance of the larva. The rostrum also aids in this

infantile expression of the larva as it is very short and blunt

and so bent down between the eyes that it cannot be seen

except from a ventral view. Both antennae and rostrum

appear useless, or at least to be poorly placed or perfected to be

of such uses as they seem to have in the adult. .

This larva is so beautifully translucent that the grossly granu-

lar blood corpuscles may be seen projected rapidly along the

blood spaces in the thorax, abdomen, legs and antennz. These

corpuscles have a long tail-like process behind and by this they

often remain fixed till dragged loose into the current again.

Scattered over the thorax and abdomen but not upon the telson

are large, branching pigment cells of a crimson color and some

are found also upon the antennules and antennze, upon the fourth

segments of the legs and upon the big claws as indicated in

Fig. 9. |

The creature remains fixed but moves its legs and this causes

movements amongst the gills which may be seen through the

carapace. There are also at times slight jerking movements of

the abdominal appendages.

Respiration seems to be carried on very actively, for the very

large scaphognathite may be seen through the carapace, as

indicated in the above figure, bailing water out of the gill

chamber at the rate of perhaps 180 per minute. The establish-

ment of this rhythmic motion so characteristic of many Crustacea

is here a slow process. Before hatching the only rhythmic

motion seen was the beat of the heart, but after the creature had

194 THE AMERICAN NATURALIST. [Vor. XXXVIII.

emerged into the water this respiratory rhythm was gradually

perfected. Two minutes after the larva had first straightened

itself out in the water the scaphognathite was seen to swing |

back and forth but with some stops. Before that at the time of

straightening of the body, there were only a few jerky contrac-

tions and then, after a minute or more, one or two swings.

Five minutes after the body had straightened out in the water

the scaphognathite was not in regular swing and even ten

minutes was not long enough to acquire an uninterrupted, regular

beat.

In some unobserved manner this first or preliminary larva

frees its abdomen from the skin within the egg-case, but still

remains holding fast by its claws. This larva is about 4 mm.

long and lives this life of restricted freedom for only about 48

hours and then moults into a second stage.

Second larval stage.— As the first larva is attached only by

its claws we would expect that when it moulted it would become

loose and no longer connected with the parent, but such is not

the case since it again possesses an anal fastening and later holds

by its claws anew. When the moulting takes place the trans-

parent skin of the larva breaks along the back and the back and

head of the larva slowly push out through the rent. Then the

antennules and antennz are pulled out of the old skin and for

the first time thrust forward into the permanent anterior posi-

tion instead of being carried backward in the embryonic position

retained in the first stage. The legs are pulled out of their old

skins and kick about in the water; then the abdomen is bent

free, all but the tip, where the anal region remains fast to that

region of the cast skin and does not break loose even when it flaps

vigorously. As the shed skin still has its claws fast locked

in the egg stalk the larva though it has drawn its hands out of

its gloves, as it were, and come out of its old clothes, still

remains indirectly fastened to the mother since its telson is fast

to its old suit and that is not broken but continuous with the

gloves, or claw skins. At first the movements of the legs make

the larva seem to be trying to crawl upon the cast skin but

after a few minutes the large claws are opened and reaching

about take hold of the shed or of the egg case. After many

No. 447.] BREEDING HABITS OF CRAYFISH. 195

minutes the claws get fastened to the egg stalk again so that the

larva is a second time moored fore and aft; by the claws to the

egg stalk and by the telson to the old shed which in turn is also

fastened by its empty claw skins to the egg stalk. There are

thus two real claws and two empty casts of claws fastened to

the egg stalk close together. The new claws are still recurved

at the tip but it did not appear that the larvae were always so

firmly fixed that they could not get away or at least be rather

easily pulled off during the six days that this stage continues.

This moulting was found to take place also when the first

larvae were removed from the mother and kept 24 hours in

running water so that the usual protection of the mother is not

absolutely necessary for the future development.

The young in this second stage are about 44 mm. long and

have the form represented in Fig. ro. The head thorax is

more elongated and crayfish-like in form. The rostrum though

bent down between the eyes is much more prominent and when

seen from above it has the characteristic lateral spines. The

large eyes are prominently stalked. The antennules and antennae

which have some thirty segments, are carried out in front of the

animal and moved about as if of use. The abdomen still ends

in a bluntly rounded telson with no sixth pleopods as yet free,

but these wide lateral appendages may be seen in bags or cases

within the lateral part of the larval telson. The whole body is

darkened by very numerous golden red pigment cells which are

so crowded at the tips of the big claws that these are conspicu-

ously crimson tipped. The yolk-colored area is reduced in

extent and dorsally divided into separate right and left areas

posterior to end of which is a large green area. On either side

of the stomach a blue area adds to the complexity of the

above region. The animal still remains somewhat translucent

and blood corpuscles can be seen in places. In the antenne the

corpuscles pass from end to end, a distance of about 2 mm. in a

few seconds. The legs bear conspicuous hairs not shown in the

figure.

When these second larvz are taken away from the mother

they scramble over one another in heaps and do not walk alone,

though if forced apart they can stand upon their legs and walk

ATURALIST. [Vor. XXXVIII.

MERICAN N.

THE A

196

Sz X

“UMOYS JOU sy[e1s-33o 0j VAIL] jo jusurqo?ejje : popuajxe MOU a'uugjue : PAIP[J

No. 447.] BREEDING HABITS OF CRAYFISH. 197

in a feeble manner. Those that hatched in McDonald jars and

moulted to the second stage floated about attached to their sheds

and to the moulded egg cases. When placed upon narrow

strips of cloth they became entangled so that they could be

hung up in the water away from sediment and with some such

substitute for the maternal pleopods they could be artificially

reared, if need be. Larva so suspended probably take food for

they were seen to pass out long cylindrical greenish fæcal

masses 48 hours after moulting. These larve were seen to

swim only when thrown into Perenyi's liquid when they darted

backward with jerks of the abdomen much as do the adults.

They live attached to the mother some six days in this second

stage and during this time the yolk areas dwindle away to a

large extent while the green areas remained and the blue areas

stood out as conspicuous organs in the position of the “ crabs

eyes " on the sides of the stomach.

Third stage.— As seen in a watch glass the moulting of the

larva from the second to the third stage takes but a few minutes.

The head and thorax break out first, then the legs are pulled

out, the abdomen flaps several times and is freed from the old

- skin entirely. There is now no anal connection of larva and

shed and the free telson is found to have the sixth pleopods

fully expanded by its sides so that a very wide tail fan is formed

and its efficiency as a swimming organ is enhanced by very long

plumose hairs. At length the larva has cut loose from its

mother’s apron strings. Yet it still remains upon the pleopods

of its mother crawling about freely over the old sheds and egg

cases. When removed from the mother these young walk about

upon their long slender legs much like shrimp and when dis-

turbed spring backward with widely expanded tail fan again

suggesting shrimp.

These active third larvae are 8 mm. long from tip of rostrum

to end of telson, while the claws 'stretch out about 2 mm. in

advance of the rostrum and the antennz are 5 mm. long.

The head-thorax is most decidedly elongated and crayfish like.

The eyes remain large and conspicuous and add to the shrimp

like appearance. The antennz still have exceedingly large

exopodite scales. The rostrum is now straight out in front,

198 THE AMERICAN NATURALIST. [Vor. XXXVIII.

sharply pointed and of the characteristic gothic style of the

adult.

The coloring of the larva has been altered by the addition

of blue pigment cells over the carapace and legs. The tips of

the big claws are no longer crimson but dark red owing to the

fineness of the pigment reticulum there. The yolk coloring is

gone but the long green hepato-pancreatic areas and the blue

* crab's eyé " regions show through the carapace.

When these young escape from their connection to the

mother they leave behind upon her pleopods the old egg cases

and stalks, and the first and second larval skins. These, how-

ever, all disappear before very long, but it was not observed

whether the female removed them or whether the young may

have taken part in tearing them off. The young continue to

crawl about upon the pleopods for about a week and before the

end of that time the pleopods are cleared of remnants of egg

cases.

These free larvae do not, however, remain continuously upon

the mother but gradually make longer and wider excursions into

the outer world, returning again and again to the mother.

Three hours after moulting some larve were found walking

about on the bottom of the aquarium and when disturbed they

leaped backward several inches with great speed. When the

female was lifted up all the young clung fast to the pleopods,

but when she was left quiet a few minutes several of the young

came off and walk about. The female gave no sign of knowing

of the existence of the young but when she walked over a loose

larva it turned upon its back, when touched by the pendent mass

of young and quickly climbed up amongst the others. But

attempts to make the young climb up on to bits of cloth or even

shed skins held in a forceps were not successful.

In occasionally swinging the pleopods back and forth the

mother was seen to wipe off against the bottom of the tank one

of.the great number of scrambling young that crowd the

pleopods. Such a one at once turned over from its back on to

its feet and walked, though it would appear that it probably had

had no experience of horizontal surfaces.

The young that walk on the bottom of the aquarium do not

No. 447.-] BREEDING HABITS OF CRAYFISH. 199

stay away long but return to the mother. Thus two days after

moulting some 60 young were shaken loose from a female and

scattered upon the floor of the aquarium, but in 20 minutes all

but 12 were back again upon the mother’s pleopods. Four days

after moulting the young crawl all over the mother not only

over the dorsal side of the abdomen but over the thorax and the

head. As many as a hundred may be off at once walking about

in the aquarium and climbing up into the chara plants while a

little later only a dozen may be away from the mother.: When

no screen was used and the water ran strongly the young were

carried away now and then and in such manner it may be that

city reservoirs, such as those in Baltimore, get stocked with

C. affinis. Eight days after moulting the young had all left the

parents though sometimes a few went back to the maternal

pleopods during the next few days. Thus by the fifth of June

the young are most all self supporting and independent individu-

als with no other protection than that they find away from the

parent. ;

How far there: is any special recognition between the

young and the mother was not determined; but when eight

- young were taken away from the parent and put in a dish with

a female from which the annulus had been removed and which

had finally lost all her dead eggs without hatching any they

climbed up on to her abdomen and claws but soon got off again.

The female did not act as did the real mother but moved about

restlessly and wiped her mouth parts as if irritated by the young

crawling over them. A few hours later there were no young

upon her and four of them had disappeared. Again when

several females were kept in the same tank there was some

evidence that larvae sometimes crawled up on to the wrong

mother. Moreover in 1900 it was noticed that when a dead

female was lifted from the water the young that dropped off

returned again to crawl upon the dead crayfish and remained

even when the parent was far advanced in decay. When the

abdomen was broken open the young crawled upon the exposed

muscles for a time but later were upon all parts of the abdomen

except the broken surfaces. | :

Later stages.— By keeping the young of a single mother in a

200 THE AMERICAN NATURALIST. [Vor. XXXVIII. :

tank by themselves some facts were made out as to the sub-

sequent moultings during the rest of May, June and the first

half of July. These young had a large aquarium with good

supply of mud, Chara, anodons and various plants and insects

with running water, sunshine and shallow water for good aération.

The young remained in the third stage for about 18 days and

their color gradually changed from red to greenish owing to the

great increase in the blue pigment. The legs also developed

bands of dark color across them though otherwise almost

invisible, yet the big claws stood out as reddish objects with

blue at the base. The fourth stage had about the same coloring

but being so much larger it was much more easily seen. The

rostrum-telson length was 12 mm.: the claws reached out

2 mm, in advance of the rostrum: the antennz were about 8

mm. long: the thorax was 24 mm. wide, and the tail-fan expanded

54mm. One of these larve had short, simple papillae upon the

first abdominal segment to represent the pleopods, apparently of

the female. These young were still translucent enough to show

the beat of the heart, at a rate of about three beats to the

second, and the circulation of blood corpuscles in the antenna

and legs, though the muscles were now much more conspicuous.

The effect of the red pigment cells was everywhere toned down

by the widely diffused blue cells.

In about 17 days, that is July 1st, many of the young had

passed into a fifth stage. The length was now 15 to 18 mm.,

with the thorax about 3 mm. wide and the tail-fan 8 mm. The

color of this fifth stage was white on the ventral side and else-

where this was overcast by greenish. The legs were now white

with dim transverse bands of gray while the large claws had

lost the red and showed brilliant blue at the base of both finger

and thumb.

The young move about actively, in walking they move the

fore pairs of pleopods back and forth very rapidly. They swim

readily and are still shrimp like in movement and in appearance,

asthey have long slender legs, protuberant — and hes tail-

fans. `

Within a week some larvze were found in a sixth stage with a

length of 21 mm., a thorax width of 5 mm., a tail-fan — of

II mm. and antiin about 19 mm. long.

No. 447.] BREEDING HABITS OF. CRAVFISH. 20I

Some eleven days later a very large larva was found with a

length of 29 mm., a width of thorax of 7 mm., claws projecting 8

mm. beyond the rostrum, antennae 25 mm. long and a tail-fan

expanse of r4 mm. This creature was then a crawling crayfish

with less tendency to keep high up amongst the water plants : it

also was dark greenish, finely speckled, and there was blue on |

the legs. Little translucency was left and the animal looked

much like a diminutive adult. The large claws were still marked

with blue and had dark red at their tips.

This specimen was a male with two long, scroll-like pleopods

carried forward in the groove under the thorax as in the adult

and the following four pairs of pleopods were not modified: thus

the male now has but one pair of modified pleopods as is the case

in the lobster.

We may summarize the growth and moultings of the young of

a single female during two months as follows. The eggs laid the

night of March 28th, hatched into a first stage May 18th. With

a length of 4 mm. they lived two days and moulted into a second

stage. This second stage was 44 mm., long and lived six days

from May 20th. The third stage was 8 mm. long and from May

26th lived 18 days. The fourth stage was 12 mm. long and

from June 13th lasted 17 days. The 5th stage was 15-18 mm.

long and lived from July 1st five days. The sixth stage was 21

mm. long and from July 6th, lived eleven days. The seventh

stage was 29 mm. long, and from July 17th lived an undeter- |

mined time as observations were then discontinued. The first

and second stages remained attached to the mother for about one

week and were not more than 44 mm. long. The third stage, 8

mm. long, kept on or near the mother for about a week and then

lived an independent existence for a week. The following stages

were all independent and gradually took on all the adult propor-

tions and colorings. ae

Growth of young crayfish.— The young were left in a tank

with running water, mud, water plants, anodons and frequent

feedings with raw hen’s egg till October 6th. Then only eight

survivors from a hundred or so were found. These were then

measured and as seen in Table I they varied greatly in the

amounts they had grown, though all were hatched from the

eggs of the same mother.

202 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Table I. Measurements of eight brothers and sisters I41

days after hatching.

TABLE I.

1 2 3 4 5 6 7 8

Length: telson rostrum . . . 62.] $8 | 83 |. 49 | 50 | 4$ | 43 | at

" c uM. - 178155102 | 6s | 60 5$ toda tt u^

s amenna: s . 315148 al 34. | 43. bee Lo 5

Wah: monr =- o s. f$ | 15 | 34 |. 233. | 12. bee ee to

Mg eg 505 S5 «1 46 121/34 132 138 20 | 20 | 19

o EE B X ALI EE. d d 9 á d 9

Length: anterior male organs. 9 9 9 8

M posterior * Nu

IO | I0:| 30 9 r

Ten other young from various females hatched about the same

date as the above and measured October 7th gave the following

results, Table II.

Tastre II.

1 2 5 14 5 6 7 8 Spe

Length: telson rostrum . . . . |6o 59 | 52) 52) 45 |39| 40 | 33 | 30| 22

4 Chel-telon . . . . . | 74/65| 65/68) ss |so| so | 41| 35| 27

IM SIO Vos «4 «s « asl olar [40 29M — 122 — 90 110

Width: thorax *- c.c dimitti S1 dio 34018 8 18

2o 2 BRI... a [o6 laci side ob lan] 18 1108 4 10

SOR VIUTTSS o d dig d |d ?

Length: first male organs . . . IO 919 te |B| 6 £84

" second male organs . . | 11 1011018 121 T | 6 14

Taking the two tables together the largest crayfish was 62

mm. and the smallest 22 mm. long ; the average length was 46

mm. During these two months and three weeks of summer

some few seem not to have grown at all, while about one third of

them have almost doubled their length. The great difference

between the largest and smallest is but imperfectly represented

by the above figures : thus the specimens, 10 of II and 1 of I,

being in length as 31 to r1 seemed in bulk much more widely

apart and in weight they were found to be as 13 to 7. Only 5

of the 18 were females but it is not known when this preponder-

No. 447] | BREEDING HABITS OF CRAYFISH. 203

ance of males arose, whether before or after hatching. The

males have now two pairs of very conspicuous and well formed

sexual pleopods of great size; the exterior pair are a little less

and the posterior pair a little more than 1 the length of the cray-

fish, but this ratio is the same in the adult. In these young

males the antenna often look longer and more tapering than in

the females and that this may now be a secondary sexual char-

acter would seem to follow from the measurements. The eight

males with perfect antennae averaged 50.75 mm. in length and

their antenne were 42.875 mm., or 84% of the length of the

body. The five females averaged 47 mm. in length and their

antennz 31.4 mm., or hardly 67% of the length of the body.

Under such conditions the young of C. affinis may then attain

a length of two inches in their first summer. Additional data

for knowing their rate of growth were obtained from some young

hatched about June 1st, 1901, from eggs laid April 11. These

young were kept in a large sink with running water, mud, water

plants and anodons which dying from time to time probably

gave them food material. The twenty survivors were measured

February r1th, 1902, or 83 months after hatching and the

lengths and widths as well as the sex are given in Table III.

The sexes could be readily determined as the females had con-

spicuous annuli and the males well developed sexual pleopods.

Taste III.

I 213 | 415 ery 8|9 ]|19|11| 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20

É 6 32 34 3 3 243127 32

Length| 41 | 38 | 24 | 56 | 32 | 44 | 43 | 39 | 39 3351331 5 é 7i

i siete Ss | ark S|? j 7 7

Width | 10 3 : I 1r | 10 9 1913F]? g| 2 à

Number 4 may have been introduced by mistake : the others

range in length from 44 to 23 mm. with an average of 34 mm.

None are as long as two inches : one half are about 3 3 mm. and

80% above 29 mm. The sexes were about equally distributed,

10 males and 9 females. i

These young did not attain to as large a size as those in the

204 THE AMERICAN NATURALIST. [Vor. XXXVIII.

preceding tables though they were four months older. But this

additional time was autumn and winter and it would seem

probable that the crayfish of Table III not being as well fed as

the others had not grown quite as fast during the summer and

had then remained stationary, Though they were kept in a

warm room they were in running cold water.

The above twenty crayfish of table III were then kept in the

same conditions during the rest of the Winter, Spring, Summer

and Autumn and until December 5, 1903. The only two sur-

vivors were both females and at that time, being 18 months old,

gave the following measurements. One was 70 mm. long and

16 mm. wide : the other 79 mm. long and 20 mm. wide. Since

last measured, a space of ten months including a summer, each

had probably doubled its length.

These two were left in the same surroundings, except when

removed for breeding as below described, and measured again

July rst, 1903, when 25 months old. One was 79 mm. long and

20 wide, and the other 76 mm. long and 20 wide. The latter

was clean and bright and the former dark and dirty as in winter

so that we infer the smaller one had recently moulted and added

6 mm. or about 44 to its length and } to its width.

During their third summer these two females were kept with

the young of Table I and thus had better food conditions. Only

one survived till October 6th and then had a length of 9o mm. and

a width of 24 mm., while its tail-fan expanded 43 mm. It had

thus gained more than: 10 and perhaps as much as 14 mm. in

length or added $3 to nearly 1 to its length.

Summarizing the above data we see that the young of

C. affinis reared in the above conditions of captivity hatch in

May from eggs 2 mm. in diameter as larve 4 mm. long and

there pass through stages of 43, 8, 12, 15—18, 21 and 29 mm.

during the first two months. During the next three summer

months they increase to an average length of 40 mm. but may

grow as long as 62 mm. During their first winter they may,

probably, not grow at all; but in the second summer they may

reach a length of 70-80 mm. In the third summer the length

may become 9o mm. as indicated M the sole survivor at

28 months.

No. 447.] BREEDING HABITS OF CRAYFISH. 205

That this crayfish grows in the open about at the rate above

indicated seems not improbable from the measurements made

upon 80 specimens of C. affinis taken early in October, 1903,

from a small pond in Baltimore that had been stocked a few

years before. Five of these were 85-100 mm. in length and

obviously belonged to a different period of life as there were

none between these and 65 mm. lengths. The remaining 75

ranged from 65 to 32 mm. There were only 6 below 40 and

12 above 56; the remaining 57 were from 40 to 56 mm, long.

Thus 76% were from 40 to 56 mm. long and we may believe

these were hatched that same Spring and were about 4 months

old; the same would apply to the few smaller ones. The 12, or

16%, that were above 56 and not above 65 mm. long are too

small for young that had passed a second summer and probably

were of the same age as the majority. The five largest speci-

mens were too large for second summer crayfish and probably

were at the end of their third summer. The absence of any in

what we would regard as second year lengths may be perhaps

connected with the draining and drying that the pond suffered

some winters.

Sexual maturity.— In rearing crayfish it was found that their

sexual instincts and organs mature long before the maximum size

was attained. Some of the young represented in Tables I and

II were found to unite in, pairs in October when but 43 months,

or I41 days old. These very small crayfish, two inches long,

thus had their sexual instincts developed. Moreover the sperm

taken from the male is then, apparently, just like that of the full

grown males. The females 1 and 5 of the first table had sperm

plugs, though the other females did not. These females were 62

and 50 mm. long. The female No. 2 of the second table was

clasped by male No. 3 and subsequently had a sperm plug : this

female was 52 mm. long. Whether these females of 50, 52 and

62 mm. in length will lay eggs next spring remains to be seen.

The males seen to unite with females were 55 and 59 mm. long.

The two females described in the last section when 70 and 79

mm. long and 22 months old were put with males April ro,

1903. Though they had been alone for some months they

at once were united with the males and the annulus of each

206 THE AMERICAN NATURALIST. [Vor. XXXVIII.

showed a sperm plug as it did not before. When they were 23 ,

months old they both laid eggs.

Amongst the females that laid eggs in captivity there were

both long specimens 120 mm. long and smaller ones down

to even 75 mm. From their size we infer these small ones

were about to begin their third summer. Thus though the

sexual unions take place at the end of the first summer we

have, as yet, no evidence of eggs being laid till the begin-

ning of the third summer. However the autumnal union in

the first year was found not only in the specimens reared in

confinement but among those eighty above mentioned as taken

in the open. And here it occurred botn in the laboratory and

in the pond and not only between the larger, third year sizes

but between those of 56 mm. which, probably, were in their first

year.

We see then that little specimens of C. affinis may have

mature sexual instincts and unite in pairs in the autumn when

they are but 4 months old and 50-60 mm. long, but as yet it is

not known if they lay before they are 23 months old and

75 mm. long.

Finally— In connection with the development of sexual

maturity we may here refer again to the apparent sexual dif-

ference in lengths of antennz noticed in connection with tables

I and II. When the above 80 crayfish taken in the Autumn

and thought to be most all of the first year, like those of the

tables, were measured little difference between the lengths of

the male.and female antennze was found. Throwing out the

specimens with broken antennz there were 37 males with an

average length of antenna of 42 mm. and 36 females with an

average length of antenna of 41 mm. Again the average male

antenna was 81.3% the average length of the male body while

the same for the female was 78.9%. The differences are so

slight that, considering the errors in measurement, there seems

here not enough evidence that the females have the shorter

antennz as a class.

BALTIMORE, November 5, 1903.

NATURAL HISTORY OF HAMINEA SOLITARIA

W. M. SMALLWOOD.

Tue shell-bearing species of the family Bullidae have been

found freely distributed in the rocks since the Tertiary period.

A large number of species have been found as fossils. Ludwig

in Leunis Synopsis der Thierkunde states that there are between

two and three hundred recent species. They have a wide dis-

tribution in both European and American waters, being com-

monly found in the sandy and muddy bays of the temperate

regions.

The Bullidae belong to the general order of Opisthobranchia

and to the suborder Tectibranchia.

Thomas Say read before the Academy of Natural Sciences at

Philadelphia on July 24th, 1821, a paper in which he gives ** An

account of some of the marine shells of the United States." In

this article we have the first reference to Bulla solitaria. His

description is as follows: “Bulla solitaria. Shell remarkably

thin and fragile, pellucid, oval, narrowed at base, with numerous

impressed, revolving lines, and transverse very obtuse wrinkles ;

apertures surpassing the tip of the shell ; spire none, substituted

by an umbilicus ; umbilicus of the base none, less than half an

inch. Inhabits the southern coast of the United States. -

July 4th, 1835, J. G. Totten described a species of Bulla,

variety znsculpta which he dredged in about fifteen feet of water

from the muddy bottom of the harbor at Newport,.R. I. He

maintains that zusceu/pta is distinct from solitaria because of

some slight variations in the color and shape of the shell. He

says, “ Thus (insculpta) can hardly be Say’s Bulla solitaria. It

is not umbilicated at the top as that species is ; having merely a

shallow pit in which nothing of the interior whirls can be seen.

The solitaria is described as being narrowed at the base; but

though our shell is regularly rounded in the passage, below, of

207

208 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the right into the left margin, it is widely rounded; and the

widest part of the shell is below the middle."

A. A. Gould, in his Z»vertebrata of Massachusetts (41), con-

siders Bulla insculpta and Bulla solitaria identical : he concludes

his description of Bula insculpta with the following paragraph : —

* The differences between solitaria and insculpta, if there be

any, must be very slight. Nor do I see that the two descrip-

tions (Totten and Say) are at all inconsistent with each other. -

Still it is true that the shells from Martha's Vineyard are pre-

cisely like.those from Charleston, S. C., and accord with Mr.

Say's solitaria ; and those from Roxbury are precisely like those -

found by Col. Totten at Newport, R. I., and described by him.

The observable differences are, that the first are of a more dead

white, are more cylindrical, the summit has a more square

appearance, the revolving lines are less distinct, and there is

always a perceptible opening in the region of the spire. These

differences may be ascribed to age or locality."

Verrill (73) describes soZitaria as occurring along the Atlantic

coast from Massachusetts Bay to South Carolina and makes no

mention of zzscu/pta except to assume that it is the same species

as solitaria. The shells of insculpta and solitaria as figured by

Totten, Gould, and Verrill seem to be identical.

The size of the shells which I have found at Wood's Holl,

Mass., varies to a considerable degree ; one shell found was only

one-fourth as large as the average, the shape of a large number

of shells which were examined varies in regard to the width and

length of the lip. All of the shells examined were dextral.

The distinctness of the revolving lines seems to depend on

the age, the larger ones being the

more distinctly marked. The state-

ment that the shells of ¢nsculpta

seem to be more of a dead white

color than that of solitaria is easily

conceal the anterior part of the explained. The shell of -the living

due ll MALE E animal is a golden gray ; the golden

tinge being given by the thin struc-

tureless membrane, the periostracum, which completely covers

the shell (Fig. 1). When this membrane has suffered disinte-

G. win in

the proboscidial snout and tentacu-

No. 447.] HAMINEA SOLITARIA SAY. 209

gration the shell has the dead white color mentioned by Gould.

In view of these variations I think that the shell as originally

described by Say is identical with the zzscz^za of Totten and is

the soZzaria found in the vicinity of Woods Holl, Mass., and of

" Long Island, N. Y.

Pilsbry has decided to limit the family Bullidz to the single

genus Bulla. Formerly the members of the Akeridz were

included with the Bullidae but the character of the radula, form

of the shell, and relation of the animal to it' enables one. to

sharply differentiate the two families.

The Bullidz have in the radula a few longitudinal rows of

teeth (formula I. 2. I. 2. IL). The animal is capable of complete

retraction into the shell. There are no epipodal or parapodal

lobes; the foot is long and tapering behind. The shell has a

mottled color pattern.

The animal described by Say, Totten and others and referred

toin this article is placed by Pilsbry! in the family Akeridze

and in the genus Haminea. “The Akeridz are a much lower

stock of Tectibranchs than Bullide, retaining the primitive

multi-dentate radula, and the PED lobes, and having a thin,

fragile unicolored shell. :

The following is the specific description of ZZamznea solitaria :

* Shell thin, subcylindrical, with gently convex sides, truncate

vertex and rounded base ; color horny or light brown. Surface

shining, having irregular growth wrinkles and (under a lens)

Jine, deeply impressed spiral grooves, much narrower than their

intervals, sometimes with smaller ones intercalated. Vertex

white, somewhat impressed in the middle, subperforate. Lip

arising to the right of the center, slightly -

thickened; outer lip. gently arched forward.

Columella thin, concave. Alt. 10, diam.

63 mill” (Fig. 2).

Habitat. — Until "b enact of Verrill, pole Bg mA of

(71-72) nothing was said of the animal or .

its natural history. All writers cogfined themselves to a con-

chological description of the species. Verrill says that so/taria

! Manual of Conchology.

210 THE AMERICAN NATURALIST. (Vor. XXXVIII.

“is restricted to muddy shores and bottoms, in sheltered situa-

tions and is found also in muddy ponds and estuaries.” Smith

and Prime state that the animal is rare, found in mud at a depth

of one or two fathoms. Balch ('99) states that the animal,

* Haminea solitaria, Bulla solitaria, is rather uncommon alive,

sometimes occurs on marsh grass, top of sea walls." In regard

to the general distribution of this species Verrill says, “From

Mass. Bay to South Carolina it is common in muddy lagoons and

salt ponds, in shallow water where not too brackish, along the

shores of Vineyard Sound, Buzzard's Bay, and Long Island

Sound. Abundant in a small pond near Holme's Hole, in New

Haven Harbor, in ditches near Fort Hale."

Haminea solitaria is found rather commonly in the Eel Pond

and Big Harbor at Wood's Holl, at Hadley Harbor, and at the

bathing beach on Buzzard's Bay at Wood's Holl. During the

laying season the animal migrates into shallow water and may

occasionally be seen upon the eel grass and algz, but usually

not in water less than two feet deep at low tide. If the eel

grass be disturbed they drop at once into the water and fall to

the bottom, where it is very difficult to see them, owing to their

form and color being so similar to the ordinary weeds and mud

at the bottom ; thus it is practically impossible to gather animals

by looking for them on the weeds or bottom. They have been

secured in two ways : first, by taking an ordinary fine-mesh dip

net and skimming off the sea weeds and half an inch of mud

from the bottom, then carefully washing out the mud; in this

way à number have been secured in the shallow ponds and

lagoons about Wood's Holl; secondly, by dredging; this

method is necessary in deep water. Bulla may be found, even

in the breeding season, in water thirty feet deep in the Big Har-

bor at Wood's Holl, — the greatest depth where they have been

found. This peculiarity may be due to the fact that their for-

mer laying spot in shallow water is now occupied by a deep sea

wall. Dr. Charles B. Wilson obtained a number of solitaria by

skimming in clear water near Edgartown about the 18th of

August. He was sure that the net did not pass over any eel

grass or sink beneath the surface. I feel sure that this is the

real explanation of the occurrence of Bulla in great abundance

No. 447.] HAMINEA SOLITARIA SAY. 211

just before the egg-laying time, and their equally sudden dis-

appearance after the egg-laying time. They simply rise to the

surface and swim away.

` In the summers of 1897 and 1898 most of the specimens

obtained by me were collected in the larger of the two so-called

*oyster ponds" near the Marine Biological Laboratory.

Although a large number of capsules were taken from this

place, over one hundred capsules were gathered from another

locality and put into this lagoon in order to assure an abundance

of material in succeeding seasons. In the summer of 1899 no

capsules could be found in this lagoon, in the summer of 1900

three capsules were found here, but all of the eggs had died

before reaching the twenty-cell stage ; the tide flows freely into

the lagoon thus keeping the water comparatively fresh. The

drainage from the Marine Biological Laboratory flows into an

adjacent lagoon and it is possible that the water seeping through

may have some poisonous effect which kills the eggs and has

destroyed or driven out the animals themselves.

The Bulla found on the sandy bathing beach at Buzzard's Bay

occurs about half an inch below the surface of the sand ; usually

near the capsule, which is the only indication of the presence of

the animal. |

General Morphology.) —1t is not my purpose in this section

to go into the details of the various systems of organs except in

so far as they are directly related to the development of the

egg; I purpose, however, to give a brief sketch of the general

form of the animal.

Probably the most striking feature of this species is its color,

which is a golden gray thickly mottled with dark brown and

occasional orange spots; one might almost say that the gray

background looks as though sprinkled with fine sand. The

shell is translucent and slightly striated spirally ; it is not at all

glossy or shiny. One would hardly expect to find much of a

shell in a tectibranch, but in so/taria the reduction of the shell

A For a foll acai of the Morphology of the family Haminea (Bullacea) see

M. Vayssiere, Recherches Anatomique sur Les Mollusques de La Familles des

Bullides Ann. Sci. Nat. Tome 9. 1879. R. Bergh, Bullacea. Reisen in Archipel

d. Philippinen, pp. 211-312, 1901.

212 THE AMERICAN NATURALIST. [Vor. XXXVIII.

has only just begun. The most noticeable feature in this process

is the fact that the shell is thin and fragile, so much so

that it must be handled very carefully. The second feature is

that in the mature animal the shell is not large enough to shelter

the whole body in its contracted state. The shell certainly

cannot afford particular protection against enemies, but when we

remember where the animal lives and its habit of crawling along

the mud and sand, we can readily see that it could have no

better protection than its color. It imitates the color of the

roots of eel grass most closely ; at first sight I have often mis-

taken the detached fragments of these roots for Haminea.

The size of the animal depends largely upon age; the smallest

shell measured was three mm. in width, and five mm. in length ;

the average size of the mature shell is 7 mm. in width, and 10

mm. in length. The average length of the extended animal

including the shell is 2 cm.; the smallest animal was 8 mm. in

length. I can make no definite statement as to the age or size

at which Æ. solitaria begins to lay, but can say that I have

never known the smaller ones to lay. In the aquaria the animals

die after laying, but I do not believe that this fact can be used

as proof that they always do.

The foot of Haminea possesses, though not clearly distin-

guished, the three divisions characteristic of the typical mollus-

can foot. From the mesopodium arise the parapodia; these

are lateral lobes, or folds, like extrusions of the edges of the

foot. In some of the Opisthobranchia the parapodia (epipodia

of Pilsbry) are highly developed and unite upon the back, com-

pletely covering the shell. In our species however the "NE

podia cover only the anterior part of the shell.

The young animal moves by stretching forward the head and

foot, and then drawing the visceral mass forward, so that the

progress results from an alternate stretching and contraction of

the anterior flexible part of the body. The posterior part being

drawn ahead at each contraction. In the mature animals there

is no such noticeable division in the movement; with them

locomotion is apparently a continuous gliding process.

The anterior part of the animal terminates in a broad thin

proboscidial snout. Running back from the end of the snout

No. 447.] HAMINEA SOLITARIA SAY. 213

there are two tentacular disks, which are divided only part way.

There is very little movement in these disks, except as they

shift about over the anterior portion of theshell. In this species

there seem to be no special sense organs in the tentacular disks.

So far, I have been unable to explain their function. The

tactile sense is usually located in the tentacles, but in our

animal the anterior portion of the snout performs this function,

as is very evident from its movements. In many of the Opis-

thobranchia the mantle folds over part of the shell, but in Æ.

Solitaria the only external evidence of the mantle is a thick fold

which occupies the lip of the shell and extends back of it for

about four mm., it is here extended and affords a surface equal

in width to the shell. At first sight one would think because of

this arrangement that the foot extended from the snout to the

posterior portion of the animal, but closer inspection shows a

sharp demarcation which indicates the boundary between the

posterior portion of the meso-podium and the mantle. All of

the parts of the body that are exposed are covered with colum-

nar, ciliated epithelium. A great quantity of mucus is secreted

by the animal, so that in crawling about the dish they often

leave a nearly perfect tube of mucus. This is secreted by

numerous typical single-celled glands, which are especially abun-

dant at the edges and tip of the snout, and the outer portion of

the tentacular disks. :

The especial characters aside from the shell which ip arae

H. solitaria are the stomach plates and zu

the form of the radula.

There are three stomach plates so

arranged that the food is trituated by

them (Fig. 3). The plates are com-

posed of chitin, having their bases

firmly imbeded in the strong muscles :

of the stomach. The form of a single pis. 3.—showing the ipea

plate is shown by the camera drawings ment of the PEOP UR S

(Figs. 4, 5). The portion of the plate int of muscle. The free sur-

that comes in contact with the food is zte perfectly the food.

differentiated into a number of ridges

which are larger and more pronounced at the anterior end. The

214 THE AMERICAN NATURALIST. [Vor. XXXVIII.

largest teeth are found on the anterior part of the plate, having

a triangular form. The size

and prominence of the teeth

gradually decreases toward

the posterior part of the

plate. This gradual increase

in size of the teeth and the

ridges indicates how both are

continually forming to take

the place of the more anterior from a profile view.

ones as they become worn *%?*

out. The youngest teeth are those found on the smallest

ridges. In some instances it is difficult to be certain that teeth

are even present.

The radula is an interesting structure but one that is difficult

to represent in a drawing. When this organ is removed from

the animal, it is somewhat triangular in shape (Fig. 6). In the

center of the anterior part there is a single row of teeth, having

a broad free margin and ter-

minating at each side in a

rather broad blunt process

(Fig. 7). There are an in-

definite number of long, slen-

der, sharp teeth arranged in

rows which correspond to the

teeth of the median ridge.

The bases of these lateral

teeth are imbeded in the

muscles of the radula (Fig.

8). Each lateral tooth is

bent at an angle of about 90

degrees. This arrangement

of the teeth would give the

following formula for H. soZi-

i 6.— The radula seen asa transparent object. /@77@ œ. 1 oc.

The anterior end is uppermost in the drawing. . :

The formula is infinity. one. infinity. x 72. The genital organs — This

system of organs is fully dis-

cussed by Lang; I have been able to demonstrate of all the

free surface. X 24

No. 447.] HAMINEA SOLITARIA SAY. 1 2.

parts as given by.him and will, therefore, quote freely from his

description. The Opisthobranchia are all hermaphrodites.

Haminea is no exception to the general rule and comes under

the first type as described by Lang. The

germinal gland consists of numerous “con-

verging diverticula.” The eggs and sperm-

atozoa arise from the same part of the

gland and are intermingled. During their

development they become detached and

lie free in the cavity of the gland. The

ducts of Z. solitaria are considerably com-

plicated, because of the development of

the accessory organs. The ovotestis lies Fic. 7.—Three of the median

between the lobes of the liver and the pos- M dimid ze:

terior part of. the body; this gland has

humerous branches, which finally collect into the common

hermaphroditic duct ; the duct empties into the common genital

cloaca. The genital aperture opens

into the extreme right anterior part

of the mantle cavity, and from it

there is, continued forward, an

open ciliated furrow, which carries

the spermatozoa to a gland called

the * prostate "; this opens into

Fic. 8— Three of the lateral teeth from the the penis. “The penis itself lies

a SP in the right, on the boundary

between the head and foot. When it is at rest its sheath lies

in the cephalic cavity near the buccal mass." Two important

glands open into the genital cloaca by a common duct; first,

the albumen gland, which is comparatively small and lies upon

the surface of a second, known as the nidamental gland; the

latter is much larger than the former and yields the outer pro-

tective envelope of the egg. The albumen gland supplies the

albumen for the egg capsule.

The * receptaculum seminis " is connected by a short duct

With the genital cloaca and receives the spermatozoa at the time

of copulation. Of course the size of this vesicle varies according

to the season and according to whether or not it is full of

Spermatozoa.

216 a THE AMERICAN NATURALIST. [Vor. XXXVIII.

Breeding habits—In general the breeding season of our

species extends from the last of June to the first of September.

There seems to be considerable variation in the time when the

egg laying begins, in 1897 the capsules were first found July

9th. In 1898 none were found until July 25th. Then three

were found which had been laid within thirty-six hours. In

1900 collecting was begun July roth and capsules were found

which must have been layed as early as June 25th, judging from

the fact that the embryos had already left some of the capsules

and were leaving others. However, the animals lay most

abundantly between July 15th and August 15th. The egg

capsules may be found scattered through the eel grass and algz

on the edges of the pond, or lagoon, but each capsule is attached

to grass or algze from two to six feet below the surface of the

water. Apparently the animals congregate in favorable spots to

lay their eggs. In one place in the eel pond, where the water

is about three feet deep at low tide, over five hundred capsules

were collected in ten days within an area about ten feet square.

During the same period repeated trips were made to various

parts of the pond, but only a very few capsules were found.

On the sandy bathing beach previously mentioned the animals

lay in the same place from year to year. Here there is a small

patch of eel grass in about three feet of water; in and around

this patch the eggs are laid in great numbers. In this locality -

the capsules are attached either to the eel grass or, as in most

cases, simply to the sand. The sandy bottom slopes out grad-

uall for about one-fourth of a mile, merging finally into a

muddy bottom. I have never dredged here, either before or

after the breeding season, so I do not know where Haminea

stays during other seasons ; but I have dug down into the sand,

both before and after the laying season, and have never been

able to find any of the animals. It is difficult to ascertain where

Haminea lives at other times than the breeding season, but

the fact that it cannot be found in shallow water except at

this time seems to show that it migrates into deeper water.

This idea is also supported by the occurrence of Haminea in

thirty feet of water in the Big Harbor, and by the statement of

Verrill (72) to the effect that, * A specimen of winter flounder

No. 447.] HAMINEA SOLITARIA SAY. 217

(Pseudopleuronactes americanus) caught at Wood’s Holl in

August, contained a large number of the shells of Bulla solitaria."

The winter flounder is dredged in the vicinity of Wood's Holl at

a depth of about fifteen to thirty feet, and is mentioned by

Verrill as devouring Bulla in large numbers. Haminea has been

dredged in May in Hadley Harbor at a depth of twenty-five feet.

These facts would tend to show that its natural habitat for the

most of the year is the deeper water. A further proof that

H. solitaria lives in deep water except during the breeding sea-

son is supplied by the following facts: August 28, 1900, Dr. C.

V. Wilson while skimming the surface water with a fine mesh net

near Edgartown, secured a large number of H. solitaria. They

all died during the first night although they were placed in a

small aquarium. The time that these individuals were collected

was after the usual egg laying periods for the forms that live in

the vicinity of Wood's Holl. When taken, they were evidently

migrating into the deep water. The observations of Verrill

and those of Wilson seem to show conclusively that H. solitaria

is a deep sea dweller except for about six weeks, when it takes

up tempofary quarters in some favorable place in shallow water.

During the first two summers that H. solitaria was under my

observation I was unable to discover any evidences of copulation,

owing to the fact that early in the season I did not have in the

laboratory any considerable number of animals ; but in the sum-

mer of 1900 I had in the laboratory about forty animals at one

time and was able to make observations on copulation. The

habit which these molluscs have of crawling over one another

and of collecting into a pile, concealed the fact and method of

copulation for a long time. One day, however, I noticed two

animals apparently copulating, the genital grooves were slightly

extended and came together, from one the penis protruded into

the genital groove of the other. The animal receiving the penis

laid a mass of eggs eight hours after the copulation, the other

one did not lay. In a second case of copulation the animal laid

after the lapse of thirty-six hours, but this was an unusually long

time and probably due to the unnatural conditions surrounding

it In the time that elapses between the period of copulation

and deposition, Æ. solitaria is similar to many of the nudi-

branchs (Smallwood 1:03).

218 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The question arose as to whether or not copulation took place

after laying ; in the three cases that I was able to observe copu-

lation did take place after one of the animals had laid but I was

unable to observe it between animals both of whom had already

laid. Fresh Haminea were collected, put into separate dishes,

and kept there for three days; during this period they did not

lay; then specimens which had copulated and laid were put in

with these, copulation followed in each case in about an hour

and the animals laid in about twelve hours. Sections of the

ovotestis before and after copulation prove the correctness of

the above observations.

In copulation the animals do not uniformly assume any

definite position in regard to each other. The genital groove

opens on the right side just anterior to the lateral fold of the

parapodium, thus obviating the necessity for an exact position.

Copulation continues for about fifteen minutes.

This species lays a single gelatinous mass (Fig. 9) which is

spherical, about three-quarters of an inch

in diameter. Its contents are chiefly

composed of albumen, which is secreted

by the albumen gland. As soon as the

albumen comes in contact with the

water it swells by the rapid absorption

of water, and thus affords a gelatinous

protection for the egg. When the eggs

first leave the genital groove they are in

Fic. 9.— The eggs of H. solitaria

are laid in a gelatinous mass,

drawing is natural size.

would be very difficult

strings; in a few hours the strings lose

their continuity and the eggs are scat-

tered throughout the egg mass. It

to count the eggs in a single mass. The

size of the capsule varies considerably; as a rule those found

on the eel-grass are about a third less in diameter than those

laid on the bottom. The egg masses laid in the laboratory were

often irregular in shape and much smaller than those collected

from the pond. The specimens in confinement that laid small

and irregular masses, often laid a second time without a second

copulation. It takes from 40 to 50 minutes for an animal to

lay a complete normal egg mass.

No. 447.] HAMINEA SOLITARIA SAY. 219

The living egg of H. solitaria is so small and so richly sup-

plied with deutoplasm that satisfactory observations on the

segmentation are impossible except in the early stages. The

egg is spherical, enclosed within a thin structureless membrane.

The size of the egg varies, the average is about O6 mm,

being smaller than the eggs of Umbrella (Heymons, 93),

Crepidula (Conklin, '97), Nucula (Drew, :o1) and that of

most molluscs that have been studied.

Before segmentation the polar differentiation of the egg is but

slightly indicated, the yolk being almost uniformly distributed

except in the region of the polar bodies. It has already been

Fic. ro.— The four celled stage which shows the relation between the four blastomeres

characteristic of Mollusca. X 275.

stated that it takes forty minutes for the animal to lay a mass

of eggs.. Within ten or fifteen minutes after each egg is laid

the first polar body appears at the animal pole and thirty

minutes later the second polar body can be seen. It happens

occasionally that the first polar body is very large and may

€ven contain yolk spheres.

The egg segments into two cells a half hour after the second

polar body has appeared. In about thirty per cent. of the eggs

Observed, the first division of the egg did not divide it into two

equal blastomeres, one being noticeably larger, a variation which

220 THE AMERICAN NATURALIST. [Vor. XXXVIII.

is similar to Umbrella (Heymons '93). Within thirty or forty

minutes after the formation of the two celled stage, the four

Fic. 11.—The four-celled stage preparatory to the formation of thefirst quartette of micro-

meres. X 275.

celled stage is formed. Just prior to the formation of the four

celled stage, the spindles do not lie parallel which is an agree-

ment with other molluscs

and indicates a spiral divi-

sion ! (Fig. 10).

After not more than thirty

minutes, the third cleavage

separates the egg into two

conspicuous parts, the proto-

plasmic micromeres and the

ini the cgh and weird ones deutoplasmic ^ macromeres.

of the spindles when the micromeres are formed. These micromeres are con-

ses siderably larger in compari-

son with the size of the macromeres than in many molluscs.

The same is true for the two following quartettes of micromeres

‘For a complete discussion of the method and significance of segmentation in

Mollusca see Mark, E. L., seriei. pisos and d =e Limax

campestris. Bull, Mus. Comp. Zool. v 1881. Conklin. E. G., Embry-

ology of Crepidula. Your. Morph. vol. 1 F he Holmes, S. w The Rs POLL

ment of Planorbis, Your. Morph. vol. 16, 1900.

No. 447.] HAMINEA SOLITARIA SAY. 221

which results in the complete envelopment of the macromeres

at an early stage. The third cleavage is dexiotropic (Figs. 11,

12; 13, 14.

The time that intervenes between the formation of the second

and third quartettes of micromeres is the same as that for the

second and third cleavage. From this time on it was impracti-

cable to follow the further cleavage stages on the living egg.

Stained preparations confirmed the observations made on the

living egg.

The second quartette of micromeres is formed by the fourth

cleavage which takes place in an anti-clockwise direction, the

Fic he f. i f the fi f micromeres. The movement

. 13.—The telop

is dexiotropic.

cells taking a position alternating with the cells of the first

quartette (Fig. 15). The next cells to undergo segmentation

are the first micromeres formed, segmentation takes place ina

leotropic direction giving rise to the turret (trochoblast) cells

(Figs. 15, 16); immediately after, indications of division in the

second quartette of micromeres and the macromeres are evident.

The egg sketched shows eight complete spindles in the meta-

phase. The spindles occurring in the macromeres participate in

the formation of the last quartette of micromeres, the movement

being in a right handed spiral. Soon after the above described

222 THE AMERICAN NATURALIST. [Vor. XXXVIII.

changes have taken place the macromere which is designated

by the letter D divides independently of the three other macro-

meres into two cells of unequal size but both containing yolk

spheres. The smaller cell is concerned in giving rise to the

mesoblastic bands and is entirely covered above by the micro-

meres.

It can be seen from the brief description of the early segmen-

tation stages and the accompanying sketches that Hamznea soli-

taria does not exhibit any segmentation phenomena other than

those characteristic of mollusca. The subsequent cleavage

iiL-.----.1b

Fic. 14.— The eight celled stage fully formed showing the position of the micromeres above

the furrows of the macromeres. x 278.

stages have been followed sufficiently to indicate that they are

in agreement with related forms that have been described

heretofore.

The embryo usually begins to move in the egg capsule at the

end of the seventh day ; the cilia on the mantle border are well

differentiated and can be seen in motion. It is difficult to

ascertain the exact length of time the embryo remains in the

egg capsule but I have known it to continue there for a week.

In most of the egg masses that have been under observation the

embryos have died before becoming free swimming individuals

although some were immediately placed in aquaria or in the

No. 447] HAMINEA SOLITARIA SAY. 223

eel pond in bottles closed with several thicknesses of cheese

cloth. In placing the embryos in their natural environment, it

was hoped that it would be possible to determine the changes

taking place between the embryo and the adult. Although

repeated experiments were made in various parts of the eel pond,

no satisfactory results were obtained and I am unable to state

how the transition from the embryo to the adult takes place.

During the summer of 1899 a number of pressure experi-

ments were tried for the purpose of determining the effect on

segmentation and the subsequent history of the embryo. The

fact that the eggs are surrounded by a great mass of albuminous

Fic, 15.— The € celled stage. The second quartette of micromeres was fo

laeotropic movement. One cell, 1b, is Ub uiniat ds segmentation which will iiem in the

formation of a eiit cel. X 275.

material made it easy to apply light pressure. A small number

of eggs were placed on a slide and covered by a second slide,

the amount of pressure was regulated by passing a small rubber

band around the slides. The eggs were taken in the one celled

stage after the polar bodies had formed; they were left under

pressure two hours and the changes which took place during the

period were carefully noted. I took pains to see that all of the

eggs were forced to segment in an irregular manner. After the

pressure was removed the eggs were placed in the aquarium in

a bottle which was stoppered with cheese cloth. Although a

224 THE AMERICAN NATURALIST. [Vor. XXXVIII.

number of experiments of this kind were tried with different

degrees of pressure I was not able to get any normal embryos.

A few abnormal embryos were reared but they lived but a few

days. It hardly seems that it will be possible to get any fruitful

results from pressure experiments on the eggs of H. solitaria

for these further reasons: eggs in the egg masses broken during

collection develop abnormally, probably because the sea water

gains access to them ; eggs laid in imperfect egg masses in the

laboratory frequently develop abnormally; occasionally I have

Fic, 16.— The sixteen celled stage. The first il LE i

turret cells, rb?. The third 3 F

i (dodi oy M ebinuR a EA.

The second quartette of micromeres is the process of division. X 275.

collected what appeared to be perfect egg masses and found that

the segmentation was very irregular, keeping these eggs under

observation, I found that they invariably died in a short time ;

some apparently normal egg masses have been found to contain

eggs in all stages of segmentation from the one celled to the

thirty celled stage.

In brief then to summarize: The Techtibranch mollusc, first

described by Say, and subsequently by Totten, Verrill, and

others, should, according to Pilsbry, properly be regarded as

Haminea solitaria. The dental formula is œ. 1 œ. The egg

No. 447.] HAMINEA SOLITARIA SAY. 225

laying period extends from the middle of June to the last of

August during which time the adults migrate from the deep

water into shallow ponds and lagoons. The eggs are laid in a

gelatinous mass, spherical in form, attached to eel grass, algze,

stones, sticks, etc. The eggs pass from the one celled stage to

the free swimming embryo in seven days. The method of seg-

mentation of /Zamnea solitaria is in close agreement with the

other mollusca. No positive results were obtained from attempts

to produce abnormal segmentation.

Zo6LOGICAL LABORATORY, SYRACUSE UNIVERSITY,

November rs

BIBLIOGRAPH Y.

BALCH, F. N.

'99. List of Marine Mollusca of Cold Spring Harbor, Long Island,

with description of one new Genus and two new species of

Nudibranchs, Proc. poe Soc. Nat. Hist. vol. 29, no. 7.

CONKLIN, E. G.

'97. The Embryology of Crepidula, Jour. Morph. vol. 13, no. 1.

Drew, G. A.

:01. The Life-History of Nucula delphinodonta, Quart. Jour. Mic. Sci.

"41. invertebraies of Massachusetts.

Heymons, R.

'83. Zur Entwicklingsgeschichte von Umbrella mediterranea, Zeitschr,

wiss. Zool. vol. 56, no. 2.

Say, T.

‘21. An account of some of the Marine shells of the United States,

Jour. Acad. Nat. Sci. vol. 2.

SMALLWOOD, W. M.

:03. Notes on the Natural History of some of the Nudibranchs.

Syracuse Univ. Series IV, no. 1.

SMITH, S. & PRIME, T.

70. gases on Mulises of Long Island, N. Y., Ann. Lyce. Nat. Hist.

Bull.

TorTEN, i G.

'85. Description of some shells belonging to the coast of New England

Silliman's Jour. vol. 1.

VERRILL & SMITH

72 ihe Invertebrates of ashes Sound, U. S. Fish Com. Report.

p M

SR rea

ERU

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Mendelism and Cytology.'— Guyer's doctor's thesis written in

1900 but printed and distributed in the latter half of 1903 is remark-

able for its “anticipation” of Mendel's law of purity of the germ

cells, the outcome, unlike Mendel's results, of a cytological study.

He first describes the course of spermatogenesis in normal pigeons.

The spermatogonia (the ancestral sperm cells immediately pre-

ceding the reduction stages) contain 16 chromosomes which are

split in the cell division that forms the primary spermatocyte. As

the primary spermatocyte grows, synapsis, ora fusion of the chromo-

somes in pairs, occurs; and this Guyer interprets as the conjugation

of maternal and paternal chromosomes. Eight thick rings are formed

which break equatorially in the division by which the secondary

spermatocytes are produced. When the secondary spermatocytes

divide to form the young spermatozoa only four chromosomes are to

be seen. These chromosomes Guyer regards as quadrivalent and he

suggests that “reduction” takes place when they divide.

He suggests that the reduction division of the four-fold chromo-

somes may be in any plane and so varying combinations of maternal

and paternal qualities will go to each spermatozoon. Thus it might

happen that in the division the maternal and paternal qualities were

segregated (and this he regards as the prevalent result in hybrids) ;

or it might happen that some of both the die and paternal

chromatin went to each spermatozoon. The “purpose” of the

formation of the quadrivalent chromosomes is to give greater

variability.

In respect to hybrid pigeons, Guyer notes that the offspring of

the common brown ring dove mated with a white ring dove are

brown. The offspring of these brown hybrids are either white or

brown and the latter color predominates. The author says (p. 36):

“This points to the conclusion that in the brown birds we may have

* Guyer, Michael > Spermatogenesis of Normal and of Hybrid Pigeons.

Dissertation, etc. Univers ity of Chicago, Chicago; 1900. 61 pp., 2 satis

plates. Dies (and printed?) 1905.]

227

228 THE AMERICAN NATURALIST. (Vor. XXXVIII.

both intermediate forms like the hybrids of the second generation and

forms which have reverted to the brown grandparent, as the white

doves have seemingly returned to the white grandparent.” Here we

have a clear recognition of what Mendel calls dominance! Also, this,

(p. 48) : *If a spermatozoón and an egg containing characteristics

of the same species unite, then the reversion will be to that of the

species; if a sperm cell containing the characteristics of one species

happens to unite with an ovum containing characteristics of the other

species, then the offspring will be of the mixed type again. By the

law of probability the latter will be the more prevalent occurrence,

because there are four combinations possible, and two of the four

would result in the production of mixed offspring, while only one

combination could result in a return to one of the ancestral species."

Here we have even the quantitative part of Mendel's law expressed in

1900!

The foregoing Mendelian generalizations are suggested by the

behavior of the hybrid germ cells in the spermatogenic stages. The

mitoses are frequently abnormal — two spindles lying side by side,

owing to the fact that the chromosomes are segregated in different

parts of the cell. This segregation suggests an incompatibility

between the chromosomes of the two species — and it results in

“pure " germ cells — with the parental qualities segregated.

Finally, the all too brief chapter of suggestions will repay careful

study. It is regrettable that so notable a contribution to the mech-

anism of heredity should have been so long delayed in appearing.

C B

Inheritance of Acquired Mental Characteristics.'— A Chicago

solicitor of patents has written a book on heredity that is bold and in

many respects crude, but which presents so many facts that it war-

rants respectful consideration. The subject is the control of the

intellectual quality of the offspring by the intellectual activity of the

parents. The thesis is that the descendants of intellectually active

parents inheret the latter's activity so that, within limits, the more

active during a given time the parents have been, or the longer the

time of their activity, the more intellectually active the offspring, the

greater their chance of achieving eminence. It is nothing new, of

course, that the offspring of intellectual or successfully active people

are especially apt to have eminent progeny, but it is rather new to

'Redfield, C. L. Control of Heredity. A Study of the Genesis of Evolution

and Degeneracy. Chicago, A. C. Clark, 1903. 8vo. 343 pp., illustrated.

No. 447] NOTES AND LITERATURE. 229

be told that of the progeny of such eminent people the younger sons

are more likely to be eminent than the older sons; or conversely,

that eminent men, in general, particularly whén not sons of eminent

men are sons of old men.

To prove the thesis stated in the last paragraph it would be

necessary first of all to find the average age a large random sample

of mothers and of fathers of a given race and time at the birth

of all their children and then to show that eminent people (using as

a measure of eminence some arbitrary standard such as the average

number of lines in the biographical descriptions in a number of

encyclopedias) were born of parents clearly older than the average

of parents of that race and time. But even this would not be wholly

satisfactory. It would be better to compare the average eminence of

the earlier and the later born of pairs of brothers. If the average

eminence of the later born brothers exceeded that of the earlier

born by several times the probable error then the greater chance of

eminence of younger sons.could be said to be demonstrated. But

even if the younger sons showed a clearly greater eminence, still we

could not assert that this greater eminence was due to inheritance of

acquired intellectual activity of the parents rather than to the possible

superior training of later sons! Now Mr. Redfield has not treated

his statistics of eminence in relation to birth rank with sufficient care ;

he is convinced of the truth of his theory; and he uses all of the

art of a skillful lawyer to prove it.

Mr. Redfield got a standard average age of Caucasian parents in

general from the Redfield genealogy, which indicates that 50% of

children are born from fathers under 33 years and mothers under 29

— these ages are taken as his standard although he thinks them

a trifle high for Caucasians in general He compares with this

standard the father's age of eminent men at the time of the latter's

birth, gleaning his facts from encyclopedias, and finds many cases

of sons of old men. He devotes one chapter to “ The Hall of

Fame" men. He finds among these many cases of exceedingly old

parentage. For instance when Franklin was born his father was 51 ;

and the total interval in three generations is 51 + 57 + 70 = 178

years. On the other hand the average birth rank of Eli Whitney's

male ancestors was 3o and for 25 Hall of Fame men the median

paternal birth rank is 35.5 years, not much above Redfield's standard.

1To iab the possible influence of superior training of younger children records

of trotting horses or milk-cattle would be superior to records of men. Red-

field thinks his theory confirmed in trotting horses.

230 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The argument is weakened by including the birth ranks of Joseph,

Moses, David and Solomon! We have little reliable information con-

cerning the ages of the ancestors of these men. Also, our confidence

in Mr. Redfield’s critical ability is terribly shaken by his comparison

of maternal impressions to mimicry, and by his attempting to account

for the intellectual inferiority of the lower animals solely on the

ground of their shorter generations.

Despite, however, all the crudities of the book we cannot deny that

it contains suggestions and that many of the conclusions cannot, in

our present state of knowledge, be refuted. The work should incite

to further and more careful investigation to confirm or refute Mr.

Redfield's theory, or, rather, to see if statistical evidence supports

the hypothesis of the inheritance of acquired dynamical qualities.

CRD,

BOTANY.

Notes.— Lieforung 29—30 of Ascherson and Grábner's Synopsis der

mitteleuropaischen Flora deals with Cyperacez;, Aracee and Palme,

— among the latter characterizing American and other foreign

species that are hardy in cultivation.

No. 26 of the new series of “Contributions from the Gray Hérba-

rium of Harvard University,” forming Vol. 39, No. 11, of the Pro-

ceedings of the American Academy of Arts and Sciences, is a revision

of the genus Flaveria, by J. R. Johnston.

Under the title Arkiv för Botanik, a new serial has been launched

by the K. Svenska Vetenskaps-Akademie. Several of the papers

of the opening number are of interest.to American botanists.

After a long interval, parts 3 and 4 of Muhlenbergia has appeared,

and contain descriptions of a number of western phanerogams by

Heller and Congdon.

The supplement to the /ndex Kewensis, in fascicle 3, reaches

Physaria. :

The embryology, etc., of Seguoia sempervirens are discussed by

Lawson in 44zza/s of Botany for January, which also contains a

historical account of the structure and morphology of ovules, by -

Worsdell.

No. 447] NOTES AND LITERATURE. 231

“The Flow of Maple Sap” is the subject of Bulletin No. 103 of

the Vermont Agricultural Experiment Station. |

The influence of lime on plant growth is discussed by Wheeler and

Adams in Buletin 96 of the Rhode Island Agricultural Experiment

Station.

Vol. 4, no. 3 of the West Indian Bulletin is devoted to cotton.

The extent of variability in Eucalyptus is considered by Maiden

in Vol. 36 of the Journal and Proceedings of the Royal Society of New

South Wales.

A systematic-anatomical study of the leaf in Acer, with special

reference to the late elements, by Warsow, has been issued from the

Fischer press, of Jena.

An anatomico-biological thesis on seeds of Podalyriez, by Lindin-

ger, has been issued from the Fischer press, of Jena.

An interesting thesis on the anatomy and biology of the fruit and

seed of certain aquatics, by Fauth, has been issued from the Fischer

press, of Jena.

The principal species of wood and their characteristic properties

are described by Snow in an illustrated volume recently issued from

the press of John Wiley and Sons of New York

An illustrated account of Persian dates and their introduction into

. America, by Fairchild, forms Bulletin No. 54 of the Bureau of Plant

Industry of the U. S. Department of Agriculture.

Germinating spores in a fossil fern sporangium are described by

Scott in Zhe New Phytologist of January 27.

A preliminary notice on fertilization, alternation of generations

and general cytology of Uredinez is published by Blackman in 77e

New Phytologist of January 27.

New or unrecorded Australian fungi are being published by

McAlpine in the current Proceedings of the Linnean Society of New

South Wales.

An article on the genus Harpochytrium in the United States, by

Atkinson, is published in the Annales Mycologici, for November.

A paper on Italian Hypogaez, by Mattirolo is separately issued

by the Accademia Reale delle Scienze di Torino.

232 THE AMERICAN NATURALIST. (VoL. XXXVIII.

Bulletin 21 of the Boston Mycological Club is devoted to Agaricus

subrufescens.

Accounts are given, in the January Journal of the New York

Botanical Garden, of the laboratories of the institution, including

that in Jamaica, and of the Carnegie desert laboratory at Tucson.

A well illustrated account of the Desert Botanical Laboratory of

the Carnegie Institution, and of the desert regions of the Southwest,

by Coville and MacDougal, constitutes Publication No. 6 of the

Institution.

A note in the Journal of the Kew Guild for 1903, shows that in

1902 1,323,376 persons visited the famous botanical gardens at

Kew, the average for each of the previous ten years being 1,355,503.

Ramírez, in Anales del Instituto Médico nacional, vol. 6, no. 2,

publishes notes on some of the manuscript icones of Sessé and

Mociño.

“The Book of Herbs,” by Lady Rosalind Northcote (John Lane,

London and New York, 1903), is a tasty and interesting little book,

well illustrated, and with a portrait of Parkinson for frontispiece.

Those who care for old books will find interest in a supplementary

catalogue of the Sturtevant Prelinnean Library of the Missouri

Botanical Garden, by Hutchings, published in the 14th Aor? of

that institution.

An account of the botanical work that has been done in the

Philippines, with a bibliography, is given by Merrill in Bulletin no. 4,

of the Bureau of Agriculture of the islands.

The third part of vol. 2 of Wood's * Natal Plants," devoted to

grasses, and the first part of vol. 4 of the same work, containing

gamopetalz, have recently been issued.

A popular Sketch of Hawaiian botany, by Morrison, is contained

in Floral Life for November.

Some views of the vegetation of the Dismal Swamp. accompany

an article on the proposed ship canal through it, in Zhe American

inventor of December 1.

Dr. Holm contributes some Notes on Canadian Species of Viola

to The Ottawa Naturalist, for December.

No. 447.] NOTES AND LITERATURE. 233

Perrot and Guérin publish an account of the Didiereas of Mada-

gascar, with habit illustrations, in the Journal de Botanigue of

August-September.

A critical revision of Gossypium, by Aliotta, has been separately

printed from vol. 5 of the Annali della R. Scuola Sup. d Agricoltura

in Portici.

Andean cacti of interest are being described by Schumann in

current numbers of the MJonatsschrift für Kakteenkunde.

The Indian Species of Polygonum and reviewed by Gage in vol.

2, no. 5, of the Records of the Botanical Survey of India, dated

Sept. 14.

Part XX of Holm's Studies in the Cyperacez is contained in Zhe

American Journal of Science for December.

The comparative anatomy and phylogeny of Sequoia are discussed

by Jeffrey in vol. 5, no. 10, of the Memoirs of the Boston Society of

Natural History, issued in November.

A new Alstonia, yielding rubber, is described from New Caledonia

by Schlechter in Der Tropenpflanzer, for November.

Shade trees, etc, adapted to New Mexico are considered by

García in Bulletin no. 47 of the Agricultural Experiment Station of

New Mexico. :

An article on Conservation and Cultivation of Medicinal Plants,

by Kraemer, is contained in the December number of the American

Journal of Pharmacy.

An account of the grape-growing industry of the United States is

contained in Zhe National Geographic Magazine, for December.

Symbiosis of Volvox and Azotobacter is discussed by Reinke in

the Berichte der deutschen botanischen Gesellschaft, vol. 21, Heft 8.

A discussion of soil temperatures and vegetation by MacDougal,

is reprinted from the Monthly Weather Review, for August.

School Science, for December, contains a description of a new and

cheap form of Auxanometer, by Lloyd.

An account of a seemingly bacterial disease of tobacco, by

Stevens and Sackett, is given in Bulletin no. 188 of the North Caro-

lina Agricultural Experiment Station.

234 THE AMERICAN NATURALIST. (Vor. XXXVIII.

Some oddly grown trees are figured by Newcomb in oral Life,

for December.

President Seward’s address before the Botanical Section of the

British Association, at its recent meeting, is printed in /Vazure of

October 8, and deals with the composition and distribution of the

floras of the past, with reference to the corresponding phases of the

flora of to-day.

Harshberger contributes to part 2 of the current volume of Proceed-

ings of the Academy of Natural Sciences of Philadelphia a paper on

Mutations of Hibiscus Moscheutos and one on Form and Structure of

the Mycodomatia of Myrica cerifera.

A spontaneous rapid vibratory movement of certain shoots of

Eucalyptus is recorded by Tavares in Broteria of October 30, —

which contains a number of other articles of botanical interest.

The propagation of plants is discussed by Corbett in Farmers’

Bulletin No. 157, of the U. S. Department of Agriculture.

Data on autumnal coloring. as correlated with leaf deterioration,

are contributed by Keegan to ature of Nov. 12.

Anatomical studies of Potamogeton are applied to the classifica-

tion of the species of this difficult genus by Raunkiær in a paper

reprinted from Heft 3 of the current volume of Botanisk Tidsskrift.

An article by Fritsch on the Use of Anatomical Characters for

Systematic Purposes is contained in Zhe Mew Phylologist for

October.

Fendlera rupicola is illustrated in Curtis’s Botanical Magazine for `

November.

Ostenfeld and Raunkiaer, in Heft 3 of the current volume of

Botanisk Tidsskrift, show that Hieracium, like Taraxacum, appears

to be apogamic,—a peculiarity not shared by other genera of

Cichoriacez experimented on.

The distinctive marks of Catalpa speciosa are figured and described

by the Editor in Arboriculture for October.

A paper on Asplenium Ruta-muraria, by Christ, in Hedwigia of

October 7, includes an analysis of North American forms.

No.447] >` NOTES AND LITERATURE. 235

Numbers of Lloyd's Mycological Notes published during the current

year include studies of Catastoma, Mitremyces, Tylostoma, Secotium,

etc., accompanied by a good many photograms.

A third supplementary list of the parasitic fungi of Wisconsin, by

Davis, has been printed in advance from Vol. 14 of the Zransactions

of the Wisconsin Academy. A total of 661 species is now recorded.

Fascicle 4 of Sydow's Monographia Uredinearum carries Puccinia

to species No. 1094.

A paper by Holden and Harper, on nuclear division and fusion in

Coleosporium, is separately printed from Vol. r4, part I, of the

Transactions of the Wisconsin Academy.

Taphria (or Taphrina) cwrulescens forms the subject of Bulletin

No. 126 of the Alabama Experiment Station, by Wilcox.

Part 9 of Koorders and Valeton's Additamenta ad Cognitionem

Flore Arbore Javanice, forms No. 61 of the Mededeelingen uit

's Lands Plantentuin, published by Kolff of Batavia.

A primer of forestry, by Pinchot, constitutes Farmers Bulletin No.

173, of the U. S. Department of Agriculture.

A popular account of broom corn, Sorghum vulgare, by Hartley,

forms Farmers’ Bulletin No. 174, of the U. S. Department of Agri-

culture.

A reprint of Schópf's “ Materia Medica Americana” (1787) con-

stitutes Bulletin No. 6 (Reproduction Series No. 3) of the Lioyd

Library.

Palaguium Supfianum, a new Gutta-Percha plant of New Guinea,

is described and figured by Schlechter in Der Tropenpflanzer for

October.

A popular illustrated account of the pulque and mescal Agaves of

Mexico is published by Dodge in the Scientific American of Septem-

ber 19.

The Journals.— American Journal of Pharmacy, January : —

Lloyd, History of Echinacea angustifolia; Schneider, Gardens of

Medicinal Plants.

The Botanical Gazette, December: — Transeau, On the vagi

graphic Distribution and Ecological Relation of the Bog | nt

Societies of Northern North America; Berry, Aralia in American

236 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Paleobotany ; Ganong,,Vegetation of the Bay of Fundy Salt and

Diked Marshes (concluded); Eastwood, Notes on Garrya with

Descriptions of New Species and Key; Bergen, Transpiration of

Spartium junceum and other Xerophytic Shrubs; and Atkinson,

Geaster leptospermus — a correction.

Botanical Gazette, January: — Wylie, Morphology of Elodea cana-

densis; Newcombe and Rhodes, Chemotropism of Roots; Weld,

Botanical Survey of the Huron River Valley — II, A Peat Bog and

Morainal Lake; Godding, Southwestern Plants; Coker, Selected

Notes — III.; and Farmer, On the Interpretation of the Quadripolar

Spindle in the Hepaticz.

The Bryologist, January :— Harris, Lichens—Peltigera; Miller,

Pogonatum urnigerum; Grout, Notes on Vermont Mosses; Hol-

zinger, The Genus Hymenostomum in North America; and E. G.

Britton, Papillaria nigrescens.

Bulletin of the Torrey Botanical Club, January : — Arthur, New

Species of Uredinez , Harper, Explorations in the Coastal Plain of

Georgia during the Season of 1902; Murrill, The Polyporacez of

North America — VI, The Genus Polyporus; Robinson, the Spines

of Fouquieria.

Journal of Mycology, December:— Morgan, A New Species of

Berlesiella; Whetzel, New Method of Mounting Superficial Fungi ;

Bates, Puccinia Phragmitis in Nebraska; Stevens, Poisoning by

Lepiota Morgani; Elis and Everhart, New Species of Fungi;

Kellerman, Uredineous Infection Experiments in 1903, Minor

Mycological Notes — II, and Notes from Mycological Literature —

VII. A portrait of Atkinson forms the frontispiece.

Journal of the New York Botanical Garden, January : — Britton,

The Tropical Station at Cinchona, Jamaica; MacDougal, Research

Work in the Garden; and The Desert Botanical ance: 3h of the

Carnegie Institution.

The Ohio Naturalist, January : — Kellerman and Jennings, Report

for 1902 on the State Herbarium, including Additions to the State

Plant List; Schaffner, Poisonous and other Injurious Plants of

Ohio (concluded).

The Plant World, December: — Baum, The Breadfruit — III;

Safford, Extracts from the Note-Book of a Naturalist on the Island

of Guam — XIII ; Simpson, Effects on Vegetation of the Hurricane

No. 447] NOTES AND LITERATURE. 237

in Florida; Crawford, Some interesting Plants formerly abundant

near Germantown, Pa.; and Thompson, Boniato — a Tree or a

Yam?

The Plant World, January : — Safford, Extracts from the Note-

Book of a Naturalist on the Island of Guam — XIV ; Roosevelt, Our

Forest Policy; Tullsen, Notes from Pine Ridge Agency, S. Dak. ;

Dobbins, The Parsleys; Barrett, Correction and Comment; and

Gorman, Oregon Wild Flowers in Need of Protection.

Rhodora, December:— Fernald, Pursh's Report of Dryas from

New Hampshire; Ames, Lodelia X syphilitico-cardinalis; Collins,

Woodsia glabella in Maine; Robinson, Records of Wolffia in Mass. ;

Rand, Matricaria discoidea in N. H.; Chamberlain, New Stations fot

Maine Plants; Pease, Trisetum in Andover, Mass.; Freeman,

Lycopodium selago on Mt. Holyoke; Osmun, Cuscuta trifolia in

Mass. ; Harger, New Station for Phaseolus perennis; and Collins,

On Corallorhiza and Taraxacum.

Rhodora, January :— Fernald, Two Allies of Salix lucida;

Brainerd, Notes on New England Violets; Rehder, Pseudo-mono-

clinism of Chionanthus virginica; Wiegand, Some Notes on Galium ;

Seymour, A Newly Introduced Galium ; and Hervey, Plants new to

the Flora of New Bedford.

Torreya, December: — Britton, Cornelius Van Brunt (with por-

trait); Cavers, Explosive Discharge of Antherozoid in Hepatice ;

Earle, Key to the North American Species of Inocybe — II ; Maxon,

A Fern New to the United States; and Holm, Linnzus’ Work on

Ferns.

Torreya, January : — Stone, Physiological Appliances — I; Banker,

Observations on Phallus ravenelii; Canby, Joseph Hinson Melli-

champ; Berry, Primary Venation in Cinnamomum.

The Zransactions of the American Microscopical Society, Vol. 24,

contains the following articles of botanical interest: — Bessey,

Evolution in Microscopic Plants; Hollis, Two Growths of Chlamy-

domonas in Ct. ; Seawell, Method of Concentrating Plankton without

Net or Filter; and Bessey, Structure and Classification of the

Phycomycetes, with a Revision of the Families and a Rearrangement

of the North American Genera.

The American Botanist, of September, contains the following —

Bailey, The Defences of Plants; Bradshaw, Collecting Seeds;

238 THE. AMERICAN NATURALLIST. [VoL. XXXVIII.

Saunders, Poison Ivy and its Extermination ; Buchheister, Variations

in the Common Polypody ; Gilbert, The Jewel Weeds; and part 6

of the Editor’s Botany for Beginners.

The Botanical Gazette for October contains the following articles :

— Harshberger, An Ecological Study of the Flora of Mountainous

North Carolina; Parish, Sketch of the Flora of Southern California

(concluded) ; Ganong, Vegetation of the Bay of Fundy Salt and

Diked Marshes, An Ecological study (continued) ; Atkinson, A New

species of Geaster ; Davis, Tilletia in the Capsule of Bryophytes ; and

Lyon, Two Megasporangia in Selaginella.

Lhe Botanical Gazette, for November, contains the following : —

Evans, Odontoschisma Macounii and its North American Allies ;

Ganong, Vegetation of the Bay of Fundy Salt and Diked Marshes;

Harshberger, An Ecologic Study of the Flora of Mountainous North

Carolina (concluded) ; Moore, Mitoses in the Spore Mother-cell of

Pallavicinia ; and Haug, Is Detmer’s Experiment to show the Need

of Light in Starch-making reliable?

Volume 10 of the Proceedings of the Iowa Academy of Sciences con-

tains the following botanical articles : — Pammel, Ecological Notes on

the Vegetation of the Uintah Mountains; Weems and Hess, Chem-

ical Composition of Nuts used as Food; and Fitzpatrick, The Scro-

_phulariacee of Iowa.

The Proceedings of the second meeting of the Jowa Park and For-

estry Association, recently distributed, contain a number of articles of

botanical interest.

The Journal of the New York Botanical Garden, for November,

contains the following : — Britton, Report on Cuban Exploration;

Lloyd, Report of a Botanical Expedition to the Island of Dominica;

and Nash, Report on Exploration in Hayti.

A new publication, “Leaflets of Botanical Observation and Criti-

cism,” has been launched by Professor E. L. Greene. The first

signature is dated Nov. 24, 1903.

The Plant World, for November, contains the following : — Mac-

Dougal, Some Aspects of Desert Vegetation ; Safford, Extracts from

the Note-Book of a Naturalist on the Island of Guam — XII; Barrett,

A Forgotten Fruit; and Waters, Field Notes. .

Rhodora for October contains the following articles : — Robinson,

Insecticides used at the Gray Herbarium ; Fernald, A new Kobresia

in the Aroostook Valley; Cushman, Notes on New England Desmids

No. 447] NOTES AND LITERATURE. 239

— II; Wentworth, Two Plants new to the Flora of Lynn, Mass.;

Woolson, New Station for Asplenium ebeneum Hortonae; B. M

Britton, A New England Station for Buxbaumia indusiata ; Rand,

Galinsoga in Maine; and A Leaflet of the Seal Harbor Village

Improvement Society.

Part 3 of Zrees and Shrubs, issued Nov. 14, adds still further to

the number of American species of Crataegus.

- The Bulletin of the Torrey Botanical Club for October contains the

following articles : — Cannon, Studies in Plant Hybrids — The

Spermatogenesis of Hybrid Peas; and Evans, Hepatice of Puerto

Rico — III.

The Bryologist, for November, contains the following : — Britton,

The Splachnums; Holzinger, On Some Fossil Mosses ; Grant, Some

Moss Societies; Miller, Buxbaumia aphylla; and Clarke, Mounting

Mosses.

In addition to an important paper on Crassulacex, by Britton

and Rose, vol. 3, no. 9, of the Bulletin of the New York Botanical

Garden contains a paper by Berry on the Flora of the Matawan

Formation, one by Williams on Bolivian Mosses, and one by Zeleny

on the Dimensional Relations of the Members of Compound Leaves.

The Bulletin of the Torrey Botanical Club, for December, contains

the following articles: — Griggs, On Some Species of Heliconia ;

Underwood, Summary of our Present Knowledge of the Ferns of the

Philippines; and Kupfer, Anatomy and Physiology of Baccharis

genistelloides.

The Fern Bulletin, for October, contains the following articles : —

Gilbert, The Fern Flora of New York ; Clute, Fernwort Notes —IV ;

House, Scolopendrium from Canada; Eaton, The Genus Equisetum

in North America— XV; Clute, The Species-conception among the

Ternate Botrychiums ; and Druery, New Forms of Ferns. A portrait

of Mr. Maxon forms the frontispiece to the number.

Part II of the botanical portion of the, /n/ernational Catalogue of

Scientific Literature has been issued under date of November, 1903.

It is of nearly double the size of the first part (626 pages), and is

essentially on the same lines as the earlier part.

The Journal of Mycology, for October, contains the following

articles: — Morgan, Some Western Specimens; Morgan, Note on

Corticium leucothrix ; I. Kellerman, The Accentuation of Mycological

240 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Compound Names; Ellis and Everhart, New Species of Fungi from

Various Localities; Kellerman, Minor Mycological Notes — I ; Ohio

Fungi, Fascicle VIII [descriptions and annotations]; Index to

North American Mycology (continued) ; and Notes from Mycological

Literature — VI.

The Journal of the Royal Horticultural Society, for October, con-

tains a number of botanically interesting articles.

The fourteenth Aor? of the Missouri Botanical Garden, in addi-

tion to the customary administrative reports, contains a Supple-

mentary Catalogue of the prelinnean library of the Garden, and a

revision of the genus Lonicera, by Rehder.

Rhodora, for November, contains the following : — Ames, Hybrids

in Spiranthes and Habenaria; Leavitt, Reversionary Stages in

Drosera intermedia; Waters, Asplenium ebenium proliferum; Eaton,

Notes on Botrychium tenebrosum; Deane, Gaylussacia in New

Hampshire — a Correction ; and Eaton, New Varieties of Isoetes.

Torreya, for October, contains the following: — Underwood, The

Early Writers on Ferns and their Collections — 1, Linnzus; Howe,

Note on the *Flowering" of the Lakes in the Adirondacks ; and

MacKenzie, A New Genus of North American Umbelliferz

[ Pseudoteenidia].

Torreya, for November, contains the following articles: Wooton,

Ferns of the Organ Mountains; House, Notes on the Flora of

Oneida Lake and Vicinity; Earle, Key to the North American

Species of Inocybe— :; Berry, A Question for Morphologists ;

Stone, Arisema pusillum in Pennsylvania and New Jersey; and

Nash, A New Bamboo from Cuba.

CORRESPONDENCE.

To the Editor of the American Naturalist:

SIR:— In a suggestive article in the American Naturalist for

August (XXXVII, 551-555), on * Vernacular names of animals,”

Dr. Edwin W. Doran remarks, * I believe that I am the first who

has thought it necessary to prepare a synonymy of the vernacular

names of animals" and announces that *the author has in prep-

aration a synonymy of all the vernacular names of vertebrates.”

It is evident from this, and from interviews had with other natural-

ists, that a great work on the subject published more than a century

ago is unknown, to most who are especially interested in its subject-

matter. Indeed, I have found that none of the naturalists of

Washington whom I had asked knew about it, so completely has it

been forgotten. Nevertheless, it is a work in four large quarto

Volumes of about 2400 pages with 4783 numbered columns and some

additional leaves (preliminary) as noted below. I give the title of

the work and analysis of the volumes.

General title.

Allgemeines | Polyglotten-Lexicon | der | Naturgeschichte | von

| Philip Andreas | Nemnich, J. U. L. | mut. mut. See below. | Zu

finden | Hamburg, bey Licentiat Nemnich | [etc.]

The title page of the first part is printed from a plate; those of

the other parts from regular type. They are paged for or were

bound in four volumes as follows :—

[Erste Lieferung. A—Canus.] Eng. title page + 8 p. l. + col. 1-

540. 1793.

Zweyte Lieferung. | — | Cap. — Fus. | title page + col. 841—1684.

1793.

. 241

242 THE AMERICAN NATURALIST. (Vor. XXXVIII.

II.

Dritte Lieferung. | G—N. | title page + col. 1-740. 1794.

Virete Lieferung. | O—Z. | title page + col. 741-1592. 1795.

Ill.

Fünfte Lieferung. | — | 1) Deutsches Worterbuch der Naturge-

schichte. | 2) Englisches Wórterbuch der Naturgeschichte. | title page

+col. 1-676 (= Fünfte Lief. Deutsches W.) + col. 677-1056 (=

Sechste Lief.). [No date.]

Siebente Lieferung. | — | 1) Frazósisches Wörterbuch der Natur-

geschichte | 2) Italienisches Worterbuch der Naturgeschichte. | title

page+col. 1057—1376 (= Französisches W.)--col 1377-1507 (=

Italienisches W.) [No date.]

Achte und letzte Lieferung. | — | 1) Spanisches Wórterbuch der

Naturgeschichte. | 2) Hollaindisches Wörterbuch der Naturgeschichte.

| 3) Schwedisches Wórterbuch der Naturgeschichte. | 4) Dànisches,

Norwegisches und Islandisches Wórterbuch der Naturgeschichte. |

5) Einige Nachtrüge. | title page + col. 1509—2108. 1798.

This bibliographical description will suffice to give some idea of

the nature and extent of the work. It is only necessary to add that

in the first 2 volumes, under each Linnzan genus, alphabetically

arranged, the species are enumerated under their Latin names in

alphabetical order and the vernacular names in different languages

added under the Latin ones. Thus, under Anas boschas domestica,

as “ Engl" 12 names applied to the subspecies as a whole in

English as well as Gaelic, Welsh and Cornish, 3 of the drake, and

3 of the young 1 A given; under Anas boschas fera 4 names are

given as “ Engl.

It cannot ab claimed that the English part is well done but the

author at least “ thought it necessary to prepare a synonymy of

the vernacular names of animals" and carried the idea into execution

in a very voluminous work.

Another noteworthy work carrying out the same idea for certain

North American birds is Gurdon Trumbull's volume on * Names and

Portraits of Birds which interest gunners," published in 1888.

Numerous minor contributions to the same subject have been

published.

THEO. GILL.

(Vo. 446 was mailed April 28, 1904).

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Vou. XXXVII, April, 1904. No. 448.

THE ANATOMY OF THE NORTH AMERICAN

CONIFERALES TOGETHER WITH CERTAIN

EXOTIC SPECIES FROM JAPAN AND

AUSTRALASIA. PART T

D. P. PENHALLOW.

INTRODUCTORY.

THE present work had its origin in 1880, in an attempt to

construct a system of classification for the North American

Coniferze, based upon the anatomy of the vascular cylinder of

the mature stem. The fundamental idea was that such a classi-

fication would prove of great value in the identification of

material used for structural purposes, but investigations had not

been carried very far when it became manifest that some such

arrangement was imperatively demanded in other directions and

for purposes of a more strictly scientific character. In entering

upon the study of fossil plants it was recognized that the most

fruitful source of reliable data must be found in the stem struc-

ture. At that time there was little in the way of an adequate

basis for further study of this sort, inasmuch as the current

diagnoses of the vascular structure were found in most cases to be

singularly inadequate, and often so incorrect as to require exten-

243

244 THE AMERICAN NATURALIST. [Vor. XXXVIII.

sive revisions. It was furthermore found that, in order to reach

correct conclusions in the case of stems which must often pre-

sent marked structural alterations arising through the influence

of decay and other conditions attending fossilization in its various

forms, it was indispensable that there should be a trustworthy

means of comparison with existing types, whereby sources of

error arising from eliminated structures might be definitely

excluded, and the fossil referred with certainty to its nearest

relative. The original intention was therefore modified with a

view to meeting the requirements of palzeobotanical research.

During the twenty odd years these investigations have been in

progress, there has been much change in the views held by

botanists respecting the significance of anatomical features as

affording evidence of descent, and our own studies brought fcrth

facts which gave repeated emphasis of the most positive kind, to

the idea that questions of phylogeny cannot be settled either by

the morphologist in the narrower sense or the physiologist,

either separately or combined, and that a proper historical point

of view can be gained only when to such labors we join the data

derived from a critical study of the stem structure in all its

details. As the greater may always be held to include the less,

the present discussion is to be regarded primarily from the bio-

logical point of view, and questions of descent will take preced-

ence over those of mere taxonomy.

The original intention was to make a complete study of all the

North American woods, comprising, as enumerated by Sargent

in his report in the Tenth Census of the United States, some four

hundred and nineteen species and varieties ; but the great impor-

tance of the Coniferze from an economic point of view, their fre-

quent representation in the fossil state and their relatively more

simple structure, eventually led to their selection as the one

group in which initial studies might be prosecuted with the most

immediate and profitable results. While the North American

species constituted the original basis, various exotic species were

added from time to time with the result that our studies as now

completed, comprise ninety-two species from North America,

twenty-one. species from Japan, and four species from Austra-

lasia. This extension has proved of great value, not only froma

No. 448.] NORTH AMERICAN CONIFERALES. 245

paleeontological point of view, but also because of the important

bearing such exotic types have had in the solution of questions

relating to descent. The methods adopted in these investiga-

tions have already been fully explained (38, p. 33 eż seg.) and

further details at this time are uncalled for.

In 1896 the work had reached such a point that it was

deemed desirable to make a preliminary statement of results.

This appeared in a paper published in the Transactions of the

Royal Society of Canada entitled the “Generic Characters of

the North American Taxaceæ and Coniferz," in which it was

shown that generic differentiations were possible. The diag-

noses and artificial key to the genera thus published, have been

in constant use since that time for the determination of fossil

species, with the most gratifying results ; and after seven years

of severe testing, and in the light of more extended studies, they

are found to be substantially correct and reliable.

The question of specific differentiation presented a far more

difficult problem, the solution of which has now been reached as

embodied in the present work.

In the prosecution of these studies I have been under deep

and often constant obligations to helpful friends working along

other lines. To Prof. C. S. Sargent of the Arnold Arboretum,

as also to Mr. Morris K. Jesup, President of the American

Museum of Natural History; Dr. N. L. Britton, Director of

the New York Botanical Gardens; the late Baron Ferd. von

Mueller of Melbourne, Australia; Sir W. T. Thistleton-Dyer,

Director of the Royal Gardens, Kew; and to Mr. E. J. Max-

well of Montreal I am indebted for much valuable type

material. To Dr. B. E. Fernow, formerly Chief of the Bureau

of Forestry of the United States Department of Agriculture, I

am under obligations for a large amount of material specially

selected with reference to testing the accuracy of diagnoses and

details of the key. To Mr. J. G. Jack of the Arnold Arboretum,

I am under particular obligations for the readiness with which he

has frequently responded to requests for material of a trust-

worthy character, and his care in selecting a large series of

specimens for testing purposes, which have contributed very

largely to the success of the final results. To my assistant,

246 THE AMERICAN NATURALIST: (VoL. XXXVIII.

Miss C. M. Derick, I wish to extend my appreciation of the

frequent and valuable assistance she has rendered.

SPIRAL TRACHEIDS.

In the genesis of the vascular system, the elements first

differentiated from the generative tissue, constitute the primitive

elements now generally recognized in accordance with the desig-

nation of Russow as protoxylem. These elements are tracheids,

and in the Coniferales as also in the Ginkgoales, they always

occupy a position immediately external to the pith and therefore

on the inner face of the zone representing the growth of the

first year, but they are not repeated in the formation of xylem

structure in subsequent years. The primitive character of such

tracheids is therefore indicated, not only by their position and

relation to development of other parts, but it is also exhibited by

their occurrence in plants where the vascular system is of a far

more simplified type and of which they constitute relatively more

prominent features. In the Equisetinez, both fossil and recent,

they are conspicuous elements of the vascular structure, being

found within the limits of the carinal canal. They similarly

occur in the Ophioglossaceze and elsewhere among the more

primitive of the vascular plants. The general evidence, then,

which may be derived from a comparative study of various types,

tends to enforce the idea that, originating as a primitive form of

the wood structure, and more or less common to all the vascular

plants, they are relatively predominant in the lower forms, with a

tendency to obliteration through replacement or modification in

the higher types, where their presence may be held to represent

a survival of ancestral characters, This view gains additional

support from a study of the peculiar structural variations which

characterize such tracheids, and the progressive modifications

which they have been found to undergo in relation to the

development of the secondary xylem.

The protoxylem elements are distinguished by the presence of

thin, spiral bands disposed upon the inner surface of the primary

wall in such a manner as to afford a substantial measure of

mechanical support. These ribbon-like bands represent second-

No. 448.] NORTH AMERICAN CONIFERALES. 247

ary growth in thickness of a local nature — the localization

being determined with reference to the requirements of such

support in the first instance. De Bary (9, p. 57) has shown

that they exhibit considerable variety in the number of fibres

and the direction and steepness of the coils. Their number is

often only 1—2 in narrow tubes which are first formed when the

differentiation of tissues begins, in others 4 or more, and it rises

in many cases in the angiosperms to 16-20. He has furthermore

pointed out that the steepness of the coils is greatest in those

tubes which are developed earliest, before the extension of the

part to which they belong has ceased; since in these the coils

are separated from one another by the extension which the tube

itself undergoes. These facts appear to suggest that the more

typical the spiral tracheid is, the more fully does it emphasize

the idea of a primitive structure ; but that as the spirals become

more dense or closer, there is a tendency toward more uniform

and less localized secondary growth of the wall, as expressed in

the structure of the higher types of plants or the secondary

xylem elements of the Conifere. In confirmation of such a

view, reference may be made to the commonly observed fact

that the spirals tend to a more compact arrangement at the ends

of the elements, becoming correspondingly more open in the

central region; and likewise to the well known transitional

forms which these structures exhibit, whereby their original

characteristics are completely lost as they merge ultimately into

tracheids devoid of spirals, but characterized by the presence of

pits of various forms. In 1840, Don ( 52) pointed out that the

tracheids of Cycas revoluta exhibit scalariform structure at one

end and bordered pits at the other. This fact has more recently

been commented upon by Williamson who observed the same

fact independently, and drew from it the inference that a defi-

nite relation exists between the scalariform markings and the

pit structures of such a nature that the one is the natural

successor of the other. In Ginkgo biloba which is now gener-

ally conceded to represent a much more primitive type than the

Conifere, though more advanced than the cycads, precisely

similar transformations are to be met with. The evidence of

fossil plants is quite as convincing as that derived from,existing

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250 THE AMERICAN NATURALIST. (VoL. XXXVIII.

species, while it is often of distinctly greater value because

affording data derived from presumably more primitive types.

Thus such transitions are well known, though of a relatively sim-

plified form in the structure of the calamitean stem, and an

excellent example of this kind is afforded by a figure given by

Scott (43, p. 22). In 1869 Williamson (52, p. 69) directed

attention to the occurrence of such transformations in the tra-

cheids of Cordaites (Dadoxylon). A more recent study of this

genus (39, p. 57 and 43, p. 418) has shown that this feature is

well exhibited in C. brandlingii, where a suitable radial section

(Figs. 1-4) will present a more or less graduated series of transi-

tions from the typical spiral tracheid of the protoxylem, through

scalariform structures to the multiseriate bordered pits of the

tracheids in the secondary xylem ; while within the limits of the

same tracheid, such transitions may be observed as it were, in

process of development, affording the most conclusive proof in

this respect. These transitions as observed in Cordaites brand-

lingit show the following phases :

In the successive radial development of new tracheids, there

is a constant tendency to a more uniform thickening of the cell

wall by secondary growth. This at first finds expression in the

more Compact arrangement of the spirals which later coalesce at

various points, thus giving rise to more localized areas devoid

of secondary: growth, and hence to a scalariform structure in

which the general lines conform more or less exactly, to the

direction of the original spirals. By a further modification the

elongated, thin areas become converted into shorter, often isodi-

ametric areas substantially by a process of division. A further

tendency to general thickening of the walls causes the margins

of the scalariform structure to project from all sides and extend

over the area of arrested growth as a lip which never completely

closes at the centre, where there is left a usually circular, some-

times oval or again lenticular or even oblong opening, and in this

manner the bordered pit is formed.

ion se which these changes occur, or the “ transi-

see a boda e c io great variation whereby the change from

radial za o rs may arise very gradually through a broad

, ordaites brandlingii or it may occur very

No. 448.] NORTH AMERICAN CONIFERALES. 251

abruptly as in the modern Coniferee. The general tendency of

such evidence is to show that with a higher type of organization,

there is a corresponding diminution in the transition zone and

increased abruptness in the structural alterations. The logical

result of an extension of this process would be the reduction of

the bordered pit to the condition of a simple pit, and ultimately,

its complete obliteration. In the Coniferze the reduction of the

bordered pit to the condition of a simple pit sometimes occurs in

the case of medullary rays or even in the case of tracheids with

very thick walls, but it becomes most prominent in the angio-

sperms where it is a characteristic feature. Instances also occur

in some of the hard pines, in which the pit is completely obliter-

ated. This applies in particular to tracheids of the summer

wood, the walls of which have become unusually thickened.

The relations to which attention has thus been directed some-

what in detail, have been expressed in more general terms by

De Bary (9, p. 321) in the statement that “ Outside the primi-

tive elements, wider trachz follow. Their development takes

place successively, advancing from the inner edge of the bundle

outwards, and as a rule at a time when the elongation of the

entire part to which they belong is nearly at an end. The

thickenings on their walls therefore have a successively denser

arrangement: dense spiral and annular trache, then reticulated

and pitted trachz follow one another in succession from within

outwards, with gradual transitions, or with the omission of one or

the other immediate form." It is probably a justifiable inference

from the preceding facts that, the relation which exists between

the spiral tracheids of the protoxylem and the pitted tracheids

of the secondary xylem in the Coniferze, is, in general terms and

from the standpoint of development, the same as that exhibited

between the lower and higher types of vascular plants. sab

Accepting the general principle which appears to be justified

by the foregoing facts, that the transition from spirals to bor-

dered pits is a feature in development which bears a direct rela-

tion to the evolution of higher types of organization, we may

utilize it for the purpose of determining the general phylogeny of

the Coniferz so far as they may show a sur vival of such ——

ters. Out of a total of 117 investigated species of indigenous

252 THE AMERICAN NATURALIST. [Vor. XXXVIII.

and exotic Coniferz, 9.4% show a more or less permanent sur-

vival of the spiral structure within the limits of the secondary

xylem. Of these 6 % fall within the Taxaceæ (Torreya 3.45 f,

Taxus 2.5%), while in the Conifera the remainder is divided

between Pseudotsuga (1.7%), Larix (0.86%) and Pinus (0.86 5).

In the genus Torreya the spirals are, on the whole, rather open

and distinguished by being 2—4 seriate. They are typical

throughout the spring wood, but in the thin summer wood they

quickly become vestigial and ultimately disappear altogether.

In 7. taxifolia there is also a marked condensation whereby the

spirals are all brought into a more compact series within the ear-

lier tracheids of the summer wood. All of these changes appear

to be directly related to a progressive increase in the thickness

of the tracheid wall. ]

In all investigated species of Torreya, there is a rather wide

variation in the angle which the spirals make with the axis of

growth, and this becomes most pronounced in T. californica,

which gives the lowest angle for any species of either Torreya

or Taxus. Usually the spiral has an angle quite distinct from

that of the lines of striation in the cell wall, but in 7. taxifolia

the two often coincide. The following will show the various

details derived from the average of ten measurements for each

species.

Average Highest Lowest Extreme

angle. angle. angle.

Torreya nucifera Gard cmt es o. 87.0 57.0 30-0

í00 08i olia . 70.4 77.0 61.0 16.0

californica . 63.0 30.0 33.9

Means . 62.3 75.7 49.3 26.3

In the genus Taxus, the spirals are rather close and in 2,

rarely 3 series. Asin Torreya they are typical throughout the

spring wood, and show a pronounced tendency to obliteration in

the summer wood. This tendency is subject to considerable

variation in different species. In 7. canadensis the spirals are

conspicuous throughout. In 7. floridana they usually disappear

No. 448.] NORTH AMERICAN CONIFERALES. 253

in the later growth and are wholly wanting in the two or three,

last formed tracheids. In 7. brevifolia they become very imper-

fect in the outer summer wood and tend to disappear completely,

only vestiges remaining in the last formed tracheids. In 7. cus-

pidata the spirals are generally absent from the summer wood,

or when present, they are merely vestigial. The angle is some-

what greater — about 7 deg.— than in Torreya, and this fact is

apparent with respect to certain species without special measure-

ment. The four species appear to be paired off in such a way

as to représent a mean difference of about 10.9 deg. as between

T. canadensis and T. floridana on the one hand, T. brevifolia and

T. cuspidata on the other. In all cases the angles of the spirals

are quite distinct from those of the lines of striation. The fol-

lowing details are based upon the average of ten determinations.

avenge | Hie | done | Bonet | Moan ot

Taxus canadensis . 72.4 88.0 66.0 22.0

"e ornidand . . . 78.4 90.0 72.0 18.0 75:4

Ho Mya o n 63.0 76.0 55.0 21.0

* cuspidata 66.1 87.0 45-0 42.0 64.5

Meats >: 69.9 85.2 59-5 25.7

A comparison of these results in detail emphasizes the fact

that the distribution of the spirals, as between spring and sum-

mer wood, is in direct harmony with the principles already stated,

and furthermore, that the angles at which the spirals develop do

not afford an adequate basis for generic differentiation. It is

nevertheless possible to recognize sub-generic groups in such

wise that in both genera a general line of division may be estab-

lished at 70 degrees. In the case of Torreya californica, the

very low angle of 46.2 degrees may be regarded as a differential

character of specific value.

In the genus Pseudotsuga, spirals are confined to the tra-

cheids of the spring wood. This has a partial exception in

P. macrocarpa, in which vestigial spirals may be observed in the

earlier tracheids of the summer wood. In this species the mean

254 THE AMERICAN NATURALIST. [VoL. XXXVIII.

angle is 70 degrees, but the spirals are always characterized by

lack of prominence, they are often widely distant, and the some-

what extended areas within which they are wholly wanting or

fragmentary, suggests a process of obliteration. In P. douglasst

the average angle is 82 degrees ; the spirals are characterized by

considerable prominence and they are also, on the whole, close.

In this genus these two factors obviously possess a well defined

differential value with respect to the two species.

Among the higher genera only two cases are known in which

spirals occur, but in each the character is of a very sporadic

nature. In Larix americana spirals are frequently found in the

summer wood, but they are so inconstant in their occurrence,

and they present such varying aspects, that the angle cannot be

determined. In Pinus téda, where the walls of the summer

tracheids are very thick, rudiments of spirals may sometimes be

seen. Here also it is manifestly impossible to determine the

angle.

Viewing these five genera collectively, their spirals conform

fully, in their occurrence and relation to progressive develop-

ment, to the general principles already stated, and especially as

formulated by De Bary. They possess no differential value of

generic rank with respect to Pinus and Larix, but they do have

such value with respect to Torreya and Taxus on the one hand,

and Pseudotsuga on the other, the differentiation resting upon

their occurrence in the summer wood in the former, and their

exclusion from that region in the latter. Were any question to

arise in this connection, it could be authoritatively decided by

the definite association of resin passages and fusiform rays in

Pseudotsuga.

It only remains for us to ascertain how far such structural

features may be employed as a basis upon which to determine

the general phylogeny of the genera. As between Torreya and

— there is very little upon which to base conclusions

respecting sequence in development, and it is apparent that both

of these genera have attained to nearly the same level. Such

differences as do exist, however, seem to point to the relatively,

though slightly, inferior position of Torreya as indicated by (1)

the smaller angle in that genus and (2) the generally more com-

No. 448.1 NORTH AMERICAN CONIFERALES. 255

pact spirals of Taxus. This fact, so far as it possesses phylo-

genetic value, appears to confirm the conclusions as to the rela-

tive positions of these two genera, already determined upon the

basis of external morphology as stated by Eichler (11, p. 108).

It has already been made clear from the preceding facts, as

well as from former discussions (40, p. 56) that in the case of the

Taxacez and also of Pseudotsuga, the spirals must be regarded

as a survival of primitive structures. On a former occasion

(40, p. 57) I was inclined to consider that their occurrence in

Larix and Pinus teda was atavistic, but in the light of more

recent evidence as now set forth, this opinion requires modifica-

tion in so far as to include the idea that they do not express

mere parallelisms in development, but that they afford evidence

of a common ancestral type at some point far anterior to the

evolution of the Taxaceze. We must therefore consider that

Torreya, Taxus, Pseudotsuga, Larix and Pinus represent differ-

ent branches of a general phylum — undoubtedly including also,

other closely related genera in which the spirals have been

wholly obliterated — which had its origin at a point anterior

even to such types as Cordaites, and therefore, in all probability

in what Coulter (7) has very happily designated as “the great

Cordaitean plexus " arising from the eusporangiate ferns, or

what, according to our more recent knowledge, and Coulter's

more recently expressed view (8, p. 172) would be designated as

the Cycadofilices. :

BORDERED Pits DISTRIBUTION AND STRUCTURE.

In the preceding pages, the derivation of the bordered pit from

the spiral tracheid, and its obviously more intimate relation to a

higher type of development have been made clear. Our present

purpose is to discuss these structures with special reference to

(1) occurrence, (2) distribution, (3) structural modifications, (4)

taxonomic value and (5) their value as evidences of descent.

At the outset, reference may be made to the occurrence of

bordered pits on the radial walls of the ray cells as exhibited

typically in Sequoia and Taxodium. Their location in such situa-

256 THE AMERICAN NATURALIST. [VOL. XXXVIII:

tions, as also upon the radial, terminal, upper and lower walls of

the ray tracheids may be held to represent a feature somewhat

distinct from their presence on the walls of the wood tracheids,

and their consideration properly belongs to a discussion of the

medullary ray as a whole; but it may be observed that they

constitute a characteristic feature in the structure of the ray

elements in the great majority of the Coniferales,

The occurrence of bordered pits on the walls — especially the

radial walls — of the wood tracheids in the Ginkgoales and Coni-

ferales, is'much too familiar a fact to call for special discussion

at this time, but reference may be made to the additional fact

that their characteristic structure is such as to permit of their

use for the general purpose of tracing possible lines of descent -

through such extinct types as Cordaites and the Cycadifilices.

It is true that similar bordered pits originating in modifications

of spiral structures, are to be met with, often in great numbers,

in the higher angiosperms, but in such cases the associated

structures permit of a clear and definite differentiation of all

such woods from the Coniferales.

Radial walls.— The characteristic situation of the bordered

pits is on the radial walls where, as was shown many years since

by De Bary (9, p. 160), “ the pits of contiguous tracheids always

correspond to one another in such a way that on each limiting

surface, all the cavities of the pits of one fit exactly over those

of the other. The plano-convex cavities are thus applied to one

another in pairs so as to form the lens-shaped pit cavities " as seen

in tangential section. But on surfaces abutting on elements of

another order, e. ¢., parenchyma cells, the bordered pits of the

tracheids correspond to non-bordered pits, or they are opposite

an unpitted wall. Four typical variations of the bordered pits

may be recognized:— (1) the multiseriate, when they are dis-

posed in any number of rows more than two, (2) the 2-seriate,

(3) the uniseriate with occasional pairs of pits, and (4) the

strictly uni-seriate. The general sequence thus presented will

be found to be in direct accord with the evolution of higher types

of structure and organization.

The most primitive type of gymnosperm presenting a multi-

seriate arrangement, is the genus Cordaites. Among eleven

No.448] NORTH AMERICAN CONIFERALES. 257

species of this genus which have been critically studied within

recent years (39) there is a general agreement in the constancy

of this character which thereby becomes of generic value. In

all the species the pits are disposed in such a compact manner

throughout the entire extent of the tracheid, as to present a hex-

agonal outline. In Cordattes acadiamum, they are 2—5-seriate

(Fig. 5). In other species they vary from 2-seriate in C. kamil-

tonense and C. newberryi (Fig. 6) to occasionally 4-seriate in C.

clarkei. In the majority of species, the rows are not constant,

but show a varying number from 1 to 3, or 2 to 5, this variation

"e! 2,

CEE Ir IXLIXE

[ E.

LARA

* -ow

ENSIJIXIT

ww wow

wrtiag ett dBtir

"JG ow» ww EP

&seee ee

ate eve

ELLE

ear

v S

Y "n

Ae VAAL

ptr RRO A es

as x LL.

e ww»

gr’

“A

evtt

wee Sad YEY

oe " N

LIFT

(LL

LES

4 €

/ e*.

^ yYY

FIG. 5.— Cordaites acadi

Radial section showing disposition of the bordered pits. X 180.

being exhibited by adjacent tracheids in accordance with the

variation of the latter in radial diameter ; and viewing this dis-

tribution as a whole, it cannot be doubted that it represents

corresponding differences in: development. One of the most

striking features of the genus is to be met with in C. newberryi

(Fig. 6), which is unique in the segregation of the pits into groups

of 6-13. !

In Araucarioxylon Knowlton has shown (25, p. 614) that,

while conforming to the characteristic form and compact arrange-

ment presented in Cordaites, the pits exhibit far less constancy

in their serial arrangement, and in this respect they are at once

258 THE AMERICAN NATURALIST. [Vor. XXXVIII.

comparable with those of the existing Araucarias. Among the

latter, A. cunninghamii shows a I-3-seriate disposition; A.

excelsa is 1—2-seriate while A. bidwillii is strictly I-seriate. All

Fic. 6.— Cordaites newberryi. Radial section showing disposition of the bordered pits. X 180.

of the extinct species as comprised in the genus Araucarioxylon

not only show similar variations, but such variations are found

to cover a much wider range. A comparison of all the species,

both recent and extinct, now available for that purpose, is of

interest in this connection.

A. n: :

wurtember: riani

sch iden um

hugelianum

‘i

e. cce sa

E T | 2-ser. 3-ser. 4-Ser-

H o xXx

"EN M E M E

1 å

ars eilliacola :

Aeerü. " c à : A : : |

cunninghamii i ue | | |

robertianu Ss |

No. 448.] NORTH AMERICAN CONIFERALES. 259

Such a comparison brings into strong relief the fact that the

Araucarias, both past and present, constitute a transitional `

group with a somewhat wide range of variations, and in this

respect they may be said to stand between the more stable Cord-

aites and Agathis on the one hand, and the far more variable

Coniferz on the other, since in Agathis australis we find essenti-

ally the same features of structure and distribution as in Cord-

Fic. 7.— Pinus cubensis. Radial section showing the form and disposition of the bordered pits. X

280.

aites, the pits being 1—3-seriate. The sequence presented above

may be held to be in the inverse order of development, and A :

robertianum must therefore be held to represent the most primi-

tive form. i

It is apparent that in Cordaites, Araucaria (including Arauca-

rioxylon) and Agathis, the pits are invariably distinguished by

two constant features; (1) their hexagonal form and (2) their

very compact disposition throughout the entire extent of the

tracheid. "They often deviate from the multiseriate arrangement

typical of the group as a whole, in that in individual cases they

are reduced to a t-seriate arrangement. They thus tend to

overlap the next group which is distinguished by a 2-seriate dis-

position, but any confusion which might arise from this cause,

may be overcome by reference to the special form and dis-

260 THE AMERICAN NATURALIST. (Vor. XXXVIII.

. position of the pits as will more fully appear in the following

lines.

Among the remaining Coniferales, 20 species of various gen-

era, or 17.2 % in all, show a 2-seriate arrangement, and to this

group we must also add the Ginkgoales and various fossil spe-

cies. Here the multiseriate disposition of the pits involves

features which at once distinguish the group as a whole from

the preceding, clearly placing.it upon a higher plane of develop-

ment. The pits are never hexagonal but generally elliptical or

round, while they also show a strong tendency to extreme seg-

regation. When brought into a compact arrangement as in

Cupressoxylon, Sequoia or various species of Pinus, they are

. flattened only along the lines of limited contact, which are

usually confined to one end (Fig. 7). A very characteristic fea-

ture of this group is the further fact that the 2-seriate arrange-

ment is not constant, either in the same section or in the same

tracheid. Both Pinus teda and P. cubensis, as also Larix ameri-

cana and Tsuga canadensis afford illustrations that while typi-

cally 2-seriate, a given section may show a strictly I-seriate

arrangement, and this difference also obtains as between con-

tiguous cells. In all such cases examination will show that the

variation is directly related to the relative size of the tracheids

in such a way that the narrower tracheids, or those arising from

a less vigorous growth, are r-seriate. Within the individual

tracheid there is a strong tendency to a r-seriate arrangement

in the central region, while it is 2-seriate at the extremities ;

and this law holds so true that in those species which are excep-

tienally 2-seriate, judgment should be reserved until it is seen

that the r-seriate form holds throughout.

The antithesis of the multiseriate type is found in the I-ser-

ite form. This is typical of 50 % of all the species included in

the present studies. In such cases the form of the pit is never

hexagonal or specially flattened. When the disposition is some-

what compact, as in Pinus strobus (Fig. 8), the outline becomes

more or less strongly elliptical, but as the segregation is more

pronounced, a definitely circular form prevails (Fig. 9). Within

the limits of the individual tracheid the same law of distribu-

tion obtains as in the 2-seriate type, whereby segregation is

always most pronounced in the central region. !

No. 448.] NORTH AMERICAN CONIFERALES. 261

Between species of the strictly r-seriate, and those of the

strictly 2-seriate type, there is an intermediate or transition

group comprising 34 species, or 29.3 % of the investigated

species, into which members of the first

two groups may occasionally be pro-

jected. The distinguishing feature of

this group is the occurrence of pits in

pairs which are usually distant, and in

no case so numerous as to distinguish

a 2-seriate disposition. They give un-

doubted proof of the passage from one

type to the other. Like the 2-seriate

type, this feature is not confined to any

one genus or to any particular group of

genera, but it applies with equal force to

any genus, the members of which may

therefore represent any or all of the three (o)

types here specified.

Viewing the distribution of the bor- | ;

dered pits from the standpoint of zonal eot i. Hague :

development, it is found to be universally disposition of the bordered pits.

true that, in the earlier spring wood there

is the strongest tendency to a multiseriate

O) (9) arrangement. With a radial increase of the

xylem, this tendency constantly diminishes, with

Q the general result that the pits become more

o summer wood in which they are sometimes

wholly obliterated — this being the case when

(0) (0)||| the cell wall acquires unusual thickness.

Upon careful examination, the foregoing facts

©) Q will be found to be in exact accord with the law

formulated by De Bary with reference to varia-

o tions in the structure of spiral tracheids and the

genesis of bordered pits as already stated. In

FLR inus strobus. a cordance with this law it is possible to con-

ing the bordered pits clude that relatively rapid growth is coördinated

er wood: with a primitive development, while the converse

DIXIT

262 " THE AMERICAN NATURALIST. [VorL. XXXVIII.

is true of a slow rate of growth which is again convertible into

terms of maturity. On this basis we may present the following

general outline of sequence in development, as preliminary to

further and more detailed discussion of phylogeny.

Cordaites. 2—5 seriate, hexagonal pits.

Araucarioxylon. I-4 “ « « | Compact through-

Araucaria. I-3 “# e « . 1 out the tracheid.

Agathis. 1-3 Tm &é m J

Ginkgo

nkgo. I-2 “ round or oval pits.

Higher Coniferales. 1-2 — « E

“a t

] More or less, often

strongly segre-

[14 LI [13

I-ser & pairs «€ « « «

M ss I—seriate. « a “ gate

Tangential Walls. —The occurrence of bordered pits on the

tangential walls is a well-known and characteristic feature of the

Coniferales. In the case of fossil forms, to which Araucarioxylon

offers a partial exception, it cannot be satisfac-

torially demonstrated because of the peculiar

| alterations of the cell wall, but that it is present

we are permitted to infer from analogy with

existing species upon which dependence must be

placed for an elucidation of the general law.

The typical position for such pits is upon the

tangential walls of the summer wood, where

they are most satisfactorily seen in radial sec-

Fats. OF ANN meee VO

tion, inasmuch as they are always readily observ-

| able when present, and their most essential

| Wi features are displayed in a manner not possible

L» | in a tangential section (Fig. 10). Pits occur in

this position in 71. of all the investigated

VU WIM po 714 f

FA species. and their absence in 28.3 % poire to

gantea, Radial sec- SOME special features in development which may

and gis a be assumed to have a general bearing upon the

question of descent and relationship. In Agathis,

as represented by the one species, A. australis,

Such pits are a prominent and characteristic

feature, but in the nearly related Araucaria, they are remarkable

for their uniform absence. In the primitive Ginkgoales they

No. 448.] NORTH AMERICAN CONIFERALES. 263

are also present, but among the Taxacez, while generally pres-

ent, they are occasionally wanting as in Tor-

reya taxifolia and T. nucifera or 66.6 % of the

investigated species of that genus. Nowhere

else among the Coniferales do we find such a

feature until we reach the genus Pinus, the

second and higher section of which is invari-

ably characterized by their absence, thus pre-

senting an exceptional feature to the extent

of 68.3 % of that genus. That such absence

represents a process of obliteration conform-

able to De Bary's law cannot be doubted,

while the sporadic recurrence of this feature

in often widely separated genera, or in partic-

ular species of a given genus, must be held to

have a more or less direct bearing upon the

general course of development. This is em- Ft: rr Sees gigant

phasized by the observation that in Larix A QAM fte de

americana and L. leptolepis as also in Picea ern —

bicolor, there is a more or less pronounced

tendency to an obliteration which is never fully developed. This

is expressed in the somewhat remote position of the pits and

V „ their very small size, which ren-

í | ders them obscure and often

(o) difficult to discover. In this

; respect these species represent

(o) transitional forms.

(] As an exceptional feature,

bordered pits may sometimes be.

() found upon the tangential walls

of the spring wood. This is

| especially noticeable at the ends

of tracheids, and in rare cases it

may apply to the entire extent

(0) | of the wall. The most notable

~ H instance of this kind, because

ia gigantea. Radial section practically unique, is to be met

quo. Rad

icine ached pits on the tangential walls

of the spring wood. X 280. , with in Sequoia gigantea (Figs.

264 THE AMERICAN NATURALIST. (VoL. XXXVIII.

11, 12). Those spring tracheids which lie in direct contact with

the summer wood of the previous year, often exhibit this feature

with great prominence, but it may also extend radially through

several successive tracheids. This is undoubtedly a primitive

character, and in the one case cited it possesses some value for

the purpose of specific differentiation, but in general terms, the

occurrence of bordered pits in such positions is of so sporadic a

nature as to give this feature no well defined value, either for

taxonomic or phylogenetic purposes. It may, nevertheless, be

stated with respect to the pits on the tangential walls of the

tracheids in general, that in their distribution they distinctly

conform to the law governing similar structures on the radial

walls.

Reference to Cordaites acadianum shows that in the multise-

riate pits of the hexagonal form, these structures always preserve

the spiral arrangement characteristic of the structures from

which they were derived (Fig. 4), and this conformity also

extends to the direction of the spirals which generally ascend from:

left to right. The general law in this respect has already been

formulated so fully by De Bary (9, p. 163), as to make it

unnecessary at this time to enter upon its consideration more in

detail, beyond a reference to one or two special features and

some apparently exceptional cases. While the spiral arrange-

ment is always typical in such genera as Cordaites, Agathis,

Araucaria, etc., it is not obvious in those cases where the pits are

strictly uniseriate and often remote from one another. Nor is

it apparent at first sight in those cases of 2-seriate pits where,

as in Cupressoxylon dawsoni from the Cretaceous, Larix ameri-

cana, Sequoia and various species of Pinus, the pits are always

paired off in such a way that the axis of each pair is at right

angles to the axis of the cell (Fig. 7). Two explanations are here

possible: (1) the spirals are in reality 2-seriate, and they are

projected through the alternate members of the two rows of pits,

or (2) the disposition of the pits represents an extreme phase in

the flattening of the original spirals conformably to a higher

type of development. This latter view, which seems the more

reasonable, is in direct harmony with De Bary’s law, while it

eun additional support from the form and direction of the pit

orifice.

No. 448.] NORTH AMERICAN CONIFERALES. 265

The orifice of the pit is variable, at different times being

round, when the pits are also round and more

or less distant; oval or oblong, when the pits

assume corresponding forms; or in the summer

wood, lenticular or oblong. The transversely

elliptical pits of Pinus strobus (Fig. 8), the ori-

fice of which is also transversely oblong, as also

the similar pits of Pinus cubensis (Fig. 7), afford

substantial proof in confirmation of the probable

correctness of this view. In the summer wood,

the pit orifice commonly assumes a position

which appears to offer a direct contradiction of

this conclusion. In Pinus strobus (Fig. 9), the

orifice is oblong and parallel with the tracheid

axis. In Pinus pungens, as in many others of

the same genus (Fig. 13), the narrow orifice is

extended above and below into a diagonal slit of ee I

great length, forming a narrow angle with the pits on the radial

tracheid axis. At first sight this would seem to ee

imply that these features represent primitive

spirals, the original direction of which has not been greatly if at

all modified, but one or two considerations will assist us toa

correct interpretation of this feature. In the first place it is to

be observed that such positions and modifications of the orifice

are invariably associated with the summer wood, or where they

occur in the spring wood, it is the result of maceration and com-

monly occurs in fossil plants, or woods in process of decay, and

they are always most conspicuous in those tracheids which have

experienced the most profound modifications with respect to the

growth in thickness of the secondary walls. It has already been

shown in the case of Taxus and Torreya, that there is no neces-

sary connection between the spiral bands and the spiral lines of

striation — that, as a matter of fact, as particularly illustrated

by Torreya taxifolia, the two are quite distinct from one another

under ordinary conditions of development. But in cases where

the wall experiences extreme growth in thickness, the oblitera-

tion of the original spiral structure is complete, and at the same

' time it is replaced by the normal striation of the wall which,

266 THE AMERICAN NATURALIST. [Vor. XXXVIII.

in such cases becomes most pronounced. Instances such as

those afforded by Pinus strobus and P.

insignis may, according to this interpreta-

(e in the obliteration of the original spirals,

O © indicative of the highest type of develop-

ment. In a few cases the structure of the

Io bordered pit presents exceptional forms.

Q In Cupressus nootkatensis the pit orifice

| shows either unusual want of regularity

(e in outline and marked eccentricity of posi-

Q o tion, or it is so enlarged as to leave only a

narrow border to the round or oval pit

(Fig. 14). Similar features occur occasion-

F16. 14.— Cupressus nootkaten- Ally in other genera, and they are generally

ieee HE Sowing conspicuous in Pinus feda. De Bary (9,

p. 159) has directed attention to the same

feature in Ephedra and Pinus sylvestris, and he

correctly interprets it as a form of arrested devel- «4

opment. Alterations also arise as a feature of O

secondary growth in those cases in which the wall

acquires unusual thickness. This is typically the e»

case in Pinus cubensis where in plan (Fig. 15), the

orifice is extended vertically to a P

length often twice the diameter of p, Pix

the original pit. In tangential sec- ^"** Radial

tion, according to the particular deformed bor-

direction of the plane of section

(Fig. 16), the orifice is either of

uniform width, or it enlarges constantly through

the entire thickness of the later growth, from

Fic. 16.— Pinus cuen. Within outwards. That such unusual forms are

sis. Tangential sec- features of extreme, secondary growth of the

tion of bordered pits a

asin Fig. 15. xix Wall, and that they may be anticipated in all

cases where such modifications of the walls

occur, is a reasonable deduction from the observed facts.

No. 448.] NORTH AMERICAN CONIFERALES. 267

TAXONOMIC AND PHYLOGENETIC.

For taxonomic purposes, the bordered pits possess a definite

though often limited value. In the genus Cordaites, as also in

Araucarioxylon, Araucaria and Agathis, this is expressed in the

hexagonal form together with their very compact, chiefly

multiseriate arrangement throughout the entire extent of the

tracheids, — characters which are of generic value and at once

serve to separate these genera from all others. The contrasting

differential feature is then to be found in the pits of the oval or

round form, together with their 2-seriate or I-seriate dispo-

sition with a more or less marked tendency to segregation.

This is characteristic of the Ginkgoales and all the Coniferales,

both fossil and recent.

As a different character of sub-generic value, the occurrence

of bordered pits on the tangential walls of the summer wood

of the first section of Pines — the soft pines — and their invari-

able absence from the same structural region in the second

section — the hard pines, — is one which may be always relied

upon.

For the purposes of specific differentiations, the pits on the

tangential walls possess a distinctly inferior value which must be

confirmed in most cases by the evidence of other factors. Their

utility in this respect is made sufficiently clear in the various

diagnoses and in the artificial key, without further discussion at

this time.!

In the genus Cordaites, according to the provisional specific

differentiations of fossil forms as at present generally employed,

the number of rows of pits, or their segregation into definite

groups, are characters of well defined, specific value, since they

are among the few features which may be utilized with cer-

tainty for this purpose. Thus C. acadianum with its 2-5

rows; C. materiarium with 2—4 rarely 3-4 rows ; C. hamiltonense

with two rows and C. mewberryi with two rows in gr x dis

6—13 pits, rest upon a basis which is not only easy of recognition,

but which may be applied with full assurance, as has been done

1 This paper will appear in the Trans. Royal Society of Canada for 1904.

268 THE AMERICAN NATURALIST. [VoL. XXXVIII.

on former occasions (38, p. 51 et seq.) In Araucaria, the three

species investigated may be similarly differentiated from one

another. The same rule is applicable to Torreya taxifolia which

is thereby separable from the other species ; likewise to Cupress-

oxylon dawsoni, Tsuga canadensis, Larix americana, and, among

the pines, P. lambertiana, P. clausa, P. sabiniana, P. teda, P.

palustris, and P. cubensis. It is to be observed, however, that

the constancy which characterizes this feature in Cordaites and

Araucaria, is wanting in the higher Abietinez. In Larix there

is such variation that very careful scrutiny is required, while in

the genus Pinus, the number of exceptions to the typical charac-

ter increases greatly, and is liable to cause some difficulty in -

the final determinations unless much care is exercised. Pinus

4eda offers a conspicuous illustration of this fact, as may be seen.

by reference to the analytical key. It is therefore manifest that

the value of the bordered pits for taxonomic purposes is most

clearly defined in the lower types of the Coniferales, and that

their value diminishes steadily, with an advance toward higher

forms of organization and development. In all cases where

exceptional forms introduce diagnostic difficulties, these may be

overcome by the controlling effect of associated characters.

We are now in a position to examime the data at hand with a

view to determining the bearing of the bordered pits upon ques-

tions of phylogeny.

. Having reference to the origin of the bordered pit and the

various modifications it presents in the course of development, it

cannot be doubted that the hexagonal, multiseriate pits of Cor-

daites, Araucarioxylon, Araucaria and Agathis place those gen-

era in a relatively inferior position, a view which gains a large

measure of support from the well known and extensively multi-

seriate disposition shown in Heterangium grievii (49, p. 341).

but the facts so far discussed have not as yet thrown any special

light upon the relative positions of the separate genera.

An examination of twelve species of Cordaites shows that the

bordered pits exhibit a much wider range of serial variation than

any other genus covered by the present studies. If then we

accept the genera] principle with respect to the development of

the bordered pits as already illustrated, it cannot be doubted that

No. 448.] NORTH AMERICAN CONIFERALES. 269

the 2—5-seriate pits stand much nearer to the primitive form of

the tracheid than do the r-seriate. From this point of view it

is then evident that in C. recentium, the name of which is

thereby seen to be fully justified, the r-seriate pits place it at

the upper end of a series which has its inferior termination in

the 2—5-seriate C. acadianum, while between the two, interme-

diate forms appear as members of a series of nine variants, and

it is possible to arrange these in such a manner as to exhibit

the probable sequence in development as follows:—

Serial variations in the Bordered Pits of Cordattes.

2-5 ser. - 2-4 3-4 2-

d SS. 2.|chiefly 3. chiefly 2 rarel

3 2-3 1- 2-ser. | 1-ser.

chiedy 2

A. chiefiy 2

- J

C. acadianum X

ohioense x

ouangondianus) x

matertarium x

clarkii

annulatum

brandlingii x

aterioide : x

tllinoisense x

hamiltonense

newberryt

recentium

The wide range of variations here shown, especially when com-

pared with other genera, at once serves to suggest that Cordaites

was in this respect somewhat of the nature of a transition group

from which others were given off, or else that it epitomized the

collective changes through which a number of genera must have

passed. And inasmuch as this genus exhibits a more highly

developed, multiseriate arrangement than any other within the

general phylum, we must concede that it is, with respect to this

Character, the most primitive of all.

The genus Araucaria shows à much more restricted range

of variations, there being only four variants pretty uniformly

distributed among fourteen species, both recent and fossil.

While the most highly developed members, four in number, are

represented by r-seriate pits, the most primitive form of 4-seri-

270 THE AMERICAN NATURAL/ST. [Vor. XXXVIII.

ate pits occurs in only one case — A. robertianum. It is there-

fore manifest that this genus is obviously of a more advanced

type than Cordaites from which it undoubtedly originated.

Agathis being represented by only one species, it is not possible

to locate it more definitely than to say that the 1—3-seriate dis-

position of its pits would place it in a position equivalent to that

occupied by Araucaria cunninghami and therefore about three-

fourths way down the scale for that genus. This fact points

with much force to the idea that of the two genera, Agathis is

of relatively lower type. |

The Ginkgoales and the Coniferales as a ‘whole, exhibit an

obviously higher type of development than the preceding group,

in consequence of the more pronounced tendency to segregation

of the pits which are now either elliptical or round, and never

hexagonal. This distinction is so clearly defined and constant

as to support the idea which gains force in other ways, that

Cordaites, Araucaria and Agathis are clearly related members of

a principal branch of the original stock, and that they therefore

diverge considerably from the particular line of descent within

which we find both the Ginkgoales and the Coniferales.

The observations so far made apply altogether to the pits on

the radial walls. We may now pass to a consideration of their

relation to the tangential walls, a factor which does not call for

very extended discussion. This feature is found to apply to

71.7 % of all investigated species exclusive of fossils. It is

wanting in three species of Araucaria, representing 2.58 fo; in

Torreya, 2 species or 1.72 %, and in the entire second section of

Pinus to the extent of 28 species or 24.1 %. But the occurrence

of pits on the tangential walls, in common with those on the

radial walls, is a well known feature of the Sigillarias (49, P-

198), where their primitive character is well established, and we

can hardly doubt that their ultimate elimination in the higher

pines is the expression of a final phase in development, consist-

ent with the position usually assigned those plants. The

absence of pits from the tangential walls of certain Araucarias

and Torreyas, is to be interpreted as one of those sporadic ten-

dencies toward a higher type of development which never

become permanent in the same line, but which are to be met

with as one of the invariable features of evolution.

No. 448.] NORTH AMERICAN CONIFERALES. 271

Ginkgo, being the unique representative of an ancient line,

cannot very well be brought into the present discussion very

much in detail. On other grounds it is known to be a primitive

form representing a group distinctly inferior to the Coniferales,

and this view is supported by the disposition of the pits in two

series, a character which, if taken alone, would give the genus

rank with Torreya taxifolia among the Taxacez, but when

regarded collectively, would place the genus distinctly below the

Coniferales as a whole. This evidence, then, indicates that the

Ginkgoales must have arisen as a side line at some point inferior

to the Coniferales, but superior to the Cordaitales.

In the Taxaceze the bordered pits do not in themselves

afford very conclusive evidence as to the relative position of the

family. Among the eight investigated species, representative of

three genera, only three, and chiefly two variants occur. Taken

alone, the disposition of the pits would lead to no final conclu-

sion, but other factors permit of placing this family in the

inferior position usually assigned to it. Within the genus three

| variants are found — the 1—2 rows of 7. taxifolia, the one row

or pairs of 7. californica and the strictly r-seriate form of

T. uucifera. In Taxus only two variants appear — the one row

or pairs of 7. floridana and the 1-seriate disposition as found in

the remaining three species. The one representative of Podocar-

pus shows but one variant, and that is r-seriate. From this

itis obvious that the generic sequence must be in the order

given, and that the sequence of species must be approximately

as given in the table of anatomical data to follow.!

The remaining genera of the Coniferales present so few devi-

ations from a typical form, that they cannot be differentiated

fully on the basis of the bordered pits. This character never-

theless has a definite value in association with others, as in the

. genus Sequoia or some of the hard pines, Larix americana, etc.

The general sequence of genera may be recognized by the bor-

dered pits only in so far as these structures serve to confirm

and emphasize the conclusions reached in other ways, and this

will become apparent from an inspection of the table of anatom-

1 This table will appear in the last number of this series.

272 THE AMERICAN NATURALIST. [Vor. XXXVIII.

ical data. It will nevertheless serve a useful purpose at the

present moment, to ascertain the general sequence based upon

the percentage distribution of the principal variants as follows.

Comparison of the principal variations in the serial arrange-

ment of bordered pits, by percentages.

Total | i+

var. | 2-5 | 2-4 2-3 1-3 2 1-2 pairs. I

Cordaites 9 |83]| 25.0 | 16.6 | 25.0 | 16.6

gathi. I D 660 0.0 40.0 33-3

Araucaria 4 6.6 | 20.0 40.0

Ginkgo I 1u0.0

Sequoia I 100.0

ari 3 323] 3331) 33:3

Taxodiu 2 25-0 5.0

Libocedrus I 100.0

id 1. I 100.0

Pseudotsuga 2 $0.0| 50.0

Pinus. 3 17.1 41.5 41.4

Abies 3 13.6| 27.3, 59-1

Taxus 2 25.0| 75.0

Tsuga . 3 16.7 | 16.7 | 66.6

cea 2 0.0| 90.0

Podocarpus I 100.0

Thujopsis I 100.0

Cryptomeria I 100.0

With respect to specific differentiations, it has already

appeared that the bordered pits may be employed with success

in Taxus and Torreya. In Cupressus this rule also applies to

C. pisifera, and C. macrocarpa, both of which are distinguished by

having their pits in one row or pairs, while the remaining seven

species have strictly uniseriate pits. An instructive example is

afforded by Cupressoxylon dawsoni. In this species, which is of

early Tertiary age (Lignite Tertiary), the pits are typically 2-

seriate, being disposed in a very compact manner similar to that |

found in existing Sequoias. But in a series of eleven speci-

mens, it is clearly seen that two variants are represented — the

second being a r-seriate form. These variations are also found,

as in the other Coniferales, to be directly related to variations in

the size and rate of growth of the tracheid. It cannot be

doubted then, that C. dawsoni is a more primitive representative

No. 448.] NORTH AMERICAN CONIFERALES. 273

than any species now existing, and that it is substantially the

ancestral form of the genus, so far as we know.

In Larix the four investigated species may be differentiated

pretty fully, and this rule applies with particular force to Li

americana, and L. occidentalis, both of which are distinguished by

a 2-seriate form, Among the pines, P. /ambertiana, P. clausa,

P. sabiniana, P. teda, P. palustris, and P. cubensis are readily

differentiated from the others by the 2-seriate pits. In all other

cases than those specifically indicated, the bordered pits afford

an inadequate basis for specific differentiation.

It is now apparent that segregated, round or oval pits in one

row müst be taken as representing the highest type of develop-

ment in the Coniferales, and any deviation from this must be

taken to indicate the sürvival of more primitive conditions,

pointing to derivation from a type like that of Araucaria or Cor-

daites. From this point of view, the occurrence of pits in 1-2

rows in Larix americana, Torreya taxifolia, Sequoia, Tanga can-

adensis and various species of Pinus, indicates the survival

of ancestral characters which are partial to the extent of 7.2 %,

and complete to the extent of 10.8 %. That such deviations from

the usual type of structure are either survivals or reversions

which serve to indicate a common origin, cannot be doubted,

more especially as they do not occur ata fixed point near the

original type, but they arise sporadically in widely separated

genera. The tendency of such evidence then, is to show a com-

mon ancestry for the various genera of the Taxaceæ and

Coniferæ, a view which is greatly strengthened by the testimony

afforded by the spiral tracheids of Larix americana, Pseudotsuga

and Pinus teda. The provisional conclusions which we now

reach are, that there were probably four main lines of descent

from the original stock represented by Cordaites :

Araucaria and Agathis.

Ginkgoales.

Taxacez.

Conifere.

bet

4 Co N

( To be continued.)

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L. MARK, DIRECTOR.— No. 151.

THE SENSE OF HEARING IN THE GOLDFISH

CARASSIUS AURATUS L.

HENRY B. BIGELOW.

I. INTRODUCTION.

To THE older investigators, such as Hunter (:82) Müller

(48), and Owen ('66), the presence of internal ears in fishes was

sufficient evidence that these animals had a sense of hearing.

Kreidl (95), however, was the first to seek experimental evidence

on this point. He tested normal goldfishes, and others from

which he had removed the semicircular canals with the attached

portions of the ears. Since both classes of fishes reacted sim-

ilaly to sounds, he concluded that the skin, not the ear, is

stimulated by sound, and that, therefore, it can be said that

goldfishes do not hear. This conclusion has since gained consid-

erable recognition, and has been accepted by Mulertt (:02) in

his recent book Zhe Goldfish. Kreidl ('96) also made obser-

vations on the trout in the fish basins of the Benedictine

Monastery at Krems, Austria, and gathered. ample evidence to

prove that the belief that the fish assembled for food at the ring-

ing of a bell to be unfounded. These observations confirmed him

in the opinion that fishes do not hear, a conclusion further sup-

ported by the subsequent observations of Lee (98) on other

species, particularly the dogfish.

In spite of this important evidence, Lang (:03) concludes a

popular discussion of the question ** Do acquatic animals hear ?"

with the statement that fishes and other like forms should be

tested more extensively before a decisive answer can be given.

Tullberg (:03), whose experiments led him to conclude that the

fish ear receives stimuli from currents in the surrounding water,

275

276 THE AMERICAN NATURALIST. [Vor. XXXVIII.

admits the possibility that the ear may also be in some degree an

organ of hearing. The fact that certain fishes do respond to

what are beyond question sound vibrations in water was demon-

strated independently by Zenneck (:03) for three fresh-water

species, Leuciscus rutilus, L. dobula, and Alburnus lucidus, and

by Parker (:03*, :03") for a salt-water species, Fundulus hetero-

clitus. Parker further showed that this response was dependent

on the presence of a functional internal ear, and that, therefore,

this species could be said to hear. But he could not get any

response to sound from the smooth dogfish, and so was led to

conclude that different fishes might differ widely in this respect,

some possessing, some lacking, a true sense of hearing. Since

none of the recent investigators who favor the view that fishes

hear, have tested the species which, in Kreidl's hands, yielded

negative results, it seemed desirable, in the light of recent work,

to examine the goldfish again ; and this I have undertaken to do.

This problem was suggested to me by Professor G. H. Parker,

and I am indebted to him for help, suggestions, and criticism

throughout the series of experiments.

II. METHODS.

In determining whether or not a fish has a sense of hearing

it must be borne in mind that the responses to sound may be

very slight and easily confused with responses to other stimuli ;

and, further, that, although the absence of response does not

necessarily mean absence of hearing, the consistent occurrence

of a response is fair evidence of the presence of this sense. To

test normal goldfishes satisfactorily, it was necessary to isolate

them from all other disturbances and to apply the sounding

instrument as directly as possible to the water, but without caus-

ing any gross mechanical vibration. The apparatus which ] used

was an aquarium, some 30 cm. wide, 35 cm. long, and 15 cm.

deep. The two sides were of glass, and the ends and bottom of

clear white pine. The aquarium was placed on a wooden table

which Stood with its feet on many thicknesses of soft paper; thus

isolating it from vibrations which might .otherwise reach it

through the floor. The bottom of the aquarium was covered on

No. 448.] CARASSIUS AURATUS L. 277

the inside with a thick layer of cotton wool, which, in turn, was

held in place by a sheet of thin cotton cloth to form a deadened

floor. On trial, I found that the light entering through the glass

sides of the aquarium was a source of disturbance to’ the fishes.

Hence I later made these sides opaque by a covering of dark

paper. My method of producing sound was by an electric tuning

fork, which was run by a small storage battery and had a pitch

of 100 vibrations per second. This was set up on another table,

very close to the first one, but not in contact with it. The foot

of this table also stood on many thicknesses of soft paper. The

fork itself rested on a deadened support, and was so arranged

that after it had been set in vibration, it could readily be moved

till its base came in contact with the wooden end of the aquarium.

This could easily be accomplished without observable jar to the

water in the aquarium, and certainly in itself had no effect on

the fishes. For when I made the fork, not in vibration, touch

the aquarium in the usual way, the fishes gave no reaction,

although to the vibrating fork they were very responsive ; I tried

this many times.

With this apparatus, I tested ‘three classes of goldfishes ; (1)

normal ones ; (2) fishes the greater part of whose integument

had been made insensitive by cutting the fifth and seventh

nerves, the lateral line nerves, and -the spinal cord close to the

medulla; and (3) fishes in which the eighth nerves had been

cut.

III. NorMAL FISHES.

Goldfishes appear to be much more irregular in their responses

to sound than some other fishes. When one is first. placed

in the aquarium, it swims about vigorously, darting from side to

side in a very restless fashion. This extreme excitability lasts

for a considerable time, often an hour; but finally the fish

becomes more quiet, sinks to the bottom, and remains nearly

motionless except for an almost incessant movement of the

pectoral fins. In a few cases, the fishes lay motionless with

these fins folded closely to their sides. If now the tuning fork

was set in vibration, and brought into contact with the wooden

278 THE AMERICAN NATURALIST. [Vor. XXXVIII.

end of the aquarium, the fishes almost always responded by one

of a number of reactions. The most important of these were:

a rapid vibration of the tail without locomotion ; sudden jerks of

the tail from side to side, often so vigorous as to cause a swift

dart forward ; normal locomotion, forward, backward, or to one

side; or in those fishes which lay at rest with the pectoral fins

folded, a vigorous spreading of the pectorals. It was evident

that individual goldfishes differed from one another in their

reactions much more than the individuals of Fundulus as

recorded by Parker (:03*, p. 51). When a fish was tested with

the vibrating fork, it might respond by any one of these reac-

tions, but every fish had one distinct reaction characteristic of

it, which the application of the fork rarely failed to elicit.

Thus, the more active fishes usually responded by vigorous

locomotion, the more quiescent ones, by tail or fin movements.

Although fishes on which no operation had been performed,

usualy responded to the sound, one, an albino, and a very

sluggish individual, gave no response. The fact that this one

died soon after my observations on it were made, is perhaps an

explanation of its unusual condition. As an example of the

characters of the responses, the following record of ten tests

from my laboratory note-book may serve. This record relates

to a fish which was subsequently operated upon by cutting the

eighth nerves.

I. Tailjerks, followed by forward swimming.

2. Tailjerks, then forward swimming after an interval of

half a second. ,

3. Sudden tail- and trunk-jerks, followed by forward swim-

ming.

Tail-jerks, but without locomotion.

Tail-vibration, but without locomotion.

Strong tail-vibration followed by a turn to one side.

Tail- and trunk-jerks, followed by forward swimming.

Tail-vibration, followed by a sudden turn to one side.

Tail- and trunk-jerks, followed by a turn to one side.

. Tail-jerks, and sudden jump forward.

all, I made 193 tests on 18 normal fishes, and observed

150 responses, about 78%, of the whole. Of the 43 failures, to

sO eM AMS

—

No. 448.) CARASSIUS AURATUS L. 279

respond, 12 were in the case of the albino fish above mentioned,

and many of the remaining 31 were probably due to faulty

observations caused by the extreme activity of the fishes. In

this respect, there is a great difference between different individ-

uals, some being in such continual movement that it is difficult

to test them, while others are more generally quiet.

IV. FISHES WITH INSENSITIVE SKINS.

The second class of fishes tested were those in which the

greater part of the skin had been rendered insensitive by cutting

the spinal cord just posterior to the pectoral fins, as well as

both lateral branches of the tenth, and the fifth and seventh

nerves on both sides of the body. The fifth and seventh nerves

were cut at a point just above the dorsal end of the opercular

opening, where they come close to the skin. The fishes were

etherized before the operation, and usually recovered and lived

many weeks. The individuals selected for this operation were

those which showed good pectoral-fin reactions when tested with

the tuning fork. Such fishes after recovery lay on their sides

on the cloth bottom of the aquarium, and were perfectly quiet

unless stimulated. That their skins were practically insensitive

was shown by the fact that they were quite indifferent to touch

with a bristle or the like.

In 65 tests on 6 such fishes, I observed 52 responses (80%)

to the vibrating fork, a condition essentially the same as that of

normal fishes. These experiments demonstrate then, that with

an almost insensitive skin, a goldfish will respond to sound as a

normal fish does.

V. FISHES WITH INSENSITIVE Ears.

The third class of fishes tested were those in which the eighth

nerves had been cut on both sides. In preparing for this opera-

tion a number of fishes were tested and only such as showed a

Clear reaction to sound were operated on. They were etherized,

and their eighth nerves were cut by piercing the skull in an

appropriate position and cutting downward with a small chisel-

280 THE AMERICAN NATURALIST. (VoL. XXXVIII.

like knife. The chief danger in this operation is in cutting too

deep, in which case excessive bleeding may follow. When this

was avoided, the fishes usually recovered, and the success of the

operation could be judged by their subsequent movements.

When they first recovered from the ether they seemed to have

lost all power of equilibration, swimming now one side up, and

now the other, or resting with their long axes vertical. After

about a day, however, they usually acquired and kept their

normal position, at least while resting or swimming slowly ; and

this ability increases, until after two or three weeks they were,

in all their ordinary movements, indistinguishable from normal

fishes. If, however, such a fish be placed in a large body of

open water and made to swim rapidly, it soon loses all power of

orientation and darts about, turning over and over until

exhausted. This condition, so far as I know, is permanent,

for in the case of one fish which lived for over three months

after the operation these reactions showed no tendency to dis-

appear, but persisted till death. The partial recovery of equili-

brium noticeable soon after the operation is probably due to a

successful attempt on the part of the fish to retain its normal

position through sight.

Earless fishes are usually more quiet than normal ones, and

hence they can very easily be tested. In all, I made 73 tests

on 7 fishes, and in no instance did I get an undoubted response

to sound. This is in strong contrast with the reactions of the

same fishes before their eighth nerves had been cut, and points

beyond question to the ear as an organ of hearing.

I supplemented the foregoing experiments by another series

in which two normal fishes that I found to respond well to

sound, were etherized, and the eighth nerve of each cut on the

right side. After an interval of twenty-four hours, they were

both tested again, and found to respond about as well as they

did before the operation. In 20 tests on the two, there were

I9 responses. They were then etherized again, and the eighth

nerves of the left sides were cut. After recovery, they were

tested once more, and, although the experiments were con-

ducted with the greatest possible care, not a single response

Was observed in 20 trials. This experiment shows that the

No. 448.] CARASSIUS AURATUS L. 281

operation of cutting the eighth nerve, severe as it is, is not

sufficiently so to account for. even a small part of the reduc-

tion in the number of responses which follows the elimination of

both ears as sense organs.

In another instance, a fish which before any operation

responded vigorously to sound by movements of the pectoral

fins, was prepared by cutting two holes in the top of the skull,

through which the eighth nerves could conveniently be reached,

and the following parts were severed : the spinal cord, the lateral

line branches of the tenth nerves, and the superficial portions

of the fifth and seventh nerves of both sides. After recovery,

the fish gave ten vigorous pectoral fin responses to as many

trials with the sounding apparatus. The eighth nerves were

then cut, and in twelve tests only one response, and that of

a doubtful character, was observed. It, therefore, seems incred-

ible that nervous shock can account for the almost complete loss

of response to sound, after cutting the eighth nerves, and I am

firmly convinced from the foregoing experiments, that the ear in

the goldfish is an organ of hearing, and that it is the loss of

this which is accountable for the difference of reaction between

fishes in which the ears were intact, and those in which the

eighth nerves had been cut.

VI. Discussion OF RESULTS.

The results of my experiments differ so essentially from

those obtained by Kreidl (’95) that a further discussion of

these differences is necessary. So far as reactions to sounds

were concerned, Kreidl was unable to distinguish between

normal goldfishes and individuals from which the semicircular

canals with the attached parts of the ears had been removed.

In my experiments, however, while normal fishes responded to

sound in about 78% of the trials, those in which the eighth

nerves had been cut, scarcely responded at all. The difference

between these two sets of results was so great that I determined

to repeat, with as much precision of detail as possible, Kreidl’s

experiments.

I tésted several goldfishes, and, having found that ey

282 THE AMERICAN NATURALIST. [Vor. XXXVIII.

responded well to the tuhing fork, I operated on them in the

following way. After etherizing the animals, I cut off the top

of the skull, exposing the brain, and the vertical semicircular

canals. I then seized the canals with forceps and drew them

out bodily with the attached sacs and their otoliths, as Kreidl

had done. I operated thus on four fishes, three of which recov-

ered. After recovery, I tested them again with the tuning fork,

and found that one responded to the sound about as well as

before the operation, and that the two others responded some-

what less regularly than before, though in .a still perfectly

definite and unmistakable manner. Thus, since these fishes

responded like normal individuals, my results confirmed in all

essential respects those of Kreidl, and I came to the conclusion

that there must be some fundamental difference between Kreidl's

methods for the elimination of the ear, and mine. The method

I generally used, cutting the eighth nerves, seemed to me a per-

fectly secure means of excluding the action of the ear. On

the other hand, the withdrawal of the semicircular canals with

the attached parts of the ear, as practiced by Kreidl, might well

leave behind and intact parts of that organ, and thus be inefficient

as a method for completely excluding the ear. To settle this

matter, I made careful dissections of the ears of goldfishes.

The ear of the goldfish is in all essential respects similar to that

of Cyprinus, as described and figured by Retzius ('8r, p. 78). `

The semicircular canals are of large size; the two vertical

canals lie free in the brain cavity, while the horizontal canal

is partially imbedded in the skull. The sac into which these

canals open, the utriculus, is of medium size, and contains a large

lenticular otolith. The utriculus, with its otolith lies free in the

brain cavity and is the structure which is removed in connection

with the semicircular canals in fishes which are operated on by

Kreidl’s method. But ventral to these parts, and largely imbed-

ded in bone is another portion of the internal ear, which probably

represents the combined sacculus and lagena. This is not

removed, nor even seriously disturbed by the Kreidl operation.

This deeper sac extends posteriorly and ventrally until that

of the right ear nearly meets that of the left in the base of

the cranium. Each sac contains two otoliths, one long and

No. 448.) CARASSIUS AURATUS L. 283

rod like, lying for the most part in the saccular portion, the

other lenticular in outline, and lodged in the lagena proper.

The wall of this sac is supplied with branches from the eighth

nerve and is so surrounded by bone that all attempts to remove

it by pulling out the semicircular canals were complete failures.

In the four fishes on which I operated by Kreidl's method,

subsequent dissection showed these structures intact. Thus, his

operation leaves uninjured a large part of the internal ear, in

fact, just that part which, from comparison with the ears of

higher vertebrates, would be expected to be concerned with

hearing. I believe, therefore, that Kreidl’s method of operating

is defective, and the reason that the fishes upon which he had

operated responded to sounds much as normal ones did, was not

because in both cases the skin was stimulated, as he believed,

but because his so-called “earless” fishes still retained intact

a part of the ear which, as I have already shown, acts as an

organ of hearing. That it is such an organ follows from the

fact that when its nerve connections are cut, the responses to

sound cease.

VII. SUMMARY.

t. Normal goldfishes usually respond in a definite manner to

sound-vibrations in water.

2. Goldfishes in which most of the skin has been rendered

` insensitive by cutting the nerves, and specimens from which

/- the ears, except the saccular portion, have been removed

still respond in an essentially normal way to sound vibra-

tions in water.

3. Goldfishes in which the eighth nerves have been cut on both

sides, thus eliminating the sacculi and lagena as well as the

rest of the ear, seldom or never respond to sound vibrations

in water.

4. Goldfishes possess the sense of hearing, and the portion of

the ear concerned with this sense is the sac which probably

represents the sacculus and the lagena of higher verte-

brates.

284 THE AMERICAN NATURALIST. [Vor. XXXVIII.

BIBLIOGRAPH Y.

HUNTER, J.

‘82. Account of the Organ of Hearing in Fish. Phil. Trans. Roy. Soc.

London, vol. 72, pp. 379-383. 1782.

KREIDL, A.

'95. Ueber die Perception der Schallwellen bei den Fischen. Arch. f.

ges. Physiol., Bd. 61, Heft 7-8, pp. 450-464.

KREIDL, A.

'96. Ein weiterer Versuch über das angebliche Hóren eines Glocken-

zeichens durch die Fische. Arch. f. ges. Physiol, Bd. 63, Heft

11—12, pp. 581-586.

LANG, A.

:03. Ob die Wassertiere héren? Reprint from Mitth. naturwiss. Gesell.

Winterthur, 1903, Heft 4, 55 pp.

LEE, F. S

'98. The Functions of the Ear and the Lateral Line in Fishes. Amer.

Journ. Physiol., vol. 1, no. 1, pp. 128-144.

MÜLLER, J.

'48. The Physiology of the Senses, Voice, and Muscular Motions, with

the Mental Faculties. Translated by W. Baly. London, 8°, xvi!

+ 849-1419 + 32 + 22 pp.

MULERTT, H.

:02. The Goldfish and its Systematic Culture. Third ed. Brooklyn,

12°, 150 pp., 19 pls.

OwEN, R.

'66. Onthe Anatomy of Vertebrates. Vol.1. London, 8?, xlii + 650 pP-

PARKER, G. H.

:03". Hearing and Allied Senses in Fishes. U.S. Fish Comm. Bull.

for 1902, pp. 45-64, pl. 9.

PARKER, G. H.

The Sense of Hearing in Fishes. Amer. Naturalist, vol. 37, 2°

435, pp. 185-204.

` RETZIUS, G.

'81. Das Gehórorgan der Wirbelthiere. Morphologisch-histologische

Studien. L Das Gehórorgan der Fische und Amphibien. Stock-

holm, 4°, xii + 222 pp., 35 Taf.

TULLBERG, T.

‘08. Das Labyrinth der Fische, ein Organ zur Empfindung der Wan

Reed Bihang till K. Svenska Vet-Akad. Handlingar,

. 28, Afd. à

ZENNECK, J. Due mE

:03. Reagiren die Fische auf Töne. Arch. f. ges. Physiol., Bd. 9%

Heft 7-8, pp. 346-356. |

CERTAIN UNDETERMINED FACTORS IN

HEREDITY AND ENVIRONMENT.

GEORGE J. PEIRCE.

IN a paper read before the Botanical Section at the Pittsburg

meeting of the American Association for the Advancement of

Science ! I suggested that certain factors of the environment

were constantly ignored in such discussions of heredity as I had

seen or heard. This paper constitutes substantially the last

section of my book? Since certain letters which I have

recently received, make me feel that in this section I have

expressed myself so concisely that my full meaning is not alto-

gether clear, I take this opportunity to give somewhat ampler

treatment to the subject.

The word environment is used ignorantly by everyone, for no

one has ever succeeded in making a complete analysis of what

is meant by this collective term. Furthermore, although we

speak of an organism as reacting to its environment, it does not

react to its environment as a whole but to each of one of the

separate influences which are the factors of its environment. It

is, therefore, very important to know what these factors are, and

what are their effects. We know that if two of these influences

are opposite and equal, there will be no visible reaction, although

the organism will be affected by both. There may be internal

results of these influences, results which, however, may not be

perceptible. If one of these opposite influences be lessened or

eliminated, the effect of the other becomes perceptible. We

judge, then, the influence of the various factors which we are

now able to distinguish from one another as constituting the

environment, only by their perceptible effects. It is conceivable

that some effects are so long deferred that they are coincident

or at least contemporaneous with the effects of other and more

recent stimuli. For this reason these long deferred effects may

! Abstract in Science, XVI. p. 137, 1902.

? Peirce, G. J. Text book of Plant Physiology, 279-83, New York, 1903.

285

286 THE AMERICAN NATURALIST. [Vor. XXXVIII.

easily be overlooked, or they may be perceived only incompletely

and with difficulty, or they may be attributed to wrong causes.

It is also conceivable that certain influences produce effects not

because they are powerful, taking into account only short times

of operation, but because they are prolonged.

This leads us to essay an analysis of environment. The

environment of an organism is all, everything, that constitutes

the world and the universe outside of itself. One may say that

this definition is too comprehensive, that only immediate envi-

ronment is meant when the word environment is generally used.

Who can say that organisms and things are affected “only by

their immediate environment ? In fact we know that the contrary

is true in certain particulars. We know, for instance, that we

are daily affected by the sun — a remote body — quite as much

as by any part of our immediate environment. But we do not

know all the radiations and other influences from all the heav-

enly bodies in the universe, the effects of these upon our earth

as a planet, and what is upon it; and yet these radiations still

unknown and unguessed, together with other unknown and

unguessed factors of environment, may operate as regularly and

as powerfully as any of the known and recognized factors.

Among the recognized factors of the environment are some,

the effects of which are very imperfectly known, if they may be

said to be known at all.’ These will be seen in their relations to

other factors if an analysis, however imperfect, of the environ-

ment be given. Thus

(1. The earth — soil, rocks, waters, atmo-

sphere; the components, conditions, CON-

tours, etc., of these.

All other planets — whatever conditions

Environment — prevail upon them; their relative positions

all that consti- to the earth and to each other.

tutes the uni-| 3. Zhe forces emanating from and operating

verse. upon these — heat, light, electricity, a

chanical forces (gravity, wind, waves, rain,

etc.).

All living things — their parts, products,

and habits. — :

If we go through the list of factors suggested by this scheme,

No. 448.] HEREDITY AND ENVIRONMENT. 287

we shall see that, in spite of their diversity and oppositeness, we

may bring them into an orderly classification. We may arrange

them under these four headings — 1, unchanging, 2, variable, 3,

occasional, 4, periodic. Among unchanging factors the following

may be mentioned,! vs. the atmosphere, water, the force of

gravity, the earth as a whole. Though these are not all of the

unchanging influences I shall discuss only these four. *

The earth’s atmosphere, consisting of nitrogen (80%) and

oxygen (20%), with only a small fraction of 1% of carbon-di-

oxide and other gaseous matters, possesses unchanging physical

properties and exerts a pressure which varies only very slightly,

taking only short intervals of time for the comparison, and varies

not at all, taking long intervals of time. For example, the baro-

metric pressures in any given locality are the same in this cen-

tury as in the last, and though we may have “low” and “high”

barometers at different times, these variations are very slight.

The buoyancy, diversity, color, transparency, permeability, etc.,

of the earth’s atmosphere are the same to-day as a “million”

years ago, as far as we know, and though these qualities and the

pressure of the atmosphere may change and may have changed

during the lapse of the millions of years during which the earth

is likely to exist and has existed as a planet, they are unchang-

ing so far as millions of generations of living organisms, and so

far as millions of series of lifeless things, are concerned.

The composition of the earth's atmosphere has changed but

little, if at all. The proportion of c rbon-dioxide may have been

greater in an earlier time, but for uncountable ages the propor-

tions have been what they are now. Let us suppose, however,

that the proportions of nitrogen, oxygen, and carbon-dioxide have

changed somewhat, taking the whole atmosphere into account, not

a small part of it merely. If only the proportions were different,

there would still have been nitrogen, oxygen, and carbon-dioxide

in the air, and these gases themselves are unchanging. We can-

not imagine some oxygen being any different from all oxygen ;

if we have carbon-dioxide we have necessarily carbon-dioxide of

the same composition and properties since the world began.

The same of the inert gas nitrogen.

! See p. 280 of my “ Text Book.”

288 THE AMERICAN NATURALIST. [Vor. XXXVIII.

When experiments are instituted for the purpose of eliminat-

ing these substances, they fail, for it is out of the question to

remove all nitrogen, oxygen, and carbon-dioxide even if all N, O,

and Co, can be excluded at the beginning of an experiment, for

N and O will be contained in the organism and Co; will be formed

by it. Nevertheless, the living organism which is dependent

upon oxygen for respiration, upon nitrogen to dilute the other-

wise too destructive oxygen, and upon carbon-dioxide from which

to manufacture food, will succumb in any experiment of more

than brief duration in which the proportions of these gases are

greatly different from those in normal air. In other words the

physical properties of the atmosphere, its components, their com-

position and proportions, being and having for ages been what

they are, living organisms represent reactions to these qualities

and will not bear sudden change, whatever might be or may have

been the result of gradual change, if there has been any.

Water is composed of hydrogen and oxygen in the propor-

tions of two to one. It always has had the same chemical com-

position, structure, and properties, the same physical qualities.

It is an indispensable constituent of living organisms and of

many lifeless things. It is a weak acid and the most universal

solvent known. Molecules and atoms of dissolved substances

move about in it with considerable freedom, and where two

volumes of water are separated from one another only by

permeable or semi-permeable membranes, there is molecular or

atomic movement both of water and of dissolved substance from

the one volume to the other through the membrane. Water is

then a medium in which ampler molecular and atomic move-

ments are possible at ordinary temperatures, etc., than in many

other substances. But the movement of the molecules and

atoms of the solutes are independent of, though taking place

among, the water molecules. The water remains the same,

physically and chemically, whatever substances may be dissolved

in it. The temperature of a solution of small volume may be

different while solution is taking place, but ultimately the tem-

perature of the solution, or of water holding nothing in solution,

will be approximately that of its surroundings, other things being

equal. The specific gravity of the solution will differ from that

No. 448.] HEREDITY AND ENVIRONMENT. 289

of pure water, but the specific gravity of the water itself and

of. the solute itself are the same as they have always been,

the specific gravity of the solution depending only upon these

two.

With its physical and chemical properties unchanged, as an

essential constituent of all living organisms and the medium in

which necessarily the food materials and foods enter and move

about and are chemically changed in the cell, water has exerted

upon living things an influence as powerful and as persistent as

have been its relations to the physical and chemical conditions

and processes of lifeless substances since the beginning. Water

can be eliminated neither outright nor by substitution from

experiments with living organisms nor, for that matter, can it be

eliminated from the majority of experiments with lifeless sub-

stances. It is a substance of universal occurrence, of uniform

properties, of uniform action. It is truly one of the unchanging

factors of the environment, to which living organisms necessarily

react, for their composition, structure, nutrition, and activities

depend upon it.

We come now to consider the effect of gravitation. The

force of gravity acts upon every particle of ponderable matter

on the earth as a direct pull toward the center of the earth.

This pull is equal, at the surface of the earth, to 32.2 foot

seconds? 7. e. a body at the surface of the earth would fall in

à vacuum at a rate increasing 32.2 feet a second per second.

The force of gravity, operating upon every particle of ponderable

matter, constantly exerts upon it this uniform force. The force

increases or decreases inversely as the squares of the distances.

But, as Newton showed, the force of gravity is not merely the

attraction between the earth as a whole and other ponderable

matters, but every portion of matter attracts every other portion

With a force proportional to the product of their masses divided

by the square of their distances apart.’ The sum of the attrac-

tions toward the center of the earth equals the amount prevr

ously mentioned, 32.2 foot seconds?, and this sum we may for

the moment speak of as gravity without necessarily taking its

components into account.

!Watson, W. A Textbook of Physics, p. 121. New York, 1900.

290 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The attraction. of gravity upon all ponderable matters is,

however, opposed by the media in which they are. This is

implied by the statement of the value of gravity at the earth’s

surface, for this statement specifies zz vacuo. Every ponderable

body is bouyed up (or supported) by a force equal to the weight

of the fluid it displaces. This law of Archimedes concerns us

as it applies to the air and to both fresh and salt water. Besides

this we have the solidity and comparative impenetrability of.

the earth itself to reckon with, for the soil is capable of

mechanically supporting much more than the weight of the

parts of plants and animals resting upon or within it. Gravity

is then opposed, partly or wholly, according to the medium in

which the attracted object is.

The opposition to gravity varies from .0013 gr. per cc. in air

at o° C., and at ordinary atmospheric pressure, through 1.00 gr.

per cc. in pure water and 1.20 gr. in sea water, to much more

than the weight per cc. of any of the substances with which

we ordinarily have to deal. Thus if we have a plant or animal

or any other portion of matter weighing x grams in air, this

matter would have a weight in pure water equal to x minus

I gram for every cc. of volume. In sea water this would be

X — 1.2 y, in which y represents the number of cc. in the

portion of matter. To take a concrete instance, suppose Wwe

have a block of wood, occupying a space of 9 cc. and weigh-

ing IO gr. inair. This would weigh in water r0— I X 9 —I

gr; in sea water this would be ro — 1.2 x 9 — 0.8 gr. or, in

other words, the block would float. In the same way the block

would be completely supported, gravity would be completely

offset, if the block were on or in the soil. But as the soil,

the water, and the air do not come into direct contact with

and are therefore not displaced by each individual part and par-

ticle of a portion of matter, there must be enough mechanical

strength within the portion of matter to resist the force of

gravity or the body would fall to pieces. We see then that,

though a body as a whole may be bouyed up by a considerable

force which resists gravity, the component particles of the body

are not necessarily so bouyed up but are subjected to the full

attraction of gravity. Although the bouyancy of the medium

No. 448.] HEREDITY AND ENVIRONMENT. 20I

in which a plant lives makes a very great difference in. the

mechanical strength of the plant as a whole, there is no dis-

coverable difference in the structure and other properties of the

protoplasm. This fact is what the foregoing discussion has lead

us to expect; for upon the component particles of protoplasm

the force of gravity operates unopposed by the bouyancy of air

or water, except such water as is in the cell-sap; and as the

force of gravity is continuous and uniform in operation, the

living protoplasm is subjected to a continuous and uniform influ-

ence. Does it not react accordingly ?

Again, the force of gravity, regarded both as the attraction '

of the earth as a whole as well as the attraction of each

particle of matter for every other, has never been eliminated

from any experiment. If it had been, can we imagine what

"would have happened? The force of gravity acts only in one

direction. The slow revolution of,a plant upon a horizontal

axis by means of a clinostat, so that all its parts will be succes-

sively turned. in this direction, and the opposition of centrifugal

or other force to the force of gravity, yield interesting results in

experiments in which these methods are employed, but they

throw little or no light on the influence of gravity upon the

component parts of the living structure. Until gravity is elimi-

nated, not merely opposed, we cannot even guess what its influ-

ence is ; but rather than ignore it, we may guess that its influence

as a formative agent is as great as we now know its directive

influence to be. And as we know that its directive influence is

always the same, that plants of a kind under like conditions

respond in approximately like ways, like times, and with like

force to the action of gravity, sending their roots downward

into the soil and their stems upward into the air, we must infer

that in the formation of new protoplasm the component parts of

this structure are affected always in the same way and that they

respond to the constant and uniform force in constant and

uniform ways.

It remains to add a word as to the earth as a whole as one of

the continuous and uniform factors of the environment of living

' Peirce, G. J. A comparison of land and water plants. Pop. Sci. Monthly,

LXIII, p. 239, 1903.

292 THE AMERICAN NATURALIST. [Vor. XXXVIII.

organisms. The earth is a spheroid whirling in space at a

rate decreasing with inconceivable slowness. Its position with

relation to other bodies of the solar and other planetary systems

changes also with inconceivable slowness. It possesses a degree

and a distribution of heat throughout its mass which changes

also with wonderful slowness. The earth possesses size, struc-

ture, composition, compactness, and other physical and chemical

properties which change so slowly that in the lapse of ages

differences can scarcely be detected, and millions of generations

come and go under exactly like influences.

Passing now from these unchanging factors to the living

organism itself, we must see that every particle of protoplasm is

affected by the components and by the properties of the atmos-

phere, by the physical and chemical qualities. of water, by the

force of gravity, and by the earth as a whole. Every particle of

protoplasm since it came into existence as such, every molecule

of every compound in it, and every atom in the molecule, has

existed from its beginning on this earth under these conditions

and subject to these influences. None of these influences has

ever been eliminated by experiment, nor has experiment ever

resulted in accomplishing any fundamental change in a living

“organism or series of living organisms. Man as an experimenter

cannot control these influences but is controlled by them. Is he

not controlled by them in every other relation in life? Is man

any more controlled by these unchanging influences than any

other living or lifeless thing ?

In the living substance of sperm and egg the component

parts, particles, molecules, and atoms, have been subjected to

these unchanging forces, not only since coming together as the

living structure, but before; and after sperm and egg fuse the

same is true; and in the growth of the fertilized egg every

particle of new material is formed, placed, and kept in place

under these influences. From the beginning to the end of its

career every individual plant and animal is subjected to these

continuous and uniform influences. But so also is every other

thing. And as we find all common salt crystals behaving alike

and being fundamentally alike at the same time that they are

unlike the crystals of all other substances subjected to these

No. 448. HEREDITY AND ENVIRONMENT. 2

same influences, so we find the plants or animals of a kind

behaving and being in the main alike at the same time that

they are unlike the plants and animals of all other kinds, although

they are subjected to the same influences. In the diversity of

composition and adjustment to these forces we have a physical

reason for the diversity of behavior of different animals, plants,

and lifeless things. They are all influenced by these forces;

what they are represents their reactions to these and to other

forces. The fundamental likeness of parent and offspring

represents the continuity of substance and of influence; the

superficial differences represent the different influences to which

they have been subjected and to which they have reacted. Not

all common salt crystals are of exactly the same size. Not all

the puppies of a litter are exactly alike. But the salt crystals

are fundamentally alike, and so are the puppies. With their

vastly greater complexity — considered merely chemically for

the moment — one should expect puppies to vary more than salt

crystals. But neither salt crystals nor puppies vary so far as

not to be salt crystals or puppies; the continuous influences

conserve their fundamental characters.

In this discussion two things have been assumed — the con-

tinuity of substance from parent to offspring, and the irritability

the power of reaction, of this substance to the various factors of

its environment. These two facts are essential to heredity.

My contention is merely that in the continuous, unchanging

factors of the environment we have forces, influences, stimuli,

under the operation of which the living substance came into

existence, under which it continues to exist, and to which it

continuously and unchangingly reacts. These influences are

factors in the environment, but at the same time, like irritability,

they are factors in heredity. The clearer our conception and

the fuller our knowledge of the irritability and the power of

reaction of living organisms to external stimuli, the smaller the

mass of unexplained though not unexplainable phenomena for.

which we shall make heredity accountable.

STANFORD UNIVERSITY,

February, 1904.

ON THE DENTITION OF RHYNCHODUS AND

OTHER FOSSIL FISHES.

C R. EASTMAN.

AMoNcsT Palaozoic chimaeroids the complete dentition is

known in at least two species of Ptyctodus, two of Rhynchodus

and one of Paleomylus. These genera are all included in the

family Ptyctodontide of the Devonian, and present for com-

parison with recent chimzroids a single dental plate on each

side in the upper jaw, with a corresponding pair biting against

the outer side of these (as shown by marks of contact) in the

lower jaw. The question therefore arises whether the unique

dental plate on each side in the upper jaw of ptyctodonts is

homologous with the so-called * premaxillary " of Chimzera,

Callorhynchus, Rhinochimzera, etc., or with the so-called “ pala-

tine plate" of the latter, or with both premaxillary and palatine

taken together. :

This question appears to be answered conclusively in the case

of Ptyctodus, from which the modern type of dentition is derived

by pushing the low and elongate upper dental plate further back

in the mouth, and introducing a “ premaxillary ” or “ vomerine :

tooth in front of it. In all cases the lower dental plate is

vertically deeper than the upper, and rises into a prominent

beak anteriorly. It is also characterized by having a descending

process at the symphysis, which is more accentuated in the

fossil than in recent forms. This process bears a triangular

groove or excavation on its inner face, the roughened surface of

which indicates that it was occupied by cartilage, since there

was no sutural union at the symphysis. That this was the case

is self-evident, for the anterior beaks could not have closed

outside the upper dental plates when the mouth was shut unless

the lower ones were mutually separated by a slight interval.

In a recent communication by Jaekel, it is stated that “von

Jakel, O., Ueber Ramphodus etc., Sitsungsber. Ges. naturf. Freunde, Berlin,

1903, pp. 383-393- a

296 THE AMERICAN NATURALIST. [Vor. XXXVIII.

der Zahnform und dem Gesammtgebiss von Ptyctodus wissen

wir noch nichts genaueres,” and it is thought that “ vielleicht

ist Ptyctodus schon ein echter Vertreter der sechszahnigen

Holocephalen,....die wohl von den Coccosteiden abstammen

mögen.” This author's evident unfamiliarity with the Ptyetodus

type of dentition is no doubt responsible for his confusion of the

upper and lower dental plates of a species of Rhynchodus from

the Upper Devonian of Wildungen, Waldeck, and for the

impossible suggestion that the nasal capsules projected into the

triangular incisions which occur in the descending process of the

mandibular dental plates. |

That which is commonly interpreted as the /ower dental plate

z of Rhynchodus, Jækel

homologizes with the

* premaxillary tooth ” of

Chimera ; and that at-

tributed to the upper

jaw of the former, Jaekel

supposes to have func-

tioned as a mandibular

element. Referring to

the lower dental plate

from the Eifel Devonian

described by F. v. Huene

Rheni under the name of Ayz-

ynchodus major, trom the a poca sed chodus emigratus, Jackel

e hn Pita & states that he prefers to

regard it as a “Preemaxillarzahn,” and notes its close resemblance

to the Wildungen teeth called by him Ramphodus tetrodon. So

far as one may judge without having compared the original speci-

mens, no essential differences exist between these forms and the

earlier described Rhynchodus major and R. rostratus! respec-

üvely. An illustration of the Wildungen dental plates is given

in the accompanying text-figure 1, slightly modified after Jekel,

that is to say, the latter's figure is inverted, and the upper dental

plate is thrust forward so as to protrude beyond the lower.

Fic. 1.— Left

1 E 7

: Eastman, C. R., Dentition of Devonian Ptyctodontide. Amer. Nat. vol.

XXII, p. 487, 1898.— Centralblatt für Mineral., 1900, p. 177.

No. 448.] DENTITION OF RHYNCHODUS. 297

RHYNCHODUS: PERTENUIS, Sp. nov.

Dental plate narrow and elongate, with sharp and extended

cutting edge and knife-blade cross-section; anterior beak prom-

inent, no symphysial process, external surface smooth.

The unique dental plate upon which this species is founded

was obtained from the Chemung of Franklin, in Delaware

County, New York, and is preserved in the State Museum at

Albany, where the attention of the writer was called to it by

Dr. J. M. Clarke, State Palaontologist, but not in time to

include its description with other fish remains already made

known from the same locality. The general outline and pro-

portions of this form differ from those of all other species, and

the absence of a symphysial process is a very unusual feature.

But for the trenchant cutting edge and narrow cross-section, the

Fic. 2.—RAynchodus pertenuis, sp. nov. Mandibular dental plate, x ł. Chemung group;

York.

Lync:

Delaware County, New

specimen might be readily mistaken for a lower dental plate of

Ptyctodus, instead of Rhynchodus. That it is properly a man-

dibular element, and referable to the latter genus, seems to admit

of no question. The hollow along the base indicates the extent

to which the plate was buried in the supporting cartilage of the

jaws. The total length is 9 cm.

1 Ann. Rept. State Geol. N. Y. 1897 (1899), pp- 317-327.

298 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Fragments of chimzroid jaws have been previously reported

from the Chemung of New York State by Clarke, but

no specific identifications were attempted by him. At best this

class of remains appears to be very rare in the eastern province,

Ptyctodus and Rhynchodus being the only genera that are

known from the New York Devonian. An undescribed species

of the former occurs in the Corniferous limestone of Le Roy,

and P. calceolus is apparently represented in the Hamilton stage

at Eighteen Mile Creek, near Buffalo. . Detached tritors from

both of these localities are preserved in the Museum of Com-

parative Zoólogy at Cambridge.

ONcHOSAURUS Gervais.

Syn. Ischyrhiza Leidy ; Gigantichthys Dames.

A comparison of the type specimens of Gervais’ Onchosaurus

radicalis! and Leidy's [schyrhiza antiqua,? the former being pre-

served in the collection of the School of Mines at Paris, and the

latter in the American Museum of Natural History in New

York, leaves no room for doubt that they are generically, and

probably also specifically identical, in which case Leidy's title

must be abandoned. The original of Gervais’ description,

together with one or two duplicates, was derived from the

Upper Cretaceous of Meudon, near Paris, and regarded through

error as of mosasaur nature. The identical form occurs also in

the Meestricht Chalk, a remarkably fine specimen from this

locality being preserved in the Paris Museum of Natural His-

tory.

The type of Leidy’s genus, /. mira? was founded on a unique

tooth from the Cretaceous Greensand of Burlington County,

New Jersey, and supposed by the author to represent a Teleost

fish related to Sphyrena. The original has never been figured,

! Zoologie et Paléontologie Françaises, vol. 1, p. 262, pl. lix. fig. 26, 1852.

* Proc. Acad. Nat. Sci. Philad. vol VII, p. 256. 1856. — Emmons, E., Ae-

port North Carolina Geol. Surv., P. 225, figs. 47, 48, 1858. — Leidy, J, m

F. S. Holmes’ Post-Pliocene Fossils of South Carolina, p. 120, pl. xxv, figs. 3-8.

1860.

? Proc. Acad. Nat. Sci. Philad, vol. VII, p. 221. 1856.

No. 448.] DENTITION OF RHYNCHODUS. 299

and its present whereabouts are unknown: The so-called

Ischyrhiza antiqua is stated by Leidy to occur in New Jersey,

North and South Carolina, New Mexico and Mississippi, but the

differences between this and Z. mira are inappreciable, and the

two were finally pronounced identical by their author. Certain

hypural fans similar to those accompanying Protosphyrzna

in the English Greensand have been theoretically associated

with Ischyrhiza, but with questionable propriety. No reasons

have been assigned for making this association, and other con-

siderations militate against it, hence it appears advisable to

exclude these fans altogether from the same genus. *

The wide geographical distribution enjoyed by Onchosaurus is

shown by its occurrence not only in Europe and America, but

also in the Upper Cretaceous of Egypt. In 1887 a tooth

differing from the type species only in unimportant particulars

was described by Dames? from the Senonian of Gizeh under the

name of Titanichthys pharao, the generic title being subsequently

changed to Gigantichthys. Dames’ figure was copied. by Zittel

in his * Handbuch "? and the two authors agree in placing this

form in the vicinity of Enchodus, although * [schyrhiza " is

referred by Zittel in the same work to the Esocida. The latter

position was first suggested by Cope, and is likewise adopted

by O. P. Hay* It will be seen, therefore, that there is good

authority for regarding Onchosaurus as one of the early fore-

runners of the pikes.

CAMBRIDGE, MASS.

! Cope, E. D., Vertebrata of the Cretaceous Formations of the West, p. 280.

187 5.— Clark, W. B., Bull. U S. Geol. Surv. no. 141, P- 60, pl vii, fig. 2a.

1897.— Maryland Geol. Surv., Eocene, p. 112, pl. xii, fig. 8. 1901.

2 Sitzungsber. Ges. naturf. Freunde, Berlin, p. 70, with figure, and p. 137. 1887.

3 Handbuch der Paläontologie, vol. TIL, p. 269, fig. 274- 1890.

* Bull. U. S. Geol. Surv. no. 179, p. 398. 1902.

FURTHER INSTANCES OF PARIETAL

DIVISION.

ALES HRDLICKA.

Since the publication of my monograph on “ Division of the

Parietal Bone in Man and other Mammals,” ! there appeared in

print three other direct or indirect communications on that same

subject by Frasetto,? Schwalbe,* and Le Double.*

The new instances of the anomaly reported by these authors

are as follows :

A. Inhuman adults.—1. An antero-posterior complete divi-

sion of the left parietal in an “adult Savoyard ” (Le Double) ;

2. A complete vertical division, “originating from the sagit-

tal point " and “terminating in the inferior border" of the right

parietal,in the skull of an adult male (Le Double). The race

of the individual is not mentioned and the description leaves

much in doubt.

3. A division in the right parietal of a ** heavy, sclerotic,”

skull of a male Egyptian. The division runs from the coronal

to the posterior third of the inferior border, and connects with a

trace of a vertical division (Frasetto). As the author states that

the usual sutures, even the parieto-temporal, are sinostosed, there

is great uncertainty as to the character of the division; it sug

gests a traumatism more than an anomalous suture.

B. In human children and fatuses.— 1. An antero-posterior

complete division in the right parietal of a seven year old child,

sex and race unknown. Skull hydrocephalic (Schwalbe).

2. An antero-posterior complete division in the left parietal

! Bull. Amer. Mus. Nat. Hist., Vol. 19, Art. VIII, pp- 231-386, N. Y. July,

1903.

* Notes de Craniologie comparée, Ann. sci. nat., Pp- 148-187, September, 1903.

? Ueber getheilte Scheitelbeine, Zeitschr. Morph. Anthrop. 1903, PP- 1-74;

Stuttgart.

* Traité des variations des os du crane de l'homme, Paris, 1903, “Parietal,”

Pp. 101-141.

301

302 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of a new born child.

phalic (Schwalbe).

3 "

Sex and race unknown. Skull hydroce-

An antero-posterior complete division in the right parietal

in a female new born child (Frassetto).

4. Ananterio-posterior complete division in the right parietal

of a six months' foetus.

lo 2

Sex and race not given (Frassetto).

An antero-posterior complete division in the right parietal

"3 p"

dem P =

1.— Anomalous division of the left parietal bone in a rachitic human foetus.

of a four to six months’ foetus, sex and race not given (Frassetto).

6. A separation of the mastoid angle of the left parietal in

an infant. Sex, age and race not given (Frassetto).

C. In Primates.— 1. Simia sabea (Cercopithecus). Right

parietal divided into four parts. This same case, apparently, is

reported by both Frassetto and Le Double.!

! Frassetto says the skull is No. A-1341 of the galeries of comparative anatomy

of the Museum of Paris, while Le Double reports his specimen as No. A-134, 1n

the Museum of Comparative Anatomy, Paris. Neither account is a thorough one.

No. 448.] PARIETAL DIVISION. 303

2. A vertical, curving, complete suture in the left parietal

of a young Semnopithecus (Frassetto).

f a

3. A vertical, complete suture in the right parietal

Cercopithecus callitrichus (Frassetto).

4. An oblique, vertical, complete, but synostosed suture in

the right parietal of a Macacus sinicus (Frassetto).

A separation of the sphenoidal angle. of the right

parietal in a Mormon maimon (Frassetto).

D. Other Mammals.— 1. Ursus americanus, young. The

liti i ietal in the rachitic human fcetus

Fic. 2.— Abnormalities of the right parietal in the rachitic human !

left parietal is divided into four irregular pieces. On the rig

} " 1345

al angle and an oblique,

IS found a separation of the sp shenoid à;

from the

incomplete suture running downward and forward

case, too, is report

sagittal border near the lambda. This

d E. e Doul DIE.

independently and imperfectly by Frassetto a

parietal « d um aeu came tí

©.

2 y) mv

l'he new cases of complete :

attention since July, 1903, are briefly these :

304 THE AMERICAN NATURALIST. (VOL. XXXVIII.

B. i. A human foetus, of white parents, born at term,

ninth pregnancy. The child weighed three pounds and lived

four days. Whole skeleton highly rachitic.

Both of the parietals, besides other bones of the skull, show

considerable modification. (Figs. 1 and 2.)

The left parietal is divided into two by a narrow and some-

what irregular membranous space running antero-posteriorly,

very nearly parallel with the line of, the sagittal junction, which

is also membranous. The upper portion is slightly higher than

the lower, its maximum height, measured by a tape, being 4.2

cm., while that of the lower piece is 3.8 cm.

The anterior third of the dividing space is very wide, forming

a large fontanel, and this is filled with one large and one smaller

secondary bones. Posterior to the two portions of the parietal

and between these and the occipital, from the sagittal line to the

mastoid, is another space, in the mean 2.5 cm. broad, somewhat

narrower inferiorly than superiorly, filled with various sized

secondary ossicles.

The squamo-parietal junction and much of the fronto-parietal

are still membranous. |

On the right there is plainly but one parietal. This is com-

paratively small and somewhat irregular. In about the middle

of the anterior border is a V shaped defect (fontanel), corre-

sponding to that on the left, and filled with a moderate sized

secondary bone The whole fronto-parietal junction is occupied

by a row of such bones and the same is true of the sagittal,

parieto-occipital and to a less degree the squamo-parietal spaces.

One of the secondary bones occupies the antero-superior angle

of the parietal area and is of a large size, but is plainly of an

accessory character, formed from an accidental accessory focus

of ossification. Another larger bony piece occupies the asteric

angle. |

The skull has been somewhat deformed in preparation or dry-

ing and the posterior parietal region on each side is depressed,

showing on this account but imperfectly in the illustrations.

The bregma fontanel is large and partly filled with small sec-

! No. 9754, Army Med. Museum ; gift of Dr. M. D. Spackman.

No. 448.] PARIETAL DIVISION. . 305

ondary ossicles. The occipital bone shows the ordinary at this

age separation of the squama, exoccipital and basal portions.

The development of the temporal bones, particularly the squa-

mze, is much retarded.

This case is of interest in several ways. It is another instance -

where the anomaly of parietal division is associated with a pro-

nounced pathological condition of the skull. Such association,

particularly with hydrocephalus (some degree of which may have

existed even in the skull under consideration), is so common in

the children and fcetal series of the cases reported that the

causal relation of these pathological conditions with the divisions

becomes more and more firmly established. They, of course,

play the role of the exciting cause only, the fundamental condi-

tion which makes a parietal division possible being the presence

of two starting foci or centers of ossification of the bone. In

this connection one is forcibly reminded of the apparent rarity

of pathological conditions in the adult human and also in the ape

and monkey skulls with parietal divisions. Even if it be granted

that much may right itself during the growth of the skull, it

would seem that at least some of the parietal divisions in man

and most of those in lower primates must be due to other excit-

ing causes than rickets or hydrocephalus.

The second point of interest in the present case is the

presence of two large and plainly accessory bones (antero-

superiorly and postero-inferiorly on the right) which in an adult

skull could easily be taken for primary portions of the parietal.

As can be seen in the illustration the small true parietal on the

right side shows a marked cleft near the middle of the anterior `

border. This cleft, it has been amply demonstrated before, is a

remnant of the original membranous space between the upper

and lower parietal centres. We had here, then, the two normal

elementary foci of the bone and in the usual position. But the

growth of the already fused primary parietal, due to rachitis,

was retarded. Such a retardation in any of the bones of the

cranial vault and from any reason leads invariably, undoubtedly

through some trophic impulse which regulates the cran

growth, to the appearance of more or less numerous secondary

foci of ossification, from which result various sized supple-

306 THE AMERICAN NATURALIST. [Vor. XXXVIII.

mentary, compensatory bones, commonly known as the wormi-

ans. Some of these secondary centres, as a rule those in

localities where the greatest deficiencies exist, which is at the

fontanels, show often more vital strength than others, enlarge

to more striking dimensions and eventually, meeting and articu-

lating with the advancing primary parietal, seem to represent

and are mistaken for separated parts of this bone. There

is no doubt but that the great majority of the “bregma,”

“human interparietal,” and supraoccipital bones, as well as

many of the “separated angles of the parietal” belong to this

category. The difference between the compensatory bone and

one that arose from lack of fusion of the primary centres is

morphologically and particularly etiologically important.

The third point that the case at hand illustrates very hand-

somely is the possibility of a formation of a vertical parietal

suture without any division, or totally independent of a division,

of the primary parietal. Had the conditions in this skull

advanced to a full development and particularly into adult life,

before which period many of the closely packed wormians fuse,

we should have had, unless an early synostosis obliterated the

L B

Fic. 3.— Bilateral, nearly obliterated vertical parietal division in a Hapale.

feature, a case very much like that of Fusari, which posess, Iam

inclined to think, falsely as an example of vertical parietal divi-

sion in a human individual. i

C i. Hapale, male, adolescent (No. 36,222, Dept. of Biol-

ogy, U. S. Nat. Mus). Skull apparently normal, symmetrical.

The ordinary sutures all open. Each parietal shows a plain

trace of a complete, vertical, now synostosed division. On the

left the division began superiorly ọ mm. posterior to the bregma

and 13 mm. anterior to the lambda, ran, slightly curving and

nearly parallel to the coronal suture, to the temporal ridge, then

No. 448.] PARIETAL DIVISION. 307

made a slight bend backward and ended a short distance anterior

to the squamo-mastoid junction. The right division began one

mm. posterior to that on the left and running a much more

angular course terminated in the same relative position as that

in the opposite parietal (Fig. 3).

2. Cebus apella, male, adolescent (No. 59,298, Dept. of

Biol. U. S. N. M). Skull slightly asymmetrical, surface of

bones irregular (rachitis?). No injury. On the left side a

serrated, vertical-oblique suture separates a large portion of the

antero-inferior angle of the parietal. The anomalous suture

begins anteriorly ro mm. from the. point where the coronal

meets the fronto-malar suture (there is a bilateral malo-parietal

articulation), and 36 mm. from the bregma; it ends inferiorly

14 mm. posterior to the meeting of the malo-parietal with the

spheno-parietal suture and 51 mm. anterior to the asterion. On

the right side 14 mm. above the point of meeting of the coronal

and the frontal suture is a small v cleft in the parietal and from

this runs backward and slightly downward a 4.5 mm. long fissure.

On the same side exists a 13 mm. long, slightly wavy, vertical

fissure in the frontal squama. It rises vertically from the fronto-

Fic. 4.—The lef ietal of a Cebus apella, showi paration of the antero-inferior angle.

malar suture and reaches the frontal part of the crest of the

temporal muscle. There is no trace of any violence that might

account for this fissure. (Fig. 4.)

The total number of ape, monkey, and lemur skulls examined

in the U. S. National Museum was 316. A large majority of

these skulls are those of adults, but no specimen was considered

in which all the normal sutures of the cranial vault were not

308 THE AMERICAN NATURALIST. [Vor. XXXVIII,

plainly traceable. The varieties, and the parietal divisions found,

are as follows :

UA m

Lemurs . , . no division.

Galagos . ; ; i : Mf: "

Tardigradus . : : i REA ie

Propitheci ; ; ie x

Gorila . : pn y

Orang. ; i i ; Pe

Chimpanzee . ; : EON "

Gibbons . ; (CLE

sembDopitbecl . `. : ; E xe

Presbytes ; ; 3 : EE ien j

Simias concolor ; 3 ; a 2

Nasalis larvatus ; ; cH n

Cynopithecus niger . : a T

Macaques no division in 79; in one, from Siam (No. 83,-

274), there is in the right parietal above the sphenoidal

angle a moderate size, curving, antero-posterior fissure; in

one M. rhesus (No. 83,476), a vertical fissure runs on the

left from the posterior third, on the right from the poste-

rior fourth of the sagittal border towards near the middle

of each parietal; in another, M. rhesus (No. 63,379), there

is on the left parietal a trace of what was probably a com-

plete vertical suture, running from the middle of the sagit-

tal to the inferior border of the bone. -

Cynocephali no division in 14; in one (No. 22,904), young,

there is a partial vertical division in each parietal, left 10

mm. long and starting from between the anterior and mid-

dle thirds, right 19 mm. long and starting from between

the third and last fourths of the sagittal border.

Colobus . ; i . no division.

Cercocebi : : : ; P E ds

Cercopitheci no division in 18; in one (No. 36,277), there is

in the right parietal a vertical-oblique fissure which begins

superiorly between the anterior and middle thirds of the

sagittal border and runs to the parietal eminence.

Cebi no division in 32; in one (No. 59,298, described in

No. 448.] PARIETAL DIVISION. 309

detail), a suture separates the left sphenoidal angle; in one

(No. 82,779), there are two fissures, each 10 mm. long, in

the superior third of the coronal border of the left parietal.

9 Hapale no division in 8; in one (No. 36,222, described in

detail) a bilateral complete vertical division.

4 Midas ; ; : : i . n0 division.

I Aotus ; " "

Brazil monkey No. T there is a 4 mm. long, vertical fis-

sure in the superior border of the left parietal, slightly ante-

rior to its middle.

7 Chrysothrix : ; . no division.

2 Lagothrix ; ; i eae "

IO Mycetes . ; ; : : Oh M "

4 Alma : ; 2 | n Ht

I Nictipithecus viti Y. js

16 Ateles no division in 14; in one 4. poo (No. 8,974),

young, the left parietal shows two incisures, one horizontal,

10. mm. long, just above the lower third of the coronal

border, and one vertical, 15 mm. long, running from the

middle of the sagittal border. In one A. ater (No. 63,425),

there is in each parietal, a short distance anterior to the

middle of the superior border, a vertical, 9 mm. long fissure.

The main facts accentuated by the examination of this

National Museum series of monkey skulls is the relative rarity

of parietal divisions in adult specimens. The condition in all

forms should be sought for preferably in the young. As a rule,

synostosis is later at least in some, and at times in all, of the

normal parietal articulations than in the abnormal divisions.

U. S. NATIONAL MUSEUM.

COMPARISON OF THE PROVISIONAL SCHEMES

OF THE CLASSIFICATION OF BIRDS?

R. W. SHUFELDT.

Ir is not my intention in the present paper to offer any scheme

of my own for the classification of birds, although it is a question

that has long engaged my attention, and I hope soon to publish,

in another connection, a provisional scheme, presenting what I

take to be a natural taxonomy of Aves in so far as it is now

understood. My only object here is to offer a few brief remarks

upon the more prominent schemes for the classification of birds

which have been put forward within recent times, and in a way

compare the views of their sponsors. Careful recapitulation,

undertaken from time to time, is always an advantage to any

science, especially if that recapitulation is made along compara-

tive lines and according to scientific methods. In other activities

in which men engage, the benefits attaching to the occasional

calling of a halt, with the view of taking account of the progress

made; to making sure that advancement is being made along the

right lines, has always been recognized. This, too, holds true in

the domain of ornithological science. In fact, those who make

the greatest, the surest and most substantial progress in anything

are the ones who command a digested and available knowledge

of all that has been previously accomplished in the field in which

they labor.

It wil be a red-etter day for our science when any spe-

cies or subspecies of birds is and are known throughout the

world by the same name, vernacular or scientific. That is, the

opinion in regard to nomenclature will be unanimous. There

will be equal rejoicing when that day arrives, when a unanimity

of opinion exists in regard to the classification of birds. It is

quite possible that many species now existing in the world's avi-

! Read by title at the twenty-first Congress of the American Ornithologists’

Union, held at the Academy of Natural Sciences of Philadelphia, 1903-

31!

312 THE AMERICAN NATURALIST. [Vor. XXXVIII.

'fauna will, when that time comes, be extinct. With nomenclature

I have nothing todo. Names are the inventions of men, whereas

on the other hand, the relationships existing among birds in nature

are actual, and in so far as invention enters here, it can only be

in the form of printed, diagrammatic, or other device, to convey

to the eye and mind what our conceptions of those relationships

are. We may change name$ at any time and invent new ones ad

libitum, but not so real relationships. These are fixed, and it

remains for us to ascertain what they really are, and express

them in the simplest terms. This is a matter of time, and I

know of but two ways by means of which a consensus of opinion

of ornithologists can be arrived at. First, by our mastering the

morphology, geographical distribution, habits and life histories of

all existing forms, and the osteology and other remains of all

extinct ones within our ken; second, by the meeting of com-

petent ornithologists in congress to discuss anything that touches

upon the classification of the Class, and especially of the visible

means of representing digested ideas in regard to it. Much

could be accomplished by an international congress, like the

Second International Ornithological Congress which met at Buda-

pest in 189r. :

Of all the papers read at that Congress, none attracted more

attention nor has been more useful or more closely studied since,

than the paper read by Dr. R. Bowdler Sharpe, entitled *A Review

of Recent Attempts to Classify Birds." It is the best thing of the

kind extant and is so well known to ornithologists the world over

as not to need further comment. I acknowledge with pleasure

the assistance it has been to me in preparing the present paper.

Apart from the many sound suggestions made by Dr. Sharpe in

that address, and the historical lore it places at one’s command,

the main assistance I have derived from it has been the oppor-

tunity it affords me to study and to compare so many of the.

schemes of classification that have been proposed from time to

ume. To be sure, there now exist a number of other avian clas-

sifications.. I refer especially to the classification of Aves pro-

posed by Cope in 1889, entitled * Synopsis of the Families of

Vertebrata," The American Naturalist, Vol. XXIII, pp. 849-877;

and also to the taxonomic scheme brought forward by Gadow in

No. 448.] CLASSIFICATION OF BIRDS. 313

his contribution to the Proceedings of the Zoblogical Society of Lon-

don for the year 1892, entitled * On the Classification of Birds,"

(pp. 229-256). So far as I am at present aware, Cope's and

Gadow's classifications of Aves are the only two of any impor-

tance that have been published since Sharpe gave us his brochure

cited above. If this be true, there has been no completed classi-

fication of this Class of Vertebrates published for over ten years.

There have, however, been a number of such schemes partly

completed and partly published, as for example the classification

of birds as set forth in Sharpe's Hand-List of Birds, now passing

through the press, and of which but one part remains to be issued.

This admirable and most useful work will contain one of the most

elaborate classifications of birds ever published. It is especially

valuable inasmuch as Sharpe belongs to that school of ornithol-

ogists which believes in employing all available characters in

classification, in ascertaining true affinities, to the end that the

Classification shall be a natural one and express as far as possible

the real relationships of all existing families of birds, even to the

minor divisions of species and varieties.

Within the past few years there have appeared some excellent

summaries of classifications ; for example, Ridgway's admirable

presentation of the matter in his Birds of North and Middle

America, “N othing original is claimed for the classification

here given," says its author, * except as to the form in which it

is presented. It is simply the result of an elective process, the

evidently good of other systems being retained and the obviously

bad rejected, according to the author's ability to correctly inter-

pret the evidence " (p. 6). In this connection I desire also to

invite attention to the avian classification found in the Catalogue

of Osteological Specimens of the Museum of the Royal College of

Surgeons of England (Part III, Aves). This admirable piece of

work is by Sharpe, who states that “The system of Classi-

fication followed in the present work is mainly that proposed by

Henry Seebohm in his ‘Classification of Birds,’ and further

elaborated in this ‘Birds of the Japanese Empire.’ There are

Some points in his system which I have slightly modified ; but

they are of minor importance when compared with the fact that

every group of birds, as diagnosed by Seebohm, possesses à

314 THE AMERICAN NATURALIST. [Vor. XXXVIII.

combination of definite features, which are characteristic of the

group, and of that group alone, be it Order or Suborder” (p. 1).

Finally, in many of the ** Manuals " and * Keys " and * Hand-

books " and * Check Lists" published in various countries, we

have other classifications, but these, inasmuch as they do

not enter upon the subject in its entirety, are apt to be more

or less unsatisfactory and often misleading. The Check-List of

North American Birds, prepared by a Committee of the Ameri-

can Ornithologists’ Union " (Second and Revised Edition, 1895),

is a very good example, for in it we find a classification that

although it would be of great credit to a taxonomer of the Cur-

vierian epoch, it certainly can now only be regarded in the light

of a curious bit of antiquated literature which it would be diffi-

cult to fit into any modern taxonomy of the Class Aves

published since the days of such worthy pioneers as Nitzsch, Ill-

iger, and Müller. As cited above this classification appeared

in 1895, yet in r9or when Ridgway, who was a member of the

aforesaid Committee responsible for the classification in the

* A. O. U. Check-List," published his own taxonomic scheme

the latter differed so markedly from the former that to compare

them is quite like making a comparison of Wilson's old single-

barrel, flint-lock gun with the finely finished modern double-

barrelled, hammerless piece now in the hands of the present day

ornithologist.

It would seem that we at least ought to be in position at the

present time, or certainly in the very near future, to decide upon

the main groups into which the Class Aves is naturally to be

divided, yet such is by no means the case. This is the more

remarkable, inasmuch as all the important part of the develop-

ment of avian classification dates no further back than the one

proposed by Huxley in 1867. This scheme belongs to the liter-

ature of the Darwinian epoch and was influenced by what was

then known of the law of organic evolution, and consequently is

the first scheme of classification worthy of our consideration.

Huxley divided the Class into three orders, the Saururze, the

Ratitæ, and the Carinatæ, and these three orders were divided

into their suborders and certain groups.

Seven years later, or in 1874, appeared the well-known classi-

No. 448.] CLASSIFICATION OF BIRDS. 315

fication of Garrod, who it may be said, almost entirely ignored

Huxley's scheme by dividing the Class Aves into two subclasses

—the first containing four orders and the latter three, or in

other words two sub-classes and seven orders as compared with

the latter’s three. Garrod’s initial scheme of classification is not

thorough since we meet with such incongruities as the Cathartidze

being considered simply as a group in the same order with the

Steganopodes, herons and others, while the Columbidz and the

Gallinze are widely separated, and the penguins are placed as a

family among the Anseres, immediately following the Anatidze, or

the ducks, geese, swans and theirallies. Still keeping before

us, however, the main divisions of the class it is to be noted

that ornithologists had little more than fairly grasped the Garro-

dian idea of avian relationships when in 1880, six years after its

publication, Sclater proposed another scheme. In it the Saururze

of Huxley are not considered, — the class is divided into two

subclasses, the Carnatz and Ratitz, the former containing no

fewer than twenty-three orders, and the latter three others, or

twenty-six orders of birds, where Huxley only recognized three ;

and these three orders Newton considers to be so many sub-

classes, while he would divide the Ratitze into no fewer than six

orders. These classifications were almost immediately followed

by Reichenow in 1882 who divided birds into seven main groups

Which he called series, and these seven series were represented

bv seventeen orders. It is very different indeed from any of

the foregoing schemes and cannot be contrasted with them with-

out great difficulty, while its chief interest lies in the fact that

he published in connection with it a phylogenetic tree of the

Class Aves, one of the first attempts of the kind employed in

ornithological science.

Within the next ten years a number of important classifica-

tions followed, — all provisional schemes for the arrangement of

the Class, but none the less entitled to our best consideration,

Coming as they have from the pens of the ablest living ornithol-

ogists.

Stejneger’s appeared in 1885; Fürbringer's in 1888; Cope's

in 1889; Seebohm's in 1890; Sharpe’s in 1891; and Gadow's

in 1892. Of all these Fürbringer's is the most elaborate and

316 THE AMERICAN NATURALIST. [Vor. XXXVIII.

exhaustive, being accompanied by several vertical aspects and

horizontal projections of his phylogenetic tree. Indeed, the

objection brought against Furbringer's classification, principally

by Gadow, is that it is too long and too elaborate for practical

use. I do not fully concur in this opinion; moreover its author

has, in many groups carried us a long way on the road toward

determining the true relationships of birds and that, I take it, is

the real goal we seek. In fact the converse of this would be

an easy matter, that is to create a brief, artificial classification

of birds based upon our present knowledge of the class, and

adapted to the practical ends of the science. Any international

congress of ornithologists, representatives from all parts of the

world, could, in a few days, prepare such a scheme. But the

problem is not to be settled in any such manner.

As it is we find hardly any more uniformity with respect

to the schemes proposed by Stejneger, Furbringer, Cope, See-

bohm, Sharpe and Gadow, than is to be found among those of

Huxley, Newton, Garrod, Forbes, Sclater, Reichenow, and others.

The majority of these schemes carry the classification down

through the families, and, in special cases, in a few instances in

each, through the sub-families. Huxley did not give the num-

ber of families in the Passeres, Garrod omitted the group

entirely in his scheme; Sclater enumerated thirty-one of them;

Reichenow but twenty-one; Stejneger thirty-three; Fiirbringer

reduced the typical Passeres to one single family, the Passeridz ;

in 1889 the present writer recognized twenty families of the

North American Passeres, and Sharpe the following year

practically adopting the scheme, included all the old world

representatives, and by so doing admitted thirty-five families as

making up the passerine group, —and so on. -

There is little need of carrying such comparisons as these

into the higher divisions into which birds have been divided.

We would but meet with greater variance of opinions, made

the more deplorable from the fact that the wealth of coinage in

new names renders the comparisons instituted even more per-

plexing. Then this perplexity is in no way diminished when a

taxonomer takes it into his head to incorporate all the known

fossil forms of birds into his scheme, as quite a number have

done, and, very properly so.

No. 448.] CLASSIFICATION OF BIRDS. 317

It may naturally be asked, what are some of the chief reasons

why the classifiers of this group of vertebrates do not exhibit a

greater unanimity — a fact more remarkable when we come to

consider that a dozen or more of those classifications coming from

the pens of competent ornithologists appeared all within a very

few years of each other. It is certainly not due to the fact that

it has been demonstrated that birds have arisen from a prehistoric

and extremely ancient stock of animals in common with the Rep-

tilia, for knowledge of this character ought to have the tendency

to harmonize views and opinions rather than to introduce the

element of disagreement among them. We may eliminate too,

I think, any difficulty that has arisen from the discovery of the

few fossil forms of birds we have come in possession of, for many

of these belong to the minor groups of existing birds, while others

are not calculated to disturb a natural classification. Indeed, in

some instances, they shed light on the subject. Again, in that

existing birds are so completely differentiated from all other

classes of animals now living upon the surface of the earth, ought

to make them the easier to classify. They alone possess feathers

and this establishes a line of demarcation for them, standing

between the group and the nearest mammals or reptiles, quite as

clearly defined as the possession of a mainspring separates all

modern watches from an hourglass. The problem then presents

itself in this wise, — to ascertain the true relationships both near

and remote existing among all living birds, and then prepare as

simple a scheme as possible expressing these relationships in

terms that shall be in harmony with the classification schemes

adopted. in the cases of other classes of animals. In doing this,

one of the first difficulties to arise is the marked homogeneity of

the group. It is like classifying so many thousands of black,

leather-covered hand cameras ; they all look a good deal alike on

their outsides, and the task would be equally difficult were we

not permitted to examine into their interiors and ascertain the

differences in their other parts, as the different kinds of lenses,

finders, shutters, and other contrivances. Exactly the same

thing obtains with birds. The great variance of opinions in the

premises at the outstart is due to the difference in the amount

of knowledge possessed by the different classifiers, especially as

318 THE AMERICAN NATURALIST, [Vor. XXXVIII.

to the characters presented on the part of the insides of the

objects they are attempting to classify, although in saying this I

do not underrate the value of the external characters.

The facts, then, that birds are a very homogeneous group and

the knowledge of all their characters possessed by individuals

who have attempted to classify them, has differed very widely in

amount, is the first factor that will account for the great differ-

ences to be seen in the various published classificatory schemes.

These are not the only reasons, however, and another very obvious

one is the attempt made by some classifiers to ignore the homo-

geneity of birds, and to arrange them after the manner of the other

great groups of animals, such as mammals or fishes. In other

words, the attempt is made to employ the same divisional group-

ings from subspecies to class in the case of birds, where perhaps no

greater differences can be found than exist between a thrush and

an ostrich, as they do in the case of mammals, where such gaps

exist as the one separating man and the ornithorhynchus. The

objection is raised here at once, however, that an order of birds,

for example, is a very different thing from an order of mammals.

This is a knotty question, and as time forbids my discussing it

here, I can only say that it leads directly to another very obvious

reason for the differences seen in the various arrangements that

have been proposed for birds, and that is this : — although ornith-

ologists, in this country at least, may be pretty well agreed as to

what is meant by a species, it is not clear that the same apparent

unanimity of opinion exists among them in regard to what is

meant by a genus, or rather what constitute generic characters ;

and so on for families, suborders and other groups, until we

arrive at the Class, and perhaps the Subclass, — groups, for evi-

dent reasons, again understood to possess the same value in all

avian schemes of classification wherein they have been employed.

The difficulty here is, no uniform laws have been drawn up set-

ting forth for birds exactly what characters constitute specific

characters, what generic characters, family characters, and so on

up to Class characters. The consequence is that one avian

classifier will employ subclasses in his scheme, which subclasses

are designated in the scheme of another as super orders, or even

as orders or some other divisional value in the scheme of a third

No. 448.] CLASSIFICATION OF BIRDS. 319

taxonomer, and so on for all those who have engaged in this

difficult subject.

Finally, there is the great question upon which no two ornith-

ologists now entertain similar opinions, and that is upon the

various relationships of birds. Both this and the former ques-

tion, however, depend entirely upon the amount of knowledge

on the subject possessed by any particular taxonomer. The

more exact and far-reaching this is, the nearer and sooner will

he arrive at the truth.

In any event, it is very clear to me that the day is still far

away when ornithologists will be agreed in reference to all these

points. It is purely a matter of evolution, of development, and

the acquirement of the necessary knowledge. Guess work will

never attain the desired end, nor will any one man settle it. It

seems to me, however, that we are in a position to discuss and

settle one class of questions, that is in the case of birds, what

groups shall be adopted in their classification, and what charac-

ters in birds themselves shall stand for those groups. For the

rest the larger part of it depends upon substantially adding to our

present knowledge of the morphology of these forms in its widest

sense, and this to be supplemented by a very general knowledge of

the entire life histories of all existing birds. From the very nature

of things the latter advances with far greater rapidity than does

the former, and we stand in great need of the addition of

many more laborers in the fields of avian morphology. Death

has materially thinned the ranks of this part of our army within

à comparatively short space of time, and it has been principally

the great captains of whom we have been deprived, — and we

have by no means rallied from the loss of such workers in the

anatomy of birds as Huxley, two of the Parkers, Gegenbaur,

Garrod, Forbes, and other men of their calibre, power and

influence, any one of whom would have said that the solution

of the classification of birds lies in our commanding a knowledge

of their history and structure.

New York Crrv.

NOTES AND LITERATURE.

GENERAL BIOLOGY.

A Well Balanced Book on Theoretical Evolution.! — It seems to

be generally agreed that we are beginning a new era in the study of

Evolution ; an era in which analytic and experimental methods will

replace that of sharp logic. But the experimentalist works blindly

without hypotheses and these the speculative writings have provided.

It is a useful thing, at the beginning of this new era to have these

hypotheses brought together by a broad-minded investigator; and

this is the very arduous task that Plate has well done.

The immediate purpose of the work has been to stem the tide

away from Darwinism, to show that whatever limitations the theory

of natural selection may have as a complete theory of the origin of

species it remains the only satisfactory theory of adaptation. The

book, which is much increased in size over the first edition, is divided

into five chapters. The first deals with the objections, less or more

serious, that have been raised against Darwinism in the strict sense ;

the second with the different forms of selection and elimination; the

third with the complementary theories ; the fourth with the basic

elements of evolution from which the theory of selection starts,

namely, excess of births, variability, and means of isolation ; the fifth

with the range of applicability of the Darwinian and the Lamarckian

factors. Then follow a Bibliography of over 10 pages and a good

index.

The book is exceedingly satisfactory in most particulars. It is

refreshing to find an author who does not insist that there is only

one method of evolution. “Das Problem der Artbildung " he says,

page 228, “darf nicht einseitig behandelt werden, weder ausschlies-

slick von Lamarck'schen noch vom selectionistischen Standpunkte ;

nur die Vereinigung beider Principien führt zum Ziele.” Naturally

the author does not follow Weismann in rejecting the inheritance of

acquired characters and he is quite ready to accept the possibility of

‘Plate, L. Ueber die Bedentung des Darwin "schen Selectionsprincips und

Probleme der Artbildung. Zweite, vermehrte Auflage. Leipzig: Engelmann,

1903. p.

321

322 THE AMERICAN NATURALIST. [Vor. XXXVIII.

orthogenesis. In this breadth of view Plate doubtless approaches

nearer to the spirit of the great master than the. Neo-Darwinians.

Nevertheless, in so far as a criticism may be aimed at the book it

concerns Plate's unwillingness to accept more freely some of the

other subsidiary or alternative theories that have been proposed.

Thus the reviewer thinks that Plate does scant justice to the claims

of the mutation theory ; and in his discussion of means of isolation

he entirely fails to mention Mendel's law of the segregation of

parental qualities in the germ cells of mongrels. Nevertheless, this

law must be an important factor in preventing the swamping of

mutations. In his account of the different theories to explain organic

adaptation there is not included the theory proposed by the reviewer

and later by T. H. Morgan that there has been a selection by the

organism of the environment for which its structure is fitted — but as

the preface date is antecedent to the appearance of that theory such

conclusion could hardly be expected.

This brief review can give no adequate idea of the scope, sound-

ness and helpfulness of the book. It is recommended to biologists

as by far the best on the subject.

C B D

Experiments in Heredity.!— Stimulated by the rediscovery of

Mendel’s Law of crossing, Bateson and Saunders have thrown

together the results of their studies on crossing plants and poultry.

As this is the first extensive post-Mendelian account of hybridization

experiments in animals it may fairly be called epoch-making.

Miss Saunders worked with two hairy species of Lychnis and also

a glabrous variety ; with two varieties of Atropa; two species, each

with two varieties, of Datura; and various races of Matthiola. In

the first cross the hairy character is dominant and the glabrous

recessive, as shown by the fact that all the first crosses were hairy.

In the second generation, however, both dominant and recessive

forms appeared in Mendelian proportions.

The Atropa experiments were less complete, but appeared to be

Mendelian, showing dominance in the first color of the type form.

The Datura experiments, involving 12 characters, were much more

complex. Also there were exceptions to Mendelism in some Cases,

although striking adherence to it in others. Finally, the Matthiola

experiments were based on so many races mixed together that the

' Bateson, W. and Miss E. R. Saunders. Report I to the Evolution Committee,

Royal Society, London: Harrison & Sons, St. Martin's Lane, 1902. 160 pP-

No. 448.] NOTES AND LITERATURE. 323

results are hard to follow ; some were clearly Mendelian, others clearly

not; and some of these aberrant cases seem to be examples of what

Millardet has called “false hybridism " — where the second and sub-

sequent filial generations show no trace of one of the parents.

The poultry experiments were made with Indian Game, White

Leghorns, Brown Leghorns, White Dorking, and White Wyandotte.

The pea comb and single comb when crossed followed Mendel's law,

the pea being dominant. The extra (Dorking) and normal toe

followed the law approximately, the extra toe being dominant. In

other cases the results were non-Mendelian. Thus it may happen

that when a usually dominant character is crossed with a pure

recessive the first filial generation is not purely dominant, but a

mixture of dominants and recessives. It appears that a usually

recessive character may sometimes dominate. The mixed result in

the first filial generation may also be due to the fact that the “ domi-

nant" used in the cross was not a pure bred dominant but gave off

“recessive " gametes.

The last 35 pages of the work are devoted to an invaluable dis-

cussion of “The Facts of Heredity in the Light of Mendel’s Dis-

covery.” Here some new terms are introduced. In experiments in

hybridization. two forms exhibiting antagonistic characters are

crossed. There may be one pair or many pairs of these antagonistic

qualities. The antagonistic qualities are called allelomorphs.. The

zygote produced by the union of gametes with allemorphs is called

a heterozygote to distinguish if from a zygote formed of similar

gametes (homozygote). Allelomorphs may be either simple like

hairiness ‘or smoothness; or they may be compound, as the varie-

gated color of some flowers. When a compound allelomorph is

crossed with a simple the second filial generation may show not two

forms only but several — the compound allelomorph has broken up

into its constituents.

The relation of Mendel’s Law to “skipping a generation,” to pre-

potency, to sex (since elaborated by Castle) and to Galton’s Law

are discussed. The whole work closes with an eloquent “outlook”

over the future of experimental breeding.

C. B. D.

Walks in New England! is a series of lay sermons which appeared

in the Springfield Republican a year or two ago ; they are the records

. ' Whiting, C. G. Walks in New England, with illustrations from photographs.

London and New York, John Lane. 8vo., pp- 301, 24 illustrations.

324 THE AMERICAN NATURALIST. (Vor. XXXVIII.-

of a saunterer among New England's woods and fields. "They record

the aspects of the changing seasons from March to December with

eyes which, in turn, are those of a lover of plants and birds, a poet,

and a deeply religious man. For science he cares little, as compared

with “the intuition of spirit"; Emerson and Whitman are more to him

than Darwin and Wallace. The letters are not full of accurate detail

like Thoreau, nor of vivid coloring like Bolles; the style is often too

involved and the thought too mystical to suggest comparison with

Burroughs; but coming as they did from week to week, they must

have been very welcome to many who could not share the author's

rambles; they breathe the calmness, the toleration, the kindly sym-

pathy of a true lover of out-door nature.

ZOOLOGY.

Influence of Man on the Distribution of Reptiles and Mammals

in Patagonia and Fuegia.— In a very complimentary review' of my

recently published Warrative of the Princeton Patagonia Expeditions,

Mr. Barnum Brown, who, as à representative of the American Museum

of Natural History in New York accompanied me on my last expedi-

tion to that country remarks that my “ observations on lizards should

have been confined to that part of Patagonia north of the Rio Santa

Cruz, for this river forms the natural southern boundary line for liz-

ards as well as armadillos though a few have been scattered south of

it by man.” I have taken these small reptiles at Fitzroy’s Springs

on the north shore of tke Gallegos river, at various points along

the coast between Cape Fairweather and Coy Inlet, about the Salt

lagoons at the estancia of Montes and Fernandez ten miles from

Gallegos, at the Mount of Observation and at Greenwood’s estancia

sixty miles south of Santa Cruz and have observed them at many

other favorable localities in the region south of the Santa Cruz

River, while other travellers have reported them as being common

not only in this region but on the plains of Fuegia as well. See

Popper's account of Fuegia in Mulhall’s Hand-Book of the River

Plate. I see no good reason for attributing the present wide

distribution of these lizards over the region south of the Santa Cruz

River to the agency of man.

Amer. Nat., Nov. 1903, pp. 799-800.

No. 448. NOTES AND LITERATURE. 32

325

Mr. Brown’s remark that the presence of the guanaco in Fuegia

while the deer, rhea and puma are absent from that island is

attributable to the agency of man rather than to the superior powers

of self-distribution possessed by the guanaco, may be correct, but the

latter hypothesis seems to me the more reasonable one. If the

presence of the guanaco in Fuegia is due to the agency of man, why

is the rhea absent from that island? It is found quite as often in

captivity among the Indians, could have been just as readily trans-

ported and is more prolific than the guanaco. The readiness with

which the guanaco takes to water is well known in Patagonia as are

also its powers of swimming and to these characters is due I believe

its presence in Fuegia. The absence of the deer in Fuegia which,

as is well known, is also a ready swimmer is I think due to the fact

that it is a forest and mountain species and does not advance on the

plains as far as the “narrows” of the Magellan Straits. Farther

west the channels between Fuegia and the mainland are too wide to

be successfully crossed by either the deer or guanaco.

. J. B. HATCHER.

The Rat-tailed Rotifers. — Jennings has published a most

interesting and valuable monograph of the Rattulid Rotifers, which

although a part of a series entitled “ Rotatoria of the United States,”

actually includes the species of the whole world. In fact, one new

species described (Diurella dixon-nutialli) has never yet been found

in America but only in England. The species are divided into

two genera; Diure//a in which the two caudal appendages or toes

are equal or the shorter is more than a third the length of the longer,

and Rattulus,in which these organs are more unequal, one being

often quite rudimentary. The author states that these are not.

natural genera, but are justified by considerations of convenience.

He remarks that the idea that all the species of a genus must be

more related to each other than to any outside species has been

largely given up in practice; but this might be admitted and yet it

might remain true that the generic characters had not been acquired

independently. If the more primitive Diurella type has given rise

separately to two or more groups of species now included in Rattulus,

it will be necessary to either divide Rattu/us into as many genera, or

unite Diurella and Rattulus under the latter (older) name. O

‘Jennings, H. S. A Monograph of the Rattulide. Bulletin U. S. Fish Com-

mission for 1902 (1903). pp. 273-352.. Pls. I-XV.

326 THE AMERICAN NATURALIST. [Vor. XXXVIII.

course it may be impossible to demonstrate this, in which case the

present classification may very well be allowed to remain. The

genus Heterognathus, Schmarda, is applicable to the species having

the toes equal — part of the present Déure//a. The author admits

that this group may be thought worthy of separation, but he does not

notice that the name belongs properly to a genus of fishes, the latter

having five years’ priority. If the equal-toed species deserve a

generic name, a new one will have to be proposed.

Looking over the paper, one notices the absence of any records

from the region west of the great plains, as well as from other great -

regions. It is to be hoped that students will arise in some of the

neglected parts of the country, now that the study is made com-

paratively easy.

T. D. Aot

Gardiner's Reports on the Fauna and Geography of the Mal- :

dive and Laccadive Archipelagoes have now begun a second vol-

ume. The First Part contains an account of the Alcyonaria, by

Hickson and E. M. Pratt, of the nudibranchs by Sir Charles Eliot, of

Sponge crabs by Borradaile, of Lagoon Deposits by Gardiner and

on a Land Planarian by Laidlaw. The Part contains nine litho-

graphed plates.

Hickson discusses the remarkable variability of the Alcyonaria and

concludes that either they constitute a large number of closely similar

species or else one species capable of extraordinary variation in cir-

cumstances that are approximately identical. For practical purposes

the author regards those variations as species which are discontinuous.

Hickson finds that the form and mode of branching are unreliable

criteria of any species because they vary with accidental variations in

environment and the presence of gall producing Crustacea that reside

in the branches.

Eliot’s Report contains many interesting general data, concerning

swimming Hexabranchidz,, hidden but highly colored Dorididze, self-

mutilating Dicodoris, a Phyllid that secretes a liquid with disagreea-

ble smell and others.

CR D

Position of the Gordiacea. Montgomery concludes! from a

study of the adults that the Gordiacea agree with the Nematoda in

only the tubular gmitalia and their opening into the cloaca. They

! Zoolog. Jahrbücher Abth. f. anat. xviii 1903.

No. 448.] NOTES AND LITERATURE. 327

agree with the Annelids in structure and innervation of the muscles

and in dorso-ventral mesenteries bounded by epithelia. They differ

from Annelids in entire absence of true metamerism, in the absence

of a prestomial ganglion, in absence of seton and appendages and in

structure of genitalia. The group cannot be regarded as degenerate

Annelids (Vejdorvsky) or as modified Nematods, but must be con-

sidered as an isolated group (Funacher, von Siebold, Villot) until

more details concerning the development are known. ‘The per-

tinence of the peculiar genus ZVecferuma to the group is at least

questionable.

North Atlantic Invertebrata.-— Several papers in the rst Hefts of

the Bergens Museums Aarbog for 1903 have an interest to students

of the Invertebrata of our northeastern coast. Emily Amesen

catalogues the Sponges of the Norwegian coast, the present paper

containing the Halichondrina. R. C. Punnett enumerates. the

Nemertini of Norway in which thirty-four species are recognized,

of which twelve are supposed to be new. Edward T. Browne reports

upon a collection of nineteen species of Medusz, mostly from the

fiords around Bergen, four of them being new and eleven others

not previously catalogued from Norway. Among the interesting

points brought out is the fact that the peculiar sucking cups described

by Heckel in Ptychogastria polaris (Pectyllis arctica Heckel) are only

the stumps of broken off tentacles. Only four species of Leptome-

dusz are enumerated in the collection. All three papers are well

illustrated.

BOTANY.

The Desert Botanical Laboratory..—Of the occurrences of

recent date interesting to the botanists of this and other countries,

one of great importance is the establishment, by the Carnegie Institu-

tion, of a laboratory at which desert plants can be studied in their

native habitat. Messrs. Coville and MacDougal were asked to consti-

tute themselves a committee of inquiry, to determine the most suitable

place where such a laboratory might be located. We have before us

! Coville, F. V. and MacDougal, D. T. Desert Botanical Laboratory of the

Carnegie Institution. pp. 1-58, Pl. I-XXIX, fig. in text 1-4. Publication No. 6,

Carnegie Institution of Washington, Nov. 1903.

328 THE AMERICAN NATURALIST. (Vow. XXXVIII.

the report of their journey, undertaken early last year, which took

them over most of the desert regions of the western part of this con-

tinent. The place decided on for the laboratory was a site about two

miles from Tucson, Arizona, and since then the laboratory has been

built and equipped. Besides the fact that the country around Tucson

is of a distinctly desert type and the flora as varied as such a flora is

apt to be, the practical questions of accessibility and of habitability

were considered. With the two authors as advisory board, Dr. W.

A. Cannon was appointed resident investigator and is at present

engaged on various researches. Arrangements have also been made

so that a limited number of trained investigators may avail themselves

of the privileges of the laboratory. ‘ Not the least important part of

the duties of the resident investigator will be to aid visiting botanists

and others.”

This contribution besides containing a description of the trip under-

taken for the purpose of selecting a site for the laboratory, also

includes an account of the general botanical and climatic features

of the deserts of the regions visited. While the two authors had

both separately made trips to many of the same districts of the west,

the itinerary of their present journey included almost all of the arid

regions of the United States and of northern Mexico. Starting at El

Paso they first made their way down tothe sand dunes of Chihuahua,

south of Samalayuca in Mexico. The winter vegetation of these sil-

iceous sand hills is scanty, only a few forms are mentioned. The

next point of attack was the Tularosa desert lying westward of

Alamogordo, New Mexico, the most interesting feature of which is

the region known as the White Sands, composed of drifting sand that

is almost wholly gypsum. The characteristic plant of the dunes is

Rhus trilobata, the roots of which bind the sand so effectually that

clumps of the plant bring about the formation of pillars of sand when

the surrounding dunes shift. A curious relation of plants of Yucca

radiosa to the dunes was also noticed. Investigation showed that a

Yucca growing out of the top of a thirty-foot dune, must penetrate

with its trunk to the bottom. On excavating it appeared that the plant

must have grown as the dune engulfed it. Inthe bottoms among the

dunes the vegetation is much denser, a grass is plentiful and Ephedra

is frequently met with.

Tucson was next visited, there the woody vegetation of the desert

consists mainly of the creosote bush, the mesquite, joint pine, and

several Cactus forms, while higher on the foot hills occur the giant

Cereus, and species of the tree known as the palo verde. At the

No. 448.] NOTES AND LITERATURE. 329

time of the rains a variety of annual vegetation may spring up.

From Tucson the authors proceeded to the Sonora region of Mexico,

stopping at Nogales, and thence to Torres. Various interesting forms

are described from this region, among the most remarkable of which

is a cucurbitaceous tendril bearing plant, /bervillea sonorae, whose

root and stem base are enormously swollen for water storage, and

a tree-like morning-glory (/pomoca arborescens), which grows twenty

to thirty feet high. At Guaymas on the Gulf of California, a curious

mixture of plant forms was observed, the beach is lined with man-

groves, while close to them were the strictly xerophytic Cacti, for as

far as rainfall is concerned Guaymas is even more arid than Torres.

In the Colorado desert of California several types of vegetation are

found, due to differences in the soils; there are the gravel hills, the

alkali, and salt flats, the two last named showing a very restricted

growth of vegetation. The fan-leaved palm, Neowashingtonia filifera

is native to the eastern foot hills of the San Bernardino mountains

which lie in the Colorado desert district. They grow in groves form-

ing miniature oases where a clayey soil, from which oozes what water

has come from the hills, crops out to the surface. Northward lies

the Mohave desert where grows Yucca arborescens and Juniperus

californica, while in the lower altitudes the creosote bush (Covillea

tridentata), is the prevailing woody plant. From the Mohave the

authors proceeded to the grand canyon of the Colorado, but were

disappointed to find that the number of woody desert plants found

along the canyon’s sides were comparatively small. This they ascribe

to the narrowness of the canyon which probably induces abnormal

climatic conditions.

Following the account of the actual journey is a consideration of

the characteristics of deserts in general and of North American

deserts in particular. Meteorological tables are given, showing the

rain-fall for various localities from Oregon, to San Luis Potosi in

Mexico. One table of especial interest gives the mean annual pre-

cipitation as compared with the estimated annual evaporation. The

ratio in favor of the evaporation is anywhere from 6 :1 to 35.2 <1. It

is also pointed out that the distribution of rain-fall during the year is

of great importance in determining the character of the flora.

Another factor in the production of desert conditions which cannot

be disregarded is the soil constituents, a fact that is illustrated by the

conditions which exist in the gypsum containing White Sands of

Tularosa, the “white alkali” which is mainly sodium sulphate and

the *black alkali" where sodium chloride and sodium bicarbonate

are the chief soluble constituents.

330 THE AMERICAN NATURALIST. (VoL. XXXVIII.

Historically the desert regions of North America are of interest,

. that area which was known as “the great American Desert " by car-

tographers as late as 1843, does not really exist as such. The des-

erts of this continent may be designated as the Sierra-Nevada desert,

comprising portions of Utah, Idaho, Washington, Oregon, Nevada,

California, Arizona, Baja California, Sonora, and Sinaloa ; and the

Chihuahua desert, which occupies the tableland of Mexico east of

the Sierra Madre and north into Texas, Arizona, and New Mexico.

It is further stated that for the purposes of this paper the desert

lands of the Dakotas, of Montana and Wyoming may be considered

as the extreme northern arm of the last named region.

As the closing section of the paper there is a discussion of the

results of experiments by one of the authors, carried on at an earlier

date. A comparison of two desert types, Mentzelia pumila, and an

Artemisia, with two moisture loving forms, tomato and ZÆucalyptus

globosus, shows that even in their natural habitats, where the con-

ditions for transpiration are of course vastly in favor of the desert

plants, the disparity between the water evaporation of the two is

very great. As to temperature it was found that the plant-body of an

Opuntia gave a maximum in the forenoon of as much as 111.2° Fahr.,

while in specimens of Cereus temperatures of 113° to 115° Fahr., were

frequently found. ‘The fact that this is above the critical temperature

usually given for chlorophyll leads to the authors’ suggestion that the

protoplasm and the chloroplasts may have undergone changes which

adapt them for such conditions, although, they add, it is not unlikely

that the death of plants in such regions is as often the result of too

great insolation as of lack of water.

The authors make no claim that their narrative is an exhaustive

account of the regions visited, which considering the short time spent

would of course be quite impossible, but express a hope that it will

serve to show the great diversity which exists in the several floras

which go to make up our desert flora as a whole. In this they can

certainly feel that they have succeeded both in the text and in the

admirable illustrations which are profusely scattered through the

pamphlet. To any intending student of the conditions which exist

in our deserts it is bound to be of great service and the full bibli-

ography by W. A. Cannon which is appended will be an additional

aid. This is the first publication relative to the desert laboratory, We

may hope that many more will emanate from this source.

M. R.

(No. 447 was mailed May 3, 1904.)

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GINN & COMPANY, PUBLISHERS. ee ce

THE

AMERICAN NATURALIST.

Vor. XXXVIII. May, 1904. No. 449.

THE ANATOMY OF THE NORTH AMERICAN

CONIFERALES TOGETHER WITH CERTAIN

EXOTIC SPECIES FROM JAPAN AND

AUSTRALASIA.

(Continued from page 273).

D. P. PENHALLOW.

THe MepurLARY Rays.

THE medullary ray, in the various details of its structure as

presented radially and tangentially, comprises some of the

most important features for diagnostic and taxonomic purposes.

While it presents numerous variations, these are, in the main, of

such a nature as to give them very positive value for both gen-

eric and specific differentiations. Primarily the medullary rays

are to be regarded as a residue of the original fundamental

structure which has been left over in the genesis of the primary

stele, but they are capable of reproduction or extension under

the influence of the cambium in the course of secondary growth.

In all such cases, however, they are typically composed of the

same elements which are necessarily parenchymatous. Devia-

tions from this structure may arise through the introduction of

other elements, but such alterations always arise in a manner

331

332 THE AMERICAN NATURALIST. [Vor. XXXVIII.

which indicates their relation to the evolution of higher types of

organization. The ray presents few features of value in the

transverse section, and these will be sufficiently dealt with in

the systematic portion! to make further reference to them at this

time unnecessary. It only remains to point out that those rays

in the Abietineze which contain resin canals, and which present

an unusual width, possess no special diagnostic value in this

plane of section.

Radial Section.— Viewed radially, the medullary ray is seen

n el

Bo 0 Sll So

Co ce

oF OC So In

TIG =

bo c — mm

Ü e. OO

Qoo

D Ou» O O C O DOS

a Q olto joo 0o

EPI. > 2 g9 o|V^7»

a | SDAA

Fic. 17.— Pinus falustris. Radial section of a medullary ray showing characteristic pits on the

ateral walls. a, a thin wall broken out; 4, thick-walled parenchyma; and c, thin-walled paren-

chyma cells. x 280.

to be composed of a series of cells extended in a radial direction

and superimposed so as to form a muriform band from one to

many cells in height. In general terms, the higher the ray the

lower the component elements, from which it follows that in one-

celled rays the cells are usually highest; but this feature is only

of general interest since it rarely has a bearing upon the chief

questions at issue. In some cases two structural types may be

‘The systematic portion here referred to, constituting Part II of the present

series, will appear in the 7ransuctions of the Royal Society of. Canada for 1904.

No. 449] | WORTH AMERICAN CONIFERALES. 333

recognized — the one containing resin passages, the other devoid

of such structures. Where such passages occur the structure

of the ray shows a variation of detail which makes it of no

value for diagnostic purposes, and it is therefore eliminated from

the following discussion.

A feature of primary importance in the constitution of the

ray is the occurrence of two kinds of parenchyma cells. In

95% of the genera the upper and lower walls are always thick-

ened by secondary growth and more or less strongly perforated

by simple pits (Figs. 175, 21 and 23). This feature also applies

to 56.1 % of the genus Pinus. It possesses no special value for

either specific or generic differentiations except so far as it applies

to cells which are markedly different and justify the special

terms “thick-walled” and “thin-walled.” It is obvious, then,

that the thick-walled cell is to be regarded as the normal struc-

ture for the ray of the Coniferales as a whole, while the thin-

walled represents the exceptional form which is introduced in

response to some special demands. Although the thick-walled

cells occur in the genus Pinus to the extent of 56 %, they show

a diminishing frequency, eventually becoming rare and ultimately

replaced by thin-walled cells. Reference to them in the follow-

ing diagnoses is always specified by (1). In 43.9% of the genus

the upper and lower walls are thin and absolutely devoid of pits.

For diagnostic purposes such cells are always referred to as (2).

In some cases they are so undeveloped as to be obscure and

readily broken out in the process of section cutting, so that they

are often entirely wanting (Fig. 17a). Such thin-walled cells

are typically developed in P. palustris, P. teda, etc, and

it is to be observed that they are always associated with the

highest forms of development. Transition forms occur. These

are first seen in the soft pines where occasional thin-walled cells,

devoid of pits, are interspersed and often coterminous with the

thick-walled elements. In the hard pines the same relation

exists, but it is gradually reversed until the thin-walled cells

altogether predominate. Such gradations are exhibited in P.

coulteri, P. jeffreyi, P. pungens, P. teda, P. cubensis and P.

inops, and they afford evidence of value as to the sequence in

development of the species. In P. murrayana, P. cubensis and

334 THE AMERICAN NATURALIST. [Vor. XXXVIII.

P.insiguis the transition forms exhibit much more detailed

gradations by virtue of which it is often exceedingly difficult to

distinguish between the two forms of cell, since whether coter-

minous or parallel the variations in thickness change in such a

way that the one type passes gradually into the other. When

these variations are viewed collectively and taken together with

the general fact that the thin-walled cells are a feature of the

higher types of organization, we may reasonably conclude that

the'thin-walled cells have been derived from the thick-walled

through a process of arrested development. The cause of such

alterations is to be sought for, and it will doubtless be found in

connection with another component of the ray.

The terminal walls of the ray cells present three typical vari-

d e e o Qi .—

= LL 2 De

ed e Z _& ae

ws e S Sla

2] 2 e S siko

a e S Sibe

ci e£ 2 2 ———

| @ || S Ld

Q ooo 499 Lond

Fic. es Laxodium distichum. Medullary ray showing the structure and position of the

pits on the lateral walls; the straight ray cells and the thin, straight, terminal walls.

X 280,

ations: (1) thin-walled and entire; (2) thin-walled and locally

thickened, and (3) thick-walled and coarsely pitted (see table,

D of anatomical data). The first feature is a characteristic

of 52.6 % of all the genera, inclusive of Ginkgo, from Agathis

to Sequoia, while it also appears in Cupressus and Abies in part

as exceptional, and in the genus Pinus to the extent of 85.3%-

The wall presents no secondary growth in thickness, either

locally or generally. In the majority of cases it crosses the line

of the principal cell axis either at right angles or diagonally,

No.449] NORTH AMERICAN CONIFERALES. 335

features which are usually of very secondary value, although in

a few cases, as Taxodium, it may serve a useful purpose as

an associated character, for differentiation from closely allied

genera (Fig. 18). In other cases the wall is more or less

strongly curved. This feature is prominent in Thuya, Cupres-

sus, Podocarpus, Thujopsis and Cryptomeria, as also in the more

highly developed hard pines. Toa less extent it also occurs in

Taxodium, and it constitutes a character of some value for differ-

ential purposes (Fig. 19). :

The second variant differs from the first in that the otherwise

thin wall is locally thickened (Fig. 20), the secondary growth

forming one or more beaded enlargements. This is a feature

which occurs exceptionally in Abies, Pseudotsuga, Picea and

Pinus, but it is typical in Cupressus (66.0 %) and Juniperus

> S 3 /

S ~ S > Lj -

v 0

=. 2 a

ao S S * 0

LI————

e x < 3

X7————

S q & o 9 EN

t -

pi > 8 a0

^ S $ 8 g

T to Co a

Fic. 19.— T'huya gigantea. Medul he f d di

1 } 1 tion of the

Ala y tay DIL 5 F : k " e

pits on the lateral walls; the thin and curved terminal walls; the cells co

tracted at th ds. X 280.

(72.7 %) where it constitutes a diagnostic element of great value.

It is in all cases, however, to be regarded as a transitional form

between the first and the third variants, and from this point of

view it also possesses a somewhat definite phylogenetic value.

The third variant is characterized by a marked general, second-

ary growth of the wall, which thereby becomes more or less

strongly thickened and traversed by numerous simple pits

(Fig. 21). It occurs exceptionally in Juniperus and Pinus, but

it is typical in Abies (90.9 %), Tsuga (100.0 %), Larix (100.0 %)

and Picea (90.0%). In Abies and Juniperus, where transitional

forms sometimes occur, the walls in the spring wood may be

336 THE AMERICAN NATURALIST. [Vor. XXXVIII.

only locally thickened, but in such cases the typical feature

always appears in the summer wood where such secondary alter-

ations are most strongly emphasized.

For taxonomic purposes, such features possess a definite

ee aE m o ET e ee Lh

Boe

ee G S S i A O A; HH

TSERTE Ws

3 N :

LH] - US NI E eee tC—EILTL—ITI——HIEIÍ-—-E

ore

di Resta gre s ci ce MESA

(Edo camer sui ener Um P Emm EUREN HD n

m FL ——HH4———LHH————HH———ÁÉEREERLLBEE——ÁH

SS | PR |

FEST a HII

IG. 20. a. Medullary ray showing the form and position of the pits;

he thin ick 1 inal walls. X 280.

value. The thick-walled cells of Tsuga, Larix and Picea permit

“of an easy and definite segregation of these three genera in those

cases which otherwise might involve a strong element of doubt,

and the same rule holds true, though to a less extent, with

HELCELS ISI 1 14-8

E NR RII: SUR SUN

FIG. 21.— Juniperus occidentalis. Medullary ray showing the form and disposition

of the pits on the lateral walls; the thick and coarsely pitted terminal walls.

X 280.

respect to the locally thickened walls in Cupressus and related

genera.

Pits on the lateral walls of the ray cells are an invariable

feature of all investigated species of Ginkgoales and Coniferales,

including fossil representatives and the Cordaitales. They vary

very much in form, size and number. In such types as Juni-

perus, they are most diminutive (Fig. 21) and generally numer-

ous, while in many of the pines, such as P. resinosa, P. kora-

tensis or P. reffexa (Fig. 22) they attain to maximum size and

No. 449] | NORTH AMERICAN CONIFERALES. 337

occupy nearly the entire surface of the wall within the limits of a

wood tracheid, thereby becoming few in number. In Sequoia

(Fig. 23) or Taxodium (Fig. 18) they are typically oval; in

Pinus cubensis or P. teda (Fig. 24), they are variously lenticu-

lar, while in P. resinosa or P. koraiensis they are oval or oblong,

or even quadrangular. Such variations as a whole, are far more

= = eee BS a

O9 oT.

WAP 18 1L lele o [bp —

o WAF N es*|P o |b —

Fic. 22.— Pinus reflexa. Medullary ray showing (a) the form and disposition of the

pits on the lateral walls; (4) the ray tracheids. X 280.

numerous and sharply defined in Pinus than in any other genus

known. In all the investigated genera, the pit is bordered.

This finds either partial or complete exceptions in the genus

Pinus to the extent of 78.1 % of the species, in which the pits

are either simple throughout, or they exhibit a more or less

definite border in the summer wood only. That a border is a

characteristic feature of fossil representatives, is justified by

comparison with existing species, but it is not always recog-

nizable in consequence of the alterations of structure due to the

general process of petrifaction. Such obliteration not infre-

quently involves the pit orifice also. It is thus apparent that

such structures often fail in the determination of fossils. In

existing species the border is often so faintly defined as to be

difficult of recognition, and this is especially the case in rays of

a resinous character. In all such cases, however, the require-

ments of a correct diagnosis are fully met by the pit orifice.

The general law of development then, is such that all genera

except Pinus may be held to be characterized by bordered pits.

338 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Their strong tendency to obliteration in that genus is found to

coincide with the more marked development of ray tracheids.

which undoubtedly assume more completely the original func-

tions of the parenchyma cells, these latter in consequence,

suffering constant structural reduction, as in the hard pines.

In the distribution of the pits an important feature appears in

= an pp

| | je [ ee

[eal —

—

Fic. 23.— Sequoia gigantea. Medullary ray showing the form and disposition of the

pits on the lateral walls. X 280.

the numerical variation in different parts of the ray. For diag-

nostic purposes it is necessary to have reference to the number of

pits, not upon the entire surface of an individual cell, butzwithin

the limits of a spring or summer tracheid as the case may be.

They are invariably most numerous in the region of the earliest

spring tracheids, usually diminishing toward the summer wood

where the change may sometimes take place abruptly, andgin

which they are most commonly reduced to one with occasional

obliteration in the most highly modified tracheids last formed.

A similar law of distribution is applicable within the vertical

limits of the ray. When these structures are several cells in

height, the number of pits is typical, and, within certain narrow

limits, constant for all except the marginal cells. Thus if the

normal number is 1-2 for the central cells, it may sometimes

rise to 4, 6 or 8 in the marginal cells only, and such exceptions

must be noted in diagnosis. When the ray is only one cell in

height, the number of pits agrees with that for the marginal

cells. Such numerical variations possess but little value for

generic purposes, but as a specific character they may be held

to constitute the principal differential feature in the last analysis.

These relations are expressed typically in the genus Sequoia,

the two species of which may be definitely differentiated. S

gigantea is characterized by oval and commonly narrowly

No. 449.] NORTH AMERICAN CONIFERALES. 339

bordered pits, the broadly oblong orifice equal to the outer

limits of the pit and chiefly parallel with the cell axis, 1-2,

more rarely 3-4 per tracheid. In somewhat sharp and definite

contrast to this, S. sempervirens has large, oval, narrowly bor-

nre

0 2

Fic. 24.— Pinus serotina. Medullary ray showing (x) the ray tracheids with dentate

1 “4 z th n

walls, (2) ah T 4- £ eh hs a cells P

chyma cells. X 280.

dered pits, 2—6 per tracheid, the round or broadly oblong orifice

either parallel with or diagonal to the cell axis. In Libocedrus

the pits are small, narrowly bordered, oval, with a lenticular,

diagonal orifice, 1-4 per tracheid. Or again in Larzx americana,

the pits are “2-6 per tracheid becoming distinctly smaller

toward the summer wood where they are abruptly reduced to 2,

and finally 1 per tracheid." In Cupressus pisifera the pits are

“chiefly 2 in radial series, or in the marginal cells and low rays

upwards of 6 per tracheid.” In Zaxodium distichum the pits

are round, conspicuously bordered and large, with a very nar-

rowly lenticular and diagonal orifice which is often as long as

the outer limits of the pit. But in the analytical key it will be

observed that this genus is naturally brought into close relations

with Sequoia which is also distinguished by large, bordered pits.

The ultimate differentiation then rests upon the fact that in the

latter, the pits are ova/, the border often narrow, sometimes

obscure, while the olbong or lenticular, usually rather broad

orifice is generally parallel with the cell axis. Asa final illustra-

tion the four pits of Pinus monophylla, or the 1-5 throughout,

349 THE AMERICAN NATURALIST. [Vor. XXXVIII.

finally reduced to 1-2 in the summer wood of P. balfouriana,

point with much definiteness to these particular species, while

among the hard pines the occurrence of large, oval or squarish

pits, 1 or rarely 2 per tracheid, segregates a group of four

species. Detailed as these features are, they are not accidental,

but of such constancy as to admit of no hesitation in accepting

the conclusions to which they point.

The length of the ray cell is subject to considerable variation,

not only within the limits of an individual, but as between one

species and another. Our studies, however, do not permit the

formulation of a law applicable to specific differentiations, even

if such a law does exist, which present evidence leads us to

doubt; but details of length, in terms of spring tracheids, have

been incorporated in all the diagnoses, since they are often very

suggestive and thus may assist in the ultimate recognition of the

species.

The form of the cell is of more evident value, although too

much stress must not be laid upon it. The cell is either

straight, as in Juniperus, Libocedrus or Picea (Fig. 21), or it

becomes fusiform through contraction of the extremities as in

Cupressus, Sequoia, Taxodium, etc. (Fig. 19). Asa well defined

differential character its value is only one degree higher than

the length of the cell, and it has been introduced into the diag-

noses for the same reason as a controlling factor of secondary

importance.

In the higher Coniferz the medullary ray is distinguished by

the presence of an element which differs materially in its struc-

ture from the associated parenchyma cells. These elements

have been designated by De Bary (9, pp. 491—492), as “ray

tracheids.” Their structure is so peculiar, and they present

such important relations to classification and development, as to

necessitate a somewhat detailed account of them, to some extent

in recapitulation of well known observations by De Bary (9, P-

491) Hartig (19, p. 13) and Goppert (17).

As stated by De Bary, the ray tracheid resembles the paren-

chyma cells, from which they differ, however, in the presence of

bordered pits on a// their walls. Furthermore, such pits not

only differ materially in form and size from the bordered pits of

No.449] WORTH AMERICAN CONIFERALES. 341

adjacent parenchyma cells, but they are always much smaller

than the pits of those wood tracheids on which they border.

Such tracheids are invariable features of the ray in all the higher

Coniferze from Tsuga and Pseudotsuga to Pinus to the extent of

25% of the investigated genera. In Juniperus they occur very

rarely, being found, so far as I am aware, in only one species

(J. nana) out of a total of eleven, and they are so sparingly de-

veloped as to readily escape observation. In Thuya they are to

be met with in 7. japonica, likewise in a rudimentary state of

development. Out of nine species of Cupressus they occur

only in C. nootkatensis. Of the ten investigated species of

Abies, they are found only in A. dalsamea. In commenting

upon this fact many years since, De Bary (9, p. 490), also pointed

out that among European species A. excelsa is similarly excep-

tional, but no attempt has been made to interpret their signifi-

cance. In Thuya, Cupressus and Abies the tracheids are strictly

marginal in the composite rays, forming the entire structure

in rays only one or two elements high. This relation obtains

in all the higher Conifers: in the first instance; but in Larix,

Picea and Pinus, where there is a notable increase in numbers,

they also become interspersed with the parenchyma cells and

eventually predominate over them, a feature which is especially

characteristic of the hard pines. Efforts have been made to

show that in all such cases the two kinds of elements succeed

one another in a definite order from above downward — or the

reverse — but our studies have failed to show that this is capable

of practical application to the purposes of classification, or even

of phylogeny (9, p. 491). The great fact of importance for our

present purpose, however, and one which stands out with much

prominence, is that the ray tracheids are not a structural feature

of the more primitive Coniferales, but only of the higher types

such as Picea and Pinus. Furthermore, the primitive position

for these structures is in the one or two-celled rays, or corre-

spondingly in the margins of the composite rays.

In Thuya and Cupressus the tracheids appear to stand by

themselves, and they exhibit no special relations to the paren-

chyma elements which would permit of inferences as to their

possible origin. In the genus Pinus, on the other hand, where

342 THE AMERICAN NATURALIST. [Vor: XXXVIII.

the relations are somewhat complex, evidence does appear of

such a nature as to suggest their derivation. In Pinus tnops,

P. torreyana, P. pungens, P. clausa, P. teda, P. palustris and

P. cubensis, we frequently find thick-walled parenchyma cells

and characteristic ray tracheids coterminous with one another.

This does not mean a simple association, since in nearly all such

cases, as typically presented by P. palustris, they also show a

graduated structure of such a nature as to confirm the belief that

the one passes into the other by structural gradations. That

such is the case cannot be doubted, and if further confirmation

were needed it is afforded by the precisely parallel relations to

be met with in the formation of resin cells and resin canals. A

further fact of much significance from the standpoint of devel-

opment, is that such interchangeable relations are peculiar to the

highest types of the genus Pinus. But we may ask, what is

the function of these structures which make their appearance

only in the higher Coniferze ; what is the proper significance of

their appearance there, and do any other plants offer parallel

examples ?

In the so-called medullary rays of Lepidodendron selaginotdes

(49, p. 141) there are numerous reticulated or spiral elements

which are undoubtedly of the nature of tracheids, and they may

be held to represent the ancestral form of the ray tracheids in the

Coniferze, toward which they bear the same relation that exists

between the spiral protoxylem element and the characteristic

wood tracheid with bordered pits. From this it is apparent that

the ray tracheid of Pinus or Tsuga represents a primitive struc-

ture which reappears in response to conditions of growth and

structural alterations of such a nature as to demand the interpo-

sition of more simple, because more primitive, elements for the

proper performance of necessary functional activities. These

activities, in the case of Lepidodendron, are probably expressed

in the radial distribution of water (51, p. 141), and we are no

doubt correct in assuming similar activities to be carried on in

the higher Coniferz. In all those species which present the

primitive structure of the thin-walled ray cells, both fossil and

recent, there are no tracheids to be found. Asa tendency to

thickening of the wall arises, there is also developed a sporadic

No. 449.] NORTH AMERICAN CONIFERALES. 343

tendency to the development of ray tracheids as in Thuya and

Cupressus. It is also a noteworthy fact that simultaneously

with a general thickening of all the cell walls throughout the

ray, as in the genus Tsuga, ray tracheids become a constant

and prominent structural feature. This relation exists in Pseu-

dotsuga, Larix, Picea and Pinus, and it is a remarkable fact that

as the type of organization advances, and the structural modifi-

cations of the wall become more profound, the tracheids gain

steadily in numbers and importance until they finally replace the

parenchyma cells more or less completely. Such facts serve to

direct attention to the idea that by such progressive alterations

the ray cells gradually lose their normal functional powers with

respect to the radial distribution of water, and under such cir-

cumstances it is imperatively demanded that this deficiency

should be met through some other structures. Under these

circumstances two alternatives are possible. First, that the

thick-walled and useless cells should return to their primitive

condition in opposition to the general course of development,

and once more resume their appropriate functions. Such struc-

tural reductions do in reality occur in these very cases, as shown

in Pinus teda, etc., but it is to be observed that they are of

the nature of a growth which has been arrested at such an

early stage as to be devoid of many of the normal structural

features. Furthermore, it would be difficult, if not impossible,

to obtain evidence from other plants in support of a hypothesis

of this nature. It is true that in the case of girdled pines the

heart wood may resume an activity long since lost, and thus take

upon itself once more the function of the sap wood, as also to

some extent the function of the bark, but such renewed func-

tional power does not in any way involve structural modifications

of existing elements, and cases of this sort cannot be cited in

support of the hypothesis stated. Under these circumstances,

therefore, it is fair to conclude that such arrested development

expresses diversion of energy to the preponderant tracheids.

The second alternative permits us to consider that in the ordi-

nary course of development the ray cells gradually lose their

functional activity asa result of extreme structural modification,

and that this loss of power cannot be restored, even though the

344 THE AMERICAN NATURALIST. [Vor. XXXVIII.

wall may return toa primitive condition of structure through

various phases of atrophy. In accordance with this idea the

tracheid would be introduced as the most natural because the

original medium for such activities as are centered in the ray,

and it would therefore acquire additional importance both nu-

merically and functionally in direct proportion to the loss of

power experienced by the parenchyma cells. This appears to

be a reasonable interpretation, and in the light of observed facts

it would seem to be the correct one.

A structural feature of great importance in the ray tracheid

appears in certain inequalities of the upper and lower walls

which take the form of teeth-like projections into the cavity

(Fig. 24). In what may be regarded as the most highly devel-

oped tracheids the teeth project across the cell cavity until they

meet and coalesce, thereby forming a more or less definite retic-

ulation which gives to the tracheid a very characteristic appear-

ance. As seen in tangential section, such reticulations often

appear as narrow bands crossing the cavity from side to side,

thus giving the cell a varying aspect. Such dentate and reticu-

lated tracheids are absolutely confined to the second section of

“the genus Pinus, in which they constitute one of the most char-

acteristic features to the extent of 68.3 % of the species. A

more detailed analysis of this feature, as applied to the hard

pines, is desirable. In P. resinosa and P. thunbergii, the tra-

cheids are simply dentate. In six species represented by E

murrayana the teeth extend into definite reticulations confined

to the summer wood ; but in six other species represented by

P. jeffreyi, such reticulations are sparingly developed through-

out the ray. In P. zzda a transitional form appears. Typically

this species shows the tracheids to be sparingly reticulated, but

occasionally they are strongly reticulated throughout. This

brings to mind the further fact that in all species which are

sparingly reticulated there is a marked tendency to strong retic-

ulation in the summer wood. In the thirteen remaining species

the tracheids are uniformly strongly reticulated throughout the

extent of the ray, and this feature attains its highest expression

in P. palustris and P. cubensis. It is therefore manifest that

we have to deal here with a graduated development of such a

No.449] | NORTH AMERICAN CONIFERALES. 345

nature that the simply dentate tracheid is the most rudimentary,

while the strongly reticulated is of the most advanced type of

structure.

The value of the ray tracheid for taxonomic purposes depends

upon: (1) its occurrence in certain genera, and (2) its structural

peculiarities. The simple wall of the tracheid, in the great

majority of cases, affords no basis of specific differentiation, but

in the various forms of dentate and reticulated walls of the

second section of Pinus, it is of well defined value in this

respect. Pinus resinosa, P. thunbergii and P. koratensis are

all characterized by the occurrence of simple teeth which are

sometimes sparingly developed. This feature is intimately asso-

‘ciated with the occurrence of large, simple and single pits on the

lateral walls of the ray cells. From this group P. densiflora

may be differentiated by the reticulations in the tracheids of the

summer wood. Among the hard pines, P. /zda is distinguished

by ray tracheids which are typically sparingly reticulated through-

out, but on the other hand, P. palustris and P. cubensis, which

probably represent the highest types of the genus, are at once

separated from all other species by reason of the extent to which

reticulations are developed.

The relations which the tracheids bear to the parenchyma

cells in the general composition of the ray also have an impor-

tant bearing upon specific differentiations. In the genus Tsuga

the tracheids are sometimes interspersed, affording the first

instance of a relation which later becomes most prominent in the

higher genera, and the same relation is also expressed in Pseu-

dotsuga and Larix. In Picea there is a somewhat stronger

tendency to an interspersal which is only expressed fully in

Pinus. In the soft pines eleven out of thirteen species show,

as a rule, as in the previous genera, that the tracheids are rarely

interspersed, P. aris tata forming a partial exception, as shown

in a sparing interspersal. P. monophylla and P. monticola, on

the other hand, show a strong interspersal of the tracheids,

and in this respect they approach the hard pines. In the

latter. group we again find the first four species character-

ized by a rare interspersal. But passing on to the more highly

developed species, such types as P. clausa, P. palustris and P.

346 THE AMERICAN NATURALIST. [Vor. XXXVIII.

glabra show that the interspersed tracheids are not only numer-

ous, but that they eventually become conspicuously predominant

and often constitute the bulk of the ray structure. It is evident

then that such features possess an evident value for diagnostic

purposes, particularly in the genus Pinus where the variations

are numerous, well defined, and applicable to particular species

or groups of species.

Tangential Section.— As displayed in tangential section, the

medullary ray exhibits two principal forms, each of

which presents features of great taxonomic and

phylogenetic value. The type of structure which

prevails, and which may be regarded as the funda-

mental form of the ray, is that of from one to

many cells superimposed in a single series of vary-

ing height (Fig. 25). Such uniseriate rays are

characteristic features of all the investigated recent

genera. In 30 % of the genera, there is a sporadic

tendency to a multiseriate form as expressed in the

development of rays which are 2-seriate in part.

Such enlargement is not confined to any particular

portion of the structure, and within the limits of

the same section it may arise at the

centre or at either end. It is never

found in Abies, Picea or Pinus, but it

is met with in Pseudotsuga macro-

carpa, three species of Cupressus, two

of Juniperus, one each of Sequoia and

Araucaria and two of Larix (Fig. 26).

^ In Libocedrus such tendency is much

= peas more pronounced, and the rays may |

a typically seri. D€ described as 2—3-seriate in part.

tom can * This feature is of so sporadic a

nature that existing species afford no

satisfactory evidence as to its origin or significance,

but reference to Cordaites tends to throw some

light upon this somewhat obscure problem. In

fourteen species of Cordaites, three of which are

European (25, p. 606—609) it is seen that the rays

No. 449.] NORTH AMERICAN CONIFERALES. 347

present four variants ranging from the strictly uniseriate form to

1-2-, rarely 3-seriate. The distribution is in the following per-

centage proportions : |

I—2-, rarely 3-seriate : : ; i ; * 21.4 %

I-2-seriate . ; : ; : i i 14.3 %

2-seriate in part. ; ; j 50.0 ffo

I-seriate 14.3 fb

From TNI it wit appear that Cordaites as a whole,

approaches the primitive, multiseriate ray such as may be found

in the Cycads, much more nearly than any of the existing

species under consideration, and from this point of view it

becomes possible to arrange a sequence showing the relative

development in the following terms: (1) Cordaites, (2) Libo-

cedrus, (3) all other genera as

enumerated above. The evi-

dence of fossil plants, how-

ever, shows that caution must

be exercised in our estimate

of what constitutes the primi-

tive ray. The structure of

Stigmaria shows a preponder-

ance of uniseriate medullary

Tays- (51, p. 224) and that

such are primitive rays cannot

well be doubted. In general,

however, we are probably not

far from correct in the assump-

tion that the highest form of

the ray is expressed in its uni-

seriate character. Deviations

from this would then require

to be interpreted as vestigial

features which indicate a rela- a. a.

tively lower type of organiza- go So al as raten piat

tion in direct proportion to the - a owing the aon of he cl

Increase of a tendency toward opposite tracheids.

a multiseriate form.

In the majority of species, the side walls of the parenchyma

THE AMERICAN NATURALIST. [Vor. XXXVIII.

cells are thick and traversed by small pits. In the

genus Pinus the wall is commonly thin and it closes

the orifice of a very large pit on the wall of the

adjacent wood tracheid. This is notably true of

the soft pines in which the side wall either projects

as a convex membrane, or it is concave and curves

into the cell cavity. Such a feature is of very little

if any importance with the exception P. reflexa, in

which the thin side walls almost invariably project

so as to give the cells a correspondingly inflated

appearance (Fig. 27a). It is not only apparent in

a tangential section, but it is very conspicuous in

the transverse section (Fig. 274) where the inflated

walls are seen to project into the cavities of adja-

cent wood tracheids, thereby giving to the ray a

beaded appearance. As an exceptional variation it

possesses no apparent significance with respect to

questions of descent.

The second form of the ray is that which has

been designated as fusiform in reference to its

characteristic outline (39). Such rays occur in

relatively few of the existing genera to the extent

of 20 %. They occur typically in Pseudotsuga,

Larix, Picea and Pinus, and they are thus seen to

be characteristic of the most advanced types.

Among extinct species they are unknown except

in the case of Sequoia burgessii! (41, 42—46) 1n

which they present a remarkable exception to the

general course of development and structure of

that genus. The fusiform rays are peculiar "

their structural features. They vary greatly in

74 height as between different genera, and such vari-

ations also occur within a given genus, the ex-

tremes being met with in the genus Pinus, where

vae antithetic

wall oii P. palustris and P. ponderosa present the

um, but d

of thyloses.

280.

relations. In most cases they are much higher

" than the uniseriate rays with which they are asso-

! Dr. E. C. Jeffrey has recently discovered the same feature in another extinct

Sequoia now in course of publication.

No. 449.] NORTH AMERICAN CONIFERALES. 349

ciated, but this rule is subject to several exceptions. They are

always distinguished by a broad-

ening of the central tract by

two to several times the original

dimensions, thereby becoming

more or less multiseriate. These

variations depend upon the na-

ture of the included structure

which exhibits modifications di-

rectly related to progressive de-

velopment of the genus. Such

broadening arises abruptly in

Pseudotsuga, Larix and Picea,

so that the terminals above and

below consist of a single series of

cells with the general structure

of the uniseriate ray (Fig. 28).

In Pinus the broadening is less

abrupt, diminishing in both direc-

tions somewhat gradually, thus

giving rise to a region of lenti-

cular form, which occupies up-

wards of half the height of the

Tay, or in some cases constitutes

the entire structure. From this

it follows that in such types as

P. palustris (Fig. 296), the ter-

minals which are often prolonged

to great length, may be linear

and uniseriate, while in P. c/ausa

the whole ray is lenticular in

outline and the terminals consist -

9f only one or two limiting tra-

cheids (Fig. 30). Within the

region of the central tract the

cells are all thick-walled in Pseu-

dotsuga, Larix and Picea, but in

Pinus the cells are generally thin-

Sees)

Fic. 29a.—Pinus albicaulis. Tangential sec-

tion of a fusiform ray showing a typical resin

canal with (a) thyloses, and (7) rather thick

tion of a fusiform ray in part, showing thin-

walled parenchyma broken out. X 280.

35°

THE AMERICAN NATURALIST. (VoL. XXXVIII.

walled, and in the hard pines this feature is emphasized by a

degeneration of the tissue to such an extent that it is readily

FiG. 30.— Pinus clausa.

. Fusiform ray sh

two tracheids. x 280.

broken out in making sections, whence it

characteristically appears either much broken

up or entirely wanting. The principal feature

of such rays, and the one which determines

their form, is the presence of a resin canal

in each case. Such resin canals traverse the

ray continuously for its entire length. They

present the same details of structure as the

resin canals which lie within the xylem. In

Pseudotsuga, Larix and Picea, the central

canal is narrow, especial in the first two

genera, and the epithelium consists of a single

layer of thick-walled cells. In Pseudotsuga

and Larix (Fig. 28), thyloses are altogether

wanting, but in Picea they are of sporadic

occurrence. In Pinus (Figs. 29 and 30) on

the contrary the canals are always distin-

guished by their great breadth; the epithe-

lium is composed of one to several rows of

thin-walled cells which are often resinous and

often much disorganized, while thyloses are

an invariable feature of the central canal.

A comparison of different genera and spe-

cies shows that there is a somewhat striking

variation in the number of uniseriate rays

(tangential) to a given area of section. Such

variations may arise within narrow limits in

the same species according to location and

conditions of growth, but apart from this

there are somewhat constant variations be-

tween different species which may be ex

pressed by the use of the relative terms few,

many. No attempt has been made to define

such variations and more exactly, but it is quite possible that a

determination of the average number to a square centimeter, OF

other convenient unit, might disclose a somewhat greater differ-

No. 449.] NORTH AMERICAN CONIFERALES. 35I

ential value than is at present apparent. A simple illustration

will serve to afford an idea of the rather limited specific value of

this character. In Tarus cuspidata the rays are numerous, while

in the two remaining species they are relatively few. The same

feature applies to the differentiation of Torreya nucifera from the

other species of that genus. In Pinus clausa, P.

serotina, P. murrayana, etc., the same rule applies,

but in all such cases it cannot be accepted as final.

The height of the ray is subject to such varia-

. tions, even within the same species, that it cannot

be defined with sufficient accuracy to admit of its

application to classification in more than a very

general sense. It is true that the rays of Ginkgo

are always low, while those of Taxus and Torreya

are often high. In Juniperus they are commonly

low, while in Pinus they range from low to very

high. Such variations do not possess sufficient

constancy to admit of either generic or specific

application in the strict sense, though they not

infrequently serve a useful purpose as controlling

factors, and they are therefore incorporated in all

the diagnoses. Variations in breadth have a much

more definite value, since the element of constancy

is well defined. The genus Thuya (Fig. 31) may

almost invariably be differentiated by this feature.

In Cupressus, C. ¢hyoides may be distinguished by

a similar feature, while C. arizonica and C. govent-

ana are equally well indicated by the great breadth

of the rays. The same rule applies also to Juni-

perus, Sequoia (Fig. 25), Pinus, and other genera,

whence it appears that this feature is of specific

value. It is always associated with and dependent

upon the form of the component cells (tangential) which afford a

means of distinguishing genera and species with much directness.

The narrowly oblong cells of Thuya (Fig. 31) serve to separate

this genus without difficulty, since a similar feature occurs but

rarely elsewhere, and then in such association as to make the

differentiation clear. In Juniperus the genus is separable into

oblong cells.

xX 280. ,

352 THE AMERICAN NATURALIST, (VOL. XXXVIII.

four well marked divisions: (1) round to oval or. transversely

oval; (2) rays broad, the cells oval to round, chiefly round; (3)

chiefly oval, and (4) rays narrow, the cells oblong to oval, chiefly

oblong. The broadly oval and thin-walled cells of Sequoia sep-

arate it from associated genera. In Picea the genus may be

subdivided accordingly as the cells are (1) variable, round, oval

or oblong; (2) equal and uniform, oblong or oval! ^ Capressus is

similarly separable into groups. But more specifically it is not

difficult to separate C. arizonica and C. goveniana by reason of

their broad rays and very conspicuously transversely oval cells, .

from C. pisifera with its round or oval cells and C. ¢hyotdes with

its narrow, oblong, rarely oval cells. In the genus Pinus atten-

tion is at once directed to P. murrayana by the conspicuously

round or transversely oval, very unequal and variable cells.

The interspersal of the tracheids often imparts a character-

istic appearance to the tangential aspect of the ray, especially in

the genus Pinus, and more particularly among the hard pines. —

In this group the tracheids present very variable forms and

sizes. In such types as P. glabra they are small, oval or round,

and wherever they occur they give rise to a marked local con-

traction. In P. palustris and P. cubensis they are commonly

oblong and not infrequently they become several times higher

than broad. As they are almost invariably narrower than the

associated parenchyma cells, they cause a local contraction

which sometimes extends over considerable distances. In P

palustris the predominance of the tracheids is carried so far that

the rays are chiefly composed of them, and it then becomes ap-

propriate to apply the term interspersed to the few parenchyma

cells. In all of the more highly organized rays of the hard

pines the appearance of the structure is so complex and variable

that a proper diagnosis can be drawn only when we take

cognizance of the principal aspects presented, and these some-

times amount to as many as four in number.

A consideration of the various structural features thus dis-

cussed in their relations to classification will show that no other

1 The term equal here applies to cells of the same ray which are of the same

width; uniform to the cells of all rays which are pretty constantly of one form;

the contrasting terms being unequal and variable respectively.

No. 449.] NORTH AMERICAN CONIFERALES. 353

portion of the stem possesses so many elements of importance

as the medullary ray which, in consequence,'attains the highest

value in this respect and affords differential characters of wide

range, great prominence, easy recognition and of primary im-

portance in the differentiation of groups, genera and species;

and asa general summary, the utility of these characters for

such purposes is approximately indicated in the following tabu-

lation :

1 Rays (tangential) of two kinds.

Ray tracheids.

Pits on the lateral walls of the ray cells sim-

ple or bordered.

Terminal walls of the ray cells thin and} entire

or locally thickened.

Form and character of the ray cell (tang.)

Form and size of pits on the lateral walls of

ray cells. !

7 Ray tracheids dentate or reticulated. — . J

Ww N

* Generic.

8 Direction and form of orifice of pits on the lat- - Specific.

eral walls of ray cells.

9 Upper and lower walls of ray cells.

10 Ray tracheids interspersed or marginal.

II Disposition of pits (radial).

I2 The number of pits per tracheid. ;

THE RELATIONS OF MEDULLARY Rays TO DEVELOPMENT.

We are now in a position to determine the relations in which

the various structural features of the medullary ray stand to

development, and for this purpose it may be most convenient to

discuss them in that sequence which is apparently consonant

with the general order of evolution of the entire group.

It has been ascertained that bordered pits are characteristic

features of the lateral walls of the ray cell in 72.4 % of the inves-

tigated species, and that in the remaining 27.6 % among the

higher types, simple pits predominate, but a closer scrutiny of

this latter group discloses some features of more than passing

354 THE AMERICAN NATURALIST. [Vor. XXXVIII.

interest. Reference to the table of anatomical data will show

that the change from bordered to simple pits is entirely confined

tothe genus Pinus, and that it does not rise abruptly as if in

response to some unusual condition whereby a profound altera-

tion in the usual course of development was induced ; but it is

effected by stages, showing that whatever influences were

brought to bear, they operated gradually through a somewhat

prolonged period of development, while here and there strong

tendencies to reversion were manifested, and that the alteration

was finally effected in a permanent way, only in the most highly

developed pines. Commencing with P. /ambertiana it will be

observed that some species of the soft pines are characterized by

simple pits. Among the hard pines P. c/ausa and P. rigida

have bordered pits, while the six following species again show

simple pits. We next come to a group of four species, with one

exception (P. murrayana) Japanese, in which there is a mingling

of both bordered and simple pits, showing a decided persistency

of the primitive character in the face of conditions which involve

achange. Following these are two species with simple pits, one

with transitional features, five with simple pits, one with bor-

dered pits, one with the transitional form and the remaining six

species with simple pits, only. It will therefore be seen that

these changes occur in waves, and that within the limits of forty-

one species there are three complete and six incomplete recurrent

phases. If we were arguing from purely theoretical grounds, all

of these species should be arranged in such order as to show,

(1) bordered pits, (2) transitional forms and (3) wholly simple

pits, and we should thereby gain a perfect, developmental

sequence. But such a position would not be justified by other

evidence of an equally, if not more weighty character, and it is

our object to interpret the facts as they are found. It has

already been shown that the occurrence of simple pits in the

pines is consonant with a higher type of development, and that

the change is not only accompanied by sporadic reversions or

survivals as one may choose to regard them, but that the change

as a whole is a process of reduction. From this point of view

then, we must regard the occurrence of bordered pits in P.

clausa, P. rigida and P. pungens as pure survivals of a more

No. 449.] NORTH AMERICAN CONIFERALES. 355

primitive structure, a feature which is less perfectly expressed in

such transitional forms as P. koraiensis or P. inops. But a mere

mingling of the two kinds of pits in the same species is not the

only evidence in this direction. The mingling of simple and

bordered pits does not occur indiscriminately, but in accordance

with a well defined law to the effect that the former are char-

acteristic of the spring wood throughout its entire extent, while

the latter occur only in the summer wood where they might be

expected if at all, since the arrested development which might

be complete in the case of relatively thin-walled cells, could be

readily overcome in part, in walls of greater secondary growth.

This in no way conflicts with the observed fact that in the

majority of cases, the usual course of development is such that

the bordered pits of the spring wood very commonly become

reduced to simple pits in the summer wood in accordance with

De Bary's law as already stated in application to other cases.

Constancy in the structure of such pits has been found to be

characteristic of Cordaites, Ginkgo, the Taxaceze and all the

lower forms of the Coniferae, from which we may conclude that

the bordered pit is essentially a primitive character. On the

other hand variation is a well marked feature of the pit in the

genus Pinus as first expressed in the large, oval or squarish and

open pits of P. resinosa or P. thunbergii, and as later appears

with greater frequency in the smaller and very inconstant pits

of P. teda or P. palustris. Such variations then, involving

a gradual and complete transformation to the condition of

simple pits, are characteristic only of the more highly developed

pines, from which it may be concluded that it is a feature con-

sistent with a relatively high order of development in exact

accord with the principles governing parallel changes in the pits

of the wood tracheids. They are also in harmony with the well-

known principle that variation is always of a more simplified

form in primitive types, but that it tends to greater diversifica-

tion with advance in organization and general development, as a

necessary sequence to the adjustment of the organism to a wider

and more complex environment. Finally it has been shown that

the elimination of the bordered pit proceeds concurrently with

the more complete organization of the ray tracheids, in response

356 THE AMERICAN NATURALIST. [Vor. XXXVIII.

to a substitution of functional activities between these structures

and the degenerate parenchyma cells. We may therefore con-

clude that extreme variation in the character of the pit is an

expression of a higher type of development, and that from this

standpoint, such structures have a definite value in solving ques-

tions of descent.

The terminal walls of the ray cells present three variants with

respect to secondary growth. All the more primitive Cordai-

tales and Coniferales are characterized by thin walls. Cupressus

and Juniperus are chiefly distinguished by their thin walls which

are also locally thickened, a feature which has been shown to be

due to incipient secondary growth. But such alterations are

already foreshadowed in Libocedrus where the local thickening

of the wall is of a sporadic nature. In Adzes magnifica and

A. grandis there is a partial recurrence of thin and locally thick-

ened walls, which is pretty fully expressed in A. concolor. A

similar recurrence is met with in Pseudotsuga macrocarpa, in

Picea polita and in Pinus parrayana, and it is also complete in

thirteen of the most highly developed species of Pinus, where

the walls have suffered extreme degeneration. Within the

limits of Picea (1) and the soft pines (5), there are six instances

in all, of sporadic and partial survival of the thin and locally

thickened wall. The first tendency to thick and strongly pitted

walls is manifested in five species of Juniperus, and such

development is fully expressed in what may be regarded as the

three most highly developed species. Thick walls are then

fully characteristic of Abies— with a partial reversion in

A. concolor, of Tsuga, Pseudotsuga douglasii, Picea, with the

exception of P. polita, five species of soft pines and three species

of hard pines. In P. teda and P. palustris the walls are so

degenerate that their structure cannot be satisfactorily deter-

mined, but they are presumably thin-walled.

From these facts it is manifest that the progressive thickening

of the terminal walls accords with the general course of develop-

ment, and once more making use of the principles already

applied to the pits on the lateral walls, we are brought to the

natural conclusion that (1) an increase in the thickness of the

walls is evidence of a higher type of organization, and (2) that

No. 449.] NORTH AMERICAN CONIFERALES. 357

the sporadic recurrence of thin walls with local thickenings

represents the persistence of a primitive character.

Ray tracheids probably constitute one of the most valuable of

the structural elements as an indication of development. This

has its foundation (1) in the fact, previously shown, that they

arise as secondary structures from the parenchyma elements,

with which they exhibit interchangeable relations, in direct

response to the requirements of a higher degree of organization,

and (2) their general relation to progressive development. The

complete absence of ray tracheids from the Cordaitales and

Ginkgoales, as also from the Taxaceze and more primitive Con-

iferæ, while they are invariable features of the higher Coniferz

in which they attain their most complete development, admits of

only one interpretation. The fact that they are exclusive fea-

tures of the Coniferze emphasizes their inferior value for deter-

mining the derivation of that group, while it points to their

superior importance as a factor in the sequence of the various

coniferous genera. They occur sporadically in Thuya (1), Cu-

pressus (2), Juniperus (1), and Abies (1). They are prominent

features of Tsuga, Pseudotsuga, Larix, Picea and Pinus. Their

invariable absence from Sequoia would appear to suggest that

this genus is more primitive than Thuya, but there are other

reasons which serve to suggest the opposite relation. Apart

from this exception it will be seen that in accordance with the

relations exhibited in the table of anatomical data, the genera

enumerated form a continuous series, commencing with those

showing sporadic tracheids, and ending with those in which such

structures attain their highest expression. From this we are

justified in the conclusion that the rare occurrence of tracheids in

Thuya, etc., is to be interpreted as the first evidence of a tend-

ency in development which is only fully realized at a later period,

and this appears to be justified by a closer examination of the

last five genera in this respect, since it is found that in them

the tracheids not only show a progressive numerical develop-

ment, but their structure likewise becomes more complicated in

direct relation to the evolution of higher types of genera and

species. We must therefore look upon the tracheids, with their

thin, simple walls, as the primitive form, while those with the

358 THE AMERICAN NATURALIST. [Vor. XXXVIII.

strongest reticulations are of the highest type, the two being

united by a transitional form characterized by the presence of

simple teeth. The evidence at hand does not appear to justify

the idea that the various genera have been segregated into small

groups representing side lines of development, but it rather

favors the thought that each genus is in itself a complete, short

line of descent, and that among these a prominent parallelism

has arisen in the tendency toward the development of tracheids

—a tendency which has been carried to completion in the case

of only five of the series, and in such a way that in only a

portion of one of these has that completion reached its highest

expression.

The occurrence of two kinds of parenchyma ray cells is an

exclusive feature of the genus Pinus, and their value for phylo-

genetic purposes is strictly confined to the relations of the various

species of pines. The first appearance of this differentiation is

among the soft pines in P. aristata and P. edulis. It is to be

observed, however, that the thick-walled cells are always domi-

nant, the thin-walled cells being interspersed among and coter-

minous with them. No further evidences of such structural

alterations are to be noted until we reach the more highly

developed representatives of the hard pines. Among these

definite transition forms occur in P. murrayana, P. coulteri, P.

Jeffreyi, P. virginiana, P. insignis and P. cubensis, while in P.

arizonica, P. ponderosa, P. sabiniana, P. pungens, etc., the orig-

inal relations are exactly reversed and the thick-walled cells show

a diminishing frequency, until in P. glabra and P. teda they

are rarely met with. Such facts give effective proof to the

belief that structural alterations of this nature are not only evi-

dences of the highest type of development among the pines,

but in the Coniferales as a whole.

The invariable absence of the fusiform ray from all except

the four genera which attain the highest structural development,

and their constant occurrence in all the species of such genera,

presents an argument of great force as showing their relation

to the evolution of advanced types. There is here no evidence

of sporadic development, foreshadowing the general course of

evolution, but the fusiform rays with their resin canals appear

No. 449] | NORTH AMERICAN CONIFERALES. 359

abruptly and permanently. Among fossil plants — except the

genus Pityoxylon which, being essentially Pinus, falls under the

general rule — there is no instance of such structures outside of

the four genera named, save in the case of the remarkable

Sequoia burgessii, from the Lignite Tertiary (4, p. 242) and S.

penhallowii of Jeffrey. As it will be necessary to further dis-

cuss the essential structure of the fusiform ray when considering

the resin passages in particular, it will be unnecessary to deal

with it more at length at the present moment.

Errata.

Page 249. For Cordaited read Cordaites.

Page 263, Fig. 12. For Radial read Tangential.

(To be continued.)

FURTHER INSTANCES OF MALAR DIVISION.

ALES HRDLICKA.

By the courtesy of Dr. Horace Jayne, Director of the Wistar

Institute, Philadelphia, I am able to report on two skulls, one

Peruvian (Indian) and the other that of a Chinese, in both of

which is found what appears to be a bilateral, complete, er

division. The specimens and anomalies are as follows :

(1) Spec. No. 53, Wistar Institute, cranium of a male, adult

(past middle age), Peruvian Indian. The occiput shows a mod-

erate, predominantly unilateral, baby-board compression. | A

smaller Wormian bone is present in the left coronal suture and

an unusual bone of the same nature, 10 mm. long and 9 mm. in

maximum breadth, in the sagittal suture, 28 mm. posterior to

bregma. The skull shows no pathological lesion, no signs of

traumatism, and no marked anomalies besides those to be

described.

Each of the indus is completely divided by an antero-poste-

rior suture into a smaller lower and larger upper portion. The

condition is very much alike on the two sides and offers a num-

ber of points of special interest. (Fig. 1.)

The body of each malar is very narrow (antero-posterior -

diameter, dorsally, at middle, 13.5 mm. on the right and 12.5

mm. on the left). This narrowness is caused by an unusual

development and extension backward of the malar process of

the superior maxillary. The central part of the malar bone was

apparently belated in development and a compensation for the

defect by the maxillary took place.

The upper, larger portion of each malar shows in its zygo-

matic border a well marked, 5 mm. long, slightly dentated suture

corresponding to the occasional posterior incomplete suture of

the normal malar. This suture is supposed ` to be the remnant

of the embryonal separation between the parts of the malar

that develop from respectively the superior and inferior center

361

362 THE AMERICAN NATURALIST. Vot. XXXVIII.

of ossification of the bone.

Fic. 1.— Right Malar of a Peruvian Indian, s E

ing an Antero-Posterior Complete Division and

above that a Posterior Incomplete suture

If this theory is correct, which thus

far there is no reason to

doubt, then we have here

a clear case of an appear-

ance of a separate, subma-

lar center of ossification.

The case would then be

not a divided malar, but an

imperfectly developed ma-

lar, with a szó-ma/ar.

Ventrally the anomalous

suture is in every respect

very much like dorsally.

From each of the incom-

plete sutures in the zygo-

matic border of the malar

proper runs a shallow

groove, such as was de-

scribed by Gruber, forward, reaching on the left the malo-maxil-

lary suture.

The upper portion of the right malar shows one, that of the

left two nutritive foramina.

This is the second instance of a malar suture observed in an

American Indian! and in both cases the subject was a Peruvian.

Measurements of the Malars.

Right.| Left.

Height pbi Dani the ends, iem i of the malo- 33-5 | 32 mm

illar ry sutu 345| ?

ux i mid

e posteriorly (dorsally : “beë ‘lowest point of ‘malo-

zygo and most eae Ns of malo-frontal i

SUDORE 44- 44-

' The first instance was reported by me in the Amer. Vat., April, 1902 (“New

Instances of Complete Division of the Malar Bone, with Notes on Incomplete

Division," pp. 273-294)

No. 449.] MALAR DIVISION. 363

Measurements of the Malars (continued ).

Right. Left.

Breadth antero-superiorly (dorsally: bet. the superior

extremity of the malo-maxillary and the anterior

end of the malo-frontal suture) HE UNT 33. | 34. mm

" at Middle .. 2:5. 5. 5. DI LI m Ws 124 1 144 7

" inferiorly (bet. lowest points of the malo-maxil-

lary and malo-zygomatic sutures). . . . + + 25 Au M

Vertical height of the inferior malar portion, to the malar

uture; anteriorly’. s . 4. 4 M Mas 13. ?

Osteriori oo. ss 24 OR eee S08 DE Pis

(2) Spec. No. 7217, Wistar Institute, cranium of a male,

adult (below middle age)

native of China. Skull

dolichocephalic, normal.

Upper third molars rudi-

mentary; styloids very

rudimentary. A small

epipteric on left, on right

a large epipteric (41.5 mm.

long, 10 mm. high), and

several episquamous ossi-

cles more posteriorly.

Both malars consist of

a larger superior and a

smaller. inferior portion,

separated by an antero- Fic.2.— Left Malar of a Chinese, showing a Com-

posterior moderately den- plete Antero-posterior Division.

tated suture, 21 mm. long on the right, 19 mm. on the left side.

(Fig. 2.)

Ventrally the aspect is entirely different. On the left a

tapering process from the maxilla reaches and articulates with

the zygoma; on the right a similar process exists, but ends

within 3 mm. of the zygoma. Both processes cover the malar

suture anteriorly and posteriorly, but in the middle leave it, and

! The lower malar portion on the left has been lost.

364 THE AMERICAN NATURALIST. [Vor. XXXVIII.

also an island-like part of the superior portion of the malar,

uncovered.

The separate inferior portions of the malars resemble some-

what those in case I

Measurements of the Malars.

Right. Left.

Hanhi antero ere re ee 32 | 36.5 mm. | 35.5 mm.

"* ^atmiddle .. pP IP M. rl Og T CI MES

"c postenoriy x ues Zee. oov XTIABESS |

Breadth axitero-saperory.. —. =- o usos oro 1395 "^

"Hes | .. $7535) c xot DEMO. IMEET

OMNEM er le a ee A sut po s 34.6 ri

Vertical height of the inferior malar portion, to the

malar suture, :

MUNIN s uM ore n bg Y II.

DEMENS. 0 e or Q e 8.

To the above two cases I am able to add the following recent

observations of malar suture in the U. S. National Museum:

(a) Skull of a male, middle aged Chickasaw Indian (No.

227, 483, Dep't. of Anthrop, U. S. N. M). This specimen,

recently returned from the Army Med. Mus., was brought to my

attention by Dr. D. S. Lamb. Skull shows a somewhat prema-

ture closure of sagittal and coronal sutures.

Each malar is divided, in very much the same manner, by an

antero-posterior suture into two portions. The lower of these,

nearly of an equal extent throughout, is at middle on the right

10 mm., on the left 9 mm. high; the height of the upper por-

tion at middle is on the right 22 mm., on the left 21 mm.

The anomalous suture is on the right as well as on the left

dorsally 15 mm., ventrally on the right 5 mm., on the left 6 mm.

long. It is shortened by a wedge-like process of the superior

maxilla, identical in appearance and only of a slightly lesser

extent, to that in the above described, Wistar Institute Peru-

vian. The upper portion of each malar shows one moderate

size foramen, but no incisure.

There are no further anomalies on the cranium.

No. 449.] MALAR DIVISION. 365

Among 380 skulls of apes and monkeys! in the Department

of Biology of the Museum,

in which the malar sutures

are clearly traceable, there

are three, a Macaque, a Cer-

copithecus, and a Chryso-

thrix, with malar division.

(b Macacus pelops,

male, adult, No. 22,062.

The right malar shows an

oblique, serrated division,

running dorsally from a

short distance above the

lower end of the malo-

maxilary suture to the

curving superior border of

the zygoma (Fig. 3). The ventral part of the suture lies in

nearly the same position. On the left is found dorsally a 4 mm.

long suture, beginning from the malo-maxillary suture in a simi-

lar position as that on the right side. The two malars are equally

high and long. The skull shows no

other anomalies.

adolescent, No. 16, 365. The right

zygomatic process is posteriorly com-

pleted by a triangular portion, separated

from the main part by a suture. Supe-

eu riorly the border of the malar portion of

Fic. 4.— Divided zygomatic pro- the arch just touches that of the tempo-

orale de ag opithicus callitae: +4) portion, inferiorly the two are sepa-

rated by a distance of 12 mm. (Fig. 4) The supernumerary

suture is serrated and equally well distinct ventrally. It is very

plainly anomalous in character; there is not the slightest trace

Fic. 3.— ee right malar of Macacus pelops. Nat-

ural si

11 Chimpanzee, 1 Gorilla, 2 Orangs, 8 Gibbons, 12 Cynocephali, 3 Sympha-

langi, 1 Nasalis larvatus, 6 Simias concolor, 55 Presbytes, 24 Semnopitheci, 116

Mascagues, 22 Cercopitheci, 10 Colobi, 5 Cercocebi, 15 Mycetes, 6 Alonata, -

Ateles, 1 ee 40 Cebi, 6 Hapale, 4 Midas, 4 Nictipitheci, 1 Saimiri, and 7

Chrysothelx

366 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of any traumatism. The dimensions of the two malars are

almost identical, even inferiorly. The specimen shows no other

anomaly.

(d) Chrysothrix (Venezuela), male, adolescent, No. 35,800.

On the left side is present an anomalous

separation of the zygomatic process from the

body of the malar. (Fig. 5.) No trace of any

traumatism. On the right the zygomatic

arch has been lost. The specimen shows no

re a further anomaly.

matic process of Chryso- No one of the three monkey skulls here

due (Vener, Natum! mentioned was in any way pathological.

In a number of Presbytes Semnopitheci,

one Hylobates, one Symphalangus, five Macaques, and one

Ateles, a more or less pronounced marginal cleft or fissure was

seen in the superior border (frontal process), three to five mm.

externally to the edge of the orbit, running to a foramen.

There were found in the series no clearly defined anterior or

posterior partial malar sutures.

The above six cases present a number of new points and will

be of value in the eventual summing up of the whole subject of

malar divisions.

Several interesting cases of the anomaly in man and mam-

mals, including an orang, have also recently been reported by

Frassetto (Notes de Craniologie Comparée, Ann. Sc's. Natur.,

Paris, Sept. 1903). |

size

U. S. NATIONAL MUSEUM,

STUDIES ON THE PLANT CELL.—I.

BRADLEY MOORE DAVIS.

INTRODUCTION.

Tus is the first of a series of papers that will follow one

another in the pages of the American Naturalist. They will

describe the chief structures in plant cells and the most

important events in their life histories, largely from the point of

view of the morphologist and student of developmental processes.

Research upon the plant cell has entirely outrun the general

accounts that may be found in several botanical text books and

in certain works of prominent zoólogists. We shall attempt to

give a general survey of the subject in its present state with

references to the most important papers ; but this is not to be an

exhaustive account of a literature that is already very large and

which can probably be treated far more satisfactorily several

years from now when it has passed through the criticism that

time will give in a field of very active botanical investigation.

American botanists have reason to be proud of the achieve-

ments of their countrymen in research upon the morphology and

physiology of the plant cell, for much of the best work of recent

years has come from them. This in itself has been a great stim-

ulus to the writer to prepare these brief accounts which he hopes

will assist the general botanist to a clearer understanding of the:

progress in this field. They will also serve to contrast the pro-

toplasmic activities among plants with those of the animal cell

Which has been so well treated in seve

English by Wilson's The Cell in Development and Inheritance.

The author will feel especially gratified if these papers should

help to change an attitude towards investigations on the plant

cell that is unfortunately too prevalent among botanists. There

is a tendency to regard cell studies as a very special field of

botanical research with elaborate technique which the average

367

368 THE AMERICAN NATURALIST. [Vor. XXXVIII.

botanist cannot be expected to master. Those who work in this

field are considered as in a department by themselves and are

labeled cytologists which is sometimes given as an excuse for

knowing little about their results. Cell studiesare nothing more .

than morphological and physiological investigations which are

frequently so broad as to break the mould of the narrower mor-

phology and physiology of former years. Cell studies must be

the foundation of all exhaustive work in morphology and physi-

ology. Indeed among the lower plants they constitute almost all

there is to morphology and will determine the classification and

relationships of great groups. There are no better illustrations

of this fact than the effect of Prof. Harper's investigations on

the ascus and sporangium upon Brefeld's theory of the origin of

the Ascomycetes. And again the results of several investigators

upon the multinucleate gametes found among the Phycomycetes

and Ascomycetes are of the utmost importance to a correct

understanding of the phylogeny of these groups. When students

. of the plant cell refuse to accept the stamp of cytologist and

insist and show that their work is simply fundamental mor.

phology and physiology we shall break away from a past that

should be outgrown.

The material of these papers will be treated under the follow-

ing heads.

TABLE OF CONTENTS.

Introduction.

SECTION I. STRUCTURE OF THE PLANT CELL.

I. Protoplasmic Contents.

(a) The Nucleus.

(b) The Plastids.

I. Plasma Membranes.

2. Trophoplasm.

Coenocentra, Nematoplasts, Physodes.

3. Kinoplasm.

Centrospheres, Centrosomes, Asters, Filarplasm, Ble-

pharoplasts,

No. 449.] STUDIES ON THE PLANT CELL. 369

2. Non-Protoplasmic Contents.

(a) Food material and waste products.

(b) Vacuoles.

3. The Cell Wall.

- Section II. THE ACTIVITIES OF THE PLANT CELL.

I. Vegetative Activities.

2. Cell Division.

(a) The Events of Nuclear Division.

. Direct Division.

. Indirect Division (Mitosis).

Prophase, Metaphase, Anaphase, Telophase.

The Dynamics of Nuclear Division.

(b) The Segmentation of Protoplasm.

. Cleavage by Constriction.

. Cleavage by Cell Plates.

. Free Cell Formation.

Ww N &

Section III. HIGHLY SPECIALIZED PLANT CELLS AND THEIR

PECULIARITIES.

1. The Zoöspore. 2. The Sperm. 3. The Egg. 4. The

Spore Mother Cell. 5. The Coenocyte. 6. The Coeno-

- gamete.

Section IV. CELL Unions AND NUCLEAR FusioNs IN PLANTS.

Section V. CELL ACTIVITIES AT CRITICAL PERIODS OF ONTOG-

ENY IN PLANTS.

1. Gametogenesis. 2. Sporogenesis. 3. Reduction of

Chromosomes. 4. Apogamy. 5. Apospory.

SECTION VI. COMPARATIVE MORPHOLOGY AND PHYSIOLOGY OF

THE PLANT CELL.

370 THE AMERICAN NATURALIST. [Vor. XXXVIII.

LITERATURE ON THE PLANT CELL.

Reference to special papers will be given by the authors name

and the date of publication through lists presented at the end of

every section.

There is no comprehensive treatise devoted to the plant cell

but the following general accounts and reviews of the literature

are important.

I. Strasburger in the Lehrbuch der Botanik and Pfeffer

in his Physiology of Plants present the best general accounts

of the structure and activities of the plant cell.

2. Zimmerman in 1893 and '94 (“ Beihefte zum Botanischen

Centralblatt " vol. 3 and 4), reviewed the literature on the plant

cell under the title * Sammel—Referate aus dem Gesammtgebiete

der Zellenlehre" and in 1896 collected the literature dealing

with the nuclei of plants in a book entitled Die Morphologie

und Physiologie des pflanzlichen Zellkernes, Jena, 1896.

3. Dangeard discusses a number of cytological topics in the

6th series. of Le Botaniste (1898) with especial reference to

his studies on the Chlamydomonadineae.

4. Fischer, Firirung Farbung und Bau des Protoplasmas

Leipzig 1899, presents a critique of the methods of cytological

research and the justification of the conclusions based thereon.

5. The most recent analysis of conspicuous activities of the

plant cell is that of Strasburger Ueber Reductionstheilung,

Spindelbildung, Centrosomen und Cilienbildner im Pflanzen-

reich, Histologische Beiträge VI, 1900.

SECTION I. STRUCTURE oF THE PLANT CELL.

It is customary to apply the term cell in Botany not alone to

the protoplasmic units of organization but also to the enclosing

wall that generally surrounds the protoplasm. Indeed these

walls alone when entirely emptied of protoplasm in specialized

2 * To this list should be added an excellent concise review by Koernicke entitled

Der heutige stand der pflanzlichen zellforschung ” Ber. d. deut. bot. Gesell 21,

(66), 1904. This article appeared too late to be quoted in the earlier papers of

this series.

No. 449.] STUDIES ON THE PLANT CELL. 37!

regions of the plant, e. g. vascular and certain supporting and

tegumentary tissues, are still called cells. When among the

lower forms and at certain periods in the life history of many

higher plants the protoplasm is naked (e. g. zoóspores, sperms,

eggs, etc.), these structures are cells in exactly the sense used by

zodlogists. We shall consider almost entirely the protoplasmic

portion of the plant structure for any extended treatment of the

walls would lead us at once into that field of microscopic anatomy

termed histology.

I.. Protoplasmic Contents.

The most highly differentiated region of the cell is the nucleus,

a structure remarkably uniform in organization among all plants.

except the lowest Algae and some very simple Fungi. These

more primitive conditions will be considered in Section VI.

Besides the nucleus there are present plastids in all groups

except the Fungi. Plastids are likewise specialized protoplasmic

elements although much simpler in structure than the nucleus.

Nuclei and plastids lie in a protoplasmic matrix called the cyto-

plasm. Cytoplasm is more variable in structure and activity

than any other region of the cell. Thus three forms of proto-

plasm, nucleoplasm, plastidplasm and cytoplasm comprise all the

living material of the cell and may be sharply contrasted with the

non-protoplasmic contents, mostly food material and waste prod-

ucts, which will be considered under a separate head. Definite

masses of nucleate protoplasm, with or without plastids are

termed protoplasts and such are either unicellular organisms.

themselves or units of a multicellular structure.

(a) The Nucleus.

The nucleus is bounded by a delicate membrane that is.

ismod largely or wholly a modification of the surrounding cyto-

plasm. The nucleoplasm very rarely completely fills the nuclear

membrane, the remaining space being occupied by a fluid known

as the nuclear sap. - The elements in the resting nucleus consist

chiefly of material that takes the form of a net work so that the

372 THE AMERICAN NATURALIST. (Vor. XXXVIII.

effect is that of a much coiled and twisted thread whose loops are

united at intervals to form large and small meshes. The ground

substance of this thread is called linin and imbedded in it as in

a matrix are deeply staining granules of chromatin. Chromatin

is regarded as the most important substance in the nucleus,

chiefly because of its behavior during nuclear division, and in

critical periods of the life history of organisms as at sporogen-

esis, gametogenesis and fertilization (to be described in Section

V). Just before nuclear division the chromatin becomes organ-

ized into bodies named chromosomes which are remarkably uni-

form in number and definite in shape for each tissue and period

of the plant's life. They will be discussed under “ The Events

of Nuclear Division " (Section II), and in Sections IV and V.

Fic. 1.— The resting nucleus. æ, Embryo sac of lily with linin thread and two nucleoli.

4, Root of onion large nucleolus. c, Tetraspore of Corallina showing large chro-

matin body and small nucleolus. 4, Spirogyra with central body containing chroma-

tin. e, Chromatin on linin net work from egg of pine. After Mitzkewitsch and

Chamberlain.

In the meshes of the linin network or lying freely in the

nuclear sap may be found one or more bodies, generally globular

in form, called nucleoli. (See Fig. 1 a and Fig. 1 4). The

nucleolus is generally regarded as a secretion of the nucleus and

it is quite certain that its substance is utilized just previous to

and during the period of nuclear division when the spindle is

formed. (Strasburger '95 and : oo, p. 125, and from the work of

others). The structure is not always homogeneous but may

Show in the interior small vesicles or areas of a different con-

sistency from the periphery. There is often present also a

rather thick outer shell or membrane. Sometimes the chromatin

in the nucleus may be gathered into a globular body that resem-

bles superficially a nucleolus, Such chromatin bodies are gen-

No. 449.] STUDIES ON THE PLANT CELL. 373

erally transitory as in Corallina, Davis '98, where the structure

(Fig. 1 c) is only found in the young daughter nucleus and later

fragments into many smaller bodies. In Spirogyra however

(Moll '94, Mitzkewitsch '98, Van Wisselingh :00, 'o2) the

chromatin is supposed to be always ina globular mass mixed with

nucleolar substance and recalls the conditions in certain Protozoa.

These chromatic structures however should never be confused

with nucleoli, whose substance is different and which are not

permanent in the cell, since they may disappear before or during

nuclear division and be formed de novo in each daughter nucleus.

The substance of the nucleolus is not well understood. It is

frequently impossible to distinguish it from chromatin except

when favorably situated in the cell and there is much evidence

that it is closely related to that substance. In large nuclei of

higher plants the chromatin is sometimes gathered into globular

bodies without apparent relation to a linin thread and these are

readily mistaken for nucleoli and have been called such, but this

loose usage of the term should be avoided. And true nucleoli

may be so closely associated with the linin net work as to have

the appearance of chromatin. Some of these conditions have

been especially described by Cavara, '98. Chamberlain, '99, has

made a study of the egg nucleus of the Pine where masses of

chromatin may take very irregular forms on the linin threads

(Fig. 1 e) and sometimes resemble small nucleoli. But such

conditions should always be sharply distinguished from true

nucleoli which are often caught in the meshes of the linin net

work and appear to be a part of it when in reality there are no

organic attachments. Itis certain that nucleoli are of secondary

importance in the cell and probably by-products of the general

constructive activities of the nucleus. In which case they may

be secretions, perhaps closely related to chromatin, or even direct

transformations of this substance. It is well known that the

nucleus has wonderful constructive powers, when the amount of

chromatin and other nuclear substances may be immensely

increased, facts that are especially well illustrated at reproductive

periods of the plant's life as during sporogenesis and gameto-

genesis. 4

Chromatin is the only substance in the nucleus that is constant

374 THE AMERICAN NATURALIST. [Vor. XXXVIII.

in its presence throughout all periods in every cell’s history.

It passes on from cell to cell through the mechanism of nuclear

division without interruption. There are periods of cell history

when the nucleus consists only of chromosomes as in the stages

of nuclear division called mataphase and anaphase. The other

structures of the nucleus have their relation to definite condi-

tions that are in part understood. The nuclear membrane

probably results from the reaction of the cytoplasm to the secre-

tion of nuclear sap among the chromosomes (Lawson, :03 a).

It would then be strictly cytoplasmic in character and similar to

the plasma membranes around vacuoles. Nucleoli must be

regarded as temporary structures since they generally disappear

during nuclear division either dissolving or else passing out into

the cytoplasm where they may remain for long periods as deeply

staining globules (extra nuclear nucleoli). Linin is believed to

be derived from chromatin and in its turn may be transformed

into the substance of spindle fibers, which are cytoplasmic, so

that chemically it holds a position somewhat intermediate between

chromatin and cytoplasm. It seems established that the linin

, net work is a temporary structure related to the activities of

chromatin.

(b) The Plastids.

These very interesting structures, characteristic of plant cells,

have not received the degree of attention that they deserve

and much valuable work may be done in the detailed study of

their protoplasmic structure and activities at various periods of

ontogeny especially through the series of changes that are

presented during developmental processes.

The primitive types of plastids are relatively large structures,

often solitary in the cells, and generally of complex form.

These are called chromatophores and are characteristic of many

algæ especially among the lower groups but are not found above

the thallophytes (Anthoceros and Selaginella excepted).

The chromatophores of the simplest algæ are replaced in most

of the higher types of these thallophytes and in all groups

above by very much smaller structures, generally .- discoid in

No. 449.] STUDIES ON THE: PLANT: CELL. COE

form, which are called chloroplasts when green, chromoplasts

when the color is other than green or leucoplasts if colorless.

These plastids are without doubt derived from the more primitive

chromatophores.

The colors of chromatophores are various. They are believed

always to contain some chlorophyll but this green is frequently so

completely masked by other pigments that its presence can only

be determined when the additional coloring matters have been

extracted. Chloroplasts are universally green except when they

may be changing into chromoplasts. Chromoplasts generally

take their tint from the predominance of other strong pigments ;

in addition to chlorophyll as phyccerythrin in the red and phyco-

phain in the brown alge. But chromoplasts may be derived

from chloroplasts whose green has largely or wholly disap-

peared leaving other pigments present as the yellow, xanthophyll,

or the orange red, carotin.

The remaining plastids, leucoplasts, are devoid of color and

are found in embryonic regions such as eggs, growing points,

and in the various tissues of seeds, underground organs and

other structures where the cells are largely or wholly removed

from sunlight. The leucoplasts may become green upon expo-

sure tó light thus changing into chloroplasts. They are respon-

sible for the secretion of reserve starch in many structures (e. g.

potato) and in consequence have been called amyloplasts.

Leucoplasts, chloroplasts and chromoplasts are morphologically

the same structures. It is well known that they may pass one

into the other in the order indicated and that chloroplasts and

chromoplasts may lose their color and become leucoplasts. It is

generally believed that plastids are not formed de nove. They

divide by constriction and thus multiplying are passed on from

cell to cell and it is believed from generation to generation.

They are therefore usually ranked as permanent organs of the

cell. However, it is but fair to call attention to the fact that

there are some serious difficulties in the way of a complete

acceptation of these views.

The protoplasmic structure of the plastids of higher plants

is rather simple while that of the chromatophores In alge is

more complex since they contain a special organ termed the

376 THE AMERICAN NATURALIST. (VoL. XXXVII.

pyrenoid. The detailed structure of chromatophores was first

described by Schmitz (82) and of plastids by Meyer ('83).

The most complete study of plastids however is that of Schimper

(85). The body of the plastid is always denser than the sur-

rounding cytoplasm. It has a porous structure that is only

visible under high magnification and there are sometimes present

very delicate fibrils. The coloring matter, oily in consistency,

is held in the pores as minute globules. The plastid may

therefore be compared to a very fine-textured sponge saturated

with pigment. All of the coloring matter of the plastid may

be readily extracted with alcohol leaving the colorless proteid

matrix.

The pigments of plastids are then in the nature of secretions

held in these specialized regions of protoplasm. Chlorophyll is

the principal substance and, as has before been said, is almost

always present, but the amount is sometimes so small that its

green is completely hidden by the color of other pigments.

Chlorophyll itself contains greater or less amounts of two other

coloring matters that may be readily separated from the pure

green, a yellow xanthophyll and an orange red carotin, both

substances closely related to chlorophyll. The other pigments,

characteristic of the chromatophores in some groups of alge,

are however quite distinct from chlorophyll. There is phycocyan,

found in the blue green algze (Cyanophyceze), phycophzein and

phycoxanthin, characteristic of the brown (Phaophycez) and

phycoerythrin of the red (Rhodophycez).

Chloroplasts are found almost universally in green plants

above the Thallophytes and are also present in the large group

of algz the Siphonales and in the Charales. They are some-

times formed very numerously in the cell, reproducing rapidly

by fission (see Fig. 2 a 2, 3) and lie in the layer of protoplasm

just inside of the plasma membrane. They are sensitive to light

and readily shift their position in the cell. Strong illumination

results in their retreat from exposed positions to the sidewalls

and bottom of the cell where the light is less intense. If the

illumination be weak they may all gather on the side most favor-

able for the reception of light. These facts are well illustrated

by the behavior of the plastids in some of the Siphonales (e. £

No. 449.] STUDIES ON THE PLANT CELL. 377

Botrydium), in the Rhodophycez (e. g. Polysiphonia) and also

in the palisade cells of leaves. Chloroplasts after exposure to

light generally contain starch but in some plants this substance

is never formed (e. g. Vaucheria, Fig. 2 A 1), the first visible

products of photosynthesis being other substances more of the

nature of oil. It is not known whether the starch grain in the

Fic. 2.— Plastids. æ, Chloroplasts: : Vaucheria, with oi] globules; 2 Bryopsis; 3

moss (Funaria), in division and containing starc ins ; 4 Oxalis, with a grain of

b, (— à 1 Tropaeolum, epidermal cell from calyx; 2 Fucus, 3

Callithamnion. c, Chromatophores : 1 Spirogyra, with pyrenoids (7) and caryoids

(c); 2 Hydrodictyon, a forming starch; 3 Nemalion; 4

sion and semel starch. d, Leucópiasts: : Phajus, plastid and starch grain at

- side ot the ucleus ; 2 Iris, from and containing oil globules; 3 Iris, ir

t, with starch grains. After Meyer, Strasburger, P. lla, 1 sese

lake and Schimpe:

chloroplast results from the direct change of some of the pro-

teid substance or whether it is a secretion. The conditions

are somewhat different when pyrenoids are present in a chro-

matophore as will be described presently.

The chloroplasts of higher plants may change color under

various conditions and become chromoplasts. Some of the best

378 THE AMERICAN NATURALIST. [Vor. XXXVIII.

examples are found in the colored cells of certain floral parts

and fruits (Fig. 2, 61). These pigments are generally either

xanthophyll (yellowish) or carotin (orange red). . Chloroplasts

may also turn brown especially in older cells that are losing their

contents. The colors of some leaves and flower parts are due

not to the plastids but to substances dissolved or otherwise held

in the cell sap of the vacuoles. The brilliant coloration of

autumn foliage is of this character as well as some of the tints

of petals, hairs and other structures. The chromatophorés of

the higher brown Algz (Phzeophycez) and most of the red

(Rhodophyceze) have the discoid form characteristic of chloro-

plasts (Fig. 2 6 2, 3). They might be called phzoplasts and

rhodoplasts if one wished to classify plastids according to their

color.

The structure of chromatophores is frequently. complicated by

the presence of pyrenoids which may be quite numerous in the

body. These structures are denser regions of the chromatophore

with a definite boundary. They are proteid in character and

are known to vary in size with nutritive conditions and may

completely disappear if the cell is starved. They have been

regarded as masses of reserve proteid material but certain func-

tions of great importance are also associated with them. The

arrangement of starch grains in the chromatophores of many

algze is clearly around the pyrenoids as centers. For this reason

they have been called amylum centers. Timberlake (:01) has

recently shown in Hydrodictyon that segments are split off

from the pyrenoids (see Fig. 2, c 2) and changed directly into

starch grains which naturally lie for a time close to the source

of their formation and only later become distributed throughout

the chromatophore. It is probable that similar conditions will

be found in other algæ (Conjugales, Protococcales, etc.) and we

may soon have a much clearer understanding of the pyrenoid.

The indications are that the pyrenoid will prove to be a region

of the chromatophore differentiated as a metabolic center, more

or less prominent according to conditions of nutrition, and that

its most conspicuous activity is the formation of starch by the

direct transformation of portions of its substance.

Some other structures besides the pyrenoids have. been

No. 449.] STUDIES ON THE PLANT CELL. 379

described by Palla ('94) in the chromatophores of several of the

Conjugales and have been named caryoids. Caryoids (Fig. 2,

¢ i) are smaller and more numerous than pyrenoids and are

distributed irregularly in the chromatophore but chiefly along

the edge. Their function is not known.

The leucoplasts complete the list of plastid structures. They

are colorless and may be found in underground or other portions

of the plant removed from light or where there is little or no

photosynthetic activities as in embryo sacs, seeds, growing points,

‘etc. They become impregnated with chlorophyll under condi-

tions suitable for photosynthesis thus changing into chloro-

plasts. An important function of the leucoplast is the forma-

tion of reserve starch in various parts of the plant. The more

recent investigations of this process (Meyer, '95, Salter, '98)

claim that it is in the nature of a secretion within the substance

of the leucoplast.. This view is opposed to the older conceptions

(Schimper, '81, Eberdt, '91), which regarded the starch grain as

formed by the direct change of proteid material in the plastid.

In view of Timberlake's ( : o1) studies on the pyrenoid of Hydro-

dictyon we may well hesitate to fully accept the views of Meyer

and Salter and ask for further investigations of this very difficult

subject. In addition to starch leucoplasts may contain proteid

crystals and oil globules.

The reproduction of plastids and their evolutionary history

in ontogeny and phylogeny offers a very attractive field for

research. It is well known that plastids multiply by fission and

it is generally believed that they never arise de nove but are

passed from generation to generation as permanent organs of

the cell. The process of division may be very favorably studied

in the spore mother-cell of Anthoceros (Fig.2, ¢ 4). The fission

begins (Davis, '99) by a constriction at the surface as though

the bounding membrane of cytoplasm exerted pressure upon an

elongating structure. There is no evidence that the interior of

the chloroplast undergoes any changes that could assist the

process further than a possible tendency of the two separating

portions to gather their substance together as division proceeds.

' The conditions suggest that the division is a mechanical separa-

tion of material too bulky for the best advantages of the cell,

380 THE AMERICAN NATURALIST. (Vor. XXXVIII.

for the proper balance of protoplasmic elements in narrow

confines, a division prompted by the activities of the cytoplasm

rather than emanating from within the plastid.

The view of the permanence of the plastid as a cell organ has

received its strongest support from the classical work of

Schimper (’85). We are not prepared to deny it and to assert

that the plastid may arise de novo. Yet those who study the

cells of embryonic tissues and reproductive phases know that it

is extremely difficult to follow the plastids and that these

structures require other than the usual methods of cell research

to establish their presence. Several writers (Eberdt, Dangeard,

Husek and others) have expressed their belief that plastids may

arise de novo but no one has thoroughly traced the appearance

or disappearance of these structures in any cells.

The plastid in phylogeny has never received the attention that

it deserves. Beginning with the conditions among the Cyano-

phycee and the lowest Chlorophyceze (which will be further

discussed in Section VI) we find the pigment distributed so

generally throughout the cell that it is doubtful if the term

chromatophore should ever be applied to regions so indefinite in

outline. Above these groups the pigment is confined to propor-

tionally smaller areas in the cytoplasm and these become

chromatophores when their form is clear. The primitive chro-

matophores were solitary and filled a large part of the cell.

The pyrenoids arose in the chromatophores probably as the

result of the influence of metabolic centers upon the protoplasm.

It is scarcely possible that a large chromatophore should be

absolutely homogeneous throughout; there would develop one

or more centers of metabolic activity and such would exert some

influence on the form of the protoplasm.

But the large single chromatophore does not seem to be the

form best adapted to the work of a cell perhaps, if for no other

reason, because it requires a mechanical adjustment of other

cell organs to itself and would interfere with the quick circula-

tion of material and the general balance of cell activities.

It seems possible that mechanical difficulties may have led to

the division of large chromatophores and the substitution of

numerous small plastids. This change was instituted in the

No. 449.] STUDIES ON THE PLANT CELL. 381

higher members of the Phaeophycez and Rhodophycez and in

the Siphonales, Charales, Cladophoraceze and some smaller

groups of the Chlorophyceze. The Conjugales whose chromato-

phores are especially elaborate have cells essentially solitary in

their life habits and with a very remarkable adjustment of the

cell organs to one another to give almost perfect symmetry.

With the splitting up of the chromatophore came the loss of the

pyrenoid and the final result was the compact plastid so charac-

teristic of plants above the thallophytes.

There is no region of the plant cell that maintains such varied

relations to its environment and performs so many visible

activities as the cytoplasm. For this reason the accounts of its

Structure and behavior have been diverse and there has developed

a nomenclature of its parts that is confusing and somewhat

difficult to harmonize.

Strasburger has for many years (since 1892) employed the

term kinoplasm to distinguish an active portion of the cytoplasm

(concerned with the formation of spindle fibers and other

fibrille, centrospheres, centrosomes, cilia, plasma membranes,

etc.) from more passive nutritive regions which he called tropho-

plasm. Kinoplasm corresponds closely to the archoplasm of

the animal cell (Boveri, 1888). This classification has been

criticised especially by Pfeffer (:00) on the ground that it

employed names signifying physiological differences when the

distinctions as far as we know are those of morphology alone.

However the physiological behavior of kinoplasm and tropho-

plasm becomes very real to anyone who studies extensively cell

activities and the morphological characters serve to emphasize

these peculiarities. The truth seems to be that cell studies

cannot be pursued from the standpoint of physiology or mor-

phology alone but must combine these attitudes. And in the

union it is hardly possible or perhaps desirable to construct

a terminology with strict regard to either field of study. We

shall use the terms kinoplasm and trophoplasm grouping the

various cytoplasmic structures under these heads.

382 THE AMERICAN NATURALIST. (Vor. XXXVII.

Cytoplasm has surface contact with three conditions and in

each case there is present a delicate plasma membrane, colorless

and very finely granular, which is very different in structure from

the cytoplasm within. The first of these three membranes is

the outer plasma membrane, which bounding the protoplast, is

consequently just inside the cell wall. This membrane is called

the *hautschicht" by the German botanists, a word for which

we have no exact equivalent, the term ectoplast more nearly

expressing the meaning than any other but for several reasons

not being very satisfactory. Since this outer plasma membrane

lies against a moist cell wall it is virtually surrounded by a film

of water. The functions of the cell wall in land plants and its

developmental history indicate a close relation to the demands

of the outer plasma membrane for a fairly uniform environment

of moisture, a matter which will be discussed in the last section

of these papers.

The second form of plasma membrane surrounds the water

vacuoles in the cell. It is very common for the plant cell to

have a single large central vacuole containing the cell sap and

the membrane around this was named the tonoplast by DeVries

in 1885. DeVries believed that this vacuole reproduced itself

by fission with each cell division and consequently was a perma-

nent organ of the cell. It is, however, now well known that the

large central space containing cell sap-is not different from other

vacuoles, indeed is frequently formed by the flowing together of

several small vacuoles as smaller soap bubbles unite in the froth

to form a larger one A vacuolar plasma membrane is of course

bathed by water since it holds the cell sap and its relation to a

moist surface is therefore more evident than in the case of the

outer plasma membrane.

The third plasma membrane encloses the nuclear sap with the

protoplasmic nuclear elements chromatin, linin and the nucleolus.

This nuclear membrane was discussed in connection with the

nucleus of which it is generally considered a part, but as there .

stated, the evidence largely indicates that it is cytoplasmic in

Character, representing a reaction of this protoplasm to the flui

nuclear sap formed around the chromosomes in the daughter

nuclei after each division (Lawson :03?) The nuclear sap

No. 449.] STUDIES ON THE PLANT CELL. 383

necessitates the development of a vacuole which becomes

bounded by the nuclear membrane. The nuclear membrane

in some cases at least differs from a vacuolar membrane in

being easily distinguished from the surrounding cytoplasm as

a definite film.

The structure of all the plasma membranes is much the same

as far as the microscope may determine. The protoplasm is

dense, colorless and filled with very minute granules (micro-

somata). There are no large inclusions such as plastids, parti-

cles of food material (starch, proteids, oils, fats, etc.), mineral

matter or waste products. These are all held well within the

cytoplasm between the outer plasma membrane and the vacuoles.

There is good reason to believe that the substance of all plasma

membranes is much the same since they perform very similar

activities both in relation to the fluids that bathe them and also

because their substance in certain cases becomes the proto-

plasmic basis of cellulose walls. These resemblances are well

established for the outer plasma membrane and that which sur-

rounds the vacuoles. Thus, the capillitium of Myxomycetes

(Strasburger, '84) is formed from the plasma membranes around

the vacuoles after the same method as a cell wall from the outer

plasma membrane. And again, during cleavage by constriction

(see section II) in the plasmodium and sporangium of the molds

(Harper, '99 and :0o, D. Swingle, : 03), vacuoles fuse with cleav-

age furrows from the outer plasma membrane to form a common

membrane which surrounds each spore mass and secretes a wall,

thus showing identity of function and structure. The resem-

blances are less conspicuous for the kinoplasm of the nuclear

membrane, only appearing indirectly with certain events of cell

division (the formation of the cell plate) which will be discussed

in the next section of the paper. The evidence indicates that

the three plasma membranes are all kinoplasmic in character, a

generalization of some importance since- it offers explanations of

many peculiar cell activities to be described later.

Since all plasma membranes have these common characters it

may well be questioned whether an elaborate terminology is

justified for structures so closely related. The terms ectoplast

and tonoplast seem undesirable since they were meant to indi-

384 THE AMERICAN NATURALIST. [Vor. XXXVIII.

cate peculiarities of structure and a degree of permanence as

cell organs that is not actually present. It seems hardly neces-

sary to define the plasma membranes further than by their posi-

tion in the cell as the outer, vacuolar and nuclear membranes.

All of the cytoplasm bounded by the plasma membranes with

the exception of certain conditions to be described later (centro-

spheres, centrosomes, asters, filarplasm and blepharoplasts) may

be called trophoplasm since it contains structures and substances

especially concerned with nutritive functions. Trophoplasm .

presents an open organization in sharp contrast to the dense

kinoplasm. This peculiarity is due in part to numerous small

vacuoles which give a spongy appearance to the usual foam like

structure and is further complicated by the inclusion of material

not strictly a part of the protoplasm in the form of various sized

granules. There are sometimes present fibrillae that impart a

somewhat fibrous texture. We cannot discuss here the theories

of the structure of protoplasm, which has not been so extensively

studied in plants as among animals, further than to point out

that it varies considerably in different regions of the cell in

relation to peculiarities that will be described later. There is

sometimes presented very typically the foam structure of

Bütschli but the introduction of small vacuoles generally gives a

spongiose appearance. This subject is critically reviewed by

Fischer, '99, and has also been treated in several papers of

Strasburger especially in '97.

Three well differentiated organs of the cell, probably tropho-

plasmic in character, require special mention, víz., coenocentra,

nematoplasts and physodes. Cocenocentra are very interesting

protoplasmic centers found in the oogonia of certain coenocytic

fungi among the Saprolegniales and Peronosporales during

oogenesis. They appear just previous to the differentiation of

the eggs as small bodies sometimes with delicate radiations (see

Fig. 3, a and 8, f), and are found one in each egg origin. They

are apt to increase in size as the eggs mature and evidently

become the centers of the metabolic activities of the cells,

drawing the sexual nuclei into their neighborhood where the

latter increase in size (Fig. 3, « 2). The coenocentrum dis-

appears in the ripe oóspore and is consequently an evanescent

No. 449.] STUDIES ON THE PLANT CELL. 385

structure. It is probably the morphological expression of a

dynamic center in the egg. — Coenocentra have been known for

several years and have been given especial attention in the

recent investigations of Stevens, '99 and 'or, and the author

(Davis, :03). They will be further considered in our account of

Coenogametes (Section III).

Nematoplasts are exceedingly small rod or thread like

Fic. 3. — Cytoplasmic structures. æ, Coenocentra of Saprolegnia; 1, odgonium, each

oenocentrum; 2, coen ocentrum and nucleus from mature egg.

cytoplasmic radiations weed like). /, Nucleus from procambium cell of Vicia, kino-

plasmic caps. g, Polle — of Lilium, filarplasm in form of multipolar

spindle. A, "bita of sperm of Gymnogramme ; r, blepharoplast at side of

sperm nucleus ; 2, blepharoplast elongating and developing cilia; 3, mature sperm,

blepharoplast and nucleus in parallel bands, cytoplasmic vescicle below. After

Zimmermann, Hof, and Belajeff.

structures reported by Zimmermann ('93, p. 215) in the cells of

hairs of Momordica and the root of Vicia (see Fig. 3B). It is

probable that organs described by Swingle, '98, and Lagerheim,

'99, under the names of vibrioides are the same as or closely

386 THE AMERICAN NATURALLIST. [Vor. XXXVIII.

related to physodes. Swingle found them in some of the

Saprolegniales and certain Rhodophycez and Lagerheim in

Ascoidea. They are probably not uncommon. Nematoplasts

may be proteid crystals but there is evidence that they move,

bending slowly back and forth, which suggests a higher degree

of organization. They should be further studied.

Physodes are bladder like structures described by Crato, '92,

in certain brown Alga. They contain a highly refractive sub-

stance which gives them a very different appearance from

vacuoles whose structure they resemble in many respects.

Very little is known about the contents of physodes and it may

well be questioned whether they are really organs of the cell

and not vacuoles set apart to hold some fluids or substances

other than cell sap.

There are left for us a group of kinoplasmic structures that

are especially prominent and sometimes only present during the

events of nuclear division and at the times when cilia are

formed. They will be discussed in later sections of these papers

(Sections II, III, V and VI) and at this time we shall give but

a brief statement of their appearances. They are centrospheres,

centrosomes, asters, filarplasm and blepharoplasts.

Centrospheres are rather large areas of kinoplasm that some-

times lie at the poles of nuclear figures and to which are

attached the fibrille that form the spindle and also those that

may radiate into the surrounding cytoplasm. If the centro-

sphere contains a distinct central body, or if such a small

structure be present alone at the poles of the spindle it is called

a centrosome. Should either structure be accompanied by

definite fibrillar radiations the whole is termed an aster. These

latter conditions are sometimes very complex and are the most

interesting types of structures. Asters with centrosomes are

known for the brown algæ in the growing points of Sphacelaria

(Fig. 3c), Stypocaulon (Swingle, '97) and the spore mother cell

of Dictyota (Mottier, :00). They are also beautifully shown in

certain diatoms (Lauterborn, principal paper '96, Karsten, : 00).

Asters with centrospheres and occasionally but not constantly

containing centrosome-like bodies are found in the oogonium

and germinating eggs of Fucus, see Fig. 3, d (Strasburger, '97^

No. 449.] STUDIES ON THE PLANT CELL. 387

Farmer and Williams, '98). Especially well differentiated asters

with centrospheres are present during the mitoses in the ascus,

functioning at the end in the peculiar process of free cell

formation (Harper, '97). Large centrospheres accompanied by

radiations are present during the germination of the spores in

certain Hepaticee (Farmer and Reeves, '94, Davis, :o1, Cham-

berlain, :03), but are less conspicuously shown in some and are

entirely absent in other phases of the life history. Remarkably

large centrospheres with inconspicuous radiations are known in

the tetraspore mother cell of Corallina (Davis, '98). Centro-

spheres occur in the basidium (Wager, '94, Maire, :02). Cen-

trosomes have been reported during the mitoses in the

sporangium of Hydrodictyon (Timberlake, :02). Centrosomes

have also been described in other types of the thallophytes but

we are justified in asking for further work on these bodies

since they are generally without raditions and may not have at

all the significance indicated. Neither asters, centrospheres or

centrosomes seem to be normally present in groups above the

bryophytes, nuclear division taking place in these plants by

methods, not found in other organisms, which will be described

in succeeding sections.

Vegetative and embryonic tissues of plants above the thallo-

phytes present very different conditions from those described in

the foregoing paragraph. The centrosphere is replaced by a

less definite structure in the form of a kinoplasmic cap which

appears at the ends of the dividing nucleus and determines the

poles of the spindle (see Fig. 3, f). They have been described

in the cells of vegetative points of several pteridophytes and

spermatophytes by Rosen, ’93, Hof, ’98, and Nemec, '99 and : O1,

and in the seta and late divisions in the germinating spore of

the liverwort Pellia (Davis, : o1).

The most highly developed conditions of spindle formation

are found in the spore mother cells of the bryophytes, pterido- _

phytes and spermatophytes. Here the nucleus becomes

surrounded by a weft of fibrillæ which form a kinoplasmic

envelope probably derived in part from the nuclear membrane.

The fibrillæ are at first quite independent of one another or of

common centers. Most of the fibrillæ enter into the spindle

388 THE AMERICAN NATURALIST. [Vor. XXXVIII.

which may in the beginning have several poles (see Fig. 3, g),

but these generally swing at last into a common axis so that

the spindle finally becomes essentially bipolar. The term

filarplasm is applied to this free fibrillar condition of kinoplasm

without organized centers. Filarplasm is peculiar to plant cells

and its remarkable activities in connection with multipolar

spindles have only been found in groups above the thallophytes.

Centrospheres, centrosomes and asters among the lower plants

resemble in general the same structures in the animal cell. But

filarplasm presents a higher form of kinoplasmic structure with

perhaps the most complex activities known in the process of

spindle formation. We shall consider them especially in Section

III when treating the spore mother cell.

The blepharoplasts are in some respects the most complex

structures derived from kinoplasm. They are most conspicuous

in the sperm cells of higher plants (spermatophytes and

pteridophytes) but they are undoubtedly present in lower

forms and probably in zoospores. The blepharoplast develops

cilia as delicate fibrille from its surface. The origin.and homol-

ogies of the blepharoplast are uncertain. In some forms they

resemble centrosomes at the poles of the last nuclear figures in

sperm tissue. But in other cases they are entirely independent

of such spindles, a character which cannot be brought into

harmony with the activities of centrosomes. They finally lie

one at the side of each sperm nucleus, see Fig. 3, 7, and with

the development of the sperm they follow the spiral twist, when

present, as a parallel band (Fig. 3, 7, 2 and 3). This structure

will receive detailed treatment in our account of the sperm

(Section III).

2. Non Protoplasmic Contents.

It is not possible to distinguish with certainty all the non-

living material of a cell from its protoplasm. We have at one

extreme cells from which the protoplasm has almost or wholly

disappeared and which are either entirely empty or set apart

solely as receptacles for various substances, sometimes waste

products and sometimes food materials. In contrast with this

No. 449.] STUDIES ON THE PLANT CELL. 399

condition are the cells filled with cytoplasm so homogeneous in

structure that only the most delicate granules (microsomata) can

be distinguished in the clear substance.

Waste products such as mineral matter, resins, certain oils,

solutions of tannin and various poisons, such as the alkaloides,

may be easily recognized. Most food substances such as starch,

proteid grains (aleurone), albumin crystals, oils, fats, etc., are

readily separated from the protoplasm in which they lie. But

the difficulties are much greater with the smaller particles of

proteid material, which are frequently such minute granules as.

to approach the microsomata in size. These may give to the

protoplasm a granular consistency that breaks up the foam or

spongiose structure characteristic of the pure condition. These

granules are undoubtedly in most cases substances intimately

concerned with the metabolism of the cell and are members of

the chains of constructive and destructive processes that charac-

terize life phenomena.

The other non protoplasmic structures of cells are vacuoles

which are essentially bubbles of fluid lying in the denser proto-

plasmic medium and surrounded by plasma membranes. The

watery fluid of vacuoles contains various substances in solution,

carbohydrates such as the sugars glucoses and inulin, mineral

salts, asparagin, tannin, alkaloids, etc., and occasionally oil and

not infrequently crystals. Vacuoles may be formed in large

numbers in protoplasm. They tend to run together as do

bubbles in a froth and in this way the large central vacuole

becomes established in the cell, gathering to itself many smaller

vacuoles until the protoplasm is forced to lie as a relatively thin

layer next the cell wall. The fluid in the central vacuole (cell

sap) is generally thinner and more watery than that in the

smaller vacuoles. The latter are apt to be more rich in albumen

which may be transformed into proteid grains as is especially

well illustrated in the secretion of aleurone. Cell sap may be

colored by pigments in solution and the tints of flowers are

largely due to this cause alone or to the effects of its color in

combination with various plastids in the cell.

It is possible that physodes, described among the cytoplasmic

Structures, are in reality vacuoles filled with substances other

than cell sap, which are not as yet understood.

390 THE AMERICAN NATURALIST, [Vor. XXXVIII.

3. The Cell Wall.

Many of the chief peculiarities of plant organization and

activities are due to the presence of the cell wall, its influence

on structure and mode of life. The cell wall is not an excretion

from the cell like a mineral shell but is formed by the direct

change of portions of the protoplasm. The regions concerned

may be the outer plasma membrane, the vacuolar plasma mem-

brane or the substance that makes up the spindle fibers which

form the cell plate. These structures are all kinoplasmic in

character and have to do with the formation of cell walls in

various ways which will be described in Section II under the

topic “ The Segmentation of the protoplasm.” The transforma-

tion of finely granular films of kinoplasm into cellulose is not

well understood but there is an evident solution of the granules

(microsomata) and the change of the resultant substance into

the cell wall. As a chemical process this change means the

replacement of molecules of an albuminous nature by those of a

carbohydrate substance. The most complete account of the

cell wall is that of Strasburger, '98.

Cell walls are chiefly composed of cellulose, but other sub-

stances are always present, modifying the structure in various

ways to give widely different properties. These modifications

are generally due to infiltrations of foreign substances but some-

times cell walls become incrusted with mineral deposits. The

group of cellulose compounds is very large and it is extremely

difficult to identify the various substances in structures so small

as the cell walls. For a detailed treatment of the chemistry of

the cellulose group the reader is referred to Cross and Bevans,

'95, and for a general account to Pfeffer, : 00, p. 480-485. There

are microchemical tests for cellulose that give good reactions

for most tissues but which cannot be relied upon for some walls

(as in fungi and many algze) yet it is well understood that the

cell walls of these organisms are from the biological point of

view essentially, the same as for other plants. The cell walls of

some fungi are very largely composed of chitin. a8 :

Several substances known to be present in cell walls give

them marked characteristics. Their association with the cellu-

No. 449.] STUDIES ON THE PLANT CELL. 391

lose is so intimate as to resist very severe treatment and there-

fore these cell walls are essentially cellulose groups modified

chiefly in their physical properties by the presence of foreign

- substances. The most conspicuous modifications of this charac-

ter are lignification, suberization and cutinization. Lignified

walls are permeable to water and gases. Several substances

have been separated from the cellulose of lignified walls, among

them lignone, coniferin, vanillin, etc. Suberized and cutinized

walls are largely but probably never wholly impervious to water

and gases; the one is infiltrated with suberin and the other

with cutin, substances that resemble one other very closely.

Even walls that appear to be pure cellulose have other sub-

stance united with them, the most important being pectose and

callose. Cell walls frequently become gelatinous or mucilaginous,

when the outer layers swell and lose their form or they may be

transformed into gums. These changes are well illustrated in

the coats of seeds and fruits and among the alge and fungi.

The cells of alga frequently secrete gelatinous envelopes or

sheaths of substances so closely related to cellulose that were

they condensed they would form a firm cell wall.

The cell wall may grow in two directions by methods quite

different from one another. There is first surface growth

which results in a stretching of the cellulose membrane (growth

by intussusception). And second there may be growth in thick-

ness by the formation of successive layers of cellulose inside of

one another, giving the wall a striated structure (growth by

apposition). The second type of growth is chiefly interesting

since it makes possible many peculiarities of structure, because

the newly formed layers may not be deposited uniformly inside

the primary wall. In some cells the secondary thickenings

have the form of rings or spirals or a reticulate structure. The

reticulate condition passes insensibly into the pitted cell in which

the secondary layers cover the greater part of the surface leav-

ing the primary wall only exposed at the pits. Further dis-

cussion of these cells falls more within the range of histology

than the purposes of this paper.

The cell wall offers a very interesting field of research among ©

the thallophytes and especially in the lower groups where we

392 THE AMERICAN NATURALIST. (VoL. XXXVIII.

may expect to find these envelopes in a fairly primitive con-

dition and may be able to establish the steps in the origin and

differentiation of this very important accessory structure to the

plant cell.

(79 be continued.)

LITERATURE CITED FOR SECTION I * THE PLANT CELL."

CAVARA.

'98. Intorno ad alcune strutture nucleari. Atti. dell. Inst. bot. Univ.

di Pavia, II, 5, 1898.

CHAMBERLAIN.

'99. Oogenesis in Pinus laricio. Bot. Gaz. 27, 268, 1899.

'O3. Mitosis in Pellia. Bot. Gaz. 36, 27, 1903.

CRATO.

'92. Die Physode, ein Organ des Zelllenleibes. Ber. d. deut. bot.

'96. Cellulose, an outline of the chemistry of the structural elements of

plants. 1895.

Davis. E

'98. Kerntheilung in der Tetrasporenmutterzelle bei Corallina officinalis

L. var. mediterranea. Ber. d. deut. bot. Gesell. 16, 266, 1898.

'99. The spore mother cell of Anthoceros. Bot. Gaz. 28, 89, 1899.

:01. Nuclear studies on Pellia. Ann. of Bot. 15, 147, 1901.

:03. Oogenesis in Saprolegnia. Bot. Gaz. 35, 233 and 320, 1903.

DEVRIES. :

'85. Plasmolytische Studien über die Wand der Vacuolen Jahrb. f. wiss-

Bot. 16, 465, 1885.

EBERDT. :

'91. 2 zur Entstehungsgeschichte der Stärke. Jahrb. f. wiss.

: 22, 295, 1891.

FARMER AND REEVES.

'94. On the occurrence of centrospheres in Pellia epiphylla, Nees

n. of Bot. 8, 219, 1894.

FARMER AND WILLIAMS. : `

'98. Contributions to our knowledge of the Fucacez; their life history

and cytology. Phil. Trans. Roy. Soc. 190, 623, 1898.

FISCHER.

'99. Fixirung, Fárbung und Bau des Protoplasmas. Leipzig, 1899.

No. 449.] STUDIES ON THE PLANT CELL. 393

HARPER.

'97. Kerntheilung und freie Zellbildung im Ascus. Jahrb. f. wiss. Bot.

30, 249, 1897

'99. Cell division in sporangia and asci. Ann. of Bot. 13, 467, 1899.

:00a. Cell and nuclear division in FwZge varians. Bot. Gaz. 30, 217,

1900. ;

Hor.

'98. Histologische Studien an Vegatationspunkten. Bot. Centb. 76, 65,

1898.

KARSTEN.

:00. Die Auxosporenbildung der Gattungen Cocconeis, Surirella und

Cymatopleura. Flora 87, 253, 1900.

ie oet

Ueber ein neues vorkommen von Vibrioiden in der Pflanzenzelle.

K. Svenska. Vet. Akad. Forhand. No. 6, 1899.

xo geh

Untersuchungen über Bau, Kerntheilung und Bewegung der

Diatomeen. Leipzig 1896.

LAWSON.

:03a. On the relationship. of the nuclear membrane to the protoplast.

Bot. Gaz. 35, 305, 1903.

MAIRE.

:02. Recherches cytologique et taxonomique sur les Basidiomycetes.

ull. d. 1. Soc. Mycol. d. France. 18, 1902.

MEYER.

'83. Ueber Krystalloide der Trophoplasten und über die Chromoplasten

der Angiospermen. Bot. Zeit. 41, 489, 503, 525; 1883.

‘95. Untersuchungen über die Stárkerkórner. Jena 1895.

NT EWTESeH.

Ueber die Kerntheilung bei Spirogyra. Flora 85, 81, 1898.

Morr.

'94. Observations sur la caryocinese chez les Spirogyra. Arch. Neer.

d. Sci. exactes et naturelle 28, 1894.

MOTTIER.

:00. Nuclear and cell division in Dictyota dichotoma. Ann. ‘of Bot. 14,

163, 1900.

NEMEC.

'99c. Ueber die karyokinetische Kernthielung in der Wurzelspitze von

Allium cepa. Jahrb. f. wiss. Bot. 33, 313, 1899.

:01. Ueber centrosomáhnliche Gebilde in vegetativen Zellen der Gefass-

pflanzen. Ber. d. deut. bot. Gesell. 19, 301, 1901.

PALLA:

'94. Ueber ein neues Organ der Conjugaten Zelle. Ber. d. deut. bot.

Gesell. 12, 153,1

PFEFFER.

:00. The physiology of plants. Clarendon Press 1900.

394 THE AMERICAN NATURALIST. [Vor. XXXVIII.

ROSEN.

'95. Beitráge zur Kenntniss der Pilanzensellen Cohn's Beitr. z. Biol. d.

Pflan. 7, 225, 1895.

SALTER.

'98. Zur näheren Kenntniss der Stárkekórner. Jahrb. f. wiss. Bot. 32,

117, 1898.

SARGANTS.

'97. The formation of sexual nuclei in Lz/w»z Martagon, II, Sperma-

togenesis. Ann. of Bot. 11, 187, 1897.

SCHIMPER.

' "81. Untersuchungen über das Wachstum der Skene Bot. Zeit.

39, 185, 1881.

'85. Untersuchungen über die asap ian und die in ihnen

homologen Gebilde. Jahrb. f. wiss. bot. 16,

SCHMITZ.

'82. Die Chromatophoren der Algen. Bonn. 1882.

STEVENS.

'99. The compound oosphere of 4/bugo Bliti. Bot. Gaz. 28, 149, 1899.

'Olb. Gametogenesis and fertilization in Albugo. Bot. Gaz. 32, 77,

1901

gol.

STRASBURGER.

'84. Zur Entwickelungsgeschichte der Sporangien von Trichia fallax.

Bot. Zeit. 42, 305, 1884.

'95. Karyokinetische Probleme. Jahrb. f. wiss. Bot. 28, 151, 1895

97a. Kerntheilung und Befruchtung bei Fucus. Jahrb. f. wiss. Bot. 30,

351, 189

797b. Ueber Cytoplasmastructuren, Kern und Zelltheilung. Jahab. f.

wiss. Bot. 30, 375, 1897

'98. Die pflanzlichen Zellháute. Jahrb. f. wiss. Bot. 31, 511, 1898.

:00. Ueber Reductionstheilung, Spindelbildung, Centrosomen und

Cilienbildner im Pflanzenreich. Hist. Beit. 6, 1900.

SWINGLE, W. T

'97. Zur Kenntniss der Kern und Zelltheilung bei den Sphacelariaceen.

Jahrb. f. wiss. Bot. 30, 297, 1

'98. Two new organs of the plant cell. Bot. Gaz. 25, 110, 1898.

SWINGLE, D.

:08. Formation of spores in the sporangia of Rhizopus nigricans and

Phycomyces nitens. Bu. Plant Ind. U. S. Dept. Agri. Bull. 37;

190

903.

TIMBERLAKE.

:01. Starch-formation in Hydrodictyon utriculatum. Ann. of Bot. 15,

619, 1901.

:02. Development and structure of the swarmspores of rydrodietyon.

Trans. Wis. Acad. of Sci. Arts and Letters 13, 486, 1902.

No. 449.] STUDIES ON THE FLANI CELL 395

VAN WISSELINGH.

Ueber Kerntheilung bei Spirogyra II. Flora 87, 355, 1900.

:02. Untersuchungen über Spirogyra IV. Bot. Zeit. 60, 115, 1902.

WAGER.

'94. On the presence of centrospheres in fungi. Ann. of Bot. 8, 321,

1894.

ZIMMERMANN.

'93 and '94. Sammel-Referate aus dem Gesammtgebiete der Zellenlehre.

Bei. z. bot. Centb. 3 und 4, 1893-94.

(To be continued.)

NOTES AND LITERATURE.

GENERAL BIOLOGY.

Plankton of the Illinois River. — Probably the most extensive

study of the plankton of any inland waters is contained in Kofoid's

report on the organisms of the Illinois river for the years 1894 to 1899.

The period of minimum productivity of plankton is January and Feb-

ruary; this is followed by rising productivity which reaches its

maximum in April, after which there is a gradual decline to win-

ter conditions. Area and depth showed little relation to plankton

production. Young waters from springs and creeks contain little

plankton, but these waters when impounded in backwater reservoirs

develop an abundant plankton. Fluctuations in hydrographic condi-

tions, temperature, and light affect plankton production. Submerged

` vegetations tends to diminish the production of plankton. The

plankton of the Illinois River is largely autonomous and may be esti-

mated at 67,750 cubic metres. The annual variations in this and in

the river fisheries show some correlation.

Where did Life begin? ?—— A second edition of Scribner's little

book, which attempts to locate the region where life first appeared on

the globe, has just been issued. The argument, now familiar to most

readers, turns on the gradual cooling of the earth's crust. Naturally

the first parts cooled would be the polar regions and here life may

have first originated. No good reason is given for selecting the

northern rather than the southern regions as the real centre nor is the

question of land and water in these regions sufficiently considered.

Since primitive organisms were without doubt water-inhabiting, the

possibility of a land-covered surface in the region where they were

Supposed to originate is not without significance. Inconclusive as

the argument really is, the whole treatment of the subject is sugges-

tive and stimulating.

! Kofoid, C. A. The Plankton of the Illinois River, 1894-1899. Pt. I. Quan-

titative Investigation and General Results. Budi. Wl, State Lab. Nat. Hist., Vol.

7, Art. 2, pp. 95-629, so pls

* Scribner, C. H. Where did Life Begin? New York, Scribner's, 1903, 12vo.

75 pp. à

397

398 THE AMERICAN NATURALLIST. [Vor. XXXVIII.

Bermuda. — The account of the Bermuda Islands by Professor

A. E. Verrill! already issued in the Zransactions of the Connecticut

Academy of Science has been published by the author as a separate

volume. After a general description of the islands, their physiog-

raphy and meteorology are considered and this is followed by a

lengthy description of the changes in fauna and flora due to man.

The geology and marine zoólogy will appear in another volume.

The body of information thus brought together will be invaluable to

the future student of these interesting islands.

Morgan on Evolution and Adaptation.” — A new book on evolu-

tion might at first thought seem superfluous, in view of the already

enormous literature on this subject, but advancement in knowledge

calls for the presentation of fundamental principles in new lights, and

no one who examines this book will find it wanting in food for

thought. ‘The general reader will findin it a convenient summary of

the older views and discussions about evolution, with extensive quota-

tions from the classical writings of Darwin, Weismann, and others.

The new point of view, which especially interests the student and

justifies the volume in his eyes, is that taken by Bateson (1893) in

his Materials for the Study of Variation, and by deVries (1901-3)

in his Mutationstheorie. From this standpoint evolution is nota

continuous but a discontinuous process, in which advance is made by

distinct steps. New species do not arise by the slow cumulation of

fluctuating individual variations in a particular direction, but are born

full fledged. A new species thus produced, which deVries calls a

mutation, differs from the parental species at first, perhaps, in only a

single respect. It possesses some new character not seen in the

parents, or it lacks altogether some character possessed by the par-

ents. It breeds true to its own distinctive character, if separated

from the parent species, or if not so separated may interbreed

freely with it. But when such interbreeding occurs the offspring fall

into two distinct classes, one resembling each parent form. Natura

selection now comes into operation to decide, not between one indi-

vidual and another, but between the two specific forms, that one being

favored which is best adapted to its environment, the other being

eliminated, or possibly being allowed to survive in a different envi-

! Verrili, A. E. Zhe Bermuda Islands, New Haven, published by the author,

I902, 8vo, X + 548 PP., 38 pls. k

2 Morgan, T. H., Evolution and Adaptation. New York, Macmillan, 1903-

8vo, 410 pp.

No. 449.] NOTES AND LITERATURE. 399

ronment. Thus mutation may multiply species without necessitating

the extinction of any or requiring the erection of barriers spacial or

physiological between the new form and the old one to prevent the

swamping of the new form by crossbreeding. A real obstacle to

the older ideas about evolution has thus been removed by fuller

knowledge of the laws of inheritance of mutations.

Though a mutation differs from the parent species at first in a

single character only, the number of differences is likely to increase,

for one mutation leads to another, as observation clearly shows. Ac-

cordingly natural selection is presently called upon to make a choice,

not simply between wo alternative forms, but among several distinct

and mutually exclusive types, some one of which will be better

adapted to a particular environment than any other.

The adaptations of organisms are almost endless and involve the

most minute details of structure and function, yet the two principles

of mutation and of exclusive inheritance are sufficient, in Morgan’s

opinion, to account for them all. Natural selection acting merely on

the fluctuating variations of individuals fails to do this.

In justice to Darwin it should be said that the mutation or sport

forming tendency of species was distinctly recognized by him, but

he attached less importance to the process than do Bateson and

deVries. The position taken by these writers, and emphasized by

Morgan, is that mutations are the exclusive source of the material on

which natural selection acts,in the production of new species.

A serious defect of the book from the student's standpoint is the

total omission of bibliographic references.

W. E. C.

ZOOLOGY

Zoülogical Investigations in the Malay Archipelago.— Under

the auspices of the universities of Edinburgh and of Liverpool,

N. Annandale and H. C. Robinson undertook an expedition in

1901-1902 for anthropological and zoólogical investigations in the

Malay Archipelago, and some of the zoólogical results of their work

have recently appeared in two fascicles.! The first contains a report

1Annandale, N., and Robinson, H. C. Fasciculi Malayenses. Zoülogy. Parts

I, II, and Supplement. London. Longmans, Green & Co., 4^, vii + 307 PP»

14pls.; xliii pp., map. |

400 THE AMERICAN NATURALIST. (Vor. XXXVIII.

on the mammals by J. L. Bonhote. In the considerable list of species

reported there are a new species of small carnivor, a new bat, a new

squirrel, and three new species of rats. C. Swinhoe reports on some-

thing over 250 species of moths, of which seventeen are new. Two

land planarians, one of which is new, are described by F. F. Laidlaw,

who also reports on the dragonflies. Three new diptera pupipara

are recorded by Speiser. G. A. Boulenger reports 85 species of

batrachians and reptiles, six of which are new. The tiger beetles

are described by H. C. Robinson.

The second part contains nine short papers: four on insects, two

on fishes, and one each on mollusks, the mouth funnel of a tadpole,

and a fossil elephant tooth. In a report on the non-operculate pul-

monates W. E. Collinge gives a full account of the anatomy of a new

and very large species of Atropos; and in J. Johnstone's paper on

the marine fishes, an interesting description of a new species of

Periophthalmus is given. Its life on the mud flats out of water and

its burrows are fully described and illustrated. The fact that when

in the air it does not respond to the report of a gun led to the con-

clusion that it was absolutely deaf. Its eyesight both in water and

1n air was acute.

The two fascicles were accompanied by a supplement containing

a map and an itinerary.

of zoólogy and particularly to mammalian anatomy, Davison! has

prepared an account of the anatomy of the cat. The volume, which

contains some 250 pages with above roo illustrations, is unsatisfac-

torily brief and in consequence it is deficient as a description of the

anatomy of a type and as an introduction to comparative study.

Although brevity may have been the aim of the author and certain

defects may therefore have been unavoidable, others are present in

the volume for which no such excuse can be found. Thus the

description of the gastrocolic omentum as a closed sac is wholly mis-

leading, and the grouping together of the corpora quadrigemina,

optic thalami, and corpora striata as basal ganglia counteracts what

has been gained for these bodies from the standpoint of comparative

anatomy. The facial nerve is placed without qualification among

the pure motor nerves, and the circumvallate papillae of the tongue

are noted as eight to twelve in number, as in man, though the figure

Phila-

n, A. Mammalian Anatomy with special Reference to the Cat.

Bes P. Blakiston’ s Son & Co., 8°, xii +250 pp., 108 figs.

No. 443:] NOTES AND LITERATURE. 401

shows six, the usual number for the cat. Defects of this kind are

too frequent to make the book really useful in the hands of most

beginners. ‘

Notes. — The action of light on organisms and the production of

light by organisms are presented in readable form by R. Dubois in

the second volume of the Zraité de Physique Biologique. The effect

of light on the action of enzymes, on the production and destruction

of pigments, and on the circulation and respiration of the higher

animal is described at some length. The influence of light on the

movements of animals is very inadequately treated probably because

most of the researches on this subject have come from other than

French laboratories. Dermatophic vision and its relation to vision

by means of eyes is fully discussed from the standpoint of Pholas.

The production of light by organisms forms a brief, compact essay

dealing with the photogenic bacteria, the light-producing protozoa,

insects and mollusks. It contains interesting statements of the rela-

tive energy values of living and mechanical sources of light and is

illustrated by some remarkable photographs taken by light from liv-

ing organisms. It is marred by an attempt to discriminate between

chemical, light, and heat rays.

A brief account of the structure of the rudimentary eyes in the

Cuban blind snake, Typhlops lumbricalis, has been published by

E. F. Muhse in the Biological Bulletin, Vol. V, No. 5, 1903. The

eye appears as a dark spot surrounded by an unpigmented circle

and covered by a large ocular scale. Internally the usual parts

can be distinguished including a well-developed lens and a retina

in which the layers typical for snakes can be seen.

R. Dubois last year reported to the French Academy of Sciences

and to the Society of Biology the success of his experiment to

acclimatize true pearl oysters on the French coast and to produce

precious pearls by artificial means. His methods resulted in

producing small but high grade pearls in one in ten oysters whereas

under natural conditions it was necessary to open 1200 to 1500

oysters to obtain one pearl. -

C. H. Eigenmann and C. Kennedy in No. 5, Vol. 4 of the

Biological Bulletin call attention to an unusual melanic individual of

the cave salamander, Spelerpes maculicaudus, to a catfish rom

Titicaca with a branched left barbule, and to a specimen of Xiphor-

hamphus with an additional left ventral fin.

402 THE AMERICAN NATURALIST. (VoL. XXXVIIL.

BOTANY.

The Journals. — Bulletin of the Torrey Botanical Club, February :

Richards and MacDougal, “Influence of Carbon Monoxide and

other Gases upon Plants"; Berry, “ Additions to the Flora of the

Matawan Formation”; Griffiths, “Concerning some West American

Smuts "; Dandeno, * Mechanics of Seed-Dispersion in Ricinus commu-

nis”, and Howe, “Notes on Bahaman Algz."

Journal of Mycology, January : — Morgan, * A New Sirothecium ";

Hedgcock, * Proof of the Identity of Phoma and Phyllosticta on the

Sugar Beet”; Atkinson, “Notes on the Genus Harpochytrium ^;

Kellerman, *Notes from Mycological Literature" — VIII, and

“ Index to Uredineous Infection Experiments."

Journal of the New York Botanical Garden, February :— Britton,

“George Washington's Palms "; Small, “The Economic Museum";

Nash, “Interesting Plants in Iura in the Conservatories " . Britton

* Botanical Exploration of the Philippine Islands."

Notes. — Two interesting illustrated articles, by Massart, are pub-

lished in the September Buletin du Jardin Botanique de l État à

Bruxelles, respectively on how perennials maintain their subterranean

level, and how young leaves are protected against climatic excesses.

The second is illustrated by Ducos-du-Hauron plates in red and blue

which give a neutral tint stereoscopic effect when viewed through

accompanying red and blue spectacles.

An important paper by Reinke, on “ Botanisch-geologische Streif-

züge au den Küsten des Herzogtums Schleswig," is published as a

complementary heft to Vol. 8, N. F. of the Wissenschaftliche Meeresun-

tersuchungen published by the German Sea Commission of Kiel.

The first part of a study of the plankton of the Illinois River, by

Kofoid, published as Article 2 of Vol. 6 of the Bulletin of the Illinois

State Laboratory of Natural History, forms a quarto volume of xviii

+ 629 pages, with numerous diagrams and plates.

Adventive bud formation on leaves of Yucca gloriosa is noted by

Lutz in the Journal de Botanique of December.

Nasmith writes on the chemistry of wheat gluten in No. 4 of the

Physiological Series of University of Toronto Studies.

No. 449-] NOTES AND LITERATURE. 403

The association of lime-loving and lime-shunning plants is dis-

cussed by Aubert in No. 147 of the Buletin de la Société Vaudoise.

An account of the sex-conditions and hybrids in Ribes, by Janc-

zewski, is separately printed from the Buletin International de P Aca-

démie des Sciences de Cracovie, of December and January.

The rubber-fig of New Caledonia, Ficus Schlechteri, is described

and figured by Warburg in Der Tropenpflanzer, of December.

Cocoa: Its Production and Use, is the title of a paper by Marshall

in the American Journal of Pharmacy, for February.

An illustrated account of Yerba Mate — Zex Paraguayensis, by

Metzger, is contained in Der Tropenpflanzer, for January.

A popular article on California mistletoe, Phoradendron, with pho-

tograms showing its occurrence on various trees, is published by

Helen Lukens Jones in Out West for February.

An account of useful and interesting plants of the Congo region,

by de Wildeman, has been issued from the press of Spineux, of

Brussels.

A short sketch of the desert flora about Phoenix, Ariz., by Dams,

is published in the Monatsschrift fir Kakteenkunde, of January.

An account of the useful plants of the Sahara desert, by Diirkop,

forms the December number of Beihefte zum Tropenpflanzer.

An illustrated account of Samoa and its crops, by Wohltmann,

constitutes Heft 1-2 of the Beihefte zum Tropenpflanser of 1904.

A good illustration of a utilized school garden is contained in the

1902-3 Catalogue of the Boston Normal School.

A portrait of Professor Farlow forms the frontispiece to Zhe Popu-

lar Science Monthly of February.

A portrait of Professor Peck is published as frontispiece to Vol. 1

of the Ohio Mycological Bulletin.

CORRESPONDENCE.

Editor American Naturalist:

Dear Sir : — Prof. B. G. Wilder has called my attention to errors

in nomenclature and typography in my article “Oral Breathing

Valves in Teleosts," etc., in the American Naturalist for February.

The observations recorded in the paper were made two years ago in

his laboratory, under the immediate supervision of Instructor H. D.

eed. I supposed that acceptance of the results included sanction

of the names employed, whereas it appears that Prof. Wilder and

Dr. Reed intended only to commend the observations themselves,

and took for granted that, before publication, proof would be sub-

mitted to them. My removal to Louisiana rendered this inconven-

ient; I took for granted that either the manuscript or a proof would

be submitted to the ichthyological editor of the Matura/ist; finally,

my own revision had to be done hastily and without access to books.

T Ramvir their causes, the errors are deeply regretted; I realize

“the right of scientific names to be correctly written,” and avail

. myself gratefully of the opportunity to publish the seat ies list of

corrections.

E. G. MITCHELL.

Baton Rouge, La., May 28, 1904.

P sae I 53» line 5 for Macullum read Macallum.

* Amiurus “ Ameiurus.

* € E i4," Bupomotus “ Eupomotis.

" 154, lines 5 and 8, for drancheostegal read branchiostegal.

* € third line from bottom, for punctatus read gyrinus.

* * last “ for Eupomotus, read Eupomotis.

- 155 fane $ second m for Clupea, read Pomolobus.

: * fourth ^ Astrocopus, read Astroscopus.

“ 156 fifth line from bottom, and page 157, 11th line from

bottom, Amébloplytes gruniens, correctly spelled is Ambloplites grun-

niens, and should be Ap/odinotus grunniens.

405

406

CORRESPONDENCE. [Vor. XXXVIII.

Page 157, figure 7, for A. grunzens, read A. grunniens.

gulosus.

158, for y-groecum read y-graecum.

159, for Echensis naucratis, read Echeneis naucrates.

160, * predacious, read predaceous.

‘ee line 25, for Amiurus, read Ameiurus.

28, “ Noturus gyrinus, read Schilbeodes gyrinus.

a6, Noturus miurus; *"

[11 [11 [11

325

Schilbeodes miurus.

Amiurus nebulosis, read Ameiurus nebulosus.

sixth line from bottom, for Amiurus, read Ameiurus.

162, 14th “ for Opistheoma, read Opisthonema.

* 16th “ “ Clupea read Pomolobus.

* 18th “ * Clupea oestivalis, read Pomolobus aestivalis.

* 1igth * © PERCADAE, read PERCIDAE.

21st americanum, read flavescens.

* 26th “ | Etheostoma, “ Percina.

gist “ “ Ambloplytes, “ Ambloplites.

sixth line from bottom, for Eupomotus, read Eupomotis.

last line, for Afomotus, read Apomotis.

163, sixth line, for auritus, read aureus.

a Fath “ Chaenobrytes gulosis, read Chaenobryttus

[11

Page 163, 18th line Apomotus correctly spelled is Apomotis, and

should be Lepomis.

Page 163, line 26, for Eupomotus, read Eupomotis.

[11

164, " 2, * comparee, read comparée.

* lines 12 and 15, for Anatomie, read Anatomy.

(Wo. 448 was mailed June 16, 1904.)

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VOL. XXXVIII, NO. 450 JUNE, 1904

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NATURALIST

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I, Charles Emerson Beecher. .- pS ee on a R. T. JACKSON e

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THE

AMERICAN NATURALIST.

Vor. XXXVIII. June, 1904. No. 450.

CHARLES EMERSON BEECHER.

ROBERT T. JACKSON.

IN the death of Professor Beecher the science of Invertebrate

Palzeontology has lost a most brilliant and eminent leader. In

hislife so prematurely cut off, he attained a reputation for criti-

cal structural and developmental work on fossil animals and

philosophical work on the same lines that placed him in the

very front rank of his profession.

Charles Emerson Beecher, son of Moses and Emily Emerson

Beecher, was born in Dunkirk, New York, October 9th, 1856.

He died suddenly of heart disease on February 14th, 1904, in

the forty-eighth year of his age. Always a delicate man he had

recently been in exceptionally good health, so that his death

came without warning. On September 12th, 1894, Professor

Beecher married Miss Mary Salome Galligan, who with two

young daughters survive him. He left also his mother and a

brother, who reside in California.

In his early childhood the family removed to Warren, Penn-

sylvania, where he attended private and High schools. A born

naturalist and collector he in childhood began collecting fossils

from the Chemung and Waverly formations about Warren,

amassing a choice and extensive collection in that region,

407

408 THE AMERICAN NATURALIST. [Vor. XXXVIII.

especially valuable for the rare Phyllocarida and for Pelecypoda.

He also collected living molluscs and minerals and became well

versed in both lines of study. He took the science course

at the University of Michigan, receiving the degree of B. S. in

1878.

In 1878 Beecher became assistant to Professor James Hall at

Albany, where he remained for ten years. While at Albany he

acquired an intimate knowledge of palzeozoic fossils, so that with

his truly remarkable memory, it seemed that he could name at

sight any species offered to him. Always interested in micro-

scopical work he prepared some two hundred and fifty sections

of the shells of fossil brachiopods for study in connection with

the work of the New York Survey on that group. Besides

work on collections of fossils at Albany, he rendered important

assistance to Professor James Hall in the preparation of that

author'slarge quarto monographs on cephalous Mollusca,! Lam-

ellibranchiata,? corals and Bryozoa? As Beecher published

little systematic work in later life I would lay stress on what

he did as assistant to Professor Hall at this period. To quote

Professor Hall, as stated in the Prefaces of the several works

cited: “In the preparation and final revision of the descriptions

of the species of Cephalopoda I have been very ably assisted by

Mr. C. E. Beecher.” Again, “ In the final revision of the genera

and species [of Lamellibranchiata] and in the preparation of the

text and the later plates of this volume the author cheerfully

acknowledges the very valuable assistance rendered by Mr.

Charles E. Beecher.” In the second volume on the Lamelli-

branchiata Professor Hall says: “In the revision of the spe-

cies, and the publication of this volume, the author acknowl-

edges with great satisfaction the assistance rendered by Mr. C.

E. Beecher. In the last work cited on corals and Bryozoa Pro-

1 Natural History of New York. Paleontology. Vol. 5, Part 2, Gasteropoda,

Pteropoda, and Cephalopoda of the Upper Helderberg, es Portage and

Chemung Groups. Albany, 1879. p. 1—15 and 1-492, pl. 1-113.

? Ibid. Vol. 5, Part 1. Lamellibranchiata [of the e Ege Hamilton,

Portage and Chemung Groups.] 1, Albany. 1884. p. 1-18 and 1-268, pl. 1-33

and 81-92. e, Albany, 1885. p. 1-62 and 269—562, 7 figs. pl. 34-80 and 93

*/bid. Vol 6, Corals and Bryozoa.... from the Lower Helderberg, Upper

Helderberg and Hamilton Groups. AT Y, 1897. p. 1-26 and 1-298, pi. 1-66

No. 450.] CHARLES EMERSON BEECHER. 409

fessor Hall says: ‘We are ihdebted to Mr. Charles E. Beecher

for the preparation of the synopsis of the genera and sub-genera

described and illustrated in this volume ; and also for the expla-

nation of plates, with reference to the pages of the volume and

the localities of the species." Beecher also assisted Professor

Hall in the publication of other works as seen by the texts dur-

ing his Albany life.

This extensive experience in pure systematic work laid the

broadest foundation for Beecher’s later developmental and philo-

sophical studies, an experience that is markedly lacking or en-

tirely left out in many who enter into biological problems. It is

the expressed opinion of the present Palzontologist of New York

State that the influence of Beecher on the paleontology of the

State was for the best, and that during his stay of ten years he

elevated the standard greatly.

In holidays at Albany he made extensive collections of

Unionidze and other living molluscs in that vicinity. This col.

lection, with the large collections of Recent Mollusca that: he

gathered at Warren, Pennsylvania, also Michigan, and other local-

ities, previous to going to Albany, he presented to the New

York State Museum in 1886 and 1887. According to Mr. Wm.

B. Marshall! this collection included some twenty thousand

specimens and as many more which were considered duplicates,

and is almost entirely made up of land and fresh-water shells of

the United States, largely collected by Beecher himself. As

shown by collating two of Marshall's published papers,?? there

are in the Beecher collection at least one hundred and twenty-

one species of land and fresh-water Mollusca, mostly from New

York and Pennsylvania. One hundred and two localities are

represented, seventy of which are in New York State, the others

being in various parts of the United States.

Of the species some forty-three belonged to the Unionidze, a

! Marshall, Wm. B. List of Shells Inhabiting the Vicinity of Albany and Hag

N.Y. 48th Ann. Rept. N. Y. State Mus., Albany, 1895. Part 1, p. 641-64

? Marshall, Wm. B. Land and Fresh-water Shells of New York Gahan at

the World’s Columbian Exposition, Chicago, Illinois, 1903. 47th Ann. Rept. N.

Y. State Mus. Nat. Hist, Albany, 1894. p. 49-75-

? Marshall, Wm. B. The Geographical Distribution of New York Unionida.

48th Ann. Rept. N. Y. State Mus. Nat. Hist., Albany, 1895. Part 1, p. 45-99.

410 THE AMERICAN NATURALIST. (VoL: XXXVIII.

beautiful series of which is on exhibition in the Albany Muse-

um. The collection, which Marshall considered as exhaustive of

the vicinity of Albany, is not only rich in species, but very full

in series, representing specimens of the same species from

various localities and of ages from young to adult. It is there-

fore very valuable for studies of later development, variation

and geographical distribution. As a collection of fine selected

material it stands an excellent proof of Beecher's indefatigable

industry and appreciation of what a collection should be, much

of it, it should be remembered, gathered when he was a youth,

or even a child.

In the Albany Museum there are also many minerals and fos-

sils collected by him, and numerous specimens of fossils pre-

pared for exhibition. In free days at Albany he made frequent

visits to the rich localities in the Helderberg mountains near

Albany, especially Clarksville and Indian Ladder, collecting an

extensive series of the fossils to be found there.

At this period Beecher published independently a paper on the

Phyllocarida of Warren, Pennsylvania, a number of papers on

the lingual dentition of Gastropoda, besides others on recent and

fossil Pelecypoda, on geological localities, etc. His most impor-

tant work was the publishing with Prof. John M. Clarke of a

monograph on the development of Silurian Brachiopoda, based

on material washed from the clays of Waldron, Indiana. This

paper is of importance as the first work on the development of

fossil Brachiopoda and opened up new lines of inquiry previ-

ously untouched. Always skilful with his pencil, at Albany he

added to his income by making drawings for the medical men of

that city. Nearly if not all his own papers were illustrated by

his own drawings or photographs. During part of his stay at

Albany Beecher was Assistant in Palzeontology in the New York

State Museum, when he did much to develop that department.

After he left Albany he was retained for a period as Consulting

Palzontologist to the museum, visiting Albany at intervals.

Many details concerning his work and connection with the

museum will be found in the 32nd to 43rd Annual Reports

of the State Museum.

In 1888 Beecher went to New Haven as Assistant in Palaeon-

No. 450.] CHARLES EMERSON BEECHER. 4II

tology to take charge of the collections of invertebrate fossils in

the Peabody Museum. He took the degree of Ph. D. at Yale

in 1889, his thesis being a most interesting monograph on the

Brachiospongidz, of which rare fossil sponges the Yale Museum

has a superb series.

Professor Marsh recognized Beecher's ability and his advance-

ment was rapid. To state his official career at New Haven in

brief: He was appointed Instructor in Paleontology in 1892,

Assistant Professor of Palaeontology in 1892, Professor of His-

torical Geology 1897, and member of the Governing Board of

the Sheffield Scientific School in the same year. On the death

of Professor Marsh he was in 1899 appointed Curator of the

geological collections of the Peabody Museum. In 1902 his

title was changed to that of University Professor of Palaeon-

tology.

In 1898 Beecher was elected corresponding member of the

Boston Society of Natural History, in 1899 member of the

National Academy of Sciences, and foreign correspondent of the

Geological Society of London. In 1900 he was elected Presi-

dent of the Connecticut Academy of Arts and Sciences, filling

that office for two years. While at New Haven he made many

trips to western or nearer localities, collecting vertebrate and

invertebrate fossils for the museum, and made a trip to Europe

with the late Dr. George Baur, visiting museums.

Indian Ladder in the Helderberg Mountains was always a

favorite and fertile spot for him, dating back to his Albany days.

It is one of the most beautiful and picturesque regions in the

Helderbergs. He collected there slabs of limestone containing

fossils which were silicified in the most perfect condition for

development by etching with acid. From such material he

etched numerous large or minute specimens of surpassing beauty

and scientific interest. Besides adult fronds of Bryozoa, Brachi-

opoda, Crustacea, and other fossils in most perfect preservation,

he obtained minute embryos and small species in large numbers

in literally wonderful condition of perfection. Young Bilobites

i mm. in length, young Acidaspis .93 mm. in length, and Arges

1.15 mm. long, both of the latter so perfect that he figured

them from both the dorsal and ventral view ; young Pleurodic-

412 THE AMERICAN NATURALIST. [Vor. XXXVIII.

tyum, consisting of the initial cup alone, and also others with first

lateral buds, young Bryozoa showing initial chambers. Such

material selected with infinite care and patience formed the basis

of a number of papers by Beecher and others. He etched also

some very choice fossils from the Hamilton of Canandaigua

Lake, New York. One sees and hears so much of poorly pre-

served fossils, that such exceptionally fine material is refreshing.

He made a special point of seeking small and embryonic material

by sifting clay from fossiliferous regions with a stream of run-

ning water. In this way he obtained choice material from Wal-

dron, Indiana. ;

In June, 1899, Beecher gave his large and exceedingly valuable

collections’ to the Peabody Museum as he said, “in grateful

recognition of the honors and favors conferred upon me during

my connection with the University.”! These collections, made

previous to his New Haven appointment, represent the result

of twenty years' labor, they were collected wholly by himself

and comprised upwards of one hundred thousand specimens,

mostly from the Devonian and Lower Carboniferous of New

York and Pennsylvania. They contained about five hundred

type specimens, published in the Paleontology of New York,

Geological Survey of Pennsylvania, and various scientific peri-

odicals. They also contained hundreds of specimens represent-

ing series in development, rare species, and choice specimens

exquisitely prepared to show structural detail.

His specimens were always fully labeled as regards locality

and identification of genus and species, so that his collections of

land and fresh-water molluscs previously mentioned and his col-

lections of fossils represent an immense amount of labor and

experience in the field and also in systematic zodlogy and palzeo-

zoology in working up the collections, and this all as a side-

show to his regular work in official positions.

In his bachelor days at New Haven Beecher with Pirsson,

Penfield and Wells roomed in “the attic,” the top story of the

Sheffield Scientific School, which was comfortably fitted up in

true Bohemian style. One of the pleasantest recollections of

1 Science, N. S. Vol. 10. p.61. July 14, 1899.

No. 450.] CHARLES EMERSON BEECHER. 413

visits to New Haven is visits to “the attic" where, after work

hours, delightful intercourse, social and scientific was held, often

far into the night.

. Already interested in studies of the development of organisms

from his work noted on the development of Silurian Brachiopoda,

in 1889 Beecher became deeply interested in the late Professor

Hyatt's methods of work. The application of the principles of

stages in development, acceleration, parallelism, and dynamic

genesis to the unraveling of the genealogical relations of living

and fossil animals. Bringing to this work his large and intimate

knowledge of species and the structure of fossil types, Beecher

entered into this field with characteristic energy and became the

leader of the Hyatt School. Beecher's reputation as an investi-

gator will rest chiefly on the rich results he obtained in the

critical, painstaking application of these fruitful principles that

Professor Hyatt labored so long to establish. |

Already interested in Brachiopoda, Beecher’s first application

of the Hyatt methods was to this group. Combining the study

of young, adult, living and fossil types, he worked out a classifi-

cation based on the principles of development, and divided the

class into four orders. He pointed out the primitive Iphidea

from the Lower Cambrian as the archaic radical from which the

whole class could be derived. This stands to-day as perhaps the

only example in which the primitive radical of a great group

. equal in systematic value to the Brachiopoda can be pointed out

with reasonable assurance. Besides his work on the class as a

whole, Beecher wrote many papers in which he worked out the

development, structural and genealogical relations of families or

minor groups of Brachiopoda. Such are his studies of the

development of Bilobites, Terebratalia, Zygospira, and his

remarkable studies of loop-bearing Brachiopoda. It is not too

much to say that a careful study of Beecher's papers on brachio-

pods, and the same may be said of his trilobites, will give the

student a more comprehensive view of the class than any other

published source. Incidentally to his work on. the group,

Beecher accumulated a rich collection of recent species of

Brachiopoda in the Yale museum, illustrating the structure and

development, often by large series, in seventy-five species and

414 THE AMERICAN NATURALIST. (Vor. XXXVIII.

five varieties, out of the total known number of one hundred

and thirty-eight species and nine varieties of living brachiopods.

This statement is taken from a list kindly sent me by Miss

Lucy P. Bush, his assistant, who prepared it recently at Pro-

fessor Beecher's request.

In corals he made important contributions by his beautiful

studies of the development of Pleurodictyum and also the struc-

ture and development of the zodids in the colonial form in

Michelinia, Favosites and recently in Romingeria.

Thoroughly established in his adopted principles of research, in

1893 he began the publication of his brilliant papers on the struc-

ture, development and classification of the Trilobita. Favored

by the discovery by W. S. Valiant of Triarthrus in unusual con-

dition of preservation at Rome, New York, Beecher entered into

the work. The presence of antennz had been announced by

W. D. Matthew ; but Beecher with his marvelous mechanical

skill and untiring patience worked out the structure of antennae,

legs and other ventral appendages with a minuteness that had

previously been impossible on any known material. His studies

of this type made our knowledge of the Trilobita as a classa new

thing, putting them on a basis for proper comparative study with

other Crustacea. He also took up studies of the development

of Trilobita, describing the development in genera in which it

was previously unknown or partially known. In 1897 Beecher

presented a classification of Trilobita based on his critical studies

of young and adult structures together with a consideration of

the geological or time sequence, ʻa natural basis for classification

as urged by Louis Agassiz and Alpheus Hyatt and nowhere

more beautifully carried out than in these trilobite studies. The

Trilobita were divided into three orders based principally on the

development of the free cheeks. At the time of his death he

was at work on an extensive monograph on the structure of

trilobites.

In all of Beecher's later work a strong philosophical bent was

evinced. This was given full expression in his charming and

forceful papers on the origin and significance of spines. In

them he urged that spinosity is an expression of growth-force

and differential development. The spinose individual or group

No. 450.] CHARLES EMERSON BEECHER. 415

of organisms begins as a smooth form, then becomes spinose in

a progressively increasing degree until the acme or most elab-

orate spinosity is attained, then in extreme decadence there is a

loss and final disappearance of spinosity, the individual and the

group in senility tending to become smooth as in its own early

growth. It is a graphic expression of the value of a new con-

ception applied to well known facts.

The student of paleontology has reason to be grateful that

many of Beecher's more important papers were combined and

published in one volume, Studies in Evolution, in the Yale Bicen-

tennial Series, 1901. It is a model of what minute, critical,

philosophical paleeozoólogical work should be. With character-

istic modesty Beecher deplored as extravagance the republication

of papers already in print. For the student it is a valuable aid

that Beecher's views on Brachiopoda and Trilobita were incor-

porated in Eastman's translation. of Zittel’s Grundzüge der

Paleontologie. The chapter on Brachiopoda being revised by

Beecher's intimate friend, Mr. Charles Schuchert, and that on

Trilobita being revised by Professor Beecher himself.

Beecher published many other papers besides those alluded to,

on Gastropoda, Cephalopoda, and Crustacea, but although

interested in stratigraphical and descriptive palzeontology he pub-

lished little in these lines. He published nothing outside of

scientific papers as I am informed by Miss Lucy P. Bush. Ina

review of his papers I find 7 new orders, 1 new family, 2 new

subfamilies, 7 new genera, and 20 new species. Most of his

papers were brief, that on spinosity being the longest, many of

them were combined however in his large work, Studies in

Evolution. His bibliography includes some one hundred and

eight titles. In addition he wrote a paper, “ Extinction of

Species,” now in press in the Encyclopedia Americana, vol. 4.

He was for several years an associate editor of this journal.

Being skilled with his hands, ingenious and fertile in mechani-

cal resources, Beecher was an exceptional preparator, as evinced

by his remarkable preparations of brachiopods and trilobites,

and numerous specimens prepared for exhibition at the Yale

University Museum. Especially noteworthy in this direction

are the splendid slabs of Crawfordsville crinoids and Cretaceous

416 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Uintacrinus, the huge mount of Brontosaurus and the gigantic

yet life-like restoration of Claosaurus, besides numerous indi-

vidual specimens of invertebrates in all groups. Beecher was a

skilful photographer and Professor Hall published! his photo-

micrographs of brachiopod shell structure from slides also pre-

pared by him. Later he published himself photomicrographs of

the appendages of Triarthrus, exceptionally difficult subjects.

He was a skilful microscopist and made numerous preparations

of recent and fossil animals, especially noteworthy are his beau-

tiful preparations of the radulz of gastropods, sections of shells

of brachiopods, sections of corals, sponges, etc. His microscope

slides were always the perfection of neatness in finish. One

method he adopted is very valuable for appearance and conven-

ience. When sections had to be ground thin he did this on a

ground glass slide, when the cover glass was in place the balsam

filled the ground surface rendering it transparent, the ground

glass forming an attractive border to the slide. This method

obviates the necessity of transferring sections to a fresh slide,

which is usually done at some risk, for the sake of appearances.

He took great interest in preparing models to illustrate struc-

ture, as shown in his models of the development and adult charac-

ters of loop-bearing types of brachiopods, his models of Triarthrus

showing structure, from the dorsal and ventral view, and his

restoration of the huge Devonian crustacean Stylonurus, measur-

ing five feet in length. The details of museum technique also

appealed to him strongly, so that he was a most able and success-

ful museum administrator, constantly devising methods to

improve the condition and accessibility of collections under his

charge. The exhibition rooms of the Yale Museum are a

graphic expression of his skill in selecting, preparing and exhibit-

ing collections of fossils.

Beecher was fond of using mechanic's tools and as a relaxa-

tion did carpentering and cabinet work at the Museum or at his

home. Full of resources he had many interests outside of his

technical scientific work.

1 On the structure of the shell in the Genus Orthis, by James Hall. 376th Ann.

Rep't. N. V. State Mus. Nat. Hist. 1884. p. 73-75, pl. 3-4-

No. 450] CHARLES EMERSON BEECHER. 417

Quiet, unassuming, modest in a very marked degree, simple,

without affectation, entirely free from all eccentricities, conscien-

tious and painstaking in every thing he had to do. In the

words of Professor Chittenden, Director of the Sheffield Scien-

tific School! *....to those who knew Professor Beecher inti-

mately no words of appreciation will be deemed too extravagant,

for close association only brought more clearly to view the

many mental traits that testified to the strength of character

and of mind that helped to make Professor Beecher one of the

strong men of the Scientific School."

Beecher was eminently successful as a teacher, as evinced by

the devotion of his pupils and the able papers produced by stu-

dents under his charge.

Always ready to help with advice, or specimens, he was an

appreciative audience, a helpful critic, a warm friend with keen

interest in his friends and their work. He will be deeply

missed as a friend, and his untimely death deplored as a loss to

the science in which he made such a brilliant mark.?

, 1 Professor Charles E. Beecher. His Life and Work Reviewed [by Miss Lucy

P. Bush, not signed]. — Prof. Chittenden's appreciation — Mr. Schuchert's Sketch.

[Tribute from a pupil] Yale Alumni Weekly. New Haven, March 2, 1904.

Vol. 13, no. 22, p. 487-489.

? The following obituary notices of Professor Beecher have also appeared.

Obituary notices were published in the following newspaper

New Haven Morning Journal and Courier, Feb. 15, 1904. p. T:

New Haven Palladium, Feb. 15, 1904. p. 1-2.

New Haven Evening Register, Feb. 15, 1904. p. 2.

Boston Evening Transcript, Feb. 15, 1904. p. II.

New Haven Evening Register, Feb. 16, 1904. p. 2.

New Haven Morning Journal and Courier, Feb. 17, 1904

American Journal Science, March, 1904. Ser. 4, vol. 17, p. 252.

Charles Emerson Beecher. Amer. Geol., March, 1904. Vol. 33, p- 189.

Charles Emerson Beecher, by W. H. Dall. Science, March 18, 1904. N.S.

vol. 19, p. 453-455.

Obituary — Charles Emerson Beecher. Museum's Journal [London], April,

1904. Vol. 3, p. 339-340

Charles Emerson Boscha by Charles Schuchert. Amer. Jour. Sci., June, 1904.

Ser. 4, vol. 17, p. 411-422, portrait as frontispiece.

Professor Charles Emerson Beecher, Ph. D., [by H. Woodward, not signed].

Geol. Mag. [London], June, 1904. N.S. dec. 5, vol. 1, p. 284-286, portrait pl. 10.

Charles Emerson Beecher, by John M. UNE Amer. Geol., July, 1904. Vol.

34, p. I-13, portrait as pl 1.

418 THE AMERICAN NATURALIST. (VoL. XXXVIII.

THE BIBLIOGRAPHY OF CHARLES EMERSON BEECHER

1876. 1. List of Land and Fresh-water Shells Found within a Circuit of

Four Miles about Ann Arbor, Mich., [by Bryant Walker and C. E.

Beecher]. Proc. Ann Arbor Sci. Assoc. 1876. p. 43-46.

1884. 2. Ceratiocaridz from the Chemung and Waverly Groups of Penn-

sylvania. 2nd Geol. Surv. Penn. Rept. PPP. 1884. p. I-22, fig. 1,

pl. 1-2.

3. Some Abnormal and Pathologic Forms of Fresh-water Shells from

the Vicinity of Albany, New York. 36th Ann. Rept. N. Y. State Mus.

Nat. Hist. 1884. p. 51-55, pl. 1-2.

4. [The plates in James Hall’s] On the Structure of the Shell in the

Genus Orthis. 36th Ann. Rept. N. Y. State Mus. Nat. Hist. 1884.

P- 73-75, pl. 3-4.

5. List of Species of Fossils from an Exposure of the Utica Slate and

Associated Rocks, within the Limits of the City of Albany. 36th Ann.

Rept. N. Y. State Mus. Nat. Hist. 1884. p. 78.

6. A New Design for a Microscope Cabinet. Amer. Month. Mic.

Jour. July, 1884. Vol. 5, p. 126—127, fig. 23.

7. Notes on a Nevada Shell (Pyrgula nevadensis), [by R. Ellsworth

Call and C. E. Beecher]. Amer. Nat. Sept. 1884. Vol. 18, p. 851-

855, fig. 6 and fig. 1—5, pl. 25.

[Reprinted in part in R. Ellsworth Call and Harry E. Pilsbry's] On

Pyrgulopsis, a New Genus of Rissoid Mollusk, with Descriptions of

two new forms. Proc. Davenport Acad. Nat. Sci. April, 1886.

Vol. 5, p. 9-14, fig. 1-5.

1885. 8. Carnivorous Habits of the Muskrat. Science. Feb. 20, 1885.

Vol. 5, p. 144-145.

1886. 9. Field Notes on the Geology of the Mohawk Valley, with a Map

[by C. E. Beecher with C. E. Hall]. Fifth Ann. Rept. State Geologist

[New York]. Albany, 1886. p. 8-10, 3 figs. [p. 4 for evidence of

authorship].

[Reprinted, without map.] 48th Ann. Rept. N. Y. State Mus. Nat.

Hist. Albany, 1895. Part 2, P. 54—56, 3 figs.

10. Note on the Oneonta Sandstone in the Vicinity of Oxford, Che-

nango County, N. Y. [by C. E. Beecher, with J. W. Hall and C. E.

! An effort has been made to render this bibliography as complete as possible.

Obligations are due to Miss Lucy P. Bush, Professor Beecher's assistant at New

Haven, for a number of titles that would otherwise have been overlooked. All

titles have been verified.

No. 450.] CHARLES EMERSON BEECHER. 419

Hall. Fifth Ann. Rept. State Geologist [New York]. Albany, 1886.

P- 11, 1 fig. [p. 4 for evidence of authorship].

11. A Spiral Bivalve Shell from the Waverly Group of Pennsylvania.

39th Ann. Rept. N. Y. State Mus. Nat. Hist. 1886. p. 161—164, pl. 12.

12. [Lingual Dentition in Amnicola dalli, in R. Ellsworth Call's] On

Certain Recent, Quatenary, and New Fresh-water Mollusca. Proc.

Davenport Acad. Nat. Sci. April, 1886: Vol. 5, p. 1-8, fig. 1-4.

[Beecher's portion, p. 2-3, fig. 1—2.]

13. Description of a New Rissoid Mollusk [by R. Ellsworth Call and

C. E. Beecher}. Bull. Washburn Col. Lab. Nat. Hist. Dec., 1886.

Vol. 1, p. 190-192, fig. 4.

1888. 14. A Method of Preparing for Study the Radule of Small Spe-

cies of Gasteropoda. Jour. N. Y. Mic. Soc. Jan., 1888. Vol. 4, p.

-IL —

[Noticed in] Jour. Royal. Mic. Soc. [London]. June, 1888. Part 1, p.

507—508.

15. Natural History of New York. Paleontology. Vol. 7, with Sup-

plement to Vol. 5, part 2; by James Hall. [Review.] Amer. Jour.

Sci. June, 1888. Ser. 3, vol. 35, p. 499-500.

16. Synoptical Table of the Genera and Species [of corals and Bryo-

zoa) described in Vol. 6 of the Palzontology of New York. 41st Ann.

Rept. N. Y. State Mus. Nat. Hist. 1888. p. 363-375.

d 43rd Ann. Rept. N. Y. State Mus. Nat. Hist. "GRIS

1890. p. 279-291.

17. Statement of the Condition of the Work on the Brachiopoda.

Paleontology of New York. Vol. 8, 41st Ann. Rept. State Mus. Nat.

Hist. 1888. p. 383-384.

[Reprinted.] 43rd Ann. Rept. N. Y. State Mus. Nat. Hist. Albany,

1890, p. 299-300.

18. Lamellibranchiata.* List of Genera Illustrated on the Plates of the

Report of the State Geologist for 1882 [with] List of Genera not Illus-

trated on the Plates of the Geologist's Report of 1882 [etc]. 41st

Ann. Rept. N. Y. State Mus. Nat. Hist. 1888. p. 385-387.

[Reprinted.] 43rd Ann. Rept. N. Y. State Mus. Nat. Hist. Albany,

1890. p. 301-303.

1889. 19. Brachiospongide : a Memoir on a Group of Silurian Sponges.

Mem. Peabody Mus. Yale Univ. 1889. Vol. 2, part 1, 4to, p. 1-28,

fig. 1-4, pl. 1—6.

20. Note on the fossil Spider Arthrolycosa antiqua Harger. Amer.

Jour. Sci. Sept, 1889. Ser. 3, vol. 38, p. 219-223, fig. 1-3.

420 THE AMERICAN NATURALIST. [Vor. XXXVIII.

21. The Development of Some Silurian Brachiopoda, [by Charles E.

Beecher and John M. Clarke]. Mem. N. Y. State Mus. Oct, 1889.

Vol. 1, no. 1, 4to, p. 1—95, fig. 1—4, pl. 1-8. :

[Reprinted.] Studies in Evolution, 1901, p. 310-398, fig. 126—130, pl.

1890. 22. On the Lingual dentition and Systematic position of Pyrgula.

Jour. N. Y. Mic. Soc. Jan, 1890. Vol. 6, p. 1-3, pl. 21.

23. On the Development of the Shell in the genus Tornoceras,

Hyatt. Amer. Jour. Sci. July, 1890. Ser. 3, vol. 40, p. 71-75, pl. 1.

[Reprinted.] Studies in Evolution. 190o1. p. 435-440, pl. 34.

24. Koninckina and related Genera. Amer. Jour. Sci. Sept., 1890.

Ser. 5, vol. 4o, p. 211—219, fig. 1, pl. 2.

25. On Leptznisca a new genus of Brachiopod from the Lower Helder-

berg group. Amer. Jour. Sci. Sept, 1890. Ser. 3, vol. 40, p. 238-240

and 245, pl. 9.

26. North American species of Strophalosia. Amer. Jour. Sci. Sept.,

1890. Ser. 3, vol. 40, p. 240-246, pl. 9.

27. Phylogeny of the Pelecypoda, the Aviculidz and their Allies, by

Robert Tracy Jackson. [Review.] Amer. Jour. Sci. Nov., 1890. Ser.

3, vol. 40, p. 421—422.

28. Contributions to the Tertiary Fauna of Florida, by Wm. H. Dall.

[Review.] Amer. Jour. Sci. Nov., 189o. Ser. 3, vol. 40, p. 423.

29. On Syringothyris Winchell, and its American species, by Charles

Schuchert. [Review.] Amer. Jour. Sci. Nov.189o. Ser.3, vol. 40,

P. 423.

1891. 30. (1) New Types of Carboniferous Cockroaches from the Car-

Pedipalpi; (4) The Insects of the Triassic beds at Fairplay, Colo-

rado; by Samuel H. Scudder. [Review.] Amer. Jour. Sci. Jan.

1891. Ser. 3, vol. 41, p. 72.

31. Bulletin from the Laboratories of Natural History of the State

University of Iowa. Vol. 2, no. 1, p. 1-98, pl. 10-12, 1890. [Review.]

Amer. Jour. Sci. Jan. 1891. Ser. 3, vol. 41, p. 72.

32. Insecta, by Alpheus Hyatt and J. M. Arms. [Review.] Amer.

Jour. Sci. March, 1891. Ser. 3, vol. 41, p. 256-257.

33. James Croll [Obituary]. Amer. Jour. Sci. March, 1891. Ser. 3,

vol. 41, p. 258. [Not signed.] ;

. No. 450.) CHARLES EMERSON BEECHER. 421

34. Development of the Brachiopoda. Part 1. Introduction. Amer.

Jour. Sci. April, 1891. Ser. 3, vol. 41, p. 343-357, pl. 17. [For Parts

2 and 3 see numbers 42 and 74 this list.]

[German translation by Moritz Fischer.] Neues Jahrb. Mineral. Geol.

und Paleontol. 1892. 1 Bd. 3 Heft, p. 178-197, taf. 6.

[Reprinted.] Studies in Evolution. 1901. p. 229-246, pl. 11.

35. Catalogue of the Fossil Cephalopods in the British Museum, part

3, by Arthur H. Foord. [Review.] Amer. Jour. Sci. May, 189r.

Ser. 3, vol. 41, p. 438.

36. The Tertiary Insects of. North America, by Samuel H. Scudder.

[Review.] Amer. Jour. Sci. June, 189r. Ser. 3, vol. 41, p. 517.

37. Development of Bilobites. Amer. Jour. Sci. July, 1891. Ser.

3, vol. 42, p. 51—56, fig. 1—2, pl. 1.

[Reprinted.] Studies in Evolution. 1901. p. 399-495, fig. 131-132,

B 1.

38. The Development of a Paleozoic Poriferous Coral. Trans. Conn.

Acad. Arts and Sci. July, 1891. Vol. 8, p. 207-214, pl. 9-13.

[Reprinted.] Studies in Evolution. 1901. p. 421-428, pl. 27-31.

39. Symmetrical cell Development in the Favositide. Trans. Conn.

Acad. Arts and Sci. July, 1891. Vol. 8, p. 215-219, pl. 14-15.

[Reprinted.] Studies in Evolution. 1901. p. 429-434 pl. 32-33.

1892. 40. (1) Notes on the Genus Acidaspis; (2) Note on Coronura

aspectans Conrad (Sp. the Asaphus diurus Green ; (3) Observations

on the Terataspis grandis Hall, the largest known Trilobite; by

J. M. Clarke. [Review.] Amer. Jour. Sci. Feb, 1892. Ser. 3, vol.

43, p. 158—159.

41. On the Occurrence of Upper Silurian Strata near Penobscot

Bay, Maine (with map). [By William W. Dodge and Charles E.

Beecher.] Amer. Jour. Sci. May, 1892. Ser. 3, vol. 43, p- 412-418.

[416-418 by C. E. B.]

42. Development of the Brachiopoda. Part 2. Classification of the

Stages of Growth and Decline. Amer. Jour. Sci. Aug, 1892. Ser.

3, vol. 44, p. 133-155, fig. 1-36, pl. 1. [For Parts 1 and 3 see num-

bers 34 and 74 this list.] '

[Reprinted.] Studies in Evolution. 1901. p. 246-273. fig. 85-120, pl.

i2.

of the State of New York. Palaontology.

Vol.8. An Introduction to the Study of the genera of Palaozoic

Brachiopoda. Partir; by James Hall, assisted by John M. Clarke.

[Review.] Amer. Jour. Sci. Oct, 1892. Ser. 3, vol. 44, p. 330-332-

n Columbia County, New

43. Geological Survey

44. Notice of a new Lower Oriskany Fauna i

422 THE AMERICAN NATURALIST. (VoL: XXXVIII.

York; by C. E. Beecher. With an annotated list of fossils ; by J.

M. Clarke. Amer. Jour. Sci. Nov., 1892. Ser. 3, vol. 44, p. 410—414.

[p. 410-411 by C. E. B.]

1893. 45. [Bibliography of] Charles E. Beecher. Bibliographies of the

Present Officers of Yale University, New Haven. 1893. p. 19-20.

46. Revision of the Families of Loop-bearing Brachiopoda. Trans.

Conn. Acad. Arts and Sci. March, 1893. Vol. 9, p. 376-391, and 395-

398, pl. 1-2.

[Reprinted.] Studies in Evolution. 1901. p. 290-309, pl. 14 and 24.

47. The development of Terebratalia obsoleta Dall. Trans. Conn.

Acad. Arts and Sci. March, 1893. Vol. 9, p. 392-395, and 398-399,

[Reprinted.] Studies in Evolution. 1901. p. 406—409, pl. 24-25.

48. Some Correlations of Ontogeny and Phylogeny in the Brachiopoda.

Amer. Nat. July, 1893. Vol. 27, p. 599-604, pl. 15.

[Reprinted.] ‘Studies in Evolution. 1901. p. 286-289, pl. 13.

49. Development of the Brachial Supports in Dielasma and Zygo-

spira; [by Charles E. Beecher and Charles Schuchert.] Proc. Biol.

Soc. Washington. July, 1893. Vol. 8, p. 71-78, pl. 10.

[Reprinted.] Studies in Evolution. 1951. p. 410-417, pl. 26.

50. On Urnatella gracilis, by C. B. Davenport. [Review.] Amer.

Jour. Sci. July, 1893. Ser. 3, vol. 46, p. 75.

51. Larval forms of Trilobites from the Lower Helderberg Group.

Amer. Jour. Sci. Aug, 1893. Ser. 3, vol. 46, p. 142-147, pl. 2.

52. A Larval Form of Triarthrus. Amer. Jour. Sci. Nov.1893. Ser.

3, vol. 46, p. 378-379, fig. 1. [Pagination of reprint 361—362.]

53. On the Thoracic Legs of Triarthrus. Amer. Jour. Sci. Dec.

1893. Ser. 3, vol. 46, p. 467—470, fig. 1-3. :

1894. 54. On the Mode of Occurrence and the Structure and Development

of Triarthrus beckii. Amer. Geol. Jan. 1894. Vol. 13, p. 38-43, pl. 3-

[Reprinted.] Studies in Evolution. 1901. p. 197-202, pl. 6.

55. The Appendages of the Pygidium of Triarthrus. Amer. Jour.

Sci. April, 1894. Ser. 3, vol. 47, p. 298—300, fig. 1, pl. 7.

56. [Review of Hall and Clarke’s Paleozoic Brachiopoda, Palzontology

of New York. Vol. 8] Amer. Jour. Sci. April, 1894. Ser. 3, vol.

47; P- 319.

1895. 57. Further Observations on the Ventral Structure of Triarthrus.

Amer. Geol. Feb., 1895. Vol. 15, p. 91-100, pl. 4-5.

[Reprinted.] Studies i in Evolution. 1901. p. 203-212, pl. 7-8.

No. 450.] CHARLES EMERSON BBECHER. 423

58. Structure and Appendages of Trinucleus. Amer. Jour. Sci.

April, 1895. Ser. 3, vol. 49, p. 307-311, pl. 3.

[Reprinted.] Studies in Evolution. 19ot. p. 220-225, pl. 10.

59. The Larval Stages of Trilobites. Amer. Geol. Sept., 1895.

Vol. 16, p. 166—197, fig. 1-8, pl. 8-10.

[Reprinted.] Studiesin Evolution. 1gol. p. 166—196, fig. 76-84, pl.

3-5.

60. Phylogeny of an Acquired Characteristic; by Alpheus Hyatt.

[Review.] Amer. Geol. Oct., 1895. Vol. 16, p. 256-259.

1896. 61. James Dwight Dana. [Obituary.] Amer. Geol. Jan., 1896.

Vol. 17, p. 1—16, portrait, pl. 1.

62. The Morphology of Ru ro Amer. Jour. Sci. April, 1896.

Ser. 4, vol. 1, p. 251-256, p

[Reprinted.] Geol. AM nu May, 1896. N. S. Dec. 4.

Vol. 3, p. 193-197, p

[Reprinted.] e in Evolution. 1901. p. 213-219, pl.

[Extract under title] Respiration of Trilobites. Amer. Nat. May,

1896. Vol. 30, p. 409 [not signed].

63. On the Supposed Discovery of the Antenna of Trilobites by

Linnzus in 1759. Amer. Geol May, 1896. Vol. 17, p. 303-306,

fig. 1-3. |

64. The Comparative Morphology of the Galeodide, by H. M.

Bernard. [Review.] Amer. Jour. Sci. June, 1896. Ser. 4, vol. 1, p.

491—492.

65. On the Validity of the Family Bohemillidz, Barrande. Amer.

Geol June, 1896. Vol. 17, p. 360-362, fig. 1-3.

66. An Ancient Pothole. New Haven Evening Register. June

22, 1896. [Not signed.]

67. On the Occurrence of Silurian Strata in the Big Horn Mountains,

Wyoming, and in the Black Hills, South Dakota. Amer. Geol July,

1896. Vol. 18, p. 31-33.

68. The Dinosaurs of North America, by O. C. Marsh. [Review.]

Amer. Jour. Sci. Dec., 1896. Ser. 4, vol. 2, p. 458-459.

1897. 69. Outline of a Natural Classification of Trilobites. Amer. Jour.

Sci. Feb.and March, 1897. Ser. 4, vol. 3, p. 89-106 and 181-207,

pl. 3.

[Reprinted.] Studies in Evolution. Igol. p. 109-162, fig. 75, pl. 2.

70. The Geological d Natural History Survey of Minnesota. Vol.

3, part 2 of the final Report, by N. H. Winchell [and others].

[Review.] Science. March 12, 1897. N.S. vol. 5, p. 449-

424 THE AMERICAN NATURALIST. [Vor. XXXVIII.

71. Vertebrate Fossils of the Denver Basin, by O. C. Marsh.

Extract from Monographs U. S. Geol. Survey. NES Amer.

Jour. Sci. April, 1897. Ser. 4, vol. 3, p. 349.

72. Models of Trilobites. Trans. N. Y. Acad. Sci., 1896-1897, vol.

16, 1898. Fourth Annual Reception, April 5-6, 1897, p. 43-44.

[Not signed.]

73. The Systematic Position of the Trilobites, by J. S. Kingsley,

Tufts college; with Remarks [p. 38-40] by C. E. Beecher. Amer.

Geol. July, 1897. Vol. 20, p. 33-40.

[Remarks reprinted under title] The ipi Position of the

Trilobites. Studies in Evolution. 1901. p. 163-1

74. [Development of the Brachiopoda, Part 3]. Morphology of the

Brachia. [In Charles Schuchert's Synopsis of American Fossil

Brachiopoda]. Bull. 87, U.S. Geol. Surv. 1897. Chap. 4, p. 105-112,

fig. 2-6. [For Parts 1 and 2 see numbers 34 and 42 this list.]

[Reprinted.] Studiesin Evolution. 1901. p. 274-285, fig. 121-125.

75. Schuchert's Synopsis of American Fossil Brachiopoda. [Review.]

Amer. Nat. Dec., 1897. Vol. 31, p. 1053-1055.

1898. 76. Catalogue of the Tertiary Mollusca in the Department of

Geology, British Museum (Nat. Hist) Pt. 1. The Australian Tertiary

Mollusca; by George F. Harris. [Review.] Amer. Nat. Feb., 1898.

Vol. 32, p. 134-135.

77. Polish Paleozoics; by Gurich. [Review.] Amer. Nat. April,

1898. Vol. 32, p. 298.

78. The Origin and Significance of Spines: A Study in Evolution.

Am. Jour. Sci. July to Oct, 1898. Ser. 4, vol. 6, p. 1-20, 125-136,

249—268, 329-359, fig. 1-73, pl. I.

[Reprinted.] Studies in Evolution. 1901. p. 3-105, fig. 1-74, pl. 1-

1899. 79. The Cretaceous Foraminifera of New Jersey, by R. M. Bagg

Jr. [Review.] Am. Geol Feb., 1899. Vol. 23, p. 126.

80. Othniel Charles Marsh. [Obituary.] Amer. Jour. Sci. June,

gg Ser. 4, vol. 7, p. 403-428. Portrait.

[Letter.] To the Corporation of Yale University : [concerning Pro-

us Beecher’s Gift to Yale University]. Science. July 14, 1899.

N. S. vol. 10, p. 6r.

82. Othniel Charles Marsh. [Obituary] Amer. Geol. Sept, 1899.

Vol. 24, p. 135-157. Portraits as pl. 5-6. [Abridged with alterations

from Amer. Jour. Sci. June, 1899. Title 80.]

1900. 83. Othniel Charles Marsh as an Ornithologist. The Osprey. Jan.,

1900. Vol. 4, p. 74-76. Portrait.

84. Trilobita. Zittel's Text.book of Palaontology, edited by Eastman.

No. 450.] CHARLES EMERSON BEECHER. 425

MacMillan and Co., London and New York. Separate, dated Jan.,

1900. p. 607-638, fig. 1261-1331.

85. Conrad's Types of Syrian Fossils. Amer. Jour. Sci. March,

1900. Ser. 4, vol. 9, p. 176—178.

86. On a large slab of Uintacrinus from Kansas. Amer. Jour. Sci.

April, 1900. Ser. 4, vol. 9, p. 267-268, pl. 3-4.

87. [Review of Geology of the Yellowstone National Park, by Hague,

Iddings and others, Monograph, 32, U. S. Geol. Survey.] Amer. Jour.

Sci. April, 1900. Ser.4, vol. 9, p. 297-300. [Signed L. V. P. and

CE. B.]

88. Restoration of Selmi lacoanus, a Giant Arthropod from

the upper Devonian of the United States. Amer. Jour. Sci.

I90O. Ser. 4, vol. 10, p. 145-150, pl. 1

[Reprinted.] Geol. Mag. [London]. Nov., 1900. N.S.Dec.4. Vol. 7,

p. 481-485, pl. 18.

89. Zittel’s Textbook, [of Pylssoido. translated and edited by

. R. Eastman. Review]. Amer. Nat Aug., 1900. Vol. 34, p.

681-682. |

90. Othniel Charles Marsh. [Obituary.] Bull. Geol. Soc. Amer. Oct.,

1900. Vol. 11, p. 521-537. [Abridged with alterations from Amer.

Jour. Sci. June, 1899 Title 80. Differs also from notice in Amer.

Geol Sept, 1899. Title 82.]

ug.,

1901. 91. The Restoration of a Dinosaur. Yale Scientific Monthly. June,

1901. Vol. 7, p. 291—293, 1 plate. y

92. Studies in Evolution. [Reprints of twenty of Professor Beecher's

papers as noted above.] Yale Bicentennial Publications, New York:

Charles Scribner's Sons, London: Edward Arnold, Aug., 1991. p.

1-23, and 1-638, fig. 1-132, pl. 1-34.

93. Notes on the Cambrian Fossils of St. Francois County, Missouri.

Amer. Jour. Sci. Nov., 19o1. Ser. 4, vol. 12, p. 362-363.

[Reprinted.] Geol. Mag. [London] Dec., 1901. N. S., Dec. 4, vol.

8, p. 559-561.

94. Discovery of Eurypterid Remains in the Cambrian of Missouri.

Amer. Jour. Sci. Nov., 19o1. Ser. 4, vol. 12, p. 364-366, pl. 7.

[Reprinted.] Geol. Mag. [London]. Dec., 1901. N. S, Dec. 4.

Vol. 8, p. 561—564, 1 fig.

95. A Treatise on Zoólogy. Edited by E. Ray Lankester. Part 5.

The Echinoderma by F. A. Bather [etc]. [Review.] Science. Nov.

29, 1901. New Series, vol. 14, p. 844-845.

1902. 96. The Reconstruction of a Cretaceous Dinosaur, Claosaurus

426 THE AMERICAN NATURALIST. [Vor. XXXVIII.

annectens Marsh. Trans. Conn. Acad. Arts. and Sci. Jan., 1902.

Vol. 11, p. 311-324, fig. 1-6, pl. 41-45.

97. Alpheus Hyatt. [Obituary.] Amer. Jour. Sci. Feb., 1902. Ser.

4, vol. 13, p. 164. 5

98. The Ventral Integument of Trilobites. Amer. Jour. Sci. March,

1902. Ser. 4, vol. 13, p. 165—174, fig. 1, pl. 2-5.

[Reprinted.] Geol Mag. [London.] April, 1902, Dec. 4. Vol. 9,

p. 152-162, fig. 1-8, pl. 9-11.

99. Note on a New Xiphosuran from the Upper Devonian of Pennsyl-

vania. Amer. Geol. March, 1902. Vol. 29, p. 143-146, fig. 1.

100. [Review. Om de Senglaciale * spite: Niváforandringer

i Kristianiafeltet Molluskfaunan], . Brogger. Amer. Jour

Sci. April, 1902. Ser. 4, vol. 13, p. fos

101. Revision of the Phyllocarida from the Chemung and Waverly

Groups of Pennsylvania. Quart. Jour. Geol. Soc. [London] Aug.,

1902, vol. 58, p. 441—449, pl. 17-19.

102. Recent Papers on the Embryology, Structure and Habits of Liv-

ing Brachiopoda. [Review.] Science. Dec. 5, 1902, N. S. vol. 16,

p. 901—902.

103. Climbing Sunset Mountain, Arizona. [In] Grand Canyon of

Arizona. Published by the Passenger Department of the Santa Fe

[Railroad]. 1902. p. 97-100, portrait of author and 5 scenic figs.

1903. 104. Exhibition of Indian Basketry at the Yale University Museum.

Saturday Chronicle, New Haven. Jan. 17, 1903. p. 13, fig. 1—5.

105. Observations on the Genus Romingeria. Amer. Jour. Sci. July,

1903. Ser. 4, vol. 16, p. 1-11, fig. 1, pl. 1-5.

106. The Correlation of Geological Faunas, by Henry Shaler Wil-

liams. [Review.] Amer. Jour. Sci. Oct, 1903. Ser. 4, vol. 16,

P- 334-

107. Pseudoceratites of the Cretaceous; by Alpheus Hyatt. [Review.]

Amer. Jour. Sci. Oct, 1903. Ser. 4, vol. 16, p. 335.

1904. 108. Note on a New Permian Xiphosuran from Kansas. Amer.

Jour. Sci. July, 1904, Ser. 4, vol. 18, p. 23-24, fig. I.

VARIATION IN THE RAY FLOWERS OF

THE COMMON CONE FLOWER

(RUDBECKIA HIRTA).

F. C. LUCAS.

Tue following observations concerning the variation in the

ray flower of the common cone-flower or brown-eyed Susan

were made during the summer of 1902.

Lot No. 1.— 318 heads. From prairies near what is known

as the Stickney tract (Chicago, Ill). Soil clayey, ground rather

ILL

ENSZIRSESNERENSEEGL same

cane BURBZEEURESERNSARCUT ER

30 H4 40 7

F 1G. 2.—Lots 2 and 3. 468 Heads.

Lot 2—11.328 - ; Mean,

Fic. r.— Lot No. 1. 318 Heads.

8 +.

Loti-—

Mean — 14, 18

moist. Collected July 7th. These heads were picked at random

from a large patch. Fig. t. |

Lot No. 2.— 225 heads. Collected from the same region and

at the same time as those of lot No. 1. In this case, however,

56 different plants were pulled and all the flowers on each were

included in the count. Fig.2 : ;

Lot No. 3.— 240 heads. From the prairies two miles directly

427

428 THE AMERICAN NATURALIST. [Vor. XXXVIII.

East of the first locality. Ground was much less moist. 4c

plants were collected at random and all the flowers counted as

in lot No. 2. Collected July 12th. Fig. 2.

Lot. No. 4.—173 heads. From the prairies about eight

miles southeast of the preceding localities and not far from the

lake shore. Ground moist from recent and continued rains.

Plants were all rather low and much shaded by tall grass and

tall weeds. Collected July 16th. Fig. 3.

Lot No. 5.— 246 heads. From the same general region as

[LE

& 34D 34 ee 25

Fic. 3.— Lot No. 4. 173 Heads. Fic. 4.— Lot No. 5. 246 Heads.

Mean — 9.612 +. Mean— 11.813 d. `

in the case of Lot No. 4, but from portions of the prairie where

there were fewer weeds ànd from lanes in the vicinity. Collected

July 21st.

In the case of both Lot 4 and Lot 5 plants were pulled up at

random and all the flowers of each plant included in the count.

The results of the various counts is shown in the following curves.

Comparing the above curves we notice one rather interesting

fact. The curve of those heads which were growing under what

had been noticed to be especially unfavorable (curve 3) is entirely

different from the other three, curves 1 and 2 being especially

symmetrical. Iam of the opinion that the reason that curve 4

is not so symmetrical as 1 and 2 is that the heads were taken

from plants not far enough removed from those represented in

No. 450.] RUDBECKIA HIRTA. 429

curve 3. The curve in fact shows the transitional condition as

regards favorable conditions of growth. Curve 3 on the other

hand is a representation in the form of.a curve of very unfavora-

ble conditions of growth.

The characteristic mode of Rudbeckia hirta is 13. There are

also secondary modes at 8 and at 20-21: and the series thus

reminds one of Ludwig’s series for Chrysanthemum leucan-

themum. In connection with these different modes it is inter-

esting to note that the number of rays is made the basis of

classification.

Species. Gray. Britton & Brown.

A apte o x sos Rays about 14 Rays 10-20

NE CINA... . s . Rays about 12 Rays 10-15

K: Spathulala ;. . 7. Rays few Rays 8-

NANI oo os Rays numerous Rays 12-20

ENGLEWOOD HIGH SCHOOL, CHICAGO.

STUDIES ON THE PLANT CELL.— II.

BRADLEY MOORE DAVIS.

THE ACTIVITIES OF THE PLANT CELL.

1. Vegetative Activities.

Every cell passes through a history whose events repeat in a

broad way activities that have become established in the organ-

ism by the experience of its ancestors. The most important of

these events is nuclear division, which is accompanied in most

plants by cell division, the important exceptions being certain

groups whose protoplasm is multinucleate throughout all, or

almost all, vegetative conditions (e. g., ccenocytic Algae and

Fungi, plasmodia and multinucleate cells in various tissues).

Protoplasm, whose nuclei can no longer divide, becomes inca-

pable of reproducing itself and must take a dependent position

in the organism, where the length of its life will be determined

by the good fortune of its environment and its vitality. Such

protoplasm becomes strictly vegetative in its functions, and while

these activities may be very highly specialized and of the utmost

importance to the organism as:a whole, nevertheless such a cell

has lost certain of the constructive, and in consequence repro-

ductive, possibilities characteristic of living matter. The most

evident and important of these constructive activities have to do

with the increase of nuclear material (chiefly chromatin), which

leads to its distribution through nuclear division, and. the devel-

opment of a complicated mechanism (the spindle) to effect this

result.

As Weismann first pointed out, from the standpoint of cell

studies, there is a stream of germ plasm flowing with every spe-

cies, protoplasm relatively fixed in its characteristics and poten-

tially immortal. The chief peculiarities of germ plasm are its

reproductive powers and the generalized structure that enables it

43!

432 THE AMERICAN NATURALIST. [Vor. XXXVIII.

to turn to any form of activity possible to the species. Portions

of the germ plasm are constantly being detached from the main

stream and relegated to more or less special duties. Such pro-

toplasm becomes the body plasm, or soma, of the individual.

Specialized body plasm generally loses very shortly the reproduc-

tive possibilities of germ plasm, and in consequence must finally

die, for its nicely adjusted dependence upon surrounding cells

cannot last forever. Yet it has been one of the surprises of

biological science that specialized tissues may keep for a very

long time the reproductive qualities of germ plasm. Investiga-

tions on regeneration in particular have brought these facts con-

spicuously to the front. . As an extreme example among plants,

it is known that even the epidermal tissues of leaves and scales

of certain ferns (Palisa, :00) may sometimes retain the funda-

mental qualities of. germ plasm and reproduce the plant.

There are no visible characters that separate body plasm from

germ plasm, excepting,.of course, when body plasm begins to

show signs of degeneration. Germ plasm may only be distin-

guished by its potentialities of growth and reproduction, potenti-

alities that cannot be accurately determined because the stimulus

to development is, in the last analysis, an external one and the

conditions which govern it may be so intricate as to escape close

scrutiny.

Germ plasm is found in its most generalized condition in the

cells of growing points, in embryonic: and meristematic regions,

and in the reproductive tissues. These tissues are well recog-

nized as the most favorable for cell studies because they present

most clearly the details of protoplasmic activities. Almost all

that we know of cell activities have come from investigations of

such regions.

One of the first signs of that specialization which transforms

germ plasm to body plasm is the slowing up and final end

of nuclear and cell division. With this change come a great

variety of modifications (peculiarities of cell wall, plastids, cyto-

plasmic activities, etc.) which may be readily associated with the

particular work of that tissue.

The vegetative activities of germ plasm are chiefly those of

growth, which in the end mean reproduction, the embryonic cells

No. 450.] STUDIES ON THE PLANT CELL. 433

drawing upon food that has been prepared for them and is either

stored in special structures (as seeds, spores, bulbs, etc.), or manu-

factured in differentiated organs or tissues (leaves, chlorophyll

bearing tissue, phloem, etc.). The vegetative activities of body

plasm are far more specific than those of germ plasm. Their

tissues have particular and highly developed activities, some deal-

ing chiefly with photosynthetic processes, some (phlaem) distrib-

uting the organized food over the plant body, some storing the

food in large quantities. Besides these there are mechanical

functions performed by highly differentiated tissues, even though

largely composed of empty cells, as the vascular tissue, support-

ing tissues, and the external protective integuments.

It is not our purpose to discuss any of these vegetative activ-

ities in detail, but only to distinguish as sharply as possible the

characteristics of germ plasm with its generalized activities from

the specialized body plasm. These generalized characters, as

before stated, are constructive activities which mean growth and

lead to nuclear and cell division. It is probable that any tissue

which presents them has regenerative powers that under the

proper environment might be expected to reproduce parts or the

entire organism. Germ plasm is distributed more widely through-

out the organism than is generally supposed, and many highly

specialized tissues still retain the spark of regenerative possi-

bilities. The significance of these conditions is not generally

appreciated, perhaps because the environmental conditions of

regeneration are little understood and are exceedingly hard to

adjust experimentally. There is presented here a very attractive

field of botanical investigation, a union of cell studies with the

more gross anatomical methods of experimental morphology.

2. Cell Division.

Cell division takes place only after periods of growth that

have led to a multiplication of nuclei and in the tissues of plants

above the thallophytes is very generally a part of the history of

each mitosis. This is because of the structure called the cell.

plate which is essentially an organ of cell division. But the

thallophytes present other methods of cell division which bear

no especial relation to nuclear activities, and in certain groups of

434 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the thallophytes nuclear division may proceed through the entire

vegetative lite of the organism without any segmentation of the

protoplasm which only takes place during the reproductive phase

of spore formation. But fundamentally protoplasmic segmenta-

tion depends on increase in the amount of protoplasm which

demands the multiplication of nuclei so that nuclear division

always precedes cell division, and we shall consider the events in

that order. i

(a) Events of Nuclear Division.

I. Direct Division.

The nucleus divides after one or two methods, either directly

by constriction or fragmentation, or indirectly (mitosis) when

there is present a fibrillar apparatus called the spindle. Direct

division is the only form present in the simplest plants and phy-

logenetically must have preceded the elaborate mechanism de-

manded for indirect division. This topic will be given especial

attention in Section VI. Direct division is also present in cer-

tain specialized cells and tissues of higher plants. These are

generally old cells or tissues that are far removed from the gen-

eralized structure and potentialities of germ plasm. Yet some-

times direct and indirect division occur in the same cell, e. g.

Valonia (Fairchild, ’94), and such forms might be made the

subject of very interesting investigations. In some cases the

phenomenon of direct nuclear division accompanies pathological

conditions or the degeneration of cells and may take the form of

extensive fragmentation. It would be outside of our purpose to

discuss such phenomena which is obviously abnormal, and the

primitive forms of nuclear division will be taken up later (Sec-

tion VI). It is possible that direct division in higher plants is in

a sense a reversion to early ancestral conditions, a reversion that

only comes on when for some reason the normal activities of the

germ cell are in abeyance or have ceased.

2. Indirect Division (Mitosis).

Indirect nuclear division, mitosis or karyokinesis, is character-

ized by a mechanism which varies greatly among plants in its

No. 450.] STUDIES ON THE PLANT CELL. 435

method of development. The characteristic appearance of this

apparatus is a spindle like figure formed of fibrille. The poles

of the spindle may be occupied by centrosomes or centrospheres

or they may be entirely free from such organized kinoplasmic

bodies. The essential structures of the spindle are sets of con-

tracting fibers which separate the chromosomes into two groups

drawing them to the poles of the spindle where the daughter

nuclei are organized. But besides these fibers there are gen-

erally present other fibrilla which complicate the nuclear figure.

Some of these extend from pole to pole (spindle fibers) others

lie outside of the spindle and end freely in the cytoplasm or,

attach themselves to chromosomes (mantle fibers), and if centro-

somes or centrospheres be present there are likely to be fibers

radiating from these centers to form asters.

The events of mitosis are generally grouped into four periods :

(a) Prophase, to include the formation of the spindle and prep-

aration of the chromosomes; (b) Metaphase, the separation of

the daughter chromosomes ; (c) Anaphase, the gathering of the

daughter chromosomes into two groups which pass to the poles

of the spindle; (d) Telophase, the organization of the daughter

nuclei. It is almost needless to say that these periods merge so

gradually one into the other that sharp lines cannot be drawn

between them. The activities during prophase are especially

variable.

Prophase. — There are two types of spindles in plants, (1)

those that are formed within the nuclear membrane and (2) those

whose fibers originate largely or wholly from kinoplasm outside

of the nucleus. Intranuclear spindles have been reported in a

number of groups of the thallophytes. They seem to be the

rule in the mitoses of oógenesis in the Peronosporales (Wager,

'96, :00, Stevens, '99, :01 and :02, Davis, :00, Miyake, :01,

Trow, :or, Rosenberg, :03). They are present in Saprolegnia,

Fig. sa (Davis, :03). Fairchild ('94) reports them for Valonia.

Farmer and Williams ('98, p. 625) state that the spindle of

Ascophyllum is largely intranuclear. Harper (:00) has not

described them for the Myxomycetes, but very little is known

about the prophases of mitosis in that group and their presence

is quite probable. Timberlake (: 02) is not positive whether the

436 THE AMERICAN NATURALIST, (VoL. XXXVIII.

spindles of Hydrodictyon are intranuclear or not; they lie in a

clear space which, however, may be a vacuole rather than the

outline of a nuclear cavity. It seems probable in such a type

that the vacuole is really the nuclear cavity whose plasma

membrane (nuclear membrane) becomes less clearly defined.

The development of the spindle is very difficult to follow among

these lower forms because it is so small. Stevens (:03) found

an exceptionally favorable type in Synchytrium and came to

the conclusion that the spindle developed from the threads of

the spirem (linin) entirely within and independent of the

nuclear membrane.

Very remarkable intranuclear spindles have been described in

the central cell of the pollen tube of Cycas (Ikeno, '98 b) and

Zamia, Fig. 5d (Webber, :o1). Murrill (:00) found them in

the mitosis following the fusion of gamete-nuclei in the egg of

Tsuga, Ferguson (:orb) at the same period for pine, and Coker

(:03) in Taxodium. They are also reported by Strasburger (: 00)

in the cells of young anthers and nucelli of the lily and in grow-

ing points (Viscum) and possibly may be found quite generally

in cells weak in kinoplasmic cytoplasm. The development of

the spindles in the above forms has not been studied in detail,

but the fibers are probably derived from the linin. We are

given a clue to the process by the events of spindle formation in.

the spore mother cell of Passiflora (Williams, '99). In this

angiosperm the nuclear cavity becomes filled with a fibrillar

network developed from the linin, the nuclear wall becomes

transformed into a mesh connecting the intranuclear fibers with

a surrounding cytoplasmic reticulum. The fibers in the central

region of this net work develop the spindle which is consequently

very largely of intranuclear origin.

Among the thallophytes the poles of intranuclear spindles are

frequently occupied by deeply staining bodies which have been

called centrosomes; but these structures can hardly be homol-

ogous with the well-known centrosomes of other thallophytes,

e. g, Stypocaulon (Swingle, '97) and Dictyota (Mottier, : 00).

They are probably merely temporary accumulations of materia

with no morphological significance.

Spindles that arise from fibers external to thé nucleus (extra

No. 450.] STUDIES ON THE PLANT CELL. 437

nuclear spindles) are of two main types: (1) those associated

with centrosomes, centrospheres or kinoplasmic caps, and (2)

those composed of independent fibrillae developed as a mesh

around the nucleus. The latter condition is especially character-

istic of the spore mother cell and is perhaps the highest type of

spindle formation known for either animals or plants. It is very

interesting to trace the relations of this highest condition to the

lower types through certain lines of evolution to be discussed in

Section VI.

Spindles with centrosomes are known in Sphacelaria, Stypo-

caulon (Swingle, '97), Dictyota, Fig. 4 a (Mottier, : 00), the zoó-

sporangium of Hydrodictyon (Timberlake, :02), in certain

diatoms (Lauterborn, principal paper '96, Karsten, :00) and in

the basidium (Wager, '94 and Maire, :02). The best accounts

of the behavior of the centrosomes are given by Swingle and

Mottier. Indeed there is much doubt about the history and

significance of the bodies in the other forms, although the con-

stancy of their presence at the poles of the spindles indicates

that they are really centrosomes. The conditions in the diatoms

are especially complicated ; an account of Lauterborn's work has

been published in English by Rowley, :03. In Stypocaulon,

Sphacelaria (Fig. 3 c, Section I) and Dictyota (Fig. 4 4) the cells

studied have permanent asters which lie at the side of the

nucleus and which divide just previous to the mitosis and sep-

arate so that they come to lie on opposite sides of the nucleus.

Fibers develop from the centrosomes on the sides nearest the

nucleus and elongating push against the nuclear membrane and

finally enter the nuclear cavity to form the spindle.

Spindles with centrospheres are well known in Fucus (Farmer

and Williams, '96, '98, Strasburger, 972), Corallina, Fig. 5 c,

(Davis, '98), in the ascus, Fig. 5 ? (Harper, '97 and '99), and in

the germinating spore of Pellia, Fig. 4 ¢ (Farmer and Reeves,

'94, Davis, : or, Chamberlain, :03). Centrospheres have been

reported in other forms but the types mentioned above have

received the most careful study. It is probable that the centro-

sphere is but a larger, more generalized kinoplasmic center than

the centrosome, a protoplasmic region whose dynamic activities

do not focus so sharply as in the latter structure. There are

438 THE AMERICAN NATURALIST. [Vor. XXXVIII.

bodies, as in the basidium, which stand intermediate in size

between centrosomes and centrospheres and are probably only

called the former because they are very distinct in outline.

Centrospheres in Fucus (Fig. 3 d, Section I), Corallina (Fig.

4 6) and Pellia (Fig. 3 e, Section I, Fig. 4 «) are formed de novo

for each mitosis by an accumulation of kinoplasm at the poles of

the elongating nucleus. The centrospheres in the ascus divide

before each of the three successive mitoses and finally remain,

one for each nucleus, to instigate the peculiar process of free

cell formation characteristic of the ascus. Centrospheres are

frequently the centers of asters which, however, are usually not

as sharply defined as those with centrosomes, possibly because

the fibers are not grouped with the same degree of symmetry as

is shown around controsomes.

Spindle fibers from centrospheres develop in precisely the

same manner as from centrosomes, 7. e. by the growth of the

fibrillæ into the nuclear cavity through the dissolving nuclear

membrane. The activity is well shown in the oógonium of

Fucus, and Farmer ('98, p. 638) believes “that the intranuclear

part of the spindle is differentiated out of nuclear material that

is unused for chromosome formation." The entrance of spindle

fibers from centrospheres at the ends of a nucleus has been

observed by myself in Corallina, Fig. 4 @ (Davis, ’98). The

germinating spores of Pellia, Fig. 4 « (Davis, : O1, Chamberlain,

:03) furnish especially good illustrations of the entrance of spin-

dle fibers into the nuclear cavity and the development of the

spindle in this form is coincident with the dissolution of the

nucleus which, according to Strasburger's theory (/95), indicates

that the latter structure contributes material for the growth o

spindle fibers.

In connection with the centrosphere mention should be made

of the blepharoplasts of the cycads and Ginko which are remark-

able bodies with radiating fibers. They have been considered

by some as asters with centrosomes, but it is known that they

take no part in spindle formation or other.mitotic phenomena in

these forms, and consequently need not be considered at this

time. They will be treated in some detail in the account of the

sperm (Section ITI).

No. 450.] SLUDIES ON THE PLANE CELL. 439

Kinoplasmic caps which form spindles are probably an evolu-

tion from the type of centrosphere that is developed de novo

with each mitosis as in Pellia. Such centrospheres by becoming

less definite in form and lacking radiating fibers would be called.

kinoplasmic caps. Indeed the centrosphere so evident in the

early cell divisions of the germinating spore of Pellia becomes a

kinoplasmic cap in the later mitoses of the older gametophyte

(Davis, : or).

Spindles developed from kinoplasmic caps are characteristic of

/ e Ean

MP IT, C

hy) D DE.

o2 (1 "n

1 4

Soe

Fic. 4.— Prophases of Mitosis. æ. Dictyota; late prophase in spore mother cell, fibers from

t ith h | 1 cav ity g

the two asters

mosomes gathering to form the nuclear plate. 4, Corallina, early prop p

mother cell; two centrospheres, the fibers form one having entered the nuclear cavity,

chromosomes shown. c, Pellia, nucleus in germinating spore ; spindle fibers from ill

defined centrospheres entering nuclear cavity, chromosomes and a nucleolus present. 4,

Gladiolus, first mitosis in pollen mother cell; a multipolar spindle, nuclear wall breaking

down at one side and fibrilie entering the nuclear cavity, chromosomes and a nucleolus

present. After Mottier and Lawson.

the mitoses in vegetative tissues, meristematic and other embry-

onic regions. They have been especially studied in higher plants

by several investigators and for a large number of forms, those

most completely described being Psilotum (Rosen, '95), Equise-

tum, Allium and Solanum (Nemec, '98a and '98b, '99b and '99c),

Pteris, Ephedra and Vicia, (Fig. 3 /, Section I) (Hof, '98) and

Allium (McComb, :00). The polar caps first appear as accumu-

lations of kinoplasm on opposite sides of the nucleus which

generally elongates. The protoplasm is granular and althóugh

440 THE AMERICAN NATURALIST. [Vor. XXXVII.

central bodies have been reported most investigators are agreed

that they are only granules without regularity or special signifi-

cance. They are no longer believed to be centrosomes. Fibrillae

are developed from the kinoplasmic caps and grow out against

the nuclear membrane and finally enter the nuclear cavity to form

the spindle. A large part of the substance of the kinoplasmic

cap is transformed into these spindle fibers.

Papers by Schaffner (98) on Allium and Fulmer (’98) on the

seedling of the pine are the last attempts to bring the centro-

some into the history of spindle formation in vegetative tissues

of higher plants. But their results cannot stand against the

accumulation of studies which indicate that centrosomes are not

present in the cells of any plant above the thallophytes with the

possible exception of the mysterious blepharoplast and certain

structures appearing in some phases in the life history of

Hepatice. Centrospheres are unquestionably present in the

Hepaticze and centrosomes have also been reported. The

centrospheres are, however, so generalized as to approach the

kinoplasmic caps in structure and development and it seems.

quite possible that they are the forerunners of this manifesta-

tion of kinoplasm. The so-called centrosomes of the liverworts

do not exhibit the specialized structure or behavior of cen-

trosomes among the thallophytes and it is probable that they

are only smaller and somewhat more clearly defined centro-

spheres. These structures in the Hepatica seem to hold an

intermediate relation between the definite kinoplasmic bodies

(asters, centrosomes and centrospheres) of the thallophytes and

the remarkable kinoplasmic activities in higher plants which

reach their highest expression in the processes of spindle forma-

tion in the spore mother cell. These topics will be treated in

Section VI.

Structures resembling kinoplasmic caps have been reported in

several other tissues than those noted above. Thus Murrill

(1:00) finds in the formation of the ventral canal cell of Tsuga a

dense fibrous accumulation beneath the nucleus which develops

one pole of the spindle in essentially the same manner as other

polar caps. The other pole of the spindle in this case appears

to be formed differently for the fibers seem to be intranuclear.

No. 450.] STUDIES ON THE PLANT CELL. 44I

It would be interesting if two types of spindle formation were

present at opposite poles of the same nucleus and further inves-

tigation of this subject is much to be desired. The mitoses in

the central cell of Pinus (Ferguson, :orb, Chamberlain, '99,

and Blackman, '98) and Picea (Miyake :03a) show spindle for-

mation from accumulations of fibrillae outside of the nucleus but

without conspicuous polar caps. Still more striking than the

irregular spindle of Murrill in Tsuga, described above, is Miss

Ferguson's (:01a) account of the mitosis in the generative cell

of the pollen grain of Pinus. The spindle here begins to develop

as a cap-like accumulation of kinoplasm below the nucleus. The

fibers enter the nuclear cavity and in coóperation with a nuclear

reticulum form a system of fibers that extend through the

nuclear cavity to the inner side of the nuclear membrane

beyond. This portion of the nuclear membrane persists until

after metaphase so that one pole of the spindle is found wholly

within the nucleus while the other is external and of unques-

tioned cytoplasmic origin. Coker, : 03, regards the spindle which

differentiates the nucleus of the ventral canal cell in Taxodium

as almost wholly of nuclear origin and the chromosomes as

derived largely from the nucleolus. There are evidently some

interesting complications in this form which deserve further

study.

It should be noted that whenever spindles are formed in con-

nection with centrosomes, centrospheres or kinoplasmic caps that

the fibers have a definite region of attachment from which they

extend into the nuclear cavity. Such regions constitute a sort

of anchorage for the spindle fibers. In this respect the physi-

ological side of the process of spindle formation in these forms

is quite similar to that of the animal kingdom and in sharp

contrast to other methods that are found in higher plants, which

will now be considered. ~ ;

When spindles are formed after the second method, 7. e.,

by independent fibrillae making up a network around the nucleus,

there is an abrupt change in the method of development.

The kinoplasm becomes distributed around the nucleus as an

investing layer and shows no inclination to gather into d

such as kinoplasmic caps or centrospheres. There is developed

442 THE AMERICAN NATURALIST. [Vor. XXXVIII.

from this granular kinoplasm a meshwork of fibrillze that extends

into the cytoplasm more or less radially. When the nuclear

membrane becomes disorganized the fibers enter the nuclear

cavity and organize the spindle (see Fig. 4 7). In some forms,

e. g., Passiflora (Williams, : 00), many or most of the fibers are

developed in the interior of the nucleus from the linin and become

connected with the extra nuclear reticulum by the dissolution of

the nuclear membrane.

The free ends of the fibrillae that lie in the cytoplasm become

gathered into several poles which are distributed variously around

the nucleus. This condition constitutes the so-called multipolar

spindle (Fig. 3g, Fig. 4 æ), which in its highest type of develop-

ment illustrates the most complex method of spindle formation

known for animals or plants. During the later periods of

prophase the several poles of the multipolar spindle converge

and fuse with one another into two poles with a common axis,

thus forming the mature bipolar spindle of metaphase (Fig. 5 f).

The spindle is in a broad sense bipolar, but one may readily see

that each pole is made up of several groups of fibrillae which

generally remain quite independent of one another (Fig. 5 f 1).

The relation of a multipolar stage to the bipolar spindle of

metaphase was first made clear by Belajeff (’94b) for Larix, and

later was established more widely by the investigations of Oster-

hout (97) on Equisetum, Mottier (’97) for the lily, and Juel (97) -

for Hemerocallis. This type of spindle formation is now well

known in the spore mother cells of numerous spermatophytes

and several pteridophytes. The same conditions in simpler

form are found in the spore mother cells of the Hepatice,

e. g, Anthoceros (Davis, '99), Pellia, Fig. 5 e (Davis, : 01), and

Pallavicinia (Moore, :03). There are a number of very interest-

ing peculiarities in this type of spindle which presents a wide

range of variation in the details of its fibrillar organization and

development. These will receive special treatment in the account

of the spore mother cell (Section III).

The only types of thallophytes known, in which the spindle is

partly or wholly of cytoplasmic origin without centrosomes, Cen-

trospheres, or kinoplasmic caps, are Chara (Debski, '97) and

Spirogyra (Van Wisselingh, :02). The developmental history 1S

No. 450.] STUDIES ON THE PLANT CELL. 443

very difficult to follow in these forms and is not fully known, but

multipolar conditions are reported which later change into bipolar

spindles.

While the spindle is being organized by kinoplasmic activities

outside of the nucleus, some events occur within which form a

very important part of the prophases of mitosis. The linin

material, which in the resting nucleus generally has the form of

a net, becomes organized into a much looped ribbon, called the

spirem thread. The chromatin material gathers along the spirem

thread as deeply staining globular bodies. These split into halves

in the direction parallel with the axis of the spirem thread, and

the two sets of chromatic bodies lie in two rows along the edge

of the ribbon, which shortens as it grows older. Finally the

spirem thread divides transversely into a definite number of seg-

ments, and these are the chromosomes. The chromosomes are

generally fully formed at the time when the spindle fibers enter

the nuclear cavity, and they are readily moved as the fibrillae

develop the spindle. Some of the fibers become attached to the

chromosomes, carrying them to the equatorial region of the

spindle to form the structure called the nuclear plate, which

always indicates the approach of metaphase.

As the spirem thread matures the amount of chromatin is

greatly increased, so that the separate globules run together and

cannot be distinguished in the chromosome which is homogeneous

in structure. Chromatin has its greatest staining power at this

period. Whether linin is closely related to chromatin in compo-

sition and is actually changed into that substance, or whether it

dissolves and contributes its material to the growth of the spin-

dle, is a problem of some importance as yet unsolved. It is

possible that the nucleolus may furnish material for the chromo-

somes, and some nucleolar like bodies are known to be chromatic

in character, but it does not seem to be established that any of

these are genetically related to an unquestioned nucleolus in any

plant form. i

Important changes come over the nucleolus coincident with the

development of the chromosomes. The structure frequently

gives signs of internal modifications early in prophase and before

the development of the spindle. It may gradually fade away or

444 THE AMERICAN NATURALIST. (Vor. XXXVIII.

decrease in size, or, if large, it may fragment. Strasburger, in

1895, advanced the view that the spindle actually drew upon the

substance of the nucleolus for the material and energy necessary

to its development. The evidence in support of this suggestive

theory lies chiefly in the development of the spindle coincident

with the dissolution of the nucleolus. There is also some evi-

dence that the nucleolus contributes material to the developing

chromosomes. Small globules, which stain as the substance of

the nucleolus, may sometimes be found adhering to the chromo-

somes as though becoming incorporated in them. These sub-

jects are naturally very difficult of investigation because stain

reactions cannot be depended upon with certainty and are not,

of course, chemical tests. Then the behavior of the nucleolus

during mitosis is exceedingly variable, since it sometimes disap-

pears quickly and sometimes remains intact, and it becomes a

very difficult matter to determine its importance. The nucleolus

is probably not absolutely necessary at any stage in mitosis, for

both spindle fibers and chromosomes develop apart from this

structure ; but it does seem to be established that the substance

of the nucleolus is generally drawn upon by the cell, especially

during prophase, when numerous spindle fibers are organized and

the amount of chromatin is being largely increased. Experi-

ments of: Hottes, which unfortunately have never been published,

have an important bearing on these problems.

If the nucleoli are not entirely dissolved they are frequently

thrown out of the spindle into the cytoplasm, where they may

lie for long periods as deeply staining globules which are some-

times called extra nuclear nucleoli. It is probable that very many

of the bodies that pass under this cumbersome title have no

relation whatever to the nucleolus. The cytoplasm frequently

contains globules that may be coagulated or precipitated food

products, and all of these stain similarly to nucleoli.

Metaphase. — The period of mitosis termed metaphase is, to

speak precisely, the time when the two halves of the split chromo-

some separate from one another. However, this is a period of

such short duration that for practical purposes nuclei are consid-

ered in metaphase when their chromosomes are lined up at the

nuclear plate. The metaphase of mitosis is generally the most

No. 450.] STUDIES ON THE PLANT CELL. 445

conspicuous of the nuclear activities not only on account of the

position of the chromosomes (see Fig. 5), but because all kino-

plasmic structures (the fibrille and centrosomes or centrospheres,

if present) are shown to their best advantage.

The best evidence indicates that the chromosomes of plants

Lr.

Ae ey E

AUN: ES d

AUD

—

ELA

a, Saprolegnia ; intranuclear spindle in oógoníum, nucie-

Erysiphe ; mitosis i i smal. centro-

heir outer membrane about to break up. e, Pellia; first

of pollen grain ; blepharoplasts, i

mitosis in spore mother-cell; broad spindle with rounded poles, the very numerous spin-

di ing i nular kinoplasm. /, Agave; first and second mitoses in pollen

the

gr +

mother-cells ; (1), multipoiar spindle just previous to metaphase; th several independent

cones of fibrille gather more closely together to complete the spindle. (2) metaphase of

second mitoses; completed spindles showing however the several independent cones of

fibrille. (After Harper, Webber and Osterhout.)

only divide longitudinally. This matter has considerable theo-

retical interest, which will be considered in Section V, and also

in connection with the spore mother cell (Section III). The

daughter chromosomes are drawn apart by the contraction of the

fibrillæ to which they are attached.

Chromosomes may take on various forms during metaphase,

446 THE AMERICAN NATURALIST. (Vor. XXXVIII.

especially while they are being separated. Thus, if the chromo-

somes are dragged apart from the ends, their form is generally

rod shaped; but if the attachment of the fibrillæ is near the

middle of the chromosome, the structures are pulled apart as

loops or V's, and the pair of chromosomes just previous to their

separation may be ring shaped. A further complication is intro-

duced in the spore mother cell by certain premature divisions by

which each daughter chromosome becomes a pair of granddaugh-

ter chromosomes instead of remaining a single structure. The

peculiarities of the heterotypic and homotypic mitoses are due

Frc. 6.— Anaphase of Mitosis, spore mother-cell of Lilium martagon. a, immediately

itosi of two grand

after metaphase of first mitosis; each daughter chromosome consists

daughter segments, adhering at the ends, making the familiar V-shaped figures charac-

teristic of the first mitosis (heterotypic) in the spore mother-cell of higher plants. 4, late

somes, each c

spindle where they will organize the nuclei of the pollen grain; this mitosis is called

a S LN ere 1 1 f sy . . CPC Spe oe ature

homotvpi

divisions of the chromosomes. All figures after Mottier.

to this phenomenon. (See account of spore mother cell in Sec-

tion III.

Anaphase.— Anaphase begins with the separation of the

daughter chromosomes at the nuclear plate (Fig.6) and ends

with the gathering of these structures at the poles of the

spindle preparatory to the organization of the daughter nuclei.

As the chromosomes move towards the poles the fibers of the

central spindle stand out sharply (see Fig. 6 4). If a cell wall

is to be formed between the daughter nuclei one may expect to

find these fibers thickening in the equatorial region of the

No. 450] STUDIES ON THE PLANT CELL. 447

spindle where the nuclear plate formerly lay. Such thickenings

are granular accumulations formed by the contraction of the

central spindle fibers and mark the beginnings of the cell plate

(Fig. 8 d) that afterwards gives rise to the cell wall. When

the daughter chromosomes reach the poles of the spindle they

generally lie in a region of granular kinoplasm which results in

part from the contraction of spindle fibers and in some cases

from the breaking down of organized centrospheres (e. g., Coral-

lina, Pellia, Fucus, etc.). The daughter nucleus at this time

hl He

‘a

cell of Passz/f. le a, very lateanaphase ;

llected at the poles of the spindle. 4, the commence-

ogether and the daughter nucleus is

ass of chromatin. c, the presence of small lacunz

increased amount

consequent enlargement of the vacuole destined to become the nuclear cavity.

chromatin has begun to break up into small masses so that it no longer holds the nuclear

sap which has established contact with the cytoplasm and is forming the nuclear plas

membrane. /, nuclear sap in contact on all sides with the cytoplasm and a complete

nuclear membraneclearly established; chroma in i y much brol p and t leoli

(n) have been formed. g, the ing 1 with ch tin di ll masses

connected by a network of linin threads; a nucleolus (n) is shown; the zone outside the

nuclear membrane is kinoplas' its appearance indicates the approach of the second

mitosis in the pollen mother-cell. All figures after Lawson.

ft. a:

(Fig. 7 a, b) is in its simplest terms, as explained in Section I,

a group of chromosomes surrounded by granular kinoplasm and

without the nucleolus, linin network or the vacuole which later

contains the nuclear sap. i;

Telophase.—Telophase is the closing period of mitosis and

completes the organization of the daughter nuclei (see Fig. 7).

448 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The chromosomes come to lie in a vacuole (Fig. 7 c, d, e) con-

taining nuclear sap and later the chromatin becomes distributed

overalinin network and one or more nucleoli develop (Fig.

7f, &. As was stated in Section I, the nuclear membrane

probably represents the reaction of the granular kinoplasm to a

fluid secretion around the chromosomes which becomes the

nuclear sap (Lawson, :03a). However, the nuclear membrane

is generally a definitely organized film, much more sharply

defined than vacuolar membranes. The development of the

linin network is not well understood. It is readily seen that

the chromosomes become joined end to end and sometimes. elon-

gate. The amount of chromatin diminishes as the linin substance

appears, but it is not certain whether the chromatin is changed

directly into linin, or whether the latter substance is a secretion.

The best evidence rather favors the former view. Nucleoli are

also believed to hold a very close chemical relation to chromatin.

It is uncertain whether or not the chromosomes lose their

organic identity in the daughter nuclei. Investigations on this

problem are surrounded by many difficulties. It has been

claimed by Guignard ('99) for Naias and Strasburger (: 00) for

several forms that the chromosomes may be followed with cer-

tainty through the period between the first and second mitosis

in the spore mother cell. But other investigators have not been

able to trace the chromosomes after telophase and are inclined

to believe that the chromosome completely loses its identity in

the resting nucleus. One of the last investigations of Lilium

(Mottier, :03) argues strongly for the latter view, and all who

have followed nuclei from one mitosis into another know that

the resting nucleus with its linin network and the granular

chromatin present conditions that generally make the recognition

of chromosomes impossible with the instruments and technique

at our command, but this does not prove that they may not be

present.

The theory of the permanence of the chromosome has met

with much favor because it is argued that otherwise how could

the number be maintained so regularly through immense num-

bers of mitoses. But it can hardly be said that the doctrine 1$

established. It has also found favor because all the events of

No. 450.] STUDIES ON THE PLANT CELL. 449

mitosis emphasize the importance of the chromosomes which

are really the only enduring structures in the nucleus and have

led to their being considered as the probable bearers of heredi-

tary qualities.

3. The Dynamics of Nuclear Division.

Mitotic phenomena in certain plant cells present evidence that

has very direct bearing on some of the theories that deal with

mechanical and dynamical explanations of nuclear division. The

methods of spindle formation and the various forms of kinoplas-

mic structures (centrosomes, centrospheres and kinoplasmic

caps) which generally in plants seem not to be permanent

organs of the cells all tend to support Strasburger's conception

of kinoplasm, which is an outgrowth and application to plants of

Boveri's well known theory of archoplasm.

The centrosome theory is supported by very few investiga-

tions in Botany, the most notable being that of Swingle (797),

for Stypocaulon, who believes that the centrosome divides with

the aster and is maintained as a permanent organ throughout

successive cell divisions. Other examples of similar conditions

may be found among the thallophytes which, after all, have

received very little attention, and such types as Dictyota and

the diatoms offer excellent subjects for studies covering à

series of cell divisions. But in contrast to Stypocaulon it should

be noted that the conspicuous centrospheres of Fucus and Cor-

allina disappear with each mitosis to be formed anew, and the

same conditions obtain in the germinating spores of liverworts

(Pellia). There seems to be no place for the centrosome in

spindle formation as presented in the spore mother cells of all

groups above the thallophytes (see Sec. III). Neither does

mitosis in the vegetative tissues of these groups, characterized

asitis by the presence of kinoplasmic caps, conform to the

program of the centrosome theory.

The morphological manifestations of kinoplasm are so various

that we are driven to a very general conception of its organiza-

tion. Kinoplasm runs through cycles in which the structure

passes from a granular condition toa fibrillar and then back again

450 THE AMERICAN NATURALIST. [Vor. XXXVIII.

to the granular state. By the granular state we mean one in

which no fibrillae seem to be present, but instead the microsomata

are densely and homogeneously massed. It is possible that such

microsomata form a closely packed network, but no such struc-

ture is visible under the microscope. The first appearance of

kinoplasm at prophase of mitosis is frequently the granular

condition. This state is illustrated by such accumulations as

centrospheres and kinoplasmic caps and by the granular zone

that has been reported around the nuclei of some spore mother

cells.

Granular kinoplasm becomes fibrillar probably by thearrange-

ment of the microsomata into a reticulum from which fibers

extend freely into the surrounding cytoplasm. These fibers

undoubtedly elongate during prophase, extending in various

directions. Some press against the nuclear membrane and when

this breaks down grow rapidly into the nuclear cavity. Of these

a portion extend from pole to pole and form the central spindle.

Others attach themselves to the chromosomes and lie either

among the central fibers or somewhat outside of the spindle

(mantle fibers). Still others may extend freely into the cyto-

plasm as astral rays from the pole of the spindle, a very com-

mon condition when centrosomes or centrospheres are present.

A contraction of the fibrillae, beginning with metaphase, is just

as characteristic of mitosis as their elongation during prophase.

The fibers attached to the chromosomes draw the latter to the

poles of the spindle. The central fibers in higher plants draw

away from the poles and give their substance to the cell plate.

The substance of contracted mantle fibers, with other kinoplasm

at the poles of the spindle, probably become distributed around

the group of daughter chromosomes so that they finally lie sur-

rounded by a sphere of kinoplasm. -

It does not seem as if we knew much more about the struc-

ture and activities of kinoplasm during mitosis than is indicated

in this cycle of change from a granular condition through a

fibrillar state back to the granular condition, with a period when

the fibers elongate and another when they contract. This with

few exceptions is the history for every mitosis. The exceptions

deal with peculiar conditions or structures. Thus, for example,

No. 450.] STUDIES ON THE PLANT CELL. 451

the astral rays of the centrospheres in theascus instead of con-

tracting to a center or disappearing in the cytoplasm after the

last mitosis grow around the nucleus and cut out a portion of

the cytoplasm to form the spores, thus contributing their sub-

stance to a plasma membrane.

There is little doubt that kinoplasmic fibrillee actually exist as

structural elements in the protoplasm. Their growth and move-

ment in the cytoplasm and nuclear cavity, their multiplication

and shifting arrangements as the spindle develops, and their

contraction to the poles of the spindle or to a cell plate give

these fibers an individuality that cannot be explained on the

theory that they merely represent lines of force or paths of

dynamic stimuli. They apparently perform all the activities

mentioned above by virtue of their own structural organization

which is that of rows of microsomata and in this organization

resemble and are probably closely related to cilia. There is an

excellent discussion of this subject by Allen, :03, p. 302, etc.

Some authors believe that there is a streaming movement in

the astral rays (Chamberlain, :03, for Pellia) either towards or

away from the pole of the spindle. This view is founded on the

granular appearance of the radiations which are sometimes very

thick in Pellia and enlarge at the points where they join the

centrospheres or the outer plasma membrane. It is not alto-

gether clear that the larger of these structures are quite the

same as spindle fibers since they seem to be actually strands of

cytoplasm rather than fibrillae. s

It is probably safe to assume that the forms which kinoplasm

takes have relation to dynamic activities, but it is not easy to

define these. Thus centrosomes, centrospheres and kinoplasmic

caps may well be the centers from which dynamic stimuli extend,

and they may be the focal points of other energies. These

problems have been very little investigated among plants. It is

obvious that differentiated regions of kinoplasm have important

physical relations to other portions of.the protoplasm, one of the

most important being the anchorage which they give to fibrilla, `

thereby largely governing the direction of such strains as come

about through the contraction of these structures in the later

periods of mitosis.

452 THE AMERICAN NATURALIST. [Vor. XXXVIII.

But the essential characteristics of kinoplasm stand out sharply

from whatever point the phenomena of mitosis is viewed, and in

this protoplasm with its power of forming contractile fibers is

vested some of the most conspicuous activities of nuclear division

as well as the important powers given plasma membranes in

relation to the segmentation of protoplasm to be considered

presently.

The dynamic activities concerned with the spindle present only

half the story of mitosis. The other important events occur

inside of the nucleus. One of these is the dissolution of a por-

tion or the whole of the nucleolus which takes place as the

spindle develops and we have already given the views of Stras-

burger ('95 and :00), supported by the studies of other investi-

gators, that its substance in certain instances furnishes material

for the development of the spindle. But the chief events in the

interior of the nucleus deal with the accumulation of chromatin

on the spirem thread which with the disappearance of the linin

indicates that the latter substance may become converted into

the former. The splitting of the spirem ribbon longitudinally is

of the utmost significance for thereby is made possible an exact

and homogeneous distribution of the chromatic material in the

nucleus. We do not know how the spirem ribbon splits nor

have we as yet any evidence of the origin and evolution of this

peculiar activity.

(b) Segmentation of the Protoplasm.

Mitosis in the uninucleate cells of plants is generally followed

by immediate cell division, which takes place in groups above

the thallophytes through the formation and cleavage of the cell

plate in the equatorial region of the spindle between the daughter

nuclei. Among thallophytes, as so far studied, cell division is

chiefly through cleavage by constriction. There are forms among

the thallophytes and also in the spermatophytes whose nuclei

gather about themselves a portion of the cytoplasm, wherein

they lie, which becomes cut out of the general mass by a cell

wall. This is free cell formation.

Multinucleate masses of protoplasm, such as plasmodia and

No. 450.] STUDIES ON THE PLANT CELL. 453

portions of coenocytes, generally divide extensively at repro-

ductive periods and always through cleavage by constriction with,

however, the frequent codperation of vacuoles which help to cut

the protoplasm in the same manner as the cleavage furrows.

Cleavage by constriction is undoubtedly the most primitive type ;

free cell formation and cleavage by cell plates being special and

very highly developed protoplasmic activities.

1. Cleavage by Constriction.

A simple example of cleavage by constriction is presented by

such an alga as Cladophora. The process consists in the build-

ing out of a ring of cellulose from the side wall into the cell

cavity. The outer plasma membrane forms a fold, thus placing

the two surfaces opposite one another (see Fig. 8 a), and the

wall is laid down between these. Spirogyra forms its wall in

precisely the same manner as Cladophora with this peculiarity,

that the new wall finally cuts through the protoplasmic strands

that connect the daughter nuclei. These strands are said to con-

tain spindle fibers (Van Wisselingh, : 02) which may contribute to

the plasma membranes forming the cell wall, as it is completed.

Another illustration of cleavage by constriction is presented in the

formation of:gametes of moulds (Sporodinia) and the abstric-

tion of conidia (Erysiphez), both processes having been studied

by Harper, ’99, p. 506. In these cases a cleavage furrow pro-

ceeds from the surface inward and divides the protoplasm. The

partition wall of cellulose is formed later between the two free

plasma surfaces. The only differences between the processes

above described are that in the first forms the cleavage proceeds

more slowly and the wall follows the furrow as it progresses in

the interior of the cell, while in the latter types cleavage is com-

plete before the plasma membranes develop the wall. Cell

division in the red Algae (Rhodophycez) is also a process of

constriction similar to Cladophora, but the wall is not generally

formed entirely across the filament so that adjacent cells remain

connected by thick strands of protoplasm.

These processes become much more complicated when large

masses of multinucleate protoplasm are divided up into many

454 THE AMERICAN NATURALIST. (Vor. XXXVIII.

smaller bodies as during spore formation among the Myxomy-

cetes and Mucorales. Very complete studies have been made of

these conditions by Harper, '99 and :00a. In the slime mould

(Fuligo) cleavage begins by furrows on the external surface

which * cut down at all angles into the homogeneous proto-

plasm." The direction of the cleavage furrows is further com-

plicated by the fact that many of them start from the bottom

and sides of deep folds. All of the furrows may bend and

secondary cleavage planes strike off from them which in time

unite with one another until the protoplasm is divided progres-

sively into very many small masses (see Fig. 8 0) that finally

round themselves off and secrete walls, becoming spores, some-

times with one nucleus and sometimes with several.

Cleavage in the sporangium of Synchytrium and the moulds,

as described by Harper, '99, is in general similar to that in the

plasmodium with, however, the additional feature that lines or

planes of vacuoles are often utilized to assist a cleavage furrow

in effecting the segmentation of the protoplasm. The -separa-

tion of the spore plasm of the sporangium of Pilobolus from the

filament below begins with a cleavage furrow from the exterior ;

but this furrow follows and makes use of a curved plane of

flattened vacuoles with the result that a dome shaped cleft is

developed and two plasma membranes are presented face to face,

which form the columella wall between them. The segmenta-

tion of the spore plasm in Pilobolus is affected somewhat

similarly through the codperation of cleavage furrows from

the exterior with vacuoles which cut into the protoplasm at

various angles to meet one another and the cleavage furrows.

The bodies first formed in the sporangium of Pilobolus are not

the final spores. Harper suggests that they may correspond to

the zooóspores of Saprolegnia. They are generally uninucleate

and begin immediately a period of growth within the sporangium

characterized by extensive nuclear multiplication and several

divisions of the protoplasmic body by constriction.

Harper finds that the spore plasm of Sporodinia is sep rated

from the filament below by a dome-shaped plane of flattened

vacuoles which fuse together and, unlike Pilobolus, cut their way

to the surface of the sporangium. Thus the cleavage is deter-

No. 450.] STUDIES ON THE PLANT CELL. 455

mined entirely by the activity of vacuoles. Spore formation,

however, is accomplished by cleavage furrows which progress

from the exterior inwards and, without the aid of conspicuous

vacuoles, cut out multinucleate masses of protoplasm which

become the spores.

Dean Swingle (:03) has extended the studies of Harper on

spore formation in the molds to Rhizopus and Phycomyces. He

confirms Harper's account of the general processes of cleavage

by furrows coóperating with vacuoles, and notes the following

characteristics in the types studied. In Rhizopus the position

of the columella is determined by a dome-shaped series of flat-

tened vacuoles which fuse and meet a cleft that extends upward

from the outer plasma membrane at the base of the sporangium.

The. spores are formed in Rhizopus by branching systems of

curved furrows that cut the protoplasm into multinucleate masses,

and in Phycomyces by angular vacuoles that develop into furrows

which extend in various directions and unite with one another

and with clefts from the region of the columella.

Other excellent illustrations of cleavage by constriction are

presented in the sporangia of such types as Hydrodictyon, Clado-

phora and Saprolegnia. Timberlake (:02) has given an account

of Hydrodictyon, and the events are also fairly well understood

for Saprolegnia. Segmentation begins in Hydrodictyon by the

development of cleavage furrows in the outer plasma membrane,

which cut into the protoplasmic layer at right angles to the

surface and meet similar furrows that make their way from the

large central vacuole outward. These cleavage planes spread lat-

erally, uniting with one another, until the protoplasm is all

divided into uninucleate masses which become the zoóspores

(Fig.8 c). In Saprolegnia (see Davis, :03, for general account)

conspicuous cleavage furrows develop from the central vacuole

and make their way to the exterior, finally breaking through the

outer plasma membrane. When this takes place there is an

immediate escape of cell sap, which was under pressure, and a

shrinkage of the sporangium so that the zoóspore origins appear

to fuse, but this is not really the case, for cleavage is continued

and the zoóspores soon separate.

A physiological. explanation of cleavage by constriction must

456 THE AMERICAN NATURALIST. [Vor. XXXVIII.

consider two sets of factors. There is an evident contraction of

the protoplasm in many examples because water is given off.

The shrinkage of the surface would undoubtedly form furrows,

but, as Harper has pointed out, these furrows do not develop in

an accidental manner. Non-nucleated masses of protoplasm are

never separated from the nucleated, but the segmentation pro-

ceeds after a system by which the final products contain only

one nucleus or at most a limited number. So it is probable that

the nuclei are the ultimate centers controlling the segmentation

which at its commencement may be quite irregular. This

explanation of sporogenesis in the plasmodium and the spo-

rangium is not altogether satisfactory for the cell division of

Cladophora, the abstriction of conidia or the development of the

gametes of a mould. In these examples the cleavage begins at

definite regions of the plasma membrane, so that the stimulus

must be local, and the direction of the plane has a definite

relation to the axis of the plant.

It is important to note (see Harper, :00, p. 240-249) how

inadequate are some of the well-known theories of the segmen-

tation of protoplasm as explanations of cleavage by constriction.

Hofmeister's law ('67) that cell division is across the axis of

growth obviously cannot be applied to the irregular segmentation

in the plasmodium and sporangium, nor is Sachs’ well-known law

of growth in vegetative points adequate. Sachs, '94, and in the

Lectures on the Physiology of Plants, chap. XXVII, conceives

a growing point of a higher plant or an embryonic structure asa

mass of protoplasm whose cell walls are determined by principles

of rectangular intersection of perpendicular planes. The outer

form of the structure determines the angles of periclines and

anticlines and the transversals conform to these. There is not

the slightest hint of such an order in the distribution of cleavage

planes in the multinucleate masses of protoplasm just described

and Sachs’ law in so far fails of general application whether or

not it be satisfactory for the conditions with which he especially

deals. There are also explanations of cell division, applicable to

the tissues of many higher organisms, based on the position of

the nuclear figure in the cell, which determines the position of

the cell plate but these theories cannot handle the events in the

No. 450.] STUDIES ON THE PLANT CELL. 457

plasmodium or sporangium where the cleavage planes are formed

without regard to the time of nuclear division or the position of

mitotic figures.

2. Cleavage by Cell Plates.

Cleavage of the protoplasm by means of the cell plate is

almost universal in cell division of plants above the thallophytes.

It is one of the peculiarities of plant cells, having been found in

comparatively few animals and. there represented rather imper-

fectly by the so-called mid-body. The general events of the

process have been known since Treubs' studies of 1878, and

were clearly described by Strasburger in 1880. Timberlake,

: 00, in a recent paper gives an historical review of the subject.

When, after the metaphase of mitosis, the two sets of daugh-

ter chromosomes separate from one another there is left between

them the spindle, made up of the central fibers. The first

appearance of the cell platé is a line of granules in the equatorial

region of this spindle where the nuclear plate formerly lay.

But several important events proceed this condition. The con-

necting central fibers begin to thicken, first near the daughter

nuclei, and then gradually towards the equatorial region of the '

spindle. The number of fibers may increase greatly, probably

by the separation of bundles of fibrille composing the spindle

into independent elements (Timberlake, : 00, p. 94). But there

is evidence that new fibrilla are sometimes formed from the

vicinity of the daughter nuclei, some of which may enter the

spindle and coóperate with the connecting fibers. In certain

forms (e. g. Allium) there is an accumulation of a stainable sub-

stance between the connecting fibers in the equatorial region of

the spindle. The reaction of this substance to stains indicates

a carbohydrate composition.

The cell plate really begins with the thickening of the con-

necting fibers in the equatorial plane of the spindle. In some

forms these thickenings are elongated bodies, in others mere

granules. The earlier writers (Treub, '78, Zacharias, '88) did

not believe that they came from the spindle fibers, but there

seems to be now no doubt of their origin from these elements,

458 THE AMERICAN NATURALIST. [Vor. XXXVIII.

which contract and thicken as the plate develops. The bodies

composing the cell plate finally lie in a plane extending the

entire width of the spindle (Fig. 8 7) and they then broaden

and come in contact with one another to form a continuous

membrane, which, as has been said, may lie in a matrix of car-

bohydrate material. The cell plate grows rapidly as the central

spindle fibers shorten and contribute their substance to the

structure. During this contraction the surrounding cytoplasm

b, c, cleavage by constriction. 4, cleavage by

ll division in Cladophora.

rum and female gamete nucleus within. g, egg of Ephedra with four embryo cells. After

Strasburger Harper and Timberiake.

enters the region between the barrel shaped group of fibers and

the daughter nuclei (Fig. 8 7). It is probable that the cell plate

is composed entirely of the substance of spindle fibers and in

consequence is kinoplasmic in character. The cell plate widens

with the accretion of material from the central spindle, which in

some cases is assisted by the radiating fibers that, lying outside

of the spindle, contract and add their material to the edge of the

plate. The cell plate thus extends laterally and finally reaches

No. 450.] STUDIES ON THE PLANT CELL. 459

the neighboring cell walls, fusing with the outer plasma mem-

brane. There are certain mitoses, as in some spore mother cells

and in the embryo sac (see Section III) where the cell plates

are absorbed into the cytoplasm leaving the original cell with

two or more nuclei and without partition walls. It is uncertain

whether the edge of the plate is ever extended by the develop-

ment of additional peripheral fibrille (Timberlake, : 00, p. 161)

from the daughter nuclei.

Cell division is accomplished by the splitting of the cell plate

(Strasburger, '98) into two plasma membranes. The division

generally begins in the center and the cleft progresses towards

the periphery until it reaches the cell wall. During the process

the thickened rod shaped portions of the spindle fibers are pulled

apart. There are thus left two kinoplasmic membranes opposite

one another and continuous with the outer plasma membrane

surrounding the daughter cells. The cause of this cleavage is

not apparent, but there are reasons for believing that the split

is essentially a thin vacuole which, starting near the center, cuts

its way through the cell plate to the periphery after a manner

very similar to the behavior of vacuoles during the cleavage of

the plasmodium and in the sporangia of certain moulds. And

there may be shown in this activity a relationship of cleavage by

cell plate to some of the events of cleavage by constriction.

After division is complete there follows the formation of a cell

wall between the two cell surfaces after the method usual to

plasma membranes.

The new cell wall generally begins in the oldest portion of the

cell plate where the cleft first appeared and is gradually built

out peripherally until it reaches the side walls. The first indica-

tion of the wall is the appearance in the cleft of a stainable

carbohydrate substance which resembles the material that was

primarily present between the fibers of the central spindle and

which disappears with the formation of the cell plate. This

material is probably the basis of the first deposits on the surface

of the two plasma membranes, but the nature of the final sub-

stance is exceedingly various. A cell wall may be formed that

is homogeneous throughout but often the thickened wall presents

three regions, two layers of a cellulose basis formed by the

460 THE AMERICAN NATURALIST. [Vor. XXXVIII.

respective plasma membranes and between them the so-called

niiddle lamella.

The middle lamella has been the subject of much discussion.

It is not the remains of the cell plate as was once supposed.

Neither is it exactly a cement between two cell walls. Its his-

tory is undoubtedly various, for the composition shows much

plasticity. The origin of the middle lamella at the surface of

a plasma membrane indicates a morphology similar to a cell wall,

but the substance, pectic in character, shows transformations far

removed from the cellulose compounds that are formed later and

which give thickness to the cell wall. Allen (: 01) discusses the

subject in detail.

The origin of the cell plate is a subject of interest which will

be further discussed in Section VI. There are some types, espe-

cially among the thallophytes, where a cell plate is present, but

apparently in a somewhat undeveloped and rudimentary condi-

tion. These forms suggest transitional conditions between c/eav-

age by constriction with the aid of vacuoles, so general among

the thallophytes, and cleavage by the cell plate, characteristic of

higher groups. The most interesting examples are Anthoceros,

Chara, Basidiobolus, Pelvetia, Fucus, and Sphacelaria.

Cell plates are formed with each of the two successive mitoses

in the spore mother cell of Anthoceros (Van Hook, : 00; Davis,

OI, p. 158), but the structure in some species is exceedingly

small (e. g., A. devis) and can scarcely extend more than one-

tenth of the distance across the cell. It is larger in other forms,

as in the one studied by Van Hook ; but even there the nuclear

figure of the second mitosis is only one-third of the width of the

cell. The protoplasm divides simultaneously in the four spores

with the characteristic arrangement. If this division were deter-

mined entirely by cell plates there would be required an exten-

sive development of fibrillae, of which there is no evidence in the

cell. But their place seems to be taken by numerous delicate

strands of cytoplasm which connect the four protoplasmic masses,

each of which contains a large chromatophore and an accom-

panying nucleus. A film is formed in the intermediate region,

and this marks the position of the cell wall. It is, of course,

quite certain that the two cell plates of the second mitosis are

No. 450.] STUDIES ON THE PLANT CELL. 461

a part of this membrane and may start its development, but the

final structure must contain very much more material than could

possibly be contributed by the sparsely developed spindle fibers.

Thus, although the splitting of the cell plate may start the proc-

ess of segmentation, its final coursé and end is probably deter-

mined by cleavage through vacuoles, thus utilizing a method

characteristic of the thallophytes.

Chara appears to have a fairly well developed cell plate (Deb-

ski, '97) which extends almost entirely across the cell, presenting

very exceptional conditions among the thallophytes. This pecu-

liarity is in keeping with other characters of the spindle, which

begins its development outside of the nuclear membrane and,

lacking centrosomes, resembles the nuclear figures of higher

plants. It is possible that nuclear studies upon Chara through-

out ontogeny might show a variation that would be very signifi-

cant for the evolutionary problems concerned with the structure

of protoplasm.

Fairchild ('97) reports a cell plate for Basidiobolus when the

beak cells are cut off from the gametes. The structure, as fig-

ured and described, is not, however, conspicuous. He points

out general resemblances between cell division in this form and

in the Conjugales, where, as Van Wisselingh (:02) described

later for Spirogyra, spindle fibers connect the daughter nuclei

and may coóperate towards the end of cell division with a cleav-

age furrow from the side of the cell. :

The conditions in the Fucales are not altogether clear. Both

Strasburger (97a) and Farmer and Williams (98) report that

the central spindle disappears in Fucus without the formation of

the cell plate and that the wall is developed between the daugh-

ter nuclei in a region of granular cytoplasm. However, in Pel-

vetia some of the radiating fibrille from opposite sides of the

daughter nuclei bend around these structures and end in the new

wall. It is not plain that they contribute much it anything to

its formation in the way of substance, but it would seem prob-

able that they hold a directive relation to the structure (Farmer

and Williams, '98).

The Sphacelariacez seem to be somewhat similar to the Fuca-

les in their methods of cell división. The beautiful figures of

462 THE AMERICAN NATURALIST. (Vor. XXXVIII.

Swingle (97) for Stypocaulon give details of the region of the

cytoplasm that forms the partition wall between the daughter

nuclei. There is a zone of fine meshed protoplasm between

much larger vacuoles. It is possible that some very long fibrillae

may connect the daughter nuclei with this zone, but they do not

form a cell plate. Consequently the wall must be developed in

this delicate alveolar layer, which probably splits along some

plane of vacuoles. The process of cleavage is then really related

to such activities of vacuoles as occur in the sporangium of the

Mucorales and in the plasmodium. But the position of the alve-

olar layer may be determined by the fibrillae, since it is always

situated nearest to the smaller of the two daughter nuclei.

It seems likely that the process of cleavage in the Fucales will

be found to be similar to Stypocaulon when the details of struc-

ture in the internuclear cytoplasm is known. So this group, with

others, is likely to furnish conditions in which spindle fibers may

determine the position of the cell wall and exert a directive influ-

ence upon it without actually laying down a cell plate. As has

been pointed out, the splitting of the cell plate is probably a

cleavage along a very thin flat vacuole, so that the process in its

essential characters is the same as cleavage through a series of

vacuoles. Thus cleavage by the cell plate is possibly an out-

growth from that phase of cleavage by constriction in which the

extensive fusion of vacuoles determines the planes of separation.

The important advance lies in the new factors, introduced through

the activities of fibrille, which become very conspicuous as actual

contributors of material to the kinoplasmic film which is laid

down as the cell plate. This function of the fibrilla probably

developed slowly from conditions such as those in Stypocaulon

and Pelvetia, where their influence upon the position of the cell

wall, if any at all, can scarcely be more than directive.

3. Free Cell Formation.

Whenever a nucleus becomes the center around which cyto-

plasm is gathered and separated from the rest of the cell con-

tents, so that the new cell lies freely in the protoplasm of the

old, this is free cell formation. Illustrations are presented by

No. 450.] STUDIES ON THE PLANT CELL. 463

the spores of an ascus, the oóspore of the Peronosporales, the

embryo cells of Ephedra, and probably other gymnosperms, and

in some cases seemed to be exemplified in the conditions pre-

sented by the egg and synergids and the antipodals of the embryo

sac.

Spore formation in the ascus is known through the studies of

Harper ('97 and '99). After the final divisions in the ascus the

nuclei lie in the cytoplasm, each with an aster at its side (Fig.

8 e, 3). A delicate prolongation carries the aster with its cen-

trosphere away from the main body of the nucleus (e, 1). The

rays of the aster now bend over and grow around the nucleus,

presenting an umbrellalike figure (e, 2). They finally meet on

the opposite side, and thereby cut out a portion of the cytoplasm

which is included in the spore. The substance of the aster

fibers forms the basis of a kinoplasmic film which becomes the

plasma membrane of the ascospore and develops the spore wall

externally after the usual method. This peculiar activity of an

aster is unparalleled in plant or animal cells.

Oógeriesis in the Peronosporales has been described in some

detail by several authors, but the process has not generally been

called free cell formation. Yet at the end of the process the

oóspore, enveloped by periplasm, lies free in the oógonium. In

the beginning the oóplasm gathers in the center of the oógonium

as a denser alveolar region around that peculiar protoplasmic

body (generally present) the coenocentrum. This accumulation

forces the vacuoles, together with most of the nuclei, to the

periphery, where they lie in a sort of protoplasmic froth next

the cell wall and constitute the periplasm. The spore wall de-

velops at the boundary of the oóplasm, so that it lies close to the

large vacuoles (Fig. 8 f) in the periplasm. There must be an

accumulation of kinoplasm, perhaps from the plasma membranes

of numerous vacuoles, to form a delicate layer between the two

regions of the odgonium. This layer of kinoplasm probably

splits along the line of vacuoles between the oóplasm and peri-

plasm, for the primary walls are certainly established between

two plasma membranes, because the secondary layers are added

to it from both sides. Nuclei in division frequently lie very

close to the boundary of the oóplasm, but there is no evidence

464 THE AMERICAN NATURALIST. (VoL: XXXVIII,

that the kinoplasmic membrane has any relation to these mitotic

figures. That is to say, there are no fibrillae to contribute sub-

stance to the membrane, and its development must be concerned

with vacuoles alone. In this respect the process recalls the part

played by vacuoles in the plasmodium and in certain sporangia

during cleavage by constriction.

Free cell formation after the method in the egg of Ephedra

(Strasburger, '79), which is also likely to be found among other

gymnosperms, takes place during the differentiation of the em-

bryo cells. The cytoplasm collects around each nucleus, forming

a sphere (Fig. 8 g), and a wall is developed on the outside of

this body. Details of the process are not known, and it is not

clear whether the position of the membrane is determined by the

vacuoles that must border upon this region or whether there are

fibers radiating from the nucleus which might. lay down a cell

plate around the denser VUE m Ren but the evidence favors the

former possibility.

Somewhat similar conditions are presented in the egg appa-

ratus of many embryo sacs. In certain forms (e. g., the lily so

well described by Mottier, '98) the egg nucleus and synergids are

thickly invested by radiating fibers, and these, together with the

cell plates, may readily determine the position of the plasma

membrane that forms the cell wall. But fibers do not seem to

be conspicuously present in the egg apparatus of many other

embryo sacs (Excellent illustrations can be found among the

Ranunculacez). In these cases the protoplasm collects around

the nuclei as dense areas bordered by vacuolar cytoplasm, and it

is possible that the vacuoles by fusing with one another cut out

these respective regions and thus determine the plasma mem-

branes of the egg and synergids. Such processes would extend

the activities of vacuoles, which accompany cleavage by constric-

tion in the thallophytes, to the highest groups of plants.

It is curious that with all of the work upon the embryo sac

we should know less about the segmentation of the protoplasm

around the synergid, antipodal, and segmentation nuclei in this

structure than in the sporangia of the molds, the ascus, or dur-

ing spore formation in the Myxomycetes.

(To be continued).

No. 450.] SPUDIES ON IHE PLANT CELL. 465

LITERATURE CITED FOR SECTION 1I.

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CHAMBERLAIN.

'99. Odgenesis in Pinus laricio. Bot. Gaz. 27, 268.

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'99. The spore mother-cell of Anthoceros. Bot. Gaz. 28, 89.

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DEBSKI.

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FARMER AND REEVES.

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FARMER AND WILLIAMS

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466 THE AMERICAN NATURALIST. [Vor. XXXVIII.

FARMER AND WILLIAMS.

'98. Contributions to our knowledge of the Fucacez ; their life history

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FULMER.

'98. Cell division in Pine seedlings. Bot. Gaz. 26, 239.

GUIGNARD.

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HARPER.

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Hor

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HOFMEISTER.

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'98. Untersuchungen über die Entwickelung der Geschlechtsorgane und

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JUEL.

199, Le Kerntheilung in den Pollenmutterzellen von Hemerocallis fulva

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KARSTEN.

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LAUTERBORN.

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McCows.

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No. 450.] STUDIES ON THE PLANT CELL. 467

MIYAKE.

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MOORE.

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468 THE AMERICAN NATURALIST. [Vor. XXXVIII.

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No. 450.] STUDIES ON HHE PLANT CELL. 469

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NOTES AND LITERATURE.

ZOOLOGY.

Notes on Recent Fish Literature.— In the Transactions of the

Liverpool Biological Society (XVII, 1903) Mr. Walter M. Tattersall

takes up again the classification of the lancelets in connection with

the study of Professor Herdman’s collection from Ceylon.

Tattersall recognizes about 12 species, with two or three marked

“varieties.” These he arranges under two generic heads, Branchios-

toma and Asymmetron. As a synonym of Asymmetron (1893) he

places Epigonichthys (1876). The last mentioned name has of

course priority over Asymmetron, if the two are identical. * Sub-

genera" Tattersall proposes to “abolish altogether ” because most

of those proposed contain but a single species ; an inadequate reason

for those who mark divergence of character rather than number of

species, by the use of subgeneric names. Mr. Tattersall regards the

American Æ. caribeum as identical with B. /anceolatum of Europe,

which conclusion is not unlikely, as the only difference known is the

usually smaller number of post-anal muscular impressions in the

American form (9 or 10 instead of ro, 11, 12 or 13). The Japanese

Lancelet (B. japonicum =B. nakagawe) is the same as the Ceylon

species, B. belcheri, and that again is indistinguishable from the

European. It has 63 or 64 muscular impressions while B. /anceola-

tum has 60 to 63 and B. caribeum 58 to 61r.. The averages of

specimens counted indicate a slight increase in number of segments

in Asiatic specimens, a slight decrease in American.

Tattersall rejects the genus Amphioxides, based on 44. pelagicus

and characterized by the absence of oral cirri “ because the absence

of oral cirri is no doubt a result of its pelagic life.” But if pelagic

life brings about such important structural modifications, a matter by

no means proved, this is no reason why we should not regard these

modifications as of generic value. If Amphioxides is really without

cirri, as is probably the case, it is surely a valid genus.

Mr. Tattersall states that “Jordan and Snyder consider Z. naka-

gawe a new species solely on account of its geographical distribution.”

In this he seems to miss the point. B. belcheri was described from

Borneo in very unsatisfactory fashion. The Japanese lancelet showed

471

472 THE AMERICAN NATURALIST. [Vor. XXXVIII.

points of difference. As in almost every case, the shore fishes of

Japan are specifically different from their analogues in Borneo, it is

antecedently probable that the lancelets differ also. It appears that

this is not the case, as a correct account of B. belcheri agrees sub-

stantially with B. nakagawe (earlier called japonicum). Yet Mr.

Tattersall finds a certain average difference. The commonest for-

mula of myotomes in B. belcheri is 37-17—9, while in B. japonicum it

is 36-17-11. As these little creatures have so few tangible charac-

ters, minute differences which can be made out have a greater

relative importance.

In the Bulletin of the Museum of Comparative Zoólogy (Vol. 39,

No. 8) Mr. Samúel Garman gives an account of the fishes taken by

Mr. Alexander Agassiz and his party on the “ Albatross,” about the

coral reefs of Fiji and the great Barrier Reef of Australia. Fourteen

new species are described and well figured, nearly all of them from

Fiji.

In the Bulletin of the Fish Commission (1903), Dr. Oliver P.

Jenkins gives a final account of the splendid collection of fishes

made by him in Hawaii in 1889. Two hundred fifty-four species are

included in this connection, and in this and two preceding papers

ninety-three species are described as new. The fauna of Hawaii is

essentially that of the tropical Pacific. ‘The same genera occur as in

the other islands, but not all the genera. Many of the types charac-

teristic of the south seas, as Periophthalmus, Synancidium, Variola,

Terapon, Cassia, never reach Hawaii. A large percentage of the

species of Hawaii are peculiar to that archipelago. Thus although

Hawaii, like other groups of Islands, has Scari, Holocentri, Gobies,

etc., it has its own species in these groups, for the most part unlike

those found in Samoa or Tahiti. The faunal isolation of Hawaii may

be due in part to the direction of the currents, which set to the west-

ward, while at Samoa their general direction is eastward. The new

genera in Jenkins' paper are Scaridea, Cirrhitoidea, Eviota, Chlamy-

des, the last two being gobies. The excellent plates in this paper

are by Mr. William S. Atkinson. j

In the Proceedings of the U. S. National Museum (XXVI, 1903),

Dr. Gill takes up Dr. Boulenger's studies of the bones of the Opah,

Lampris luna. Dr. Gill makes a very different interpretation of the

osteology of the shoulder girdle from that of Dr. Boulenger. He

finds the so-called infraclavicle to be the hypocoracoid, and believes

that the bones of the Opah differ little from those of the related

No. 450.] NOTES AND LITERATURE. 473

mackerel-like fishes. He finds no warrant in associating the Opah

with the sticklebacks to form a group Catosteomi. The present

writer agrees fully with Dr. Gill in his view of this matter.

In the Proceedings of the U. S. National Museum (XXVI, 1903),

Dr. Gill discusses the generic names in a forgotten work of Heinrich

Friedrich Linck, 1790. “Versuch einer Eintheilung der Fische nach

der Zähnen ” in * Magazin für das Neueste aus der Physik und

Naturgeschickte,” published at Gotha. The paper is without value,

but some of its generic names antedate those in common use. ‘These

are Mustelus for Mustelus levis = Squalus mustelus. This antedates

the use of the name for Mustelus canis, for which Dr. Gill suggests

the new group name, Cynias. Pristis and Mola of Linck are equiv-

alent to the genera later so named by Latham and Cuvier. The other

new generic names are Rhinobatos (without type indicated), Cal-

lichthys (no type), Alosa (no type), Thymallus (no type), Soarus

(definition unintelligible), Barbatula (— Cobitis).

Gill further shows that,the name Macrodon Schinz (1822) was

intended for’the genus of Sciznoid fishes called Ancylodon by

Cuvier in 1817, preoccupied by Ancylodon Illiger 1811, and later

named Sagenichthys by Berg. The type of the Sciznoid genus

should, therefore, stand as Macrodon ancylodon.

For the genus of Erythrinidz called Macrodon by Müller in 1842,

Gill substitutes the name Hoplias.

He further calls attention to the fact that Oken in 1817 (in Zs/s)

gave classical names to the genera of fishes left with French names

only by Cuvier in the first edition of the Régne Animal of the same

year. From this work, the following names must date: Monacan-

thus, Alutera, Triacanthus, Curimatus, Piabucus, Cirrhinus, Bagre,

Lota, Brosme (not Brosmius,a later form), Monochirus, Aurata,

Plectropomus, Priacanthus, Stellifer, Sander (— Lucioperca), Zingel,

Otolithes and Chelmo.

In the same Proceedings, Mr. B. A. Bean figures the rare eel AA/ia

egmontis from Barbados. It has been received also from the Tor-

tugas. :

In the Mark Anniversary Volume, Dr. C. R. Eastman discusses

again the character of the extraordinary structures found in Carbon-

iferous rocks and known as Edestus, with a bibliography of the

subject. These are now believed to be coalescent whorls of teeth of

Some cestraciont shark.

474 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In the Proceedings of the Washington Academy of Sciences (Vol. V,

pp. 189-229), Messrs. Edmund Heller and Robert E. Snodgrass

give an account of the new species of fishes taken in their expedition

to the Galapagos under the auspices of the Hopkins Seaside Labora-

tory of Stanford University. The following are the new genera and

species : 5

Evolantia for Æxæoctus micropterus, Sphyrena idiastes, Apogon

atrodorsatus, Galeagra pammelas (a new genus allied to Acropoma),

Corvula eurymesops, Scena perissa, Azurina eupalama, Pomacentrus

redemptus, Pomacentrus arcifrons, Nexilosus albemarleus (a new

genus allied to Hypsypops), Scarus noyesi, Pontinus strigatus, Eleotris

tubularis, Cotylopus cocoénsis, Gobius rhizophore (a species of Cory-

phopterus), Gobius gilberti (a species certainly referable to the Japan-

ese genus Pterogobius), Arbaciosa truncata, Malacoctenus zonogaster,

Lepisoma jenkinsi, Encheliophis jordani, Petrotyx hopkinsi, Eutyx

diagrammus (Petrotyx and Eutyx are new genera of Brotulide),

Antennarius togus, Allector chelonia (Allector is a new genus allied

to Chaunax). A list of all the species constituting this splendid col-

lection will appear later.

In the Proceedings of the Biological Society of Washington (XVI,

1903), Austin H. Clark shows that the earliest name of the common

American Eel is Anguilla chrisypa Rafinesque, the name meaning

“besmeared or anointed below." Rafinesque calls it “Gold-breast.”

In the Bulletin du Museum a’ Histoire Naturelle Paris (1903), Dr.

Pellegrin describes a number of new species of Cichlide, one of

them, Heros ( Cichlasoma) labridens, being from Huasteca Potosina,

from the veteran naturalist, Dr. Alfredo Dugès of Guanajuato. It is

near Cichlasoma bartoni. Several others are from Guiana.

In the Bulletin of the U. S. Fish Commission (1902) Professor W.

J. Mcenkhaus describes a new species of Darter, Zadropterus ever-

manni from Lake Tippecanoe in Indiana.

D: 6) 9)

BOTANY.

Notes. — Professor Ganong's address on the cardinal principles of

ecology is published in Science of March 2 Ss

Daniel gives an account of a graft-hybrid between pear and quince

in the Revue Générale de Botanique of Jan. 15.

No. 450] NOTES AND LITERATURE. 475

An account of DeVries’ mutation theory, with portrait, is pub-

lished by Harris in Zhe Open Court for April.

The vegetative vigor of hybrids and mutations is considered by

Cook in a leaflet of Proceedings of the Biological Society of Washing-

ton, issued April 9.

Nelson describes a number of new flowering plants from Nevada,

and proposes new names for ten old species known under preoccu-

pied names, in leaflets of Proceedings of the Biological Society of

Washington, issued on April 9.

Pammel publishes an account of some weeds of Iowa as Bulletin

70 of the Experiment Station of that State.

Miss Perkins has issued (Gebrüder Borntreger, Leipzig, January,

1904) the first fascicle of * Fragmenta Flore Philippinz," embody-

ing the results of studies carried on at the Berlin Museum and based

on the collections of Warburg, Merrill, Ahern and others.

Merrill has published, from the Manila Bureau of Government

Laboratories, papers on new or noteworthy Philippine plants and

the American element in the Philippine flora.

A list of the plants known from Siam, by Williams, is being pub-

lished in current numbers of the Bulletin de T Herbier Boissier.

Vol. 4, Sect. 2, of the Fora Capensis, under the editorship of This-

elton-Dyer, runs from Hydrophyllacez into Scrophulariacez.

A number of views of the vegetation of Samoa accompany an arti-

cle on that group of islands, by Kellogg, in Out West for April.

Vol. 5, fascicle 1, of Coste's “ Flore descriptive et illustrée dé la

France” deals with Scrophulariacez and Labiate.

An analysis of the vegetation of Madeira, by Vahl, has been issued

from the Gyldendalske Boghandel, of Copenhagen.

A posthumous paper by Weber, published by Roland-Gosselin in

January from the Buletin de la Société Centrale d Agriculture, etc., de

Nice, deals with the restored genus Cleistocactus.

Certain Arizona cacti are illustrated in Zhe Gardener's Chronicle

of March 19.

Country Life in America for April contains an illustrated article by

Julia E. Rogers on Magnolias.

476 THE AMERICAN NATURALIST. [Vor. XXXVIII:

An account of Mexican tea (Turnera aphrodisiaca) is published by

J. U. Lloyd in the Pharmaceutical Review for April.’

A note on Rudbeckia, by Pihl, in Svenska Tradgardsforeningens

Tidskrift of January, is illustrated by colored figures of several culti-

vated species.

A series of ten illustrations, showing the development of an elm

shoot, by Richards, is published in Country Life in America for April.

A third signature of Professor Greene’s “Leaflets” continues his

study of Polygonacez.

Holm's “Studies in the Cyperacee — XXL," published in Zhe

American Journal of Science for April, deals with new or little known

species of Carex.

Parish begins a preliminary synopsis of Southern California Cy-

peracee in the Bulletin of the Southern California Academy of Sci-

ences for March.

Certain Canadian mosses are enumerated by Cufoni in No. 7-9 of

the Bullettino della Società Botanica Italiana of 1903, which also con-

tains a note by Baroni on the botanic garden of Mt. Ætna.

The necessity for reform in the nomenclature of fungi is discussed

by Earle in Science for March 25.

Copeland has an article on Californian fungi in Annales Mycologict

for January.

Rehm publishes diagnoses of a number of North American Asco-

mycetes in Annales Mycologici of January.

An account of two hundred Portuguese fungi has been distributed

by Professor d'Almeida of Lisbon.

Klebahn's “Die wirtswechselnden Rostpilze " (Gebrüder Born-

treger, Berlin, 1904) forms a large volume containing a detailed

analysis of heteroicism followed by an account of the species show-

ing this peculiarity and ample indexes to fungi and hosts.

Taphria cerulescens is the subject of Bulletin 126 of the Alabama

Experiment Station, by Wilcox.

Salmon gives an account of recent researches on the specialization

of parasitism in Erysiphacez in Annales Mycologici for January, and

The New Phytologist of Feb. 27.

An account of dry rot of the potato, caused by Fusarium oxyspo-

No. 450.] NOTES AND LITERATURE. . 477

rum, by Smith and Swingle, is published as Bulletin 55 of the Bureau

of Plant Industry of the U. S. Department of Agriculture.

Fungous diseases form the subject of Bulletin 63 of the Delaware

College Experiment Station, by Chester and Smith.

Oudemans has distributed separates from the Proceedings of Jan.

30 of the K. Akademie van Wetenschappen te Amsterdam, describing

Exosporina Laricis, a new destructive parasite of the larch.

Arthur describes and figures the three common edible species of

Coprinus in Bulletin 98 of the Purdue University Experiment Station.

A paper by Stahl on the means by which lichens are protected

against feeding animals is separately distributed from the Festschrift

commemorative of Heckel’s oth birthday, issued by the Fischer

press of Jena.

The development of lichen apothecia is considered by Baur in the

Botanische Zeitung, Abteilung I, of March tr.

Coville has recently distributed separates of a well illustrated

paper on the Indian use of Mymphea polysepala, from the Report of

the U. S. National Museum for 1902.

An account of a new African fiber-banana, Musa ulugurensis, is

given by Warburg and Moritz in Der Tropenpflanzer for March.

Van Es and Waldron give an account of stock-poisoning plants of

North Dakota in Bulletin 58 of the Experiment Station of that State.

An illustrated account of lumbering by machinery is contributed

by K. Smith to ZZe World’s Work of February.

Clement is publishing a series of illustrated articles on “the Japan-

ese floral calendar” in current numbers of Zhe Open Court.

The reports on the New York Botanical Garden for 1903, con-

tained in Vol. 3, No. to, of the Bulletin of the institution, show that

84,163 specimens were added to the herbarium, 1,656 bound volumes

were added to the library, and 1,000 species were added to the living

collection, bringing the latter up to about 11,600 species.

An illustrated account of the Arnold Arboretum, by Miller, is pub-

lished in Country Life in America for March. |

40,396 visitors to the conservatories of the St. Petersburg Botani-

cal Garden, in 1903, are reported in its Buletin of March 5.

A portrait of Behrens is published as frontispiece to Heft 79 of

the Zeitschrift für wissenschaftliche Mikroskopie.

PUBLICATIONS RECEIVED.

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ANNANDALE, N. and RoBiNsoN, H. C. Fasciculi Malayenses Antl ological

and Zoólogical Results of an Expedition to Perak and the Siamese Maylay ipa:

1901-1902. Pt. I, Zoólogy, pp. 1-200, pls. 1-10; Pt. II, Zoólogy, pp. 201-307.

pls. 11-14-vii; Supplement pp. xlii and map.— BAILEY, L. H. Plant Breeding.

Being five Lectures upon the Amelioration of Domestic Plants. Third Edition,

New York, Macmillan, 1904. Svo, xiii + 324 pp., 22 figs.— BASTIAN, H. C.

Studies in Heterogenesis. London and Oxford, Willams and Nargate, 1903.

8vo, viii + 354 + xxxiii, pp. , 815 figs. 31/6.— BERGE N,J. Y. Notebook to Accom-

pany Bergen's Text Books of Botany or for suiit use in Botanical site cap

T Secondary Schools. Boston, Ginn & Co.,1904. 4to, 144 pp» $0.7 5.—

A. Mammalian ume, with Special 1éfinínie to the Cat. Philadelphia,

Blakiston, 1903. 8vo, xi + 250 pp., 108 figs. $1 1.50.— GARDINER, J. S. he Fau

and Geography of the Maldive and Laccadive Archipelagoes, etc. Vol. II, Pu

III, pp. 589-698, pls. 35-48.— GIRARD, A. Controverses Transformistes. Paris,

C. Naud, 1904. 8vo, 178 pp., 23 figs.— LEIGHTON, G.R. The Life-History of

British Lizards and their Local Distribution in the British Isles. Edinburgh,

Geo. A. Morton, 1903. 8vo, xiv + 214 pp., many illustrations.— MOUILLEFERT, P.

Traité de Sylviculture. mires et Aménagement des Bois. Paris, Alcan,

1954. Svo, 476 pp., 10 pls., 27 figs. 6 francs.— MouNEYRAT, A. La Purine et

ses Dérivés. Meu uni Biologique, N o. 18. Paris, C. Naud, 1904. 8vo,

98 pp.— Nites, Grace G. Bog-trotting i Orchids. New York, Putnam's,

1904. 8vo, xvi + 310 pp. 72 plates, 24 of them colored.— PERRY, J. H. and

EMERSON, B. K. The Geology of Worcester, Massachusetts. ergata Nat-

ural Hist. Soc., 1903. 8vo, xii-166 pp., plates and maps.— PogTER, T. C. Flor

of Puri: Boston, Ginn & Co., 1904. 8vo, xv v + 362 pp. PAE TEA

T. C. Catalogue of the Bryophyta (Hepatic, Anthocerotes and Mosses) and

fucum (Ferns and Fern-allies) f € in Pennsylvania. Boston, Ginn &

1904. 8vo, 66 p.— REESE, A. ek n Introduction to Vertebrate Embry-

sin Based on the Study of the Frog dd the Chick. New York, Putnam's,

1904. Svo, xvii + 291 pp., 83 figs.— SckIBNER, G. H. Where did Life Begin ?

A Brief Inquiry as to the Probable courses of Migration therefrom of the Flora

and Fauna of the Earth. A Monograph. New York, Scribner’s, 1903. 8vo,

etai s portrait. $1.20.— SNYDER, H. The Chemistry of Plant and Animal

Li w York, Macmillan, 1903. 8vo, xvii + 406 pp: 102 figs. $1.40.—STEIN-

MANN, P Einführung in die Paläontologie. — a, 1903. 8vo.

ix + 466 pp., 818 figs. 12 poem gus Ew e Geometry. Boston,

Ginn & Co., 1904. 8vo, viii + 171 P ., 58 figs.

ARNOLD, R. The Paleontology id Stratigraphy of the Marine Pliocene and

Pleistocene of San Pedro, Cal. Contrib. to Biol. Hopkins Seaside Lab., Stanford

Uni o. xxxi + 420 pp., 37 pls.— BALL , E. D. Descriptions of Some New

Tree’ k open from the United States. nu. Biol. Soc. Wash., Vol. xvi, pp-

479

480 THE AMERICAN NATURALIST. [Vor. XXXVIII.

177-182.— Bancrort, F. W. and EsrTERLYy, C. O. A Case of T ALL Pol-

arization in the Ascidian Heart. Univ. Cal. Publ., ZoÓl., Vol. I, pp. 105-114.—

BARBOUR, T. A New Batrachian from Sarawak, Borneo. Proc. Biol. Soc. Wess

Vol. xvii, pp. 51-52.— BERRY, E. W. Oralia in pies Paleobotany. Bot.

Gazette. Vol. xxxvi, pp. 421-428.— BERTRAND, C. E. Les Coprolithes de

Bernissart. Mem. Mus. Roy. d' Hist. Nat. Ped Tom. I, 154 pp., 15 pls —

‘BOUDREAU, W. J. odern Rice Culture. Farmer's Bull. Rureau WR EL

0. 3, 46 pp., 16 figs.— BoULANGER, E. Les Mycelium Truffiers Blancs. Paris,

Oberthur, 1903. 4to, 23 pp., 3 pls.— BOULANGER, E. re de Liasco-

spore de la Truffe. Paris, Oberthur, 1903. 4to, 20 pp., 2 pls.— BREED, R. S.

The Changes which Dur in the Muscles of a Beetle, d mainc

Chem. during Metamorphosis. Bull. Mus. Comp. Zoöl, Vol. XL, pp. 317-382, 7

. pls.— Burrum, B. C. Wheat Growing on the on Plains. Bull. Wyo. Agr.

Exp. Sta. No. 60, 39 pp., pls.— CAPE or Goop Hope. Department of pilti

ture. Annual Report of the Geological Commission. 1901,67 pp., map. 1902,

128 pp., — Mpeg T. D. A. The Colorado Rubber Plant. Bull. Col.

Mies M No. 1, 2 pp.— COMISION DE PARASITOLOGIA AGRICOLA. Circular

— COMISION DE jc litat AGRICOLA. Las Plagas de la Agricultura.

t1.— Cook, O. F. The Vegetative Vigor of Hybrids and Mutations. Proc.

pon "Soc. eR Vol. xvii, pp. 83-90.— DALL, W. H. . Diagnoses of New Species

of Mollusks from the ese apu Channel, California. Pros. Biol. Soc. Wash.

Vol. xvi, pp. 171-176.— DALL, W. H. and Bartscu, P. Synopsis of the Genera,

Subgenera and ricis s pu Family japant Proc. Biol. Soc. Wash

Vol. xvii, pp. I — Davenport, C. B. Report on the Fresh-Water Bryozoa

of the United duis Proc. U. S. Natl. Mus. Vo He xvii, pp. 211—221, pl. 6.—

DEAN, BASHFORD. Notes on Japanese Myxinoids. Jour. Coll. Sci. Imp. Univ.

okyo. Vol xix, 23 pp., 1 pl.— DEAN, BAsHFORD. Notes on the Long-Snouted

Chimaeroid of Japan. wo (Harriotta) Pacifica (Garman) Mitsikurii.

Jour. Coll. Sci. Imp. Univ. Tokyo. Vol. xix, 15 pp., t pl.— DEAN, BASHFORD.

Notes on Chimera. Two Japanese Species, C. phantasma Jordan and Snyder,

and C. Mitsikurii N. S., and their Egg Cases. Jour. Coll. Sci. Imp. Univ. Tokyo.

Vol. xix, 9 pp., 1 pl.— Dickson, C. W. The Ore Deposits of Sudbury, Ontario.

Trans. Amer. Inst. M. Engineers, for 1903, 65 pp, 26 figs.— DonsEY, G. A.

The Arapho Sun Dance; The Ceremony of the Offering Lodge. Field Columb.

Mus Publ. Anthrop. Ser. Vol. IV, xii + 288 pp. 137 pls.— DORSEY, G A. Tr

ditions of the Osage. Field Coumb. Mus. Publ. Anthrop. Ser. Vol. vii, pp. 9-60.

— Dorsey, G. A. and KROEBER, A. L. Traditions of be Arapho fn under

the Auspices of the Field Columbian Museum and of the American Museum of

Natural History. Field Columb. Mus. Publ. ma Vol. v, x + 475 pp.—

Dorsey, C. W. Soil Conditions in the sgg pci Bull. Bureau Agr. P. 1.

No. 3, 57 pp. maps and plates.— DonsEv, C. Quien of Tobacco.

Farmer's Bull. Bureau Agr. P. 7., No. p 20 pp., 4 figs.— Dorsgy, C. W.

SER on the Agricultural Soils of Union Province, Luzon. Bull. Bureau Agr.

Actinians. Bull. Amer. Mus. Nat. Hist. Vol. xix, PP- 495-503. pls. 44-47.—

Epwarps, H. T. Maguey in the Lege Phil. Bur. Agr., Farmer's Bull.

No. 10, 14 pp.— ELrior, D. G. escriptions of Twenty-seven Apparently New

Mus. Publ., Zoól. Ser. Vol

iii, pp. 239-259.— ELLIOT, D. G. Catalogue of Mammals Coliected by C. Heller

No. 450.] PUBLICATIONS RECEIVED. . 481

in Southern California. Field Columb. Mus. Publ, ZoOl. Ser. Vol. iii, pp. 271-

321, pls. 38-50.— ELLIOT, D. G. escriptions of Apparently New Species of

Mammals of the Genera jones s and Ursus from Washington and Mexico.

Field Columb. Mus. Publ., . Ser. Vol. iii, pp. 233-237.— ELLIOT, D. G.

List of Mammals Pies. er Edmund Heller in the San Pedro Martir and

Han una Mountains and the Accompanying Coast Regions of Lower

Cons with Descriptions of Apparently New Species. Field Columb. Mus

Publ. Zool. Ser. Vol. iii, pp. 199-232, pls. 33-38.— ELLIOT, D. G. Doicilpdións

of Apparently New Species and Subspecies of Mamma b. and a New Generic

Name Proposed. pues Columb. Mus. Publ, Zool. Ser. Vol. iii, pp. 263-270.—

FARRINGTON, O. C. Catalogue of the Collection of Meteorites. reld Columé.

Mus. Publ., Geol. Ser. Vol. ii, pp. 79-123, pls. 30-39.— FELT, E. P. Grapevine

oot Wo Bull. N. Y. State Mus., No. 72, 55 pp., 13 pls T, E.P. Insects

Affecting pa orest Trees. From Seventh Ann. Rept. Forest, Fish and Game Comm.,

PP- 479-534, 66 pls., 26 figs.— aN G, S. Preliminary bipes of Field Work

in the Town of Minerva, Essex Co., Rept. N. Y. State Geol. for 1900, pp.

91-1035, pl. 8.— Gasser, H. The n Idea. Medical RM: Aug. 1903,

30 pp.— GILBERT, C. H. and SrARKs, E. C. The Fishes of Panama Bay. Contrib.

to Biol. api Seaside Laboratory, Stanford Univ., No. xxxii, » pp. 33 pls.—

GOLDMAN, E. A. Descriptions of five New Mammals from Mexico. Proc. Biol.

Soc. Wash. Vol. xvii, pp. 79-82.— GRABAU, A. W. Notes on ai Development

of the Biserial Arm in Certain Crinoids. Am. Jour. Sci. Vol. xvi, pp. 289-300,

11 figs. — GRARAU, A. W. Stratigraphy of Becraft Mountain, Columbia PRA

N.Y. Rept. N. Y. State Paleont. 1902, pp. 1030-1079, 13 figs, maps.— GRAB

A. W. Paleozoic Coral Reefs. Bull. Geol. Soc. Amer. Vol. xiv, pp. daia

pls. 47-48.— HALLOCK-GREENWALT, Mary. Prose and Rhythm. Pop. Sci.

IM Sept. 1903, pp. 425-431.— Hare, R. F. Canaigre. Bull. N. Mex. Agr.

. Sta. No. 49. 15 pp.— Hart, C. A. Synopsis of Insect Collections for Dis-

— in Illinois High Schools. ZZ. State Lab. Nat. Hist., 64 pp., 74 fi

HERRICK, C. e The Doctrine of Nerve Components and Some of its ven

tions. Jour. Comp. Neurol. Vol. xiii, pp. 301-312.— JOHNSON, D. W.. Block

Mondta i in gend Mexico. -Amer. Geol. Vol. xxxi, pp. 135-139.—JORDAN, D.

S. Notes on Fishes Collected in the Tortugas Archipelago. Bull. U. S. Fish

Com. for 1902, pp. 539-544, 2 pls.— JORDAN, D. S and SNYDER, J. O. On the

Species of White Chimera from Japan. Proc. U. S. Natl. Mus. Vol. xxvii, pp.

223-226.— JORDAN, D. S. and Starks, E. C. A Review of the Cottide or

Sculpins dies in the Waters of Japan. Proc. U. S. Natl. Mus. Vol. xxvii, pp.

231-335, 43 figs.— JoRDAN, D. S. and Starks, E. C. A Review of the Japanese

Fishes of ^ Family of Agonida. Proc. U. S. Natl. Mus. Vol. xxvii, pp. 575-

599. 13 figs.— JORDAN, D. S. and STARKS, E. C. A Review of the Scorpenoid

Fishes of Japan. Proc. U. S. Natl. Mus. Vol. xxvii, pp. em -175, pls. 1-2.—

nd. Aca

Jupay, C. The Plankton of Wina Lake. Proc. Ind. Acad. Sci. 1902,

120-133.— KANDA, M. Studien über die noc einiger Metallsalze sr

das Wachsthum höheren Pflanzen. Jour. Coll. Sci. Imp. Univ. Tokyo. Vol. xix,

37 pp. 1 pl — Kemp, J. F. A New Spheroidal came deibnee: Vol. tica pp-

Tectonique des Alpes Francaises. nn. Univ. Grenoble. Tom. xvi, 10 pp.—

Koroip, C. A. The Plankton of the Illinois River, 1894-1899, with introductory

482 i THE AMERICAN NATURALIST. (Vor. XXXVIII.

Notes upon the Hydrography of the Illinois River and its Basin. Part I.

— Q Investigations and general results. Bul. ///. State Lab. Nat. Hist.

Vo pp. xviii and 95-629, 50 pls.— LAMBE, L. M. Lower Jaw of Dryto-

saurus perae (Cope). Ottawa Nat. Vol. xvii, s 133-139, 3 pls.— Law-

SON, A; C. The os Gabbro at Dehesa San Diego Co., Cal Univ. Cal.

Publ. Bull. Dept. Geol. Vol. iii, pp. 383-396, pl. PEIEE W. m A Pri

on the Cultivation of pole Cane. Farmers Bull. Bureau Agr. P. I. No. 1, 19

p.— Lyon, W. S. Report on the Introduction and Distribution of Seeds and

Plants by the Bureau of Agriculture. Farmer’s Bull. Bureau Aor. Pads NO,

18 pp., 3 pls.— LvoN, W. S. Cacao Culture in the PURUS, Farmer's Bull.

Bureau Agr. P. I. No. 2, 25 pp.— Lucas, F. A. New Batrachian and a New

“ea from the Trias of Arizona. Proc. U. S. Natl. Mus. Vol. xxvii, pp. 193-

195, pls. 3-4. — Mason, O. T. Aboriginal American Basketry ; Studies in a Tex-

tile w sion t Mahisa Rept. U: S. Natl. Mus. for A ds pp- 171-548, 248

pls.— McGRE . C. Birds from Benguet Province, Luzon and from the

Islands of Mon Modes. ud and resin, Eu Philippine Mus.

sU. ne

23 16 p

in California. Univ. Cal. Pu = Bull, Dept. Geol. Vol. iii, -381.

M, J. C. The Pliocene and — Canidz of the Great Volar of

— MERR

Cone ei Cal. Publ., Bull. Dept. Geol. Vol. iii, pp. 277-290, pls Siren

— MERRILL, E. D. Botanical Work in the Philippe Bull. Bureau Pe

Z. No. 4, 53 pp. 1 pl— MILLER, G. S., Jr. w Hare from Greece.

Proc.

Biol. Soc. Wash. Vol. xvi, pp. SS UR D G. S., JR. Descriptions of

two New Mole Rats. Proc. Biol. Soc. Wash. Vol. xvi, pp. 161-164.— MILLER,

G. S., JR. A Second Specimen of Euderma Maculatum. Proc. Biol. Soc. Wash.

Vol. xvii, pp. 165-166.— MILLER, G. S., JR. New PO from Lower Siam.

Proc. Biol. Soc. Wash. Vol. xvi, euge s — MILLER, G. S., JR. Notes on the

Bats Collected by William Palmer in Cuba. Prac. U. S. ees Mus. Vol. xxvii,

ts o e

PP- 337-348, pl. g.— MILLspauGH, C. ntæ Yucatanza. Plan th

Insular, Costal and Plain Regions of the Peninsula of Yucatan, Mexico. Fe

P B Vol. iii, pp. 85-151, figs.— NE

Wheat-Grasses of Wyoming. Bull. Wyo. Agr. Exp. Sta. No. a 2 Abe Meg and

figs.— NELSON, A. A Decade of New Plants. Proc. Biol. Soc. Vol. xvii,

pp. 99-100.— NELSON, A. New Plants from Nevada. Proc. rs e Wash.

Vol. xvii, pp. 91-98.— NELsoN, E. W. Descriptions of New Birds from South-

xico. Proc. Biol. Soc. Wash. Vol. xvi, pP. 151-160.— NELSON, E. W. A

Revision of North American Mainland Species of Myiarchus. Proc. Biol. Soc.

ash. . Vol. xvii, pp. 21-50.— OBERHOLSER, H. C. M of a New

African Weaver Bird. Proc. U. S. Natl. Mus. Vol. xxvii, p. 68 3.— OBER-

HOLSER, H. C. A Review of the Wrens of the Genus Tooni s. Proc.

U. S. Nat. Mus. Vol. xxvii, pp. 197-210.— OBERHOLSER, H. C. A Revi-

sion of the American Great Horned Owls. "o au. M.

xxvii, pp. 177-192.— Oscoop, W Haplomylomys, a New Subgenus of

eromyscus. Proc. Biol. Soc. Wash. Vol. xvii, pp. 5 — Oscoop,

3-54: m nu

Thirty New Mice of the Genus Peromyscus from Mexico and Guat E Proc.

Biol. Soc. Wash. Vol. xvii, pp. 55-77.— PALMER, T. S. Index MES Mam-

maliu A List of the Genera and Families of Mammals. = S Dept. Agr.,

Division Biol Surv. W. A. Fauna No. 23, 984 4 pp.— Pat QE. Ihe

Ancient Canal Systems and Pueblas of the Salt River Valley, Lo. Bur.

No. 450] PUBLICATIONS RECEIVED. 483

Phoenix Free Mus. No. : IO pp. 2 pls.— PEARY, R. E. Field Work of the

Peary Arctic Club. Bull. Geog. Soc. Philad. Vol. iv, 48 pp., pls.— RABr, C.

Ueber die Züchtende ku ia fanion Reize. ‘Rektoratsrede, Leipzig,

Engelmann, 1904. 8vo, 44 pp. 0.80 marks.— REHN, J. A. G. Studies in

American Mantids or Aa Proc. U. S. Natl. Mus. ux xxvii, pp. 561-

574.— RICHARDSON, HARRIET. oe to the Natural History of the

Isopoda. Second Part. Proc. U. S. Mus ol. xxvii, pp. 657-681.—

RICHARDSON, HARRIET. Cou to d Natural o of the aae

Proc. U. S. Natl. Mus. Vol. xxvii, pp. 1-89.— RICH QE. A Rev

of the Sunfishes of the Current Genera Apomotis, Dero eh Eu upomotis, dh

els Reference to the Species found in Illinois. rudi JH. State Lab. Nat. .

251. vii, pp. 27- nbi dd R. Diagnosis of nine new forms of

American Birds. rec. l. Soc. Wash. Vol. xvi, pp. 167-170.— RicGs, E. S

Structure and viti co of Opisthocoelian Dinosaurs. Part I. Apatosaurus

Marsh. Field Columb. Mus. Publ. Geol. Ser. Vol. ii, pp. needa pls. 45-53.—

RUSSELL, xd Research in State Universities. Bul. Univ. Michigan, N. S.

Vol. v, 24 pp.— Sarro, K. Untersuchungen über die NE EEA Pilzkeime.

I. Jour. Coll. € Imp. Univ. Tokyo. Vol. xviii, 57 pp., 4 pls.— SHIMEK, B

Amer. Geol.

T La M 54-73

SHUFELDT, R. W. Osteology of the Limicole. Ann. Carnegie Mus. Vol. ii,

— SiLLoway, P. E. Additional Notes to Summer Birds of Flathead

Lake with Special Reference to rdi Lake. Bull. Univ. Montana, No. 18,

293- -368, pls. 54-57.— Simms, S. C. A Wheel-shaped Stone Monument in

yoming Ani. Anthrop. Vol. v, pp. 107-110.— SiMMs, S. C. Traditions

of the Crows. Field Colum. Mus. ML. Anthrop. Ser. Vol. ii, pp. 281-324.—

SMITH, E. F. and SwiNGLE, D. B. The Dry Rot of Potatoes due to Fusarium

oxysporum. U. S. Dept. Aer, Bureau Plant Industry. Bull. No. 55, 64 pp

pls.— SNopGRass, R. E. and H es E. Papers from the Hopkins-Stanford

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72.— SNYDER, J. O. Beers a the Shore Fishes Collected by the Stea-

mer preio about ^ Hawaian Islands in 1902. Bull. U. S. Fish Com. for

1902, Rig js P od 13 pls.— Starks, E. C. The Osteology of some Berycoid

Fis Natl. Mus. Vol. xxvii, pp. 601-619, 1o figs.— STEJNEGER,

x: Sed pode jue the Rio Grande Valley, Texas. Proc. Biol. Soc. Wash.

Vol. xvii, 17-20.— STEJNEGER, L. A New Species of Large Iguana from the

Bahama Islands. Prec. Biol. Soc. Wash. Vol. xvi, pp. 129-132.— STEWART,

J. H. and Hire, B. H. Commercial Fertilizers. Bull. W. Va. Agr. Exp. Sta.

No. 91, 56 pp.— SwarTH, H. S. Pacific Coast Avifauna. No.4. Birds of the

Huachuca Mountains, Arizona. Cooper Ornithological Club, Los Angeles, Cal.

Svo, 70 pp.— TINSLEY, J. D. and Vernon, J. J. S > Moisture Investigations

for the Season of 1903. Bull. N. Mex. Agr. Exp. Sta. No. 48,

Tu. e S. On the Fossil Echinoids of be. So Coll. Sci. Imp. Univ.

Vol. xvii, 27 pp., 4 pls.— TRUE, F. W. On Species of "uc American

Hae sete described by Dr. R. A. Philippi. Prac. Biol. Soc. l. xvi,

Pp. 133-144.— TRUE, F. W. Notes on a Killer Whale (Genus feci from

the Coast of Maine. Proc. U. S. Natl. Mus. Vol. xxvii, pp. 227-230, pls. 7-8.

— UHLER, P. R. List of Hemiptera-Heteroptera of Las Vegas Hot Springs,

New Mexico, Collected by Messrs. E. A. Schwarz and Herbert S. Barber. Proc.

U. S. Natl. Mus. Vol. xxvii, pp- 349-364.— UNIVERSITY OF Montana. Lec-

484 THE AMERICAN NATURALIST. [Vor. XXXVIII.

tures at Flathead Lake. Bul. No. 17, Biol. Ser., pp. 197—288, pls. 47-52, figs.

R. The Fi

4- Ta TRR: Oraibi Summer Snake Ceremony. Field Columb

Mus. ^, Anthrop. Ser. Vol. iii, pp. 271—358, pls. 148-209.— Voru, H. R.

The ono ni Ceremony. Field Columb. Mus. Publ., Anthrop. Ser. Vol. vi,

pp. —28.— WASHBURN, F. Eighth Annual Report of the one

aus A Minnesota. xvi +184 pp. 119 figs.— WASHBURN,

Mediterranean Flour Moth. S ten Report of the Entomologist of Minnesota.

3! pp. ! pl., 20 figs.— Weeks, F. B. Bibliography, and Index of North Ameri-

can Geology Duero Petrology and Mineralogy for the year 1902. Bull

U S Ce ; No: 200 pp.— WHEELER, H. J. and ADAMS

(Ae Bw

Further ud oF the Niel js Lime upon Plant Growth. Bull. Rhode J.

gr. Exp. Sta. No. 96, pp. 23-44, 4 figs.— WHITEAVES, J. F. Mesozoic Fossils.

Él i Pt.v. On some additional Fossils from the Vancouver Cretaceous, with

a revised List of the Species. Geol. Survey Canada Publ. pp. 309-415, pls. 40-

51.— WHuITEAVES, J. F. Bibliography of Canadian Zoóiogy for 1902 Exclusive

of Entomology. Trans. Roy. Soc. Canada. Vol. ix, pp. 163-167.— WHITEAVES,

ie The Canadian Species of Trocholites. Ottawa Naturalist. Vol. xviii, pp.

di OW ARA Vc J.F. Additional Notes on some Canadian Specimens of

Hisce Undatus. Ottawa Nat. Vol. xvii, pp. 161—163.— WIELAND, G. R

Polar Climate i in time the Lore Factor in the Evolution of Plants and Animals.

-430.— WILLISTON, S. W. On the Osteology

of AVA (Nyetodaciu with notes on American Species. Field Columb.

Mus. Publ. Geol. Ser. Vol. ii, pp. 125-163, pls. 40-44.— WILSON, C. B.

Species of Aun with a more Complete Account of two Species already

cribed. Proc. U. S. Natl. Mus. Vol. xxvii, pp. 627-6 5.

Anales del Museo Nacional de Montevideo. Tom. v.— Boletin de la Comision de

Parasitologia e [Mexico]. Tom. ii, Nos. 3-4.— Bulletin of the Johns Hop-

kins Hospital. Vol. xiv, Nos. 152-153. Vol. xv, Nos. 154-1 56.— Bulletin Musée

Oceanographique de Mic Nos. 1-3.— Field Naturalists Quarterly, The. Vol.

iii, Mch.— Higher Science Mch.-Apr., 1904.—Le Réveil Sali cole, Ostréicole et des

Péches Maritimes. Ann. iii, Nos. of — Medical Brief, The. Vol. xxxii, Jan.—

Notarisia, La Nuova. Ser. xiv, Oct. 903, Jan. 1904.— Physical Review, The,

Vol. xvii, No. 6, Dec.— Proceedings of in Natural Science Association of ‘Staten

Island, Vol. viii, Nos. 24-25, Vol. xix, Nos. 1-4.—Revista Chilena de Historia

Natural. Año vii, No. 2.—School Science. Vol. iii, Nos. 6-7.— 7it-Bits of Ani-

mal Life. Vol. iii, No. 4.— Transactions of the American Microscopical Society.

25th Ann. Meeting, Piruw. 1902.

(No. 449 was mailed August 1, 1904.)

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VOL. XXXVIII, NOS. 451-452 JULY-AUGUST, 1904

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THE

AMERICAN: NATURALIST.

VoL. XXXVIII. July-August, 1904. Nos. 451—452.

PROCEEDINGS OF THE AMERICAN SOCIETY

OF ZOÓLOGISTS.

First ANNUAL MEETING OF THE EASTERN BRANCH.

Tue first annual meeting of the Eastern Branch of the Amer-

ican Society of Zoólogists, the fourteenth annual meeting of the

Society since its establishment as the American Morphological

Society, was held in Biological Hall, University of Pennsylvania,

Philadelphia, Pa., on Tuesday and Wednesday, December 29

and 30, 1903.

In accordance with the report of the committee appointed at

the preceding meeting to consider the matter of the publication

of the proceedings of the Society, of an official organ, etc., it

was voted to be the sense of the Eastern Branch of the Ameri-

can Society of Zoülogists that the American Naturalist should

become the official organ of the Society, and the committee was

empowered to arrange for the publication of this year’s proceed-

ings of the Eastern Branch in the 4 merican Naturalist.

The committee in charge of the preparation of an invitation

to the International Zoólogical Congress to hold its meeting in

1907 in America, presented its report, which was adopted. As

the result of this action, and similar action by the Central

-485

486 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Branch, the American Society of Zoólogists will invite the

International Zoólogical Congress to meet in America in 1907.

The necessary preliminary arrangements have been left to a

committee. The committee appointed to draw up a definition

of zoólogical requirements for admission to college, conformable

with the definition of the requirements in other sciences, pub-

lished by the College Entrance Examination Board, submitted a.

report of progress that caused considerable discussion, and the

committee was continued. ;

The list of officers for 1904 is as follows:

President, E. A. Andrews ; vice-president, W. E. Castle; sec-

retary-treasurer, G. A. Drew. Additional members of the

executive committee, T. H. Montgomery, Jr., H. C. Bumpus,

H. S. Jennings.

The following are abstracts of papers presented at the meet-

ing :

The Egg Cases of Chimeroid Fishes: Bashford Dean, Colum-

bia University. — The egg cases of chimzeroid fishes present an

extreme case of adaptation, not to the egg of the time it is

placed in the capsule, but to the young fish when about to

escape. From this standpoint the capsule is adapted in the

matter of extreme size, definite modeling to the head, trunk and

tail of the young fish, durability, mode of attachment, position,

which enables the young fish to develop dorsal side upward, ori-

entation, wherein the snout of the young fish develops in the

direction of the opening valve, and the tail in the direction of

the long and narrow end of the capsule. Further adaptations

include an elaborate provision for circulation of water within the

capsule, the mechanism for which improves as the embryo devel-

ops, by means of a process of weathering, whereby the perfora-

tions of the capsule become progressively larger; and a special

exit valve which can be opened only from within, and only at the

time when the fish is ready to escape. The exit valve has then

a kind of tension resulting in “dehiscence,” comparable some-

what to that of the elements of a seed pod. Adaptation is made

still more complete by provision for the specific characters of

the adult; thus the species which have the longest tails have the

Nos. 451-452] AMERICAN SOCIETY OF ZOOLOGISTS. 487

longest tail sheaths in the egg capsules, and those having the

deepest bodies have also the deepest cavity in the main portion

of the capsule.

The foregoing characters evidently cannot be explained by the

Lamarckian hypothesis, for so perfect a case could not have been

formed primitively zz wero around the embryo; for, by the prin-

ciple of use and disuse itself, such an elaborate capsule would

not have been formed for the protection of an embryo which

was just about to be hatched, nor would the capsule be provided

for aquatic breathing. It is, moreover, a well known law that

in uterine development the secondary membranes are reduced

rather than increased. Natural selection, as it is generally

understood, is equally inadequate to explain the formation of

the present capsule ; for, from this standpoint, fluctuating varia-

tions would have had to be selected throughout a long series of

complicated changes in two entirely different lines, 7. e., in the

line of the evolution of the embryo and in that of the evolution

of the capsule. And it is more than difficult to imagine that in

each of these parallel series the corresponding favorable fluctua-

tions could have continued to be successively selected. This

series of constantly adapted stages furnishes a notable example

of development in a definite line.

The Reproductive Organs of the Limbless Lizard Anniella:

W. R. Coe and B. W. Kunkel, Yale University.— The subject

of this investigation, Anniella pulchra, lives on the barren sand

plains in certain portions of California and Arizona. It is a

slender, worm-like lizard, entirely devoid of limbs and limb girdles

except for slight rudiments of the pelvic arch. It lives beneath

the surface of the sand and feeds upon small insects and spiders.

Although described in 1852, it has never been carefully

studied except with regard to its external anatomy and skeleton.

An examination of its viscera reveals a number of interesting

anatomical peculiarities, among which those of the reproductive

organs are perhaps the most striking. These relate particularly

to the oviducts and uterus of the female and the copulatory

organs of the male.

As in many other lizards, Anniella is ovoviviparous. The

young, only two of which are produced in a season, are born in

488

THE AMERICAN NATURALIST. [Vor. XXXVIII.

September. Both of these develop in the right oviduct (Fig. 1,

Fic. 1.— Diagram of the urogenital organs of

Anniella, showing the right ovidu

wee

» kidney ; z, ureter;

AH *

«cum 5

ut, ut^), the left being very rudi-

mentary and apparently quite

functionless (Fig. 1, /. od). The

right and left ovaries are equally

well developed and a single ovum

matures in each, but both ova

enter the right oviduct.

The rudimentary left oviduct

is represented by a slender tube

usually extending forward only

about as far as the anterior ends

of the kidneys, which are situ-

ated in the most posterior por-

tion of the body cavity (Fig. 1,

L od). The anterior end of this

aborted oviduct is terminated

by a very narrow canal which

usually shows a tendency to be-

come more or less convoluted.

It is especially noteworthy that

this convoluted, terminal canal

shows great variation in length

in different individuals. In a

single specimen examined it was

about half as long as that of the

functional right oviduct. In this

single instance the rudimentary

oviduct is terminated anteriorly

by an ostium opening into the

body cavity. The posterior end

of the left oviduct is but little

smaller than that of the right,

and has a similar structure and

similar relations with the cloaca.

These peculiarities furnish a

striking example of what appears

to be a very recently acquired anatomical character — one which

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOG/STS. 489

has not reached a condition of stable equilibrium, as instanced

by the great variation of the parts concerned. The left oviduct

has apparently been eliminated as a functional organ of the body,

and yet the case mentioned where it was half as long as the

functional one suggests the possibility of further variations,

which, in an exceptional case, might show a reversion to the

primitive condition in which an embryo might develop in this

now rudimentary organ.

So far as we are aware no instance of anaborted and function-

less oviduct has been reported for any of the lizards. The

related genus, Anguis, of Europe, has both oviducts well

developed. In many reptiles, however, the left is shorter than

the right oviduct, so that Anniella presents a variation along the

same line as these other reptiles, but one which has been carried

to an extreme condition.

It should be pointed out that it is the left oviduct which is

aborted in Anniella, while it is the right which has become func-

tionless in birds. In the latter, however, the degeneration of

the parts is carried to a greater length in that the right ovary is

e of embryo of Anniella shortly before birth, showing the pair

projecting from the lateral borders of the cloacal aperture

] pits with which each phallus is provided. X 22.

Fic. 2.— Portion of ventral surfac

of large copulatory organs (phalli)

and the oblique groove and termina

likewise more or less aborted, while in Anniella both ovaries

retain their normal functions.

In the male the spermaries and the sperm ducts are developed

equally on the two sides of the body. The copulatory organs

(phalli) are formed as finger-like, external projections from the

490 THE AMERICAN NATURALIST. [Vor. XXXVIII.

lateral borders of the cloacal aperture, and nearly to the time of

birth have a curious resemblance to a pair of rudimentary limbs

(Fig. 2). Shortly before birth they are drawn into the cloacal

aperture, and by a process of invagination are converted into a

pair of slender pouches which extend backward into the caudal

region for a considerable distance behind the-cloaca. They then

lie on the ventral side of the tail and immediately internal to the

muscles of the integument.

It isa remarkable fact that these copulatory organs develop

in females as well as in males. In females, however, they cease

their development after they are converted into post-cloacal

pouches, although they are retained throughout the life of the

animal. In the males, on the other hand, they increase greatly

in size at the, time of sexual maturity. During the act of copu-

lation the large blood sinuses with which they are provided

become distended with blood. This surcharging with blood,

together with the contraction of certain muscles which surround

the pouches, causes the phalli to be everted to their original,

embryonic condition, and thus to function as a pair of true

copulatory appendages.

Some Account of the Temporary Biological Station at Bermuda

for the Season of 1903: Charles L. Bristol, New York Uni-

versity.— The following invitation was issued about May 1, 1903:

Harvard University and New York University unite with the

Bermuda Natural History Society in inviting botanists and zoól-

ogists to spend six weeks in the temporary biological station

provided for the present season at Bermuda.

By special arrangements with the Quebec S. S. Co. and the

Hotel Frascati it has been possible to make the total expense,

including transportation from New York and return, and board

and lodging for six weeks at Bermuda, one hundred dollars.

The Bermuda Natural History Society has expressed its inten-

tion to do everything in its power to make the summer's work

as profitable and pleasurable as possible, and to this end has un-

dertaken to provide, among other things, the necessary facilities

for collecting, namely, a steam launch, thirty to forty feet long,

with crew; a sail boat with fish-well and crew; three rowing

Nos. 451-452.| AMERICAN SOCIETY OF ZOOLOGISTS. 49I

boats, and a carriage with two horses, capable of carrying ten or

twelve persons. The laboratory will be equipped with all

necessary reagents and utensils except microscopes and dissect-

ing instruments, which should be brought by each investigator.

The well known richness of the sub-tropical fauna and flora,

the healthfulness and equable temperature of the islands, and

the ease with which they may be reached combine to make the

Bermudas a most attractive field for biological research.

Venerable George Tucker, Archdeacon, President Bermuda

Natural History Society.

Hon. W. Maxwell Green, Consul U. S. A., Vice-president B.

N S

F. Gòodwin Gosling, Honorary Secretary B. N. H. S.

E. L. Mark, Director of Zoölogical Laboratory, Harvard Uni-

versity. ;

C. L. Bristol, Professor of Biology, New York University.

It met with a hearty reception, and in response to it thirty-

three botanists and zoólogists from widely scattered portions of

the United States availed themselves of the opportunity offered.

The committee in charge was somewhat embarrassed at first, as

our temporary laboratory was equipped for twenty research stu-

dents, but the proprietor of the Hotel Frascati kindly gave up

two large rooms in the hotel so that all were comfortably cared

for.

The facilities provided and paid for by the Bermuda Natural

History Society enabled the party to make daily excursions

either by launch to the reefs and lagoons, by wagons to botani-

cal collecting localities, or by rowboat to the nearby places.

During the season Capt. W. E. Myers of St. George placed

his powerful sea-going tug, **Gladisfen," at the disposal of the

expedition to collect on the “ Challenger" bank. The trip occu-

pied three days and was very fruitful in results.

During the stay of the party, between June 22 and August

22, the weather was pleasant save one or two days of dead calm,

when it was disagreeably warm, though at no time did the ther-

mometer go above 85? F. The general health of the party was

excellent, and demonstrates what my experience had led me to

believe, — that the climate is favorable for college workers in

midsummer.

492 THE AMERICAN NATURALIST, [Vor. XXXVIII.

The accommodations at Hotel Frascati were excellent. Our

party so nearly filled the house that the proprietor, Mr. Alonzo

Peniston, put. it practically at our disposal, accommodating his

meals to our excursions whenever it was desired. The table

was first-class, and not to be compared with the usual summer

board at seaside laboratories. The food was well cooked, was

abundant, and well served at tables always decked with a profu-

sion of flowers. |

The party was welcomed on its arrival at the dock by the

venerable Archdeacon Tucker, Hon. W. Maxwell Greene and

Mr. F. Goodwin Gosling, and this welcome was continued by

every one — high and low — during the season. His Excellency,

the Governor, Gen. H. LeG. Geary, was kind enough to extend

a hearty welcome to the party by holding a reception at Mt.

Langton, to which nearly two hundred guests were invited.

Hospitality was extended on all sides, and almost to the embar-

rassment of the work at times. ;

The people of Bermuda became interested in the annual visits

of the New York University Biological Expeditions from 1897

to 1902, and in 1902 a committee of the Legislature was ap-

pointed to consider and report on the advisability of the estab-

lishment of a Marine Biological Station. The Colonial Secretary,

by direction of His Excellency the Governor, addressed a letter

asking for information and coóperation to the following: the

British Association for the Advancement of Science, the Carne-

gie Institution, the Royal Society, the Minister of Marine and

Fisheries, Canada, Dr. Auton Dohrn, Naples. The responses

were all heartily in favor of the idea, and this encouraged His

Excellency to ask the approval of the Imperial Government.

This was granted in a despatch from the then Secretary of State

for the Colonies, Hon. Joseph Chamberlain, and the way was

clear for further action by the Colony. In November, 1903,

Governor Geary sent a message to the House of Assembly which

has resulted in appropriate legislation without a dissenting voice.

It may be safely said now that before long a well equipped :

station will be available for biologists at Bermuda that will be

open the year round.

During the coming summer another “temporary” station

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 493

will be established and good facilities offered for advanced

workers.

A Key to the Development of Sipunculus nudus: John H.

Gerould, Dartmouth College.— The development of Phascolo-

soma shows that the embryonal envelope or serosa of Sipunculus

nudus, described by Hatchek, is a modified prototroch. In the

gastrula stage of each form the prototroch consists of two or

three rows of broad, flat cells which encircle the embryo. In

the trochophore of P4. vulgare the apical plate, the mid-dorsal

cord of ectoderm in the interruption of the prototroch and the

somatic plate are strikingly similar to the corresponding parts in

S. nudus, but in the former no sinking beneath the vitelline

membrane occurs, and no amniotic cavities are formed. The

resemblance between the trochophores extends even to the apical

rosette and to the cells which bear the postoral circlet of cilia.

The remnants of the prototroch of Sipunculus are cast off

with the vitelline membrane, the huge prototroch cells of Phas-

colosoma degenerate during metamorphosis, and their substance

in the form of yolk granules passes backward into the newly

formed coelom. Thus the ccelom of the larva becomes filled

with yolk granules, which Selenka mistook for blood corpuscles.

These gradually are absorbed. A cuticula is formed beneath

the vitelline membrane, the latter is shed, and an overgrowth of

the region vacated by the prototroch cells takes place.

Sipunculids are primitive Trochozoa, closely allied to chzeto-

pods. The adult sipunculid retains the retractor muscles and

nephridia of the trochophore; and the loss of the prototroch,

development of the ccelom and elongation of the trunk are the

only fundamental changes which the trochophore undergoes in

passing into the adult form. Resemblance of the trochophore

of Phascolosoma to that of chzetopods, e. g., Amphitrite, is

striking. Transitory metamerism of the mesoblastic bands and

1 The reader is referred, for a more complete account, to Studies on the Em-

bryology of the Fe I. The Embryonal Envelope and its Homologue.

Mark Anniversary Volume. Cambridge, Mass., 1903; and The Development of

Phascolosoma (relie. ei Archives de Zool. expér. et gén. Notes et Revue.

4e série, T. 2, No. 2, 1904. p. xvii-xxix

494 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of the fundament of the nerve cord in PA. gouldii, immediately

before metamorphosis, points definitely to an intimate relation-

ship between the sipunculids and the annelids. Sipunculus is

more highly modified than Phascolosoma.

Divergence Under the Same Environment as Seen in the

Hawatian Snails: John T. Gulick, Oberlin, Ohio.— The

Island of Oahu presents two mountain ranges; one near the

northeast coast, about 35 miles in length, and enjoying a heavy

rainfall, especially on the side of the range most exposed to the

northeast trade-winds; the other, near the southwest coast,

about half as long, and receiving much less rain. The forest

regions on these two mountain ranges are occupied by about

300 species of snails belonging to eight genera of the family

Achatinellide. A large majority of these species belong to four

genera found always on the trees or other vegetation. Auricu-

lella clings to the leaves, is about the size of a grain of wheat,

and is represented by one abundant species on the dry mountain

range, and by several species on the other range. Apex is from

half an inch to an inch in length, and occupies the dry range,

and the drier ridges of the other range. Achatinella and

Bulimella are in size not very different from Apex, and like it

are found on the trunks and branches as well as on the leaves ;

but their distribution is almost entirely on the longer mountain

range, Achatinella being more abundant on the less rainy side,

and Bulimella prevailing on the more rainy side.

Natural selection has undoubtedly been an important factor

in adapting Bulimella for the rainiest districts ; and Achatinella

and Bulimella for the shady groves of the less rainy region, and

Apex for the driest region as well as for the deficient shade on

the ridges of the intermediate region.

But we come now to species whose divergence refuses to be

explained by either natural selection or sexual selection. As an

example of divergent evolution that must depend on other con-

trolling principles, let us consider the formation of the series of

Species of Achatinella occupying the valleys on the southwest

side of the main mountain range. The temperature, the rain-

fall, the soil, and the plant and animal species surrounding these

snails are much the same.

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 495

Commencing at the most eastern of these valleys and pro-

ceeding toward the northwest, we find that the first seven or

eight miles brings us past about that number of deep valleys,

occupied by shady groves and inhabited by many varieties

belonging to fifteen or more species. These species are all

of the larger type of the genus, being near an inch in length.

With some of them, all the individuals of the species are found

in the groves of a single valley, and in the case of most, even

stragglers are not found at a distance of more than a mile from

the metropolis of the species.

Proceeding toward the northwest, we find that the genus is

represented by many species, of progressively diminishing size,

till at the other end of the mountain range the shells have

dimensions not more than half those presented by the shells

found in the valleys first explored. The habits of feeding are

also wholly different; for as we approach the northwest end of

the range we find that the larger trees have been deserted, and

the shrubs and herbs have become the homes of most of the

species. This diversity either in the inherited aptitudes or in

the acquired habits of the isolated groups of one stock, resulting

in different methods of dealing with the same environment, un-

doubtedly results in different forms of selection helping to mould

the different groups into different species. We, however, note

that, in these cases, the process is not natural selection ; for it

is not difference in the environment causing the accumulation

of divergent adaptations, but it is slightly divergent adaptations,

protected by isolation from the swamping effects of free-crossing

with the original stock, that initiates divergent methods of deal-

ing with the same environment, and so determines the difference

in the selection which emphasizes the divergence. This form

of selection, determined by the aptitudes and habitudes of the

species in dealing with the environment, I call endonomic

selection.

A point of very great interest is the important part filled by

isolation in producing divergent forms of endonomic selection.

If a pair of individuals from a plastic (or variable) species are

completely isolated from the original stock, the probability is

that in some way their method of dealing with the environment

496 THE AMERICAN NATURALIST. (Vor. XXXVIII.

will differ from the method prevailing in the original home,

though under certain circumstances it may have been tried by

some of the original stock. New relations having been estab-

lished with the environment, and the swamping effect of associa-

tion and intergeneration being removed, divergent evolution is

insured. Two genera of the Achatinellidae are found almost

exclusively on the ground among the dead leaves. The diver-

gence between these genera living on the ground and those liv-

ing on the trees and shrubs was probably gradually introduced

in some such way. In the same way may have arisen the differ-

ences between the genus of minute species feeding on the leaves

and the genera of larger forms feeding on what is found on the

trunks and branches of the trees.

The Function of the Lateral Line Organs in Fishes: G.H.

Parker, Harvard University. — This work was carried on at the

Biological Laboratory of the United States Bureau of Fisheries

at Woods Hole, Massachusetts. Tests were made on smooth

dogfish, skates, winter flounders, scup, and two species of Fun-

dulus. The action of the lateral line organs was eliminated by

cutting the branches of the fifth, seventh, and tenth nerves sup-

plied to these organs. After recovery from shock the reactions

of individuals on which these operations had been carried out

were compared with those of normal fishes. No significant dif-

ferences could be discovered in the'reactions of normal and oper-

ated fishes to the following stimuli : temperature changes from

9° C. to 30° C.; salinity changes having a range between a mix-

ture of one part sea water and four parts fresh water, and sea

water concentrated to three fourths its original volume ; reduced

oxygen supply ; increased carbon dioxide; foul water; reduction

of pressure to nine pounds; increase of pressure to twenty-two

pounds; and currents of water. On testing for equilibrium it

was found that the lateral line organs were of no more signifi-

cance in this respect than the general integument. Normal and

operated fishes did not respond differently to sound vibration of

the rate of 100 per second. Operated fishes failed to respond

to vibrations at the rate of six per second, whereas normal

fishes reacted to these with great precision. The lateral line

No. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 497

organs are, therefore, concerned with vibrations of low fre-

quency. These vibrations are often produced when large objects

move in the water or when wind blows on its surface.

The Sexual Elements of the Giant Salamander: Albert M.

Reese, Syracuse University. — Eggs and spermatozoa were ob-

tained by stripping giant salamanders (Cryptobranchus alleghen-

iensis) during the latter part of September.

Each egg is a spherical mass of yellow yolk, about 6 mm. in

diameter, surrounded by two layers of transparent jelly ; it does

not completely fill the inner envelope, but is surrounded by an

aqueous fluid.

The egg proper is filled with yolk granules of various sizes,

and is surrounded by a thin vitelline membrane.

The gelatinous envelopes are continued as slender cords from

egg to egg, forming a continuous string similar to the egg-strings

of Amphiuma. The distance between two adjacent eggs is

about four or five times the diameter of a single egg.

Some dozens of eggs were obtained from one female, all of

them apparently being derived from one ovary.

The spermatozoa, in which no motion could be seen, though

stripped with little difficulty from the males, consist of an elon-

gated, cylindrical nucleus, a sharp, gradually-tapering apical

body, and a tail of moderate length which consists of a central

fibre and a transparent envelope. No middle-piece can be made

out.

Artificial fertilization was attempted but was entirely unsuc-

cessful.

Some Recently Discovered Cases of Mutual Sexual Adapta-

tion: P. P. Calvert, University of Pennsylvania. — In studying

the Odonata of Mexico and Central America for the Biologia

Centrali-Americana it was found that a new species of Proto-

neura, P. peramans, presented a remarkable adaptation of the

sexes for mutual grasping during pairing, in that the bifid, supe-

rior, terminal, abdominal appendages of the male are received

between bifid, mesothoracic processes of the female. The other

species of this genus from this region show no (or less perfect)

adaptations in this regard.

498 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Segmentation of the Ovum of Desmognathus fusca :!. William

A. Hilton, Cornell University. — During

thelast few years several amphibian eggs

have been found which, from the few stages

of development described, seem to indicate

a meroblastic rather than a holoblastic

method of cleavage. The first of these

EDI c eua. WA reported by H. H. Wilder (1899) for

ode e stage niin Desmognathus, the second for Aaütodax,

sprees by Ritter and Miller, also 1899, and the

third by Montgomery for Plethodon in r9or. In all of these

forms only the later stages were observed.

As Desmognathus is very abundant in

central New York, advantage was taken of

the opportunity for a thorough investigation

of the earlier stages.

The eggs are about 3.5 mm. in diameter

and entirely devoid of pigment. The general F'*- Section of four

appearance is creamy white except at the and second division

animal pole, which is chalky white. In this =" ex»

chalky white area the first division furrows

make their appearance. The first and sec-

3 ond furrows are at right angles (Fig. 1),

2 but soon these lose this geometrical regu-

larity and the later division lines may be

very irregular.

In the four-cell stage the first cleavage

plane has cut somewhat deeper than the

side, z and 2, etc., first and second, but neither has yet cut deeply into

secon o etc., division planes

(about x 8). the central mass of yolk (Fig. 2). In the

— Surface view of an

1 Due to a slight GCE Sei € uem Wilder and the author have

been working on precisely the same r. Wilder's results, if not pub-

lished by the time this appears, will s oe soon. In his paper he comes

to the conclusion that the eggs of Desmognathus are holoblastic and not mero-

blastic in cleavage. His conclusions were reached by the study of the surface

aid of a few sections of early stages as well as a few surface views. I will leave

a more detailed account of the early development of this animal for a later

paper

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 499

next stage there is no distinct equatorial groove.

stage is gradually derived from the four-cell

condition by the appearance of furrows be-

tween the two original ones. In the eggs of

this stage which Wilder has studied (see foot-

note) two rather regular vertical planes of

cleavage cut the egg at nearly right angles to

the first division plane, but in a considerable

number of eggs which I have examined, some

Fic, 4.— Section o

eight-cell stage (about

X 8.)

The eight-cell

of an

of them living and others preserved, I only found one or two

with such regular vertical cleavage planes; in some there was an

approach to it, but in a number of others the eight-cell condition

IG. 5.— Surface views ofa

few-cell blastula from the

side (about X 8).

into one or more

seemed to be gradually derived from the four-

cell stage by the appearance of furrows which

were hard to recognize as parts of vertical

planes of cleavage. In the eight-cell stage

the first furrow nearly encircles the egg, but

the position of the first and the second is

often so shifted that it is difficult to believe

that they were ever at right angles (Fig. 3)-

At this time the animal pole may be divided

distinct blastomeres, but the vegetative pole

shows almost no signs of cleavage planes (Fig. 4).

As segmentation proceeds the surface appearances correspond

in general with those of other amphibian ova

(Fig. 5), but sections bring out the fact that

the segmentation is shallow, and does not in-

volve the whole thickness of the egg (Fig. 6).

Later, however, when the surface appearance

islike that of a fine celled blastula (Fig. 7),

sections show that the seg-

mentation has extended

Fic. 7.— Surface view of

a many-ell blastu

Fic. 6.—Sect

JAS

ion of a few-

cell blastula (about X

8).

through the entire egg and that the cleavage

in this stage is as truly holoblastic as in any

other of the salamanders (Fig. 8).

As the embryo increases in size the cell

outlines in the yolk are lost, and if sectioned

in this stage only, one would be justified in

Be the side (about suspecting a meroblastic segmentation. This.

500 THE AMERICAN NATURALIST. (VoL: XXXVIII.

idea is helped by the fact that the growing embryo soon comes

to occupy a position on the yolk strikingly

like that of a fish embryo.

In a word, then, the segmentation of the

ovum of Desmognathus is truly holoblastic,

but the total cleavage is rather slow in

Fig.8.—Sectionofamany- appearing, and in the later stages of devel-

cell blastula @bourX®). opment the yolk mass becomes homogeneous

by the disappearance of cell walls.

The Early Development of Dinophilus as Compared with that

of the Annelids: James A. Nelson, University of Pennsyl-

vania.— The ovum of Dinophilus segments into four cells, or

macromeres, each of which gives rise by oblique alternating or

* spiral " cleavages to five generations of smaller cells, or micro-

meres. The first three quartettes give rise to ectoderm. The

posterior left hand’ member of the fourth quartette divides

bilaterally and equally into two cells which later give rise to the

mesoderm bands. The remaining cells give rise to the ento-

derm. This method of origin of the three germ layers is

identical with that found in the Mollusca and the Polycheeta.

As in the two former groups the brain is formed from the first

quartette, and the greater part of the trunk ectoderm from the

posterior cell of the second quartette. In the details of the

cleavage there isa striking agreement with that of the poly-

cheetous annelids. Cells arising from the first quartette iden-

tical in origin with the « primary trochoblasts " of the annelids,

together with cells derived from the first and second quartettes,

make up a cell row which very probably forms at least a part of

the second ciliated band situated on the head of the adult, in a

position corresponding with that of the prototroch of the annelid

larva. Other details increase this resemblance. The sym-

metrical cell pattern known as “the cross" is formed by the

same cells and at the same divisions as in the Polycheta, while

in the divisions of the posterior cell of the second quartette

bilateral Symmetry appears at the same time and in the same

cell as in several members of this group.

Such close agreement in the details of the cleavage, as well

“Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 501

as in the fate of the corresponding cells with the conditions

obtaining among the Polychzta, speaks strongly for a genetic

relationship between this group and the Dinophilide, since

cleavage patterns are inheritable, as well as other characters.

Organ-forming Germ Regions in the Eggs of Ascidians and

Snails: Edwin G. Conklin, University of Pennsylvania.— The

early development of ascidians and snails is of such a deter-

minate character, even in the earliest cleavage stages, that it

seems probable that a definite polarity and localization of mate-

rials exists in the unsegmented egg. A study of the eggs of

these animals shows that this antecedent probability is realized

in fact.

In the living eggs of the fresh water snails, Limnza, Physa

and Planorbis, one may readily observe the segregation of unlike

materials of the egg before cleavage begins. Before maturation

the nucleus can be seen in such eggs as a large, clear vesicle,

while the rest of the egg substance is filled with yolk granules

and is yellow in color. During the maturation divisions the

nuclear vesicle is dissolved and its contents form a clear area

surrounding the polar amphiasters. After the second polar

body is formed this clear area spreads out as a disk and then as

a cap which covers the upper two-thirds of the egg. The lower

third of the egg which is not covered by this cap of clear proto-

plasm remains of a bright yellow color. These two portions

remain distinct throughout the subsequent development, the

clear area giving rise to the ectoderm and the yellow one to the

endoderm and mesoderm. In these snails, therefore, the germ

layers are visibly outlined in the unsegmented egg.

A somewhat similar localization of germinal materials, before

cleavage, occurs in the eggs of certain ascidians. In Cynthia

(Styela) partita, Ciona intestinalis and Molgula manhattensts

the central yolk mass of the immature egg is surrounded on all

sides by a peripheral layer of protoplasm. In the living eggs

of Cynthia the central yolk is gray in color while the peripheral

layer contains granules of yellow pigment ; in Ciona the yolk is

red, the peripheral layer clear and transparent ; in Molgula the

yolk is gray, the peripheral layer clear. As soon as a sperma-

502 THE AMERICAN NATURALIST. [Vor. XXXVIII,

tozoon enters the egg this peripheral layer, together with the

nuclear sap from the germinal vesicle, flows rapidly to the lower

pole, at which point the sperm always enters. The yolk is left

uncovered at the upper pole and here both polar bodies are

formed. The protoplasm at the lower pole then moves up to a

position just below the equator on the posterior side of the egg ;

and here the pronuclei meet and the first cleavage spindle

arises. At the close of the first cleavage the greater part of

the clear protoplasm moves into the upper hemisphere while the

gray, yolk-laden protoplasm is largely collected in the lower

hemisphere. In Cynthia the yellow pigment collects into a

brilliant crescent, just below the equator on the posterior side of

the egg; in Ciona and Molgula this crescent is clear and almost

colorless. Owing to its color in Cynthia the substance of this

crescent can be traced through the whole development until it

gives rise to the muscle cells and the mesenchyme of the tad-

pole. The gray substance of the unsegmented egg gives rise

chiefly to the endoderm, while the transparent protoplasm goes

principally into the ectoderm. In these eggs, therefore, there is

not only a localization of the material of the three germ layers

before cleavage, but all the axes of the future animal are clearly

established in the unsegmented egg.

An Experimental Examination of the Germ Regions of the

Molluscan Egg: Edmund B. Wilson, Columbia University.—

No abstract.

Polarity and Axial Heteromorphosis : T. H. Morgan, Bryn

Mawr College. — The reversal of the polarity that is seen in

axial heteromorphosis appears to give us an opportunity for an

examination of the phenomenon of polarity itself, for, if we

could determine the conditions that determine this reversal we

might hope to gain some insight into the factors that determine

the polarity. It is from this point of view that I offer the

following observations on two species of planarians, P/anaria

maculata and P. lugubris.

If a planarian is cut in two at almost any level a new tail

regenerates on the posterior end of the anterior piece and a new

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS 503

head on the anterior end of the posterior piece. It appears

that the same material is doubly potent, being able at nearly

every level to form a head or a tail. Something in the piece

itself determines that a head shall develop at the anterior cut

surface and a tail at a posterior cut surface. This “something”

is what we call polarity.

I have found certain exceptions in the planarians to this rule.

Occasionally a cross-piece develops a head at each end, and in

Planaria lugubris, especially, when the tip of the old head is cut

off just behind the eyes, another reversed head regenerates from

its posterior cut surface.

In the few cases in which I got double-headed forms I was

impressed by the fact that this occurred only when the cross-

pieces were very short, and never occurred in long cross-pieces.

This lead me to try making the pieces very short in order to see

if I could obtain in this way more of these double-headed forms.

This I found to be the case, and I have obtained from such

pieces double-headed forms from every part of the body of

Planaria maculata. Wt is clear, therefore, that the result can

not be connected with the presence of any particular structure

at the cut end. A number of different kinds of experiments,

mainly with pieces having obliquely cut ends, seemed to show

that the main factor in the production of the heteromorphic

head is connected with the shortness of the piece. I suggest as

a tentative hypothesis that this shortening of the piece reduces

the strength of the polarity so that it comes to have less influ-

ence on the new part than certain innate tendencies of the mate-

rial itself. There is reason to conclude, as I have stated, that

the new material that develops over the cut surface has the

power of producing either a head or a tail. If now the polarity

d, that one of these two tendencies which

is reduced, or remove

In the case of

is stronger will have a chance to assert itself.

Planaria maculata the tendency to produce a head seems

stronger than the tendency to produce a tail, hence the appear-

ance of the heteromorphic head in the short cross-pieces, in

which the polar influences fail to act. This conclusion I hold

subject to future revision when we have gained more light as to

the nature of the so-called polarity itself.

504 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In Planaria lugubris I had found that when the head ‘is cut

off just behind the eyes a heteromorphic head is produced on the

posterior end of the old head. From this result I confess that

I anticipated that in this worm it would be even easier than in

P. maculata to obtain double-headed forms from short cross-

pieces. To my surprise I found on the contrary that these

double-headed forms do not arise except in a few cases in which

the cross-pieces have come from the region immediately behind

the old head. Most of the other short cross-pieces produce a

head and a tail, but in a small number of cases the structure

.that develops at the anterior end appears to be a tail. It would

be difficult to determine with absolute certainty whether this

anterior pointed structure is a heteromorphic tail or an undevel-

oped head, unless the pharynx were to develop. In one case a

pharynx appeared in the new part at the anterior end, and

another in the new part at the posterior end. In both the new

pharynx pointed outwards, 7. e., towards the tip of the new part,

hence there can be little question that the new structure is

really a heteromorphic tail.

It appears in Planaria lugubris that while the material at

every level has the double potency of producing a head or a tail

according to which end of the piece it comes to lie at, yet in

very short pieces from the middle and posterior regions of the

body a double-tailed form may arise. We must suppose, there-

fore, on our hypothesis, that while in this species also the mate-

rial is totipotent, yet when the determining influence of the

polarity is removed the stronger tendency is to produce a tail,

while in P/azaria maculata, as we have seen, the stronger ten-

dency is in the other direction.

In Planaria lugubris it has been necessary to assume that the

most anterior part of the body has a different predisposition

than has the rest of the body, since in the former a double-

headed piece may develop and in the latter a double-tailed form.

This result finds a parallel in certain other cases that I have

observed in the earthworm and tadpole.

If an earthworm is cut in two behind the region of the gizzard

there often develops from the anterior cut surface of the poste-

rior piece of the worm a heteromorphic tail. In this case the

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 505

posterior part of the worm is not a short piece, and yet it is

worthy of note that this region has practically the same structure

throughout. This may mean that the conditions are fundamen-

tally the same as those in the short cross-pieces of the planarian,

and if so this may throw some light on the phenomenon of

polarity itself.

It has also been shown in the earthworm by Hazen that from

the posterior cut surface of a short piece of the head end a het-

eromorphic head may develop. Here we must assume that the

tissues are predisposed to form a head, while in the posterior

region they are predisposed to form a tail, and in both when the

polar influences are removed these structures appear. It has

been shown by Harrison in the tadpole that a taillike outgrowth

develops from the anterior cut surface of the tail. I have tried

to show that this outgrowth is really a heteromorphic tail.

Here again it is noticeable that the old tail has practically the

same structure throughout, hence possibly, as in the earthworm,

there may be a decrease in the polarity and the consequent

- development of a heteromorphic structure.

These considerations are tentative, and possibly premature ;

but the facts give us at least some grounds for the hope that the

problem of polarity may not be beyond the reach of further

analysis.

The Development of the Germ Layers in a Nudtbranch Mollusc:

Dana B. Casteel, University of Pennsylvania.— The cleavage

of the egg of the nudibranch, Frona marina, is of the spiral

type well known for molluscs. From the mesentomere (4 d)

arises the primary mesoblast and also enteroblasts, these latter

being concerned in the formation of the intestine. Secondary

mesoblast is formed from the third quartet of ectomeres in a

manner similar to its formation in Physa and Planorbis. Two

cells in each anterior quadrant sink below the surface into the

cleavage cavity and by division there form two bands of cells on

the sides of the anterior end of the gastrula. These elements

later take part in the formation of the muscles of the velum, a

larval organ, and are thus truly “larval” in character.

The anal excretory organ, characteristic of opisthobranch

506 THE AMERICAN NATURALIST. [Vor. XXXVIII

larvae, is undoubtedly of ectodermal origin in Fiona. It arises

from one large and several small third quartet cells of the right

posterior quadrant which, after torsion begins, are carried far-

ther to the right side and above the anal opening. The larva

also possesses additional organs of excretion, two in number,

lying bilaterally in the anterior region of the body cavity.

These, the *nephrocysts," are composed of but a few large

vacuolated cells which contain concretions and fluid excreta.

The origin of these cells has not been determined definitely, but

they are probably derived from the mesoderm.

A Pair of Giant Nerve Cells of the Squid: Leonard W. Wil-

liams, Brown University. — No abstract.

The Function of the Accessory Chromosome in the Spider:

Louise B. Wallace, University of Pennsylvania.— In the sper-

matogenesis of insects a number of authors have described a

peculiar chromosome which has been called by various names.

While these all agree in being unlike the ordinary chromosome,

there seems to be sufficient disagreement to subdivide them into

O groups. Montgomery’s “chromatin nucleolus ",and Paul-

mier's * small chromosome ” are probably homologous and these

authors have suggested that they may represent degenerating

chromatin. McClung’s « accessory chromosome ” in Orthoptera

Nos.451452] AMERICAN SOCIETY OF ZOOLOGISTS. 507

is too well developed to allow of such an explanation and since

it divides in the second maturation division only and is thus

distributed to but one-half of the spermatids, he regards it as a

sex-determinant.

In the spider Agalena nevia, the peculiar chromosome closely

resembles the one described by McClung and therefore I adopt

his term but my results differ from his in regard to its distribu-

tion. I am convinced that, although split longitudinally, it

takes part in neither of the maturation divisions and that, there-

fore, only one-fourth of the resulting spermatids possesses it.

Such a condition would hardly favor the idea of sex-determination.

Figures r, 2 and 3 show spermatocytes of the first maturation

division and Figures 4, 5 and 6, spermatocytes of the second

maturation division.

In nests of maturing spermatozoa are found cells undergoing

degeneration (Fig. 7) and these occur in great numbers in the

testis and also in the ducts (Fig. 8) together with the much

smaller number of ripe spermatozoa. This fact leads me to

suggest that possibly only those spermatids which contain the

accessory chromosome develop into functional spermatozoa and

that the remaining three-fourths never reach maturity. If this

interpretation bea true one, a parallelism could be drawn between

the development of the spermatozoon and of the ovum, since in

the latter only one of the four oótids develops into a ripe ovum,

508 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the other three being thrown off as polar bodies and later

degenerating.

The Lamellibranch Nervous System: Gilman A. Drew,

University of Maine — The nervous system of the giant scallop,

Pecten tennicostatus, is greatly specialized, and the size of the

animal makes it possible to study it by dissection as well as by

serial sections. The cerebral and pedal ‘ganglia are small, owing

no doubt to the reduction of the parts they supply, and the

visceral ganglia, supplying the large adductor muscle and sending

a large number of nerves to the borders of the mantle, are very

large and composed of. distinctly marked regions. The pallial

nerves that follow along the borders of the mantle lobes near

the bases of the tentacles and eyes, present the structure of

ganglia and supply nerves to the tentacles and eyes, which are

very abundant in this form. The need for such ganglia is

apparent, and their presence instead of a further complication of

the visceral or cerebral ganglia indicates the ease with which

such centers may be established when need arises.

The fact that the pallial nerves, that here show the structure

of ganglia, are joined at intervals for nearly their whole length

by nerves from the visceral ganglia, while only a very few nerves

are sent to them from the cerebral ganglia, and these only to

their extreme anterior ends, is taken as an indication that the

loss during development of the anterior adductor muscle is

accompanied by a suspension in the growth of other anterior

portions, and that the bulk of the animal if formed by the

excessive growth of the posterior portions.

The relation of the nervous systems of other lamellibranchs.

was also discussed.

The Early Stages in the Development of Ophiothrix Fragilis :

. W. MacBride, McGill University.— Comparatively few and

scattered notices are to be found in zodlogical literature with

reference to the early development of Ophiuridea and in a pre-

liminary notice like the present communication, it will only be

necessary to refer to one of them. In 1891 Russo! published

Le prime fasi di sviluppo nell’ "s squamata — Bollettino della Societé-

in Naturaliste in Napoli. Vol. 5, 1891

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 509

an account of the formation of the layers in the embryo of the

viviparous Amphiura squamata; and ^

in that paper he made the startling

statement that in Amphiura squamata

the endoderm was formed by delami-

nation from the inner ends of cells

forming the wall of a hollow blastula,

and that, further, the coelomic sacs

appeared as cleavage spaces in the

midst of masses of mesenchyme cells.

Such fundamental differences be- usi spacio ao amas

tween the developmental processes ‘sents! division.

in this form and those in all other. Echinodermata, were hardly

to be credited without ample

confirmation, especially when it

is recollected that Amphiura

squamata is not a favorable

object for study owing to the

difficulty of obtaining the earlier

stages in quantity.

mm For this reason I selected the

Fic. 2.— Section through embryo at the closeof common British form, Ophio-

TARTE se thrix ragilis for study, and,

during a visit to the Plymouth Biological Station in 1898 I arti-

ficially fertilized a number of the eggs and reared some of the

larvae through the entire period of

development up to and including the

metamorphoses, the whole occupying

about three weeks.

The egg of Ophiothrix fragilis is

very minute, being less than .1 mm.

in diameter, and is quite opaque so

that in order to follow the earlier

stages of development it is necessary

to use serial sections. Fic. 3.—Longitudinal section of young

gastrula arch., archenteron.

As shown in Figure 1, the result

of segmentation is not as in other Echinodermata, a hollow

blastula, but a solid morula. From a comparison with one

510 THE AMERICAN NATURALIST. [Vor. XXXVIII.

another of my preparations, it appears that the egg at first

divides into a few blastomeres, which are arranged around a

vestigial, or perhaps we might say virtual blastoccele, and that

as development proceeds this

blastocole swells but is kept

continually filled by cells pro-

duced by tangential divisions of

the peripheral blastomeres. At

the conclusion of segmentation,

the embryo strikingly recells a

ccelenterate planula, as shown in

Figure 2.

The interior mass of rounded

cells, however, does not consti-

tute the endoderm, for,in the next stage (Fig. 3) a regular

invagination has made its appearance, giving rise to an archen-

teron, which is, however, two or three cells thick on one side.

The interior cells of the former stage are seen to occupy the

space between archenteron and ectoderm, homologous with the

segmentation cavity of other forms. In the completed gastrula

Fic. 4.—Longitudinal section of older gastrula

arch., archenteron c. 2., cellular plug.

Fic. 5.— I itudinal EURE E nh

5 = g mbryo two da ld cæ., cælom ezt., enteron.

a curious plug of cells is seen projecting into the archenteron

which is no doubt the remains of the thickened side of the arch-

enteron of the younger gastrula (Fig. 4).

When the embryo has attained the age of two days, the rudi-

ment of the coclom appears as a vesicle at the anterior end of

the archenteron exactly as it does in the embryos of Asteroidea.

At the same time the embryo takes on a shape which may be

Nos. 451-452.] AMERICAN .SOCIETY OF ZOOLOGISTS. SII

described as roughly triangular. The two basal angles of a

triangle (Fig. 5) are really the rudiments of the first two arms

of thé larva, which throughout the whole larval existence are

much longer than the rest, and are the most important organs

of locomotion. The cells which form the interior mass at the

close of segmentation are.now seen to be destined to form the

supporting calcareous rods for these arms, and hence we may

attribute the leading peculiarity of the development, vés.: the

appearance of a solid morula instead of a hollow blastula, to the

extremely precocious formation of the skeletogenous. cells for

these arms.

On the Homologies of the Archenteron and Blastopore in Verte-

brates: Charles S. Minot, Harvard University.— No abstract.

Medusa Fauna of the Bahamas: Alfred Goldsborough Mayer,

Brooklyn Institute of Arts and Sciences.— The medusa fauna

of the Bahamas is poor in comparison with that of the Tortugas,

Florida, The writer secured ninety (90) species of Hydro-

medusze, Scyphomedusz, Siphonophore and Ctenophorz at the

Tortugas, while only forty-three (43) species were found at the

Bahamas. What is more interesting, the relative abundance of

specific forms is quite different in the two regions. The com-

monest medusa during the summer months at the Tortugas,

Pseudoclytia pentata, is not found at the Bahamas, and the

commonest medusa during the summer months at the Bahamas,

Lymnorea alexandri, is not found at the Tortugas.

Altogether, of the forty-three species of Bahama medusa,

twenty-three (23) are about equally abundant on the other side

of the Gulf Stream at the Tortugas. Sixteen are more abun-

dant at the Bahamas than at the Tortugas, while four are more

abundant at the Tortugas. This is accounted for by the follow-

ing facts: The Tortugas lie to leeward of the Gulf Stream and

the extraordinarily rich pelagic life of the great current is con-

stantly driven upon their shores, while the Bahamas lie to the

windward of the stream and their local fauna is not reinforced

by creatures characteristic of the great current. There is,

however, something more than a mere concentration of individ-

512 THE AMERICAN NATURALIST. (VoL: XXXVIII.

uals at the "Tortugas, for a surprising number of characteristic

species appear to be confined to this region alone. Moreover,

atthe Tortugas, we have a small land mass surrounded by pure

deep ocean water, while at the Bahamas we find a vast area of

shallow flats covered mainly by coralline mud, forming veritable

submarine deserts covered only with a sparce growth of coralline

algae and a few scattered clusters of coral reefs and gorgonians.

The water over these shallow banks is almost as deficient in life

as is that of the desert bottom itself, very few Sagitta, Salpz

or Crustacea being found,and among the Medusz only repre-

sentatives of the Gonionemidz and Lymnorea are abundant.

Indeed, the water of these shallow banks is usually charged

with a flocculent mass of silt which adheres to pelagic animals,

and appears to be rapidly fatal to them. The deep water regions

of the Bahamas, however, such as the tongue of the ocean, or

the water to the eastward of Great Abaco Island, are rich in

pelagic animals which appear to be free from silt and in good

condition. This water of the deep areas must, however, be

often driven over the shallow banks by the winds, and its life be

thus destroyed.

It is found that Olindias is closely related to Gonionemus, for

inits ontogeny it passes through a stage in which all of the

tentacles arise from the side of the bell, and are sucker-bearing

as in Gonionemus. Eucheilota paradoxica is the only Leptome-

dusa known which produces an asexual generation of Medusa

by adirect process of budding. These daughter Medusz are

derived from both ectoderm and entoderm of the gonad of the

parent, as is the case in the Sarsiadz and in Hydroids. On the

other hand, the Medusae buds found upon the manubrium of

Bougainvillia niobe are developed entirely within the ectoderm,

the entoderm remaining inert and passive during the growth of

the bud, its limiting membrane being unbroken, and no connec-

tion ever being established between the gastro-vascular cavities .

of the bud and the parent. In B. niobe, however, the ectoderm

of the manubrium is of considerable thickness, affording abun-

dant material for the formation of the bud. It is possible,

therefore, that this peculiar method of formation of medusa

buds from the ectoderm, which has been observed in Z. niobe by

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 513

us, and in Rathkea octopuncta and Lizsia clappadei by Chun,

may be due toa gradual process favored by the thickness of the

ectoderm which may have prevented the deep-lying entoderm

from taking an equal share in the formation of the bud until

finally it has come to remain passive throughout the period of

formation of the bud, as in B. zzobe.

Medusz produced from ectoderm alone may, therefore, be

phylogenitally homologous with Medusae produced by the more

primitive and universal coóperation of both ectoderm and ento-

derm.

Among the new forms discovered, Parvanemus degeneratus

is the most degenerate free-swimming hydroid medusa yet

described. It lacks tentacles, sense organs and peripheral vas-

cular system. It swims, however, with great activity, but is

short lived.

Correlation as the Basis for Selection in Lepidoptera: Henry

E. Crampton, Columbia University.— Read by title.

Exhibition of a Cat, Showing Abnormal Placement of Viscera:

C. M. Clapp, Mount Holyoke College.— No abstract.

The Origin and Function of the Medullary Sheaths of Nerve

Fibres: Porter E. Sargent, Browne and Nichols School, Cam-

bridge.— No abstract.

Demonstration of Pulsatile Anterior Lymph Hearts in Young

Tadpoles: Henry McE. Knower, Johns Hopkins University. —

No abstract.

Excretory Activities in the Nuclei of Gastropod Embryos E

O. C. Glaser, Johns Hopkins University. — The primitive uri-

nary bodies of the larva of Fasciolaria tulipa attain great size

and are very favorable objects for study. They originate as two

pear-shaped patches of highly vacuolated ectoderm at the sides

of the definitive mouth. On account of their connection with

1 The material on which this work was done was collected during three sum-

mers in North Carolina. Iam deeply indebted to the Hon. Geo. M. Bowers for

the courtesies of the Beaufort Laboratory of the Bureau of Fisheries.

514 THE AMERICAN NATURALIST. [Vor. XXXVII.

the velar fold they are later carried outwards from the wall of

the embryo and ultimately hang down from the under side of the

velum.

Between the two extremes in development mentioned, the

cells composing these bodies increase in size, in number by addi-

tion, and undergo a period of active excretion. Early in this

period the nuclei divide amitotically, and the cells become poly-

nucleated. The nuclei are granular, very irregular in outline,

and each one has at least one nucleolus surrounded by a clear

area. In addition the cells are characterized by their highly

vacuolated condition (Fig. 1).

In sections through these nuclei, before and after amitosis,

Fic. 1.— Surface view of part of external kidney.

the chromatin is arranged in an irregular network (Fig. 2

A, B, C). The nucleolus (z/) has a different staining reaction,

and Is probably a typical plasmosome. The halo surrounding it,

instead of being circular or oval in outline, as it appears to be

in whole mounts and in optical sections, is highly irregular, its

outline projecting in finely attenuated processes into the body of

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 515

the nucleus. These processes are so delicate and the granules

bounding them so small in comparison with the chromatin gran-

ules in the network, that they are invisible except in sections.

The location of large chromatin granules continuous with the

nuclear reticulum, in the indentations of the halo, explains its

circular appearance in entire preparations, because these granules

are the only parts of the outline visible. They also afford an

explanation of the irregular outline itself, for if we imagine a

sphere of liquid to proceed out from the nucleolus in all direc-

aR Y) TED

ray ASe em

OPERA

Sie

SERA

Fic. 2.— Sections of nuclei of excretory cells.

tions, any larger objects with which it might come in contact

would, after yielding as much as possible, detain the surface at

certain points, allowing it to advance only where there is no

resistance, vís., between the larger granules. In many cases

the projections of the halo end distally in bulges or vacuoles (A,

vac) which, apparently through the agency of the larger granules

of the nuclear reticulum, become constricted off and may after-

wards be found loose in the nucleus (A, B, vac’), from which

they may finally escape (C, vac"). In other cases the projec-

tions of the halo may break through the wall of the nucleus

before the vacuoles separate (B, vac").

516 THE AMERICAN NATURALIST. [Vor. XXXVIII.

These phenomena strongly suggest an excretory activity, and

this suggestion is strengthened by our knowledge of the behavior

of amitotically dividing nuclei in other animals, but especially by

the results of R. W. Hoffmann on the behavior of the nuclei and

nucleoli in the large macromeres of the embryos of Nassa muta-

bilis. Complete proof, however, that the activities described in

the sub-velar masses of Fasciolaria are really the exhibition of an

excretory process, is found in a chemical analysis of an aqueous

extract of these bodies. Such an extract analyzes like a dilute

solution of urea. |

Double Reproduction in the Medusa Hybocodon prolifer: H.

F. Perkins, University of Vermont.— Hybocodon is familiar to

American biologists as a singularly asymmetrical jelly-fish found

along our coast in the early spring. Of the four radial canals

only one is continued beyond the bell in a tentacle.

The method by which this medusa has been described as

reproducing its kind is by developing, upon the bulbous base

of this solitary tentacle, gemmiferous buds. These, when

mature, become detached from the parent and swim away as

free jelly-fish, Only the one tentacle is developed upon these

buds, and at the time of liberation the base of this tentacle

frequently carries maturing buds of a second generation.

While this is a rapid and efficient means of propagation, it is

not the only one. In addition to this asexual process, a sexual

process is also present. Eggs or sperms are produced within

the gonads which surround the manubrium, and these develop

within the tissue to the condition of tentacled actinula larve.

Thus the adult Medusz give origin to two sorts of offspring,

one by gemmation, the other by sexual production of viviparous

young. This in itself is not anything extraordinary, but it is a

matter of interest that the two processes are found taking place

simultaneously in the same individual, and that this coincidence

does not lessen the rapidity with which the embryo jelly-fish are

produced on the tentacle bulb. The fact is of interest only as

showing that the reproductive activities of Medusz are even

more unrestricted than we have realized.

The sexually produced larvee of Hybocodon mature upon the

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. RI?

wall of the parent manubrium, attached to this member, after

the earliest stages, by a slender connecting stalk. Ten tentacles

appear while the larva is still a spherical mass, and as they

lengthen assume a direction pointing away from the point of

attachment. The mouth and oral tentacles of the actinula are

developed after liberation at the pole which was, during the

attached period, next to the parent.

The Development of the Heart and Branchial Blood Vessels of

Ceratodus : William E. Kellicott, Columbia University.— No

abstract.

Regeneration in Scyphomeduse : Chas. W. Hargitt, Syracuse

University.— Notwithstanding the remarkable development and

scope of experimental zodlogy during the past decade, extending

to members of every phylum from Protozoa to Vertebrata, com-

paratively little has been done directly upon any of the Medusz,

and almost nothing upon Scyphomedusz. The incidental refer-

ence of Haeckel to such a tendency among certain of these

organisms is hardly more than a hint, with nothing whatsoever |

as todetails. Such is also the case with the later work of Eimer

and Romanes on the nervous system of the Medusz, at least as

it pertains to problems of regeneration, though painstaking and

important in its bearings on problems of locomotion and co-ordi-

nated activities. More recently Uexküll has also reviewed some

of this work, and while differing in many respects from that of

his predecessors as to conclusions, leaves untouched the subject

of regeneration. ;

The series of experiments of which this abstract furnishes

the barest outline was undertaken with a view to ascertain the

comparative capacity of Scyphomedusz to reproduce lost parts,

such as bits of the various parts of the body which might be

lost by the usual exigencies of the struggle of life. The experi-

ments comprised some three phases of regenerative energy :—

First, the general ability to recover from such injuries as rents

of the umbrella, etc.; second, regeneration of such organs as

oral appendages; third, ability to regenerate such specialized

organs as the sensory bodies.

518 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Concerning the first, it required but few experiments to deter-

mine very clearly the existence of such a capacity, though its

progress was somewhat more slow than had been anticipated.

Various rents of the umbrella were soon healed, but excisions

of the margins of the body were regenerated much more slowly.

Concerning the second class, the responses were similarly

slow. An excised oral arm required several weeks to completely

regenerate, as was also the case with excised portions of the

gastric pouches.

Concerning the third class I had entertained some doubt,

namely the power to reproduce the more specialized organs,

such as rhopalia or sensory bodies. The experiments were

an agreeable disappointment, as these organs were quite as

promptly regenerated, if not indeed more so than had been the

case with the former. In some cases the first signs of regener-

ation were noted within five days of their excision, and in

from eight to twelve days their functional power was clearly

recovered, as was proved by details of experimentation imprac-

ticable to describe here. The associated organs, such as lap-

pets, hood, etc., were also quite as fully regenerated as the other.

Histological examinations show beyond any doubt the opera-

tion of typical histogenic processes, various phases of the regen-

erating organ being easily traceable from stage to stage to perfect

maturity of form and structure.

In earlier experiments of a similar sort made upon Cyanea

and Aurelia by the writer it was found impossible to maintain

the normal vitality of the animals for sufficient time to secure

any conclusive results. Rhizostoma, however, lends itself most

remarkably to experimentation of various sorts and for continu-

ous periods of from four to six weeks without material loss of

vigor or regenerative power. It was found, however, that this

was somewhat dependent upon the size and, presumably, age of

the specimens. Those of large size, 75 to 125 mm. in diameter,

proving much less hardy under an aquarium environment than

specimens of smaller size; though very small specimens also

proved less satisfactory. Specimens of from 40 to 60 mm. in

diameter proved much the better, not only from the apparently

greater vigor, but also in convenience for experimentation and

observation.

Nos. 451-452.] AMERICAN SOCIETY OF ZOOLOGISTS. 519

The experiments clearly prove the capacity among these

organisms for regeneration of organs or parts of organs, from

the more generalized sort to the most specialized, and that with

approximately similar readiness.

A New Generic Type of Polygordide: J. Percy Moore, Uni-

versity of Pennsylvania.— The annelid referred to is known only

from a couple of fragmented specimens found in association

with marine Oligochaeta on Cape Cod. Like Polygordius it has

a slender, elongated form, an acutely conical prostomium bearing

a pair of prominent, slightly articulated, apical tentacles, a round,

open mouth, and above it a pair of deep, ciliated, sensory pits.

The segments are very obscurely indicated externally in the

anterior region but are clearly defined at the caudal end. The

body wall exhibits the same succession of layers as in Polygor-

dius; and internally are found a similar digestive tract with

eversible proboscis, divided of the ccelom by transverse inter-

segmental dissepiments and dorsal and ventral longitudinal mes-

enteries, an oblique muscle band supporting the metameric

gonads, and an epidermal nérvous system ending anteriorly in

a bilobed cephalic ganglion. In all these and other characters

the worm resembles Polygordius, but in three important respects

differs from that genus.

By an arrangement which can be described as a shortening

of the oblique muscle sheet, the thick layer of longitudinal

muscles is folded on each side in the form of a longitudinal

ventrolateral ridge in which the muscle fibres assume (in trans-

verse section) a radial arrangement and undergo certain struc-

tural changes. Thus is approached the condition found in many

of. the more primitive Polychzta and the manner in which the

segregation of the dorsal and ventral longitudinal muscle tracts

of annelids may have occurred is indicated.

At the caudal end the last ten or twelve somites are setigerous.

Two of these bear a pair of seta on each side but the others

only a single slender, vitreous, capillary seta, which arises on

each side from the place of insertion of the oblique muscle and

curves over the back for a distance nearly equalling the diameter

of this region.

520 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The anal segment or pygidium shows no enlargement nor

other peculiarities exhibited by most species of Polygordius.

There are no marginal papille or special appendages of any

sort, but the pygidium continues the general outline of the rest

of the caudal region to the anus. The caudal setze evidently

serve the same purpose as the bulbous enlargement with its

papillae of Polygordius in anchoring the posterior end of the

worm. For this worm the name of Chetogordius canaliculatus

is proposed. Its chief interest is that it somewhat bridges the

gap between Polygordius and the Polychzeta and partially con-

firms the view expressed thirty years ago by McIntosh that

Polygordius finds its nearest polychaete relations in the Ophe- .

liidæ.

The First Steps Toward Degeneration in the Parasitic Cope-

pods: Charles B. Wilson, State Normal, Westfield, Mass.— No

abstract.

Gymnandromorphous Ants; W. M. Wheeler, American

Museum of Natural History.— No abstract.

Trematode Parasites of American Frogs: HM. S. Tha

Haverford College.— Up to the present time the following

species of trematodes have been found in American frogs :—

Halipegus (Distomum) ovocaudatus Vulpian in the mouth and

eustachian recesses; Hematolechus (Distomum) | longiffexus

Staf., H. similiplexus Staf, H. breviplexus Staf., H. vario-

plexus Staf., and Ostiolum (Distomum) formosum Pratt in the

lungs ; Pleurogenes (Distomum arcanus Nickerson encapsuled

in the pylorus and the liver; Distomum quietum Staf. Cepha-

logonimus americamus Staf., D. retusum Leidy, and Holosto-

mum nitidum Leidy, in the small intestine : Diplodiscus

(Amphistomum) subclavatus Gceze in the rectum; Gorgodera

(Distomum) amplicava Looss, G. simplex Looss, G. attenuata

Staf., and G. translucida Staf., in the urinary bladder; Disto-

mum tetracystes Gal. encapsuled in the muscles of the throat ;

and Monostomum ornatum Leidy in the body-cavity.

The Structure and Development of the Compound Eye of the

Nos. 451-452. AMERICAN SOCIETY OF ZOOLOGISTS. 521

Bee: E. F. Phillips, University of Pennsylvania.— The omma-

tidia which make up the compound eye of the honey bee are

each composed of a crystalline cone of four cells and a rhabdome

surrounded by eight or occasionally nine retinular cells. Each

ommatidium is sheathed. by two kinds of pigment cells, the inner

pigment cells, two in number which surround the base of the

crystalline cone and the outer pigment cells, twelve in number

which extend the entire length of the ommatidium. Each retin-

ula cell sends in a process to the retinular ganglion which has

the property of a nerve fibre. The nerve fibres of the eye are

differentiated portions of the retinular cells which send in fine

branches to the rhabdome, which is probably the nerve ending

of the eye. There is no connection between the crystalline cone

and the rhabdome and the cone has not a sensory function.

The ommatidia develop from a single layer of thickened hypo-

dermis and the region of the compound eye is marked out in the

unhatched larva. The first indication of ommatidia is the

grouping of cells with a distinct boundary in young larve. The

cells of these groups become differentiated until each group is

composed of a spindle-shaped mass with large nuclei, surrounded

by numerous cells with smaller nuclei. The central spindle

forms the retinula and the rhabdome is early visible as a clear

space at the outer surface. The crystalline cone arises from the

sides of the retinula and the pigment cells are still farther out.

The ommatidium is formed as a morphological invagination of

the hypodermis as was held by Watase. The rhabdome and

crystalline cone are formed as intracellular secretions and the

lens is secreted by the inner pigment. cells whose nuclei in the

young pupal eye are distal to the cone and afterward migrate to

the base of the cone.

THE ANATOMY OF THE CONIFERALES.

(Continued from page 359).

D. P. PENHALLOW.

RresiNous TRACHEIDS AND RESIN CELLS.

THE investigations of Eichler (11, p. 35) show that in Ginkgo

the wood is characterized by the presence of wood parenchyma

elements which take the form of short idioblasts of a lenticular

form in longitudinal section, and are distinguished by the stor-

age of crystals of calcium oxalate. These structures are pecu-

liar to this genus in which they form a specific character of

definite value, but a more detailed account of them at this time

is not called for.

In a large proportion of the Coniferales the wood is charac-

terized by the presence of more or less numerous wood-paren-

chyma cells. These are always distinguished by their cylindrical

form and transverse terminations. They are invariably associated

with the production of resin, either as entering into the compo-

sition of resin passages, or as isolated cells. It is this latter

group with which we are most particularly concerned at the

present moment and as, with very few exceptions, they are uni-

formly characterized by the presence of resin which gives them

a distinctive appearance, I prefer to describe them as “resin

cells" rather than by the more commonly employed designation

of * wood-parenchyma," which conveys no suggestion of their

special function and most prominent feature.

Before proceeding to consider these structures more in detail,

it will be necessary to digress for a short timeand discuss certain

other elements which have been erroneously regarded as wood-

parenchyma. It has been stated by Eichler (11, p. 35) that

wood-parenchyma elements occur in Araucaria and Agathis, but

this is evidently due to a wrong interpretation of certain features

presented by species of those two genera, which, according to

5*3

524 THE AMERICAN NATURALIST. [Vor. XXXVIII.

our investigations, are wholly devoid of such structures, in the

sense defined above.

In Araucaria excelsa a transverse section shows more or less

numerous elements containing resin. These are not to be dis-

tinguished in their general structure from the surrounding tra-

cheids, and they are to be recognized solely by their contents,

which are usually somewhat prominent. Their distribution is

characteristic. They occur in small, scattered groups, or more

commonly in rows one or two elements wide, parallel with the

medullary rays and in immediate contact with them on each side.

When the plane of section passes near the position of the sup-

posed terminal walls the latter may be seen to be cut through in

various ways, but they never exhibit any structural features, and

tn they are therefore in no way compar-

able with the terminal walls of the

wood-parenchyma cells. In a radial

section they are seen to be long and

fusiform, exactly resembling the wood

tracheids, except for a reddish brown

transverse plate which occurs either

close to or exactly opposite a medul-

lary ray, a position which is more

clearly shown in a tangential section

(Fig. 35). The dark plates closely

resemble Sanio’s bands, for which

they might very readily be mistaken

upon casual observation, or they might

likewise be mistaken for terminal and

unpitted walls. In Agathis australis

these features are represented in their

typical form. The transverse section

" Modes sie, Tra shows such elements to be numerous

of the resinous tapas and disposed in radial rows on each

Puce the medullary rayatz. side of the medullary ray (Fig. 32).

In a radial section they present the

porn fibrous and fusiform Character as in Araucaria, but in

addition the wall usually experiences a marked increase in

secondary growth within a region exactly opposite the ray (Fig.

Nos.451-452]] ANATOMY OF THE CONIFERALES. 525

33). This feature is also prominent in a transverse section

(Fig. 32). Such local increase in thickness always arises in

adjacent cells in such a way that the more strongly thickened

regions are exactly opposite, and they serve to constrict the cell

cavity gradually from above and below in such a way as to leave

a channel about half the usual width of the cell cavity, which

gradually widens upward and downward (Fig. 33). It is at the

position of maximum constriction that we find a transverse

plate of variable thickness, but always of a reddish brown color.

These plates are always thinnest in their central region, and

they may be of uniform thickness for the greater part of their

extent. At the region of contact with the tracheid wall they

become thicker and thereby attain a vertical distribution to an

extent four or five times greater than the general thickness. At

such position also there is somewhat clear differentiation between

the plate and the wallin point of color. Such plates show abso-

lutely nothing of the nature of pits, and they are in no sense

comparable with the terminal walls of the wood-parenchyma

cells, except in form and position (Fig. 33).

The peculiar position of these plates, their resinous color and

their simulation of both Sanio's

bands and terminal walls, excited

a suspicion as to their true nature

and led to the belief that they

might not be structural features

atall. They were therefore sub-

jected to a series of careful tests

to determine (1) if they were

structural, (2) if they were resin-

ous, and (3) if the latter, to

what extent. It was recalled in

this connection that, although , _ a d UE Bd uds

devoid of any special secretory showing the local thickening of the tracheid

^ and the occurrence of resin plates (7. 7 )

reservoirs in the wood, Agathis éppieiie k midallary ap. ;

is nevertheless well known for

its production of the resin called gum dammar. It was sus-

pected that the plates might be local deposits of resin, and they

were therefore brought into direct comparison with gum dammar,

526 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the characteristics of which are well known and described by

Wittstein (D. 53, p. 63). Tests were applied to thin radial and

tangential sections, employing for this purpose (1) various essen-

tial and fixed oils, (2) ether, (3) alcohol, (4) ammonia, (5) potas-

sium hydrate in one and one half percentage solution. The

plates were found to be very refractory with respect to both the

fixed and essential oils, as well as towards ether, alcohol, ammo-

nia and xylol, and in all of these cases no change was to be

observed, even after an action extending over several weeks. A

partial exception applies to alcohol and ether. In the latter case

there did appear to be a certain diminution in volume, apparently

through solution, when the

reagent was first applied,

but after that there was no

further alteration. The

application of alcohol, both

in the hot and in the cold,

showed that while the resin

contained in the medullary

rays was all dissolved, the

plates were only partially

affected. The reaction of

the reagent was chiefly manifested in the

development of strong curvature, often

accompanied by fracture. This was evi-

dently due to an increase in volume, and

a tendency toward solution, and it gave

the first definite evidence that the plates

could not be of a cellulose character. Be-

yond this no further change was brought

about, even after several weeks of action,

The potassium hydrate gave the most

positive results. At first there was no

apparent change, but after an interval of

about ten days or two weeks the plates ;

a oe com dene "a i TTE

y clear channel in the cel] therelation of the resin per

petites oe proot of the resinous ? factored ped be X

ese plates is to be found in %5

-——

Fic. 34.— Agathis australis. Radial section showing

the origin of the resin plates (7. 7.). X 225.

Nos. 451452] ANATOMY OF THE CONIFERALES. 527

the ruptures which they not infrequently exhibit (Fig. 35), and

in the various developmental stages which may be seen not

infrequently (Fig. 34). These show that resin gathers locally

upon the inner face of the tracheid wall, and as its volume

increases it projects toward the center from all sides, until com:

plete coalescence occurs.

The facts thus obtained proved most conclusively that the

transverse plates are obviously resinous and not of the nature

of cellulose, even partially, and the conclusion appeared to be

justified that they consist of gum dammar, but of a highly modi-

fied and highly refractory character. The same evidence also

conclusively shows that the cells in which the plates are devel-

oped are normal wood tracheids and not wood parenchyma, which

is altogether unknown in both Araucaria and Agathis, within the

limits of the investigated species.

We are naturally led to ask what is the purpose of these resin

plates? The peculiar form in which the resin is deposited,

and the particular location of the plates, points with much force

to their connection with some functional activity, since if it were

simply a question of storage of the secreted products the latter

would hardly be disposed as found but rather after the manner

common to so many of the Cupressinez ; and this suggestion

gains strength from the fact that both in the particular form of

the resin masses and their location in the tissue, Araucaria

and Agathis are peculiar among all allied genera. No exact

comparison can be established with other plants, and it is dif-

ficult to suggest an explanation which is adequate. One thing

does seem clear, however, and that is that since these plates are

of an impervious nature and developed, in some cases at least,

in connection with a special constriction of the tracheid cavity,

they offer, and possibly they are specially designed to afford, a

definite obstruction to circulation in a vertical direction. In

this sense they may be designed to serve the same general

purpose that is accomplished by the development of thyloses in

the vessels of the angiosperms, or in the resin canals of the

higher Coniferz. It is therefore possible that they may be

connected in some way not at present clear, with a more com-

plete restriction of the circulation to a radial direction, and

528 THE AMERICAN NATURALIST. [Vor. XXXVIII.

particularly through the medium of the medullary rays as

specialized channels for that purpose.

The occurrence of such resinous tracheids is almost exclu-

sively confined to Araucaria and Agathis, in which it is a feature

ef particular species, but it is a noteworthy fact that similar

structures occur, though rarely, among the higher Coniferze. In

the genus Abies they are prominent features in both 4. fraseri

and A. grandis. In the former a transverse section shows them

to be prominent and scattering through the summer wood, more

rarely in the spring wood, while in the radial section the resin is

seen to be massive in the summer wood, forming a peripheral

layer in the spring wood. In A. grandis the resin is usually

more abundant, but otherwise the features are the same.

The taxonomic value of the resin tracheids applies exclusively

to Agathis and Araucaria, where they are of specific value, and

permits of the differentiation of at least one species in each

genus. In Abies such tracheids are so sporadic and present so

little constancy as to be of no great value.

Returning to a consideration of the resin cells, these struc-

tures are found to be entirely wanting in those species of Taxus

(4) and Torreya (3) which are included in the present studies.

They do occur, however, in Podocarpus where they present the

usual structural features, but they are there remarkable for their

number and the great abundance of massive resin which they

contain. This distribution in the Taxaceze does not altogether

accord with the conclusions of Eichler (11, p. 35) who states

that they occur very sparingly in Taxus, but he makes no

mention whatever of their presence in Podocarpus where they

are: much too prominent to escape even the most casual obser-

vation.

In the Coniferæ, resin cells are characteristic of all genera

ML Picea and Pinus, where they are completely replaced by

resin passages. They are therefore features in the wood struc-

ture of twelve genera, and they are constant characteristics of

all their species, with very few exceptions. Such exceptions

apply exclusively to the genus Abies, in which four species — 4.

f npe. A. lasiocarpa, A. veitchi, and A. balsamea — are wholly

devoid of such Structures, ;

Nos.45t-452.] ANATOMY OF THE CONIFERALES. 529

The recognition of the resin cells presents no difficulty inthe

great majority of cases because of the abundance and depth of

color of the resinous contents. This finds its most complete

expression in Taxodium, Sequoia, Cupressus, etc. In Abies, on

the other hand, where these cells have experienced extreme

numerical reduction and where there also seems to be a corre-

sponding reduction in their secretory power, it is impossible to

recognize them in this. way. In such cases it is often possible

to distinguish them by their slightly different form and some-

what thinner walls as compared with the adjacent wood tracheids ;

by their situation slightly in advance of the outermost row of

summer wood tracheids; and most particularly by their pitted

terminal walls when the latter lie near the plane of section.

This last feature may also be relied upon in all other cases when

any element of doubt is involved (Fig. 36). In longitudinal

section the characteristic form of the cell serves to distinguish

Fic, 36.— Abies amabilis. Transverse section showing the positions and structure

of the resin cells (7. c.) on the outer face of the summer wood (S.

it beyond all doubt, even in the absence of resinous contents.

Whether exposed in radial or tangential section, the cell has the

form of a narrow cylinder upwards of 300 » in length, and

always several times longer than broad, except in cases where

there is a definite tendency, through aggregation, to the forma-

tion of resin canals.

The resin cells sometimes occur in pairs, but more generally

as isolated structures separated by one or more tracheids. The

terminal walls are transverse and more or less strongly marked

with simple pits. The side walls, especially the radial, are pro-

530 THE AMERICAN NATURALIST. [Vor. XXXVIII.

vided with simple pits, though often few in number, and this

feature serves to a large extent, to assist in their differentiation

from adjacent tracheids of similar form (Figs. 36 & 38). It

nevertheless not infrequently happens that in transitional forms,

such as are met with in Seguota sempervirens (Fig. 37c),

bordered pits occur on the lateral walls.

The resin is in all cases massive and often very abundant.

Pie: ye Sequoia sempervirens. Radial sections showing (a) the form of the resin cells

and the associated parenchyma tracheids; (4) resin cells f he spring wood showing

pong of the resin; (c) resin cells showing transitional forms with bordered pits.

In such genera as Taxodium or Sequoia it completely fills the

entire cell'cavity (Figs. 39 & 40), but in Larix, Tsuga and

Pseudotsuga it takes the form of a peripheral layer in immediate

contact with the inner face of the cell wall (Fig. 42). The

reduction thus indicated is, in some species, carried to such an

Nos. 451-452.] AWA TOMY OF THE CONIFERALES. 531

extent that the resin is barely recognizable, while in Abies it is

wholly wanting.

A relation of more than ordinary interest is to be found in

the relation of the resin cells to certain forms of tracheids. In

Sequoia sempervirens it commonly happens that the resin cells

lie in immediate contact with tracheids of special form. These

structures are wholly unlike the wood tracheids among which

they are found, but they are, in all essential respects, like the

tracheids of the medullary rays. They have the form of long,

cylindrical elements with abrupt terminations, and they thus

bear an external resemblance in form to the wood parenchyma

cells with which they are associated. They differ, however, in

the distinguishing presence of bordered pits upon their side and

terminal walls (Fig. 3842). The relations of these two elements

is nevertheless a much more intimate one o

than is implied by mere association. In

Sequoia an interchangeable relation is

manifested as already pointed out, in the

occurrence of resin cells with bordered pits =a

(Fig. 37), while in Adzes amabilis (Fig. 38)

resin cells and tracheids also form a coter-

minous series. It is thus obvious that we

have here precisely the same interchange- ©

able relations that have been found to o

occur in the medullary rays, and it is evi- © V

dent the one element must arise through

modification of the other. The precise

order of this sequence is not altogether

clear from the available data, but the fact

that ray tracheids are derived from their a b

associated parenchyma cells, and that in F M d^ des apice

such types as Podocarpus, 'Taxodium, etc., de scáihus ul the parii-

the resin cells occur without tracheids, į

while the latter do occur in Sequoia and «æ and 4 being normally

. . coterminous, X 200.

especially in Abies, seem to justify the

inference that here also they are derived forms, having their

origin substantially in special modifications of the parenchyma

elements. In view of these relations, it is necessary to distin-

532 THE AMERICAN NATURALIST. [Vor. XXXVIII.

guish such elements as parenchyma tracheids in order to estab-

lish their proper identity and differentiate them from the wood

tracheids, which have a wholly different origin, as well as from

the ray tracheids, which have a wholly different location. It is

probable that the parenchyma tracheids also serve a similar pur-

pose to the ray tracheids with respect to the distribution of

nutrient fluids. The origin of the parenchyma tracheids as sug-

gested finds support in the statement of Eichler (11) that the

wood parenchyma arises through the activity of the cambium

cells, abundantly in the Cupressineze and Abietinez, forming in

exceptional cases the epithelium of the resin canals, since it at

the same time shows how the parenchyma tracheids arise and

how they may be intimately connected with the wood-paren-

chyma ; but it finds additional support in a knowledge of the

genesis and structure of the resin passage.

In Sequoia and Abies we have two genera which are remark-

able for their transitional forms of structure, affording a fairly

clear conception of the genesis of the resin passage. In each

case there is a well defined tendency toward the aggregation of

the resin cells into compact groups which take the form of

longitudinal strands, enclosed on all sides by the accompanying

parenchyma tracheids. Under such circumstances the indi-

vidual cells undergo a continual reduction in length until they

eventually become but two or three times longer than broad,

or they may even become isodiametric. This change is not

accompanied by any alteration in the thickness of the walls in

the earlier stages of development, but as a result of such a

shortening the effect is to bring about the concentration of a

greater number of simple pits within a given area. Such cells,

therefore, are always more strongly pitted than those which are

isolated and of greater length. When aggregates of this sort

have attained to a certain degree of development a line of

cleavage arises in the center of the mass and results in the for-

mation of an intercellular space which, according to Eichler (11),

always arises schizogenously. This space is short and either

isodiametric or but little longer than broad, the length coincid-

Ing with the principal axis of growth. Such cyst-like reservoirs

Or sacs represent the primitive form of the resin canal, and they

E CONIFERALES.

ATOMY OF THE

52.] AN.

Nos. 45

They

are typically developed in Sequoia, Abies and Tsuga.

always form a continuous series extending in a direction parallel

with the axis of growth; but as the type of reorganization ad-

vances they merge, forming a continuous” canal such as may be

found typically in Pseudotsuga or Pinus.

From these state-

ments, then, it is clear that the parenchymatous resin cells

undergo modification in two directions, passing into parenchyma

tracheids, on the one hand, and on the other becoming shorter

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Transverse section showing the scattering distribution of the

Fic. 39.—Thuya dolabrata.

X 55.

resin cells.

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and shorter, according to conditions of aggregation

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534 THE AMERICAN NATURALIST. [Vor. XXXVIII.

forming an epithelium and (3) the central reservoir in the form

of a cyst or canal. This structure is fully exemplified in the

genus Pinus, where the highest form of development is attained.

While the occurrence of resin cells in particular genera is a

feature of great taxonomic value, their importance in this respect

is greatly emphasized by the particular form of their distribution

and the constant tendency they exhibit toward the formation of

definite aggregates. In Thujopsis and Cryptomeria (Fig. 39)

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Fic. 40.— Sequoia sempervirens. Transverse section showing scattering resin cells in the

Spring wood, and contiguous resin cysts with aggregates of resin cells on the outer face of

a growth ring. X 55.

the resin cells are always scattered throughout the entire trans-

Verse section and they show no tendency to the formation of

aggregates. In Podocarpus, where there is a notable increase

in numbers, the same general law of segregation prevails, but

there is nevertheless a somewhat well defined tendency toward

aggregation. In Thuya 66.6 % of the species show definitely

scattering cells, 33.3% show the cells to be scattering with a

Nos. 451-452.] 4J.VA TOM Y OF THE CONIFERALES. 535

tendency toward a more compact disposition, while in 33.3 % the

cells fall into-well defined aggregates or an approximation to

such an arrangement. The genus Sequoia is characterized

chiefly by the widely scattering distribution of the resin cells

(Fig. 40), but in .S. sempervirens there are individual cases in

which there is also a definite aggregation into groups. In Cu-

pressus 53.9% of the species are distinguished by the presence

of widely scattering cells which become definitely arranged in

Fic. 41.— 7.

zone. X 55.

zones in 38.4%, and aggregated into groups in 7.7 % of the spe-

cies. It will be observed here that this feature of distribution

is, on the whole, more pronounced in the relatively primitive

genera, and that it diminishes in force in the genera of a rela-

tively high order.

In Taxodium (Fig. 41) and Libocedrus, both of which are

distinguished by the presence of very prominent resin cells, these

Structures are disposed in well defined zones which are concen-

4 TURALIST. [VOL XXXVIII.

AMERICAN N

536

tric with the growth rings and lie either in the spring or summer

wood, or in both. This is to be interpreted as a definite tend-

ency to aggregation which is nevertheless not fully expressed,

since in each case there are numbers of cells which are not

zonal in their distribution, but which conform to the law appli-

In Juniperus the cells are

cable to Thujopsis and Podocarpus.

typically zonate, being also scattering in only one species.

These are

Abies only 63.6 % of the species bear resin cells.

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‘Transverse section showing the distribution of the resin cells

F16. 43.— Tsuga pattoniana.

on the outer face of the sum

mer wood. X ss.

previous types

at in 50 % of such cases, or in 36.3 f

neither scattering nor zonate in the sense of the

but it is to be observed th

of all- species

groups as a preliminary

, they are aggregated in

On the other hand,

show the resin cells to be few, inconspicu-

resin passages.

step to the formation of

of all species

36.3

ous, non-resinous and scattered alon

mer wood.

IS to be re

c to the final

g the outer face of the sum-

se in distribution leading

This, for reasons which will appear more fully later

arded as a pha

Nos. 451-4452] ANATOMY OF THE CONIFERALES. 537

obliteration of such structures, which is fully accomplished in

36.4 % of all the species as represented by A. balsamea, A.

fraseri, A. lasiocarpa, A. veitchi. This last form of distribution

is wholly typical of Tsuga (Fig. 42), in which there are no other

resin cells than those on the outer face of the summer wood.

Finally, in Picea and Pinus, there are no separate resin cells in

any of the situations described, since they have been completely

replaced by highly organized resin passages. It thus appears

that the distribution of the resin cells presents four variants

which bear a direct relation to the organization of resin passages,

as the latter eventually replace the former. These facts will

appear somewhat more clear from the following summary :

TABLE SHOWING PERCENTAGE DISTRIBUTION OF RESIN CELLS.

lOn the outer

No. of |% of occur-| Scattering. | In zones. | Grouped. ltace of sum-

species. rence. : | er wood.

Ginkgo I 000.00 |

Agathis I 000.00 |

Araucaria 3 000.00 |

Torrey $ 000.00 |

Taxus 4 000.00 |

Thujopsis I 100.00 | 100.00 |

Cryptomeria I 100.00 100.00

Podoc [ 100.00 | 100.00 | (100.00)

Thuy 2 66.60

= 3 100.00 |. 2230

x I (33-30)

Sequoia a nen 100.00 100.00

I 50.00

Cupressus . 7 53-90

5=9 100.00 38.40

I 7-70 |

: : I 100.00 | (100.00) | 100.00 |

Libocedrus I 100.00 | (100.00) | 100.00 |

Juniperus . men 100.00 00.00 |

I I.IO j

Abies 4557 100.00 | 36 30

5 45.50

Tsuga . eak 100.00 33.30 s

5 100.00

Pseudotsuga . 2 100.00 100.00

Larix Senne 4 100.00 100.00

Picea IO 000.00

Pinus 4t 000.00 |

Figures in parentheses refer to exceptional forms of occurrence.

538 THE AMERICAN NATURALIST. [Vor. XXXVIII

From such data it is clear that the distribution of the resin

cells bears an important relation to the recognition of sub-generic

groups and even of species. But viewing these structures from

the broader standpoint of the Coniferales as a whole, it is

obvious that they must be placed among the structural elements

which belong to the first rank for taxonomic purposes.

We are now in a position to determine what relation, if any,

such resin bearing elements bear to questions of phylogeny, and

we may first of all consider the resinous tracheids. These

structures have been seen to be peculiar to Agathis, Araucaria,

and Abies, in which they occur only in certain species. In

answering this question, we cannot avail ourselves of evidence

derived from fossil plants, since it is in such cases of a nega-

tive character. Neither Cordaites nor Araucarioxylon affords

definite proof of the presence or absence of such structures,

since they do not appear in any of the published diagnoses, and

our own studies have not resulted in their recognition. If

originally present, they must have been obliterated in the course

of fossilization. We must therefore depend entirely upon such

evidence as is afforded by existing species. From this point of

view it is obvious that they furnish no evidence as to the-origin

of either of the three genera in which they occur. It is, on the

other hand, possible to determine from other data, that both

Agathis and Araucaria are much inferior to Abies in point of

structural organization and development, and from this we may

be permitted to conclude that the resin tracheids.of Abies are

vestigial forms of elements which were typically developed in

Agathis and Araucaria, and possibly characteristic also, of their

Progenitors. If such inferences are to be regarded as justifiable,

they go far to support the idea of a. common origin for all three

genera, and they thus lend force to conclusions which lead to

the same result, but upon the basis of independent data.

From a study of the distribution of the resin cells, it is

apparent that they fall into four categories in which the typically

segregated cells may be held to represent the most primitive

form of disposition. This view is greatly strengthened by the

observation that in all such cases, the resin cells are rarely if at

all accompanied by parenchyma tracheids, while the structure of

Nos. 451-452.] dAWATOMY OF THE CONIFERALES. 539

the cell is farthest removed from that which is found to enter

into the composition of resin passages, whence they are also to

be regarded as of a primitive character. This view is supported

by the observed fact that those genera and species in which such

segregations occur, are also of a relatively primitive type. With

an advance in organization, there is a tendency to the formation

of aggregates as expressed in the zonal distribution of Taxodium,

Libocedrus or Sequoia, where we also find the definite formation

of groups of cells which later exhibit the initial stages in the

formation of a definite canal. But in Sequoia, as also in Abies

where similar changes take place, the more complete aggregation

of the cells is invariably accompanied by structural alterations

whereby they become greatly shortened and more strongly

pitted, while they are always accompanied by parenchyma

tracheids with which they are interchangeable. In this connec-

tion it is also to be noted that the aggregates in Sequoia, Abies

and Larix leading to the formation of resin sacs, are always dis-

posed in a zonal manner, conformably to the zonal disposition of

the separate elements, a relation which is in direct harmony with

the view already advanced, that the zonal disposition of the

isolated cells is an advance upon the strictly segregated form,

and that it leads directly to the formation of resin passages.

Following upon the zonal distribution, a more complete aggrega-

tion results in the formation of local groups of short, resin cells

ultimately leading to the formation of a true resin canal. Such

a feature of distribution, occurring in genera which, from other

data, may be shown to be relatively high in development, is in

itself significant ; but we further find that the scattering, zonal

and grouped forms bear such relations to one another, that the

real succession is in the order already given. Thus while both

species of Sequoia are characterized by scattering cells, S. sem-

pervirens also shows them aggregated to form groups and

eventually imperfectly organized resin canals. Or in Cupressus,

the transition is expressed in a more complete form, involving

all three modes of distribution. In Tsuga there is an obvious

tendency toward the elimination of the resin cells which are now

greatly reduced in numbers and confined to the outer face of the

summer wood. In Abies a similar tendency is also manifested,

540 THE AMERICAN NATURALIST. (VoL. XXXVIII.

but it is expressed ina different way, and just here we must

note a fact of more than ordinary significance. Resin cells are

present on the outer face of the summer wood in 4. grandis,

A. concolor, A. amabilis and A. magnifica. Groups of resin

cells are present in A. nobilis, A. concolor, A. bracteata and

A. firma, but it will be seen that in only one case — A. concolor

—are the two forms of distribution presented in the same

species. This is in direct conformity with the idea that the

resin passage eventually displaces the resin cell, bringing about

an obliteration of the latter, and it goes far to support the idea

that with respect to these particular structures, the genus Abies

occupies a transitional position, standing next to Picea and

Pinus, from both of which the resin cells have completely dis-.

appeared, Furthermore, from another point of view, the grad-

ual replacement of the resin cells appears to be indicated by a

corresponding reduction in the contained resin. Nowhere is the

resin so abundant in the resin cells, as in those genera like

Podocarpus and Taxodium, which show no development of resin

passages, even in their most simple forms ; but with the develop-

ment of resin sacs, as in Abies or Sequoia, or of resin passages.

as in Larix and Pseudotsuga, there is a remarkable diminution

of the resin, apparently in direct response to its more ready

production by more specialized structures.

The genus Abies then, appears to form a transition group,

having parallelisms with Agathis and Araucaria through tbe

occurrence of resinous tracheids ; with Thuya, Cupressus, etc.,

through the survival of isolated resin cells approaching oblitera-

tion; with Tsuga, Larix and Pseudotsuga through the develop-

ment of rudimentary resin canals leading to.the formation of

definite resin passages ; and with Sequoia through the survival

of isolated resin cells and the development of rudimentary resin

canals. Through these parallelisms the connection appears to

be most direct on the one hand. with Sequoia, and on the other

with Tsuga. This relation of Sequoia to Abies has been shown

by Penhallow on former occasions (38), and has more recently

been indicated in other ways by Jeffreys (24), but so far as the

present evidence is of value, it would not permit us to infer that

Sequoia, Abies and Tsuga form a continuous and coterminous.

Nos. 451452} ANATOMY OF THE CONIFERALES. 54I

series in the order given, but rather that they represent separate,

though short, side lines of*development, between which the

general sequence is manifested.

RESIN PASSAGES.

Structural.

Our studies of the resin cell have shown how peculiar aggre-

gates of these structures lead in a natural way to the organiza-

tion of resin passages, the structure of which it is now necessary

to discuss somewhat in detail, and in doing so it will be most

profitable to have reference to (1) the primitive form, (2) the

intermediate form and (3) to the advanced or fully organized

form.

The primitive form of the resin passage is to be found in

Sequoia, Tsuga, and Abies and inasmuch as within these genera

they exhibit differences in organization which correspond ap-

proximately to the sequence given, it will be necessary to discuss

them somewhat in detail, with special reference, however, to

Sequoia. This genus possesses special interest with respect to

the occurrence and organization of secretory reservoirs, since it

is in all probability not only the most ancient genus in which

such structures occur, but it is, so far as I am aware, the only

genus affording special data with respect to important variations

of structure and mode of occurrence. Being also, on the whole, :

the most primitive of the three genera, I shall deal with it first.

In Sequoia sempervirens the secretory reservoirs occur in

rows within the initial layers of the spring wood, and they there-

fore lie exactly on the outer face of the summer wood of the

previous year. Within this row the reservoirs are contiguous

and in many cases they become confluent so as to form a more

orless extended and continuous compound reservoir lying tan-

gentially, In their most rudimentary forms they present the

aspect of simple aggregates of resin cells without any differenti-

ation of a resin sac or of an epithelium. In a more advanced

Stage of development there is produced a central cavity in the

form of an intercellular space (Fig. 43, C) which has obviously

542 THE AMERICAN NATURALIST. (Vor. XXXVIII.

originated schizogenously. About this the resin cells are gener-

ally flattened radially and disposed in such a manner as to sug- .

gest the future development of a definite, limiting layer or

epithelium. In the completed form of the structure the central

space has broadened out and taken a circular form, assuming the

character of a definite cyst bounded by as definite a limiting

epithelium in which the cells are always flattened radially and

disposed concentrically (Fig. 43, C). Externally to these cells

there may be a second layer of similar resin cells, constituting

the outer epithelium, while the whole is enclosed on three sides

by a layer of parenchyma tracheids which are exceedingly like

E: YN

NES

Fic

- 43-— Sequ pervi Transverse section showing two contigu-

ous resin cysts, C, completed and with a normal epithelium Æ; C’ an

intercellular space as rudiment of a cyst with imperfectly developed

epithelium.’ 7. R. the medullary ray; S. W. the summer wood. X

the associated tracheids ot the spring wood, but from which they

may usually be distinguished by (1) their greater size and rela-

tively thinner walls, (2) the occurrence of bordered pits on the

tangential and terminal, as also upon the radial walls. Such

parenchyma tracheids never occur in the adjacent summer wood

for very obvious reasons, but on the radially opposite side of the

reservoir they are very commonly flattened radially (Fig. 43).

and they not infrequently present the same structural aspects as

the epithelial cells. The interchangeable relation between resin

cell and parenchyma tracheids as already shown would lead us to

Suspect a substitution in the composition of the epithelium, and

Nos. 451-452.] ANATOMY OF THE CONIFERALES. 543

such substitution does actually occur, since it is often to be

noted that the second and third rows may be made up, at least

in part, of tracheids.

In a longitudinal radial section the reservoir is found to have

the form of a sac of vary-

ing form and size, but

generally elongated par-

allel with the axis of

growth and completely

closed at both ends (Fig.

44). The epithelium

which immediately de-

fines the limits of the

sac generally consists of

short, cylindrical cells,

while in the second or

outer layer the cells be-

come much elongated

and several times longer

than broad. Beyond

this, the third layer con-

sists of parenchyma tra-

cheids readily distin-

guishable whenever the

terminal walls lie near Fic. 44.— Sequoia sempervirens. Radial section of a resin

the plane of section, Or cyst, showing the epithelium (z4.) ; the central cyst (C)

. . with a thylosis (£4); parenchyma tracheids (7r. £.), and a

otherwise as already in- „acheni of the spring wood (Sf. T) X 300.

dicated. Certain devia- ;

tions from this typical structure require examination. The resin

sacs are placed in vertical series of indeterminate extent, but at

varying intervals of such a nature that they may sometimes be

separated only by a rather thick wall of short resin cells. At

other times they are somewhat distant and separated by an

extensive vertical tract of resin cells. From this it is obvious

that in any given plane of section there will be a great diversity

of aspects presented, but in the main exhibiting structural grada-

tions in the development of the reservoir as already recounted.

In some cases thick-walled cells of circular outline may be seen

544 THE AMERICAN NATURALIST. (Vor. XXXVIII.

in transverse section to stand out from the general line of the

epithelium and lie within the cavity proper. More rarely such

cells are so multiplied as to fill the entire cavity, and they may

themselves be filled with granular resin. Such features are

Fic. 45.— Sequoia sempervirens. ‘Transverse section of a resin cyst show-

ing an associated resin cell (7. c.); the epithelium (e/.); the thyloses

(/A.), one of which contains resin, and one of which is directly derived

from an epitheli ll; the parenchyma tracheids (fr. ¢.). X 300.

clearly defined (Fig. 45), and it is evident from the way in

which such cells originate from the epithelial cells that they are

of the nature of thyloses. A longitudinal section through such

a reservoir (Fig. 46) shows how such thyloses occupy the entire

cavity of the cyst, while in other cases they may be purely

local (Fig. 44). Among fossil sequoias similar thyloses form à

most characteristic feature in the resin passages of the medullary

rays in S. burgessii.

In Tsuga caroliniana there are no secretory reservoirs, but

Just in the region between the spring and summer wood of the

same growth ring there are peculiar aggregates of resin cells of

a more or less rounded outline but forming a continuous series

of considerable extent. An analysis of these aggregates shows

them to be composed of thick-walled and rounded resin cells,

among which there mày be a small central intercellular space

without any definite organization of epithelium. In such aggre-

Nos. 451.452] ANATOMY OF THE CONIFERALES. 545

gates the component cells are far less resinous than the isolated

resin cells of the same section.

The parenchyma tracheids are not

clearly distinguishable from the

associated wood tracheids. In

radial section the cells are seen to

be very variable, thick-walled and

sometimes with more or less promi-

nent intercellular spaces. Between

the rays they are several times

longer than broad, but opposite the

rays they are short, cylindrical and

more copiously pitted: while some-

times they may be seen to merge

into ray elements and thus to con-

tinue their course at right angles

to their primary direction. A care-

ful comparison of these cell aggre-

gates with those of Sequoia and

Abies leaves little room for doubt

as to their structural and functional

identity, and we cannot do other-

wise than conclude that they repre-

sent the most primitive structural

condition which is capable of di-

rectly giving rise to definite cysts

by central cleavage.

In 7suga mertensisana the secre-

tory reservoirs are disposed like section of

those of Sequoia, on the outer face ETE e Dd (CUM DM.

of the summer wood, where they areresinous. This figure corresponds to

form tangential series. They ex- "'* ^^*

hibit all the gradations from simple cell aggregates without a

central space to perfectly formed cysts with a definite epithe-

lium. This latter is in one, more rarely in two rows, and it is

composed of more or less rounded or radially flattened elements.

The parenchyma tracheids are few in number, and they are not

readily distinguishable from the adjacent wood tracheids. In

Fic. 46.— Sequoia sempervirens. Radial

i a resin cyst showing the epithe-

546 THE AMERICAN NATURALIST. [Vor. XXXVIII.

longitudinal section the reservoirs are variously rounded or

oblong cysts, contiguous or isolated, and forming a longitudinal

series. In their general form and structure they are essentially

the same as in Sequoia.

In the genus Abies secretory reservoirs occur in at least four

species where they form more or less extensive tangential series,

within which they are usually contiguous and more or less con-

fluent. They present the same general variations in structural

organization as in Tsuga and Sequoia, but in A. concolor, and

less conspicuously in A. nobilis, they are often extended in a

radial direction so as to become narrowly oval or oblong, and

several times longer than broad. The epithelium consists of a

well defined structure composed of one to three rows of cells.

The first row, immediately bordering upon the canal, consists of

rounded or oval and thick-walled cells which are much smaller

than those of Sequoia, and similar to those of Tsuga. They are

_ always characterized by an abundance of strongly defined, simple

pits, and many of them contain resin, which usually takes the

form of rounded granules of diverse sizes. The parenchyma

tracheids are so nearly like the accompanying wood tracheids as,

in some cases, to be separable with some difficulty, but they

generally surround the resin sac, at least within the limits of the

spring wood, and they not infrequently replace the parenchyma

cells of the epithelium more or less completely. Not infre-

quently they form somewhat extended radial series from the

epithelium into the spring wood, as in Picea (Fig. 48). In such

cases they are usually recognizable by their rather unusual size

and thinner walls, and in addition they commonly show bordered

pits on the tangential walls. When the terminal wall lies suffi-

ciently near to the plane of section, it shows from one to several

rather large bordered pits, and by this feature such tracheids

mined be located with much certainty. Thyloses have been defi-

nitely noted only in the case of 4. concolor, in which species

they are essentially of the same general character as in Sequoia,

They are thick-walled and either isolated or so numerous as to

a oo In one case of contiguous cysts, an epithe-

- ed und to form thyloses in both cysts — in the one

case giving rise to an isolated cell, in the other forming a tissue

which nearly filled the entire cavity.

Nos. 451452] ANATOMY OF THE CONIFERALES. 547

In radial section the reservoirs are round or oblong cysts of

variable size, and they are either contiguous or distant. In the

former case they rarely or never become confluent, but they

maintain their separate identity as in Sequoia and Tsuga. In

the latter case the intervening region is occupied by an aggre-

gate of resin cells in all essential respects like those in the same

regions of Sequoia and 7suga mertensiana, or like the aggregates

which are generally characteristic of Tsuga caroliniana. The

inner epithelium usually consists of short, cylindrical and strongly

pitted cells which, in the second and third rows, become suc-

cessively longer and less strongly pitted, so that those in the

outer row may be identical in form and markings with the

isolated resin cells. In the two outer rows the cells not infre-

quently show bordered pits on their radial walls, thus presenting

transitional forms which gradually pass over into tracheids, and

the one then replaces the other. The parenchyma tracheids,

Which are always most characteristic of the spring wood, are al-

ways distinguished by the presence of large and prominent bor-

dered pits, but in addition they are sometimes broad and thin-

walled, and lie in radial series.

From these facts it is clear that the secretory reservoirs of

the three genera in question always take the form of closed

sacs, which DeBary has already pointed out as a feature of cer-

tain Coniferz (9, p. 440), and in order to clearly differentiate

them from those which occur in the higher Coniferz, I shall

reserve for all such cases the term resin cyst. That such sac-

like reservoirs represent the primitive form of the resin passage

scarcely admits of question when we observe the various transi-

tional forms which they present and the relation they bear to

the resin passages of the higher Coniferæ — a view which is

strengthened by the observation of DeBary (9, p. 443) that

primitive forms of the secretory reservoir occur in the pith of

Ginkgo in the form of elongated sacs.

DeBary has shown (9, p.440) that the secretory passages

traverse the wood longitudinally, at first as prismatic tubes which

usually acquire a round or elliptical, transverse section. This

Statement is applicable to Pseudotsuga, Larix, Picea and Pinus,

but inasmuch as there are important differences of detail between

548 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the first three genera and the last, in such a way that the former

represent an intermediate, whilethe latter represents a completed

type, it will be necessary to examine them separately. In all of

these cases, however, the secretory reservoir is invariably char-

acterized by the presence of a definite and continuous canal

of indeterminate length, in consequence of which I reserve for

them the appropriate and long used term, “resin passage," as

distinguished from the resin cysts of the previously discussed

genera.

In Pseudotsuga the resin passages are always scattering,

though they frequently occur in tangentially extended groups of

two or four contiguous or even coalescent reservoirs. The

central canal, which is usually small and not infrequently very

narrow, is rather more generally rounded than in previous types.

The epithelium is very clearly defined and consists of one to

three rows of thick-walled parenchyma cells, sometimes con-

taining resin, the first row of which are rather small and radially

flattened, but in P. macrocarpa they are rather thin-walled. In

P. douglasii the epithelium is commonly extended on the two

Sides of the resin canal in such a way as to form a tangehtially

elongated tract which not infrequently extends beyond and

involves neighboring medullary rays. In P. macrocarpa, on the

other hand, the epithelium is concentric with the canal, thus

forming a tract of about equal thickness all around. Such a

deviation as is expressed in P. douglasii constitutes the first

evidence of a tendency in development which is fully and fre-

quently expressed in Pinus. Thyloses are of infrequent occur-

rence, and they appear to be confined to P. macrocarpa, where

they are few in number and generally rather thin-walled. Paren-

chyma tracheids are usually not apparent in a transverse sec-

ton. This results from the frequent location of the resin

passages in the summer wood, which is not favorable to their

development, and from the close resemblance which they bear

to the tracheids of the spring wood; and while such elements

form an integral part of the resin passage, their particular dis-

position cannot be exactly defined, though there is no g

reason for supposing that they differ in this respect from what

may be observed in other cases. In a longitudinal section the

Nos. 451-542.| ANATOMY OF THE CONIFERALES. 549

canal is found to be more or less continuous, though it presents

frequent constrictions, and it is thereby reduced to very narrow

dimensions. It is this feature which causes the canal to exhibit

such marked variations in size when seen in transverse section.

The epithelium cells are narrowly cylindrical and rather long

and thick-walled, as well as somewhat strongly pitted. Out-

wardly they become much longer and relatively narrower, and

they eventually merge with the surrounding parenchyma tra-

cheids by which they may also be replaced.

In Larix the same features of contiguity and coalescence

may be observed, except that in Z. occidentalis the resin pas-

sages sometimes form into

continuous zones of imper-

fectly organized structures

with the aspect presented in

Tsuga martenstana. The epi-.

thelium is always well defined

(Fig. 47) and it consists of

one, sometimes two, rows of

cells. The cells of the first

row are small, very variable

in form and size, thick-walled

and more or less strongly

flattened radially. They are

also commonly resinous and

more or less strongly pitted.

When there is a second row

of epithelium, the cells are

essentially like the wood tra-

cheids, and like the paren-

chyma tracheids from which

talis. ‘Vransverse section

Fic. 47.— Lari. ; i

from the inner spring wood showing a pair of

resin passages with the central canals (c); the

cheid at (ør. t.) and the summer wood (s. w.).

they may be separated with

difficulty. The latter, therefore, which are absent from the

summer wood, can be distinguished from the elements of the

spring wood only when the pits on the terminal walls (Fig. 47

pr.t.) are brought into view, or, more rarely, when the pits on

the tangential walls are in evidence. Thyloses rarely occur, and

so far they have been noted only in Z. occidentalis, where they

550 THE AMERICAN NATURALIST. [Vor. XXXVIII.

are infrequent and thick-walled, and in Z. americana, where they

are of rare occurrence and thin-walled. In longitudinal section

the central canal is always continuous, though constricted at

intervals, a feature in all essential respects the same as in

Pseudotsuga. Radially, the first row of epithelial cells are short

cylindrical, or in Z. occidentalis short fusiform, but there is a

graduated increase in length outwardly, so that in the second, or

in the third row if present, they become narrow and very long,

and they eventually merge with the parenchyma tracheids through

intermediate forms with bordered pits. All of the epithelial

cells are thick-walled and strongly pitted, and they thus offer a

somewhat strong contrast to the rather thin-walled parenchyma

tracheids with bordered pits.

The resin passages of Picea differ from those of Pseudotsuga

and Larix in being more strictly segregated, and in consequence

there is a conspicuous absence of contiguous structures, which

may nevertheless sometimes be seen in P. zzgra, and especially

of coalescent forms. They are usually narrow, but well rounded

or oval and there is far greater uniformity of structure and

form than in any of the preceding types. The epithelium con-

sists of one row, one to two rows, or even one to three rows of

cells, differences which apparently belong to particular species,

though no attempt has been made to define the precise limita-

tions of such features. The cells are generally small, round or

radially flattened and thick-walled, though occasionally a cell

may be thin-walled as in P. alba. In cases of thick-walled epi-

thelium, the outermost cells merge with similar tracheids from

which they are not readily distinguishable, while the general

epithelium becomes extended into a tangentially elongated tract

as in Pseudotsuga douglasii and Pinus. Thyloses have been

noted as of occasional occurrence in P. nigra, P. pungens and

P. sitchensis, but they are always thin-walled. Parenchyma

trachieds are not obvious in the summer wood, but they are

recognizable in the spring wood where they appear to replace

the resin cells, though they are apparently of much less frequent

occurrence than in the genera previously discussed. In P. alba,

however (Fig. 48, pr. 7.), we sometimes find a radial series of

tracheids which also extends laterally so as to form an enclosing

Nos.451-452.] ANATOMY OF THE CONIFERALES. 551

layer. Radially the canal is continuous, but with more or. less

frequent constrictions as in Pseudotsuga and Larix. The epi-

thelium consists of narrow, cylindrical and much pitted cells

which increase in length in the outer layers where they become

five to seven times longer than broad, and finally they merge

with the parenchyma tracheids which replace them.

While the general composition of the resin passage in

Pseudotsuga, Larix and

Picea is the same as that ( ) LOW,

of the resin cyst, it is obvi- | O% T | (9 3 prt

ous that the frequent con-

strictions in the canal indi- ?

cate a partial survival of

the cystic formation. We

must therefore regard these

structures and the three

genera to which they be-

long as forming a transi-

tion group between the

primitive resin cyst on the

one hand and the perfectly

organized resin passage of

Pinus, with its canal of uni-

form width, on the other.

In the Benus Pinus, the Fic. 48.— Picea alba. 'Vransverse section of a resin pas-

resin passages show COn- sage from the spring wood, showing the central canal

. o 4 (25 the thick-walled epithelium (e) and the paren-

siderable variation in detail, chin enata LAC 4). 3 400.

but they all conform to the

same structural type (Fig. 49). The central canal is broad and

round, often very large, and in longitudinal section it is a per-

fectly continuous passage of uniform width. The epithelium

consists of large, but very variable and thin-walled cells in one

to several rows. In the soft pines it generally forms a concen-

tric zone of uniform width, but in several of the hard pines there

is a marked tendency to extension in a tangential direction and

the formation of rather extensive eccentric tracts. In all of the

pines there is a pronounced tendency for the epithelial elements

to become so thin-walled that they are readily broken out in

552 THE AMERICAN NATURALIST. [Vor. XXXVIII.

making sections, while in the hard pines, as P. cubensis, P. teda,

P. pungens, etc., the cells are often strongly resinous. In the

outer epithelium, the thin-walled elements may be associated

with occasional thick-walled elements with which they are inter-

changeable, precisely as in the similar relations displayed by the

medullary rays of P. pungens and P. cubensis. In the same

region also there is a similar association with and transformation

HNO

FIG. 49.— Pinus refle va. Transverse section of a resin passage from the inner face

of the sping wood showing the central canal (C); the thin-walled and resinous

Pasarela og or (); the spring wood (S2. W.) and

into parenchyma tracheids, which also has its parallel in the

medullary ray. Somewhat more specifically, special reference

to two examples may serve to illustrate the general nature of -

some of the more important variations. In longitudinal section

the parenchyma tracheids are usually of much greater length

than the associated parenchyma cells, with which they are par-

allel or coterminous, and they occur in large numbers in P.

lambertiana. In p. reflexa they are coterminous with paren-

chyma cells which they finally succeed, to be replaced in turn

by thin-walled wood | tracheids. In P. lambertiana they are

Nos.451-452.]] ANATOMY OF THE CONIFERALES. 553

always to be distinguished by the bordered pits on the radial,

tangential and terminal walls, while in P. refera they are char-

acterized by the large number of bordered pits on the radial

walls, with very few on the tangential walls. In the former

situation the pits are much smaller than in adjacent wood tra-

cheids. Together with adjacent wood tracheids, the paren-

chyma tracheids may be more or less involved in bearing resin

(P. lambertiana), while finally, as exhibited in transverse sec-

tion, their numbers may be so large that they form extensive

areas about the resin passage (Fig. 49). In such a case the

sequence of elements in transverse section would be :—

1. Canal with thyloses.

2. Thin-walled epithelium.

3. Epithelium — cylindrical parenchyma tracheids.

4. Parenchyma tracheids.

5. Wood tracheids with thin walls.

Thyloses are a constant feature in the structure of the resin

passages of Pinus (Fig. 30). They are always thin-walled and

completely fill the canal. So constant are these features in

association with those previously recounted that they serve to

afford a ready means of accurately recognizing the genus under

all circumstances.

The general course of development thus outlined shows that

the parenchyma tracheid stands in such relation to the organiza-

tion of the resin passage that its more frequent occurrence is

directly correlated with a higher type of organization and devel-

opment in the plants to which they belong.

We are now ina position to present a general summary of

the relations which the resin cells bear to the organization of

the secretory reservoirs — cysts and passages — and the posi-

tion which the latter occupy in the economy of the plant, as

follows :

1. Resin cells, which are of the nature of wood parenchyma,

at first occur as isolated structures filled with resin, but

they show a definite tendency to association, and later form

definite aggregates.

2. Parenchyma tracheids become associated with such aggre-

gates for the purpose of effecting a more complete nutri-

tion of the secretory cells.

554 THE AMERICAN NATURALIST. [Vor. XXXVIII.

3. Resin cells, when aggregated beyond a certain point, develop

schizogenous, intercellular spaces which form either central,

closed cysts, or central canals of indeterminate length. — -

4. The structure of the cyst or passage always presents the

same sequence of elements, and the work of the reservoir

is then divided between

(a) the tracheids which provide nutrition for the secretory

cells ;

(b) the secretory cells or epithelium in which the formation

of the resin takes place ;

reservoir for the surplus product ;

(d) thethyloses which may impede the proper storage of

the resin, or which may individually serve the purpose of

storage.

So long as the formation of resin is not excessive, it is stored

in the cells where produced. This is true of all isolated resin

cells, as well as of many which enter into the composition of

complex cysts and passages. When the resin is excessive, how-

ever, the surplus is excreted into specialized reservoirs of the

form of closed Cysts, or of canals, and we are led to interpret

the appearance of these structures in the higher Coniferz as a

response to such needs. The development of the resin passages

wil thus be seen to stand in direct relation to the capacity of

the plant as a resin producer — a fact which is otherwise ap-

parent from our knowledge of the general capacity of the differ-

ent genera as resin producers, and a comparison of this feature

with their known position in the line of descent.

(To be concluded.)

A LIST OF BERMUDIAN BIRDS SEEN DURING

JULY AND AUGUST, 1903.1

HAROLD BOWDITCH.

Tune following list comprises those birds which fell under my

personal observation on or near the Bermuda Islands between

the sixth of July and the twenty-second of August, 1903, the

time of my stay at the Bermuda Biological Station. The names

of such birds as are included in the Check-list of the American

Ornithologists’ Union are preceded by their respective check-list

numbers, and every effort has been made to bring the scientific

names down to date. I wish here to express my thanks to Prof.

Edward L. Mark, Director of the Biological Station; Mr. John

T. Nichols; Mr. Owen Bryant; Mr. Goodwin Gosling, of Ham-

ilton ; and the Messrs. Louis L. Mowbray and R. S. McCallan,

of St. George, for their valuable aid in furthering my observa-

tions.

70. Sterna hirundo (?) Linn. Common Tern.— Terns were

seen three times off the north shore, in each case a single indi-

vidual, but not once was I near enough to make a positive iden-

tification. They were in all probability S. Azrundo.

89. Puffinus gravis (?) (O'Reilly). Greater Shearwater.—A.

Shearwater-like Petrel (family Procellariidae) was seen in sight

of Bermuda, July sixth ; it was probably a Greater Shearwater,

although certain identification of this bird and of the two others

seen on the same day would not be possible without specimens.

109. Oceanites oceanicus (?) (Kuhl). Wilson's Petrel.— A

* Mother Cary’s Chicken " was seen from the S. S. “ Gladisfen "

on the “Challenger” bank, about twelve miles southwest of

Gibbs’ Hill, on the first of August. Identification was impos-

Sible, but the bird was probably a Wilson's Petrel. Merriam

(84, p. 284) reports finding a Leach’s Petrel, dead, on the

1Contributions from the Bermuda Biological Station for Research. No. 3.

555

556 THE AMERICAN NATURALIST. [Vor. XXXVIII.

shore at ***Tuckerstown Beach, May 1, 1881,” and says :

“The bird is common enough at sea a hundred or two miles

from the Bermudas, but I never saw one near the islands.”

112. Phaéthon americanus Grant. Yellow-billed Tropic Bird ;

“ Long-tail."— The Yellow-billed Tropic Birds were perhaps the

' most conspicuous birds in Bermuda until the middle of August,

when their numbers seemed to decrease. These birds lay

one egg with brown markings, in holes in the cliffs, almost

always with no attempt at a nest; as they are very close sitters,

many opportunities are offered for study at short range. Mr.

Owen Bryant found chicks in the * natal down” on Castle Island

on June thirtieth. Eggs were found in Harrington Sound as

late as July twenty-sixth, and a downy young bird was found on

Brangman's Fort on August twelfth.

I was very much interested in observing the change in the

plumage of the young * Long-tail," as seen in many specimens.

The bird emerges from the egg covered with puffy white down,

shaded with gray on the wings, and has a pale grayish bill, gray

feet, and gray, unfeathered lores. This « natal down ” is changed

fora plumage of white, with black spots on the head, wings and

back, and one black spot on the end of each tail-feather. The

bill is yellowish flesh-color and grayish ; the lores are feathered,

and the tail-feathers are all of the same length. During the

persistence of this plumage, which in being speckled resembles

that of the Red-billed Tropic Bird, the bill becomes yellowish

orange, the tint which persists throughout life. The fact that I

never saw a speckled bird on the wing, but always on the “ nest,” `

leads me to suppose that the speckled plumage is exchanged for

a plumage indistinguishable from that of the adult bird, before

the bird-of-the-year takes flight ; and it seems as if the change

could not be due to wear, as the bars and spots of black are

almost half an inch from the ends of the feathers.

194. Ardea herodias Linn. Great Blue Heron.— I saw one

of these birds in Tucker's Town Bay, July fifteenth, and another

was seen on an island near Hamilton Harbor on the eighth of

August. Capt. Savile G. Reid ('84, p. 242), quoting Major

edderburn, says: «In 1846 the nest of this bird, containing

two eggs, was found amongst the mangrove trees at Hungry

Bay.” This is said to be the only known nest-record.

Nos. 451-452] LIST OF BERMUDIAN BIRDS. 557

234. Tringa canutus (? Linn. Knot; Robin Snipe.— A

single bird, probably of this species, was seen on July twenty-

seventh on Cooper's Island, in company with three other waders,

a Turnstone, a Spotted Sandpiper, and a supposed Sanderling.

All were rather shy and hard to approach.

248. Calidris arenaria (2) (Linn.). Sanderling.— One instance,

not a positive identification; Cooper’s Island, July twenty-

seventh.

263. Actitis macularia (Linn.). Spotted Sandpiper.—I have

only two days’ positive records for the Spotted Sandpiper, namely

July twenty-seventh, when I saw two or three on Cooper’s

Island ; and July twenty-ninth, two at Hungry Bay. However,

Sandpipers were seen on one or two other days, and were prob-

ably the Spotted.

274. Ægialitis semipalmata Bonap. Semipalmated Plover :

| Ringneck.— One instance, a single bird on Cooper's Island,

August twelfth. ,

283.1. Arenaria morinella (Linn.). Ruddy Turnstone.— One

. record, Cooper's Island, July twenty-seventh.

289. Colinus virginianus (Linn.). Bob-white ; “ Quail."— Ac-

cording to Capt. Savile G. Reid (84, p. 227) these birds were

imported from the United States in 1858 or 1859. My records

for them extend from July seventh to August twentieth, seldom

more than one in a day. I heard the spring call as late as Au-

gust tenth, and was told that a brood of young was seen near the

Hotel Frascati, where the Biological Station was located, shortly

before my arrival in Bermuda.

320b. Columbigallina passerina bermudiana (Bangs and Brad-

lee). Bermuda Ground Dove.— This is one of the common

Bermuda birds, being found especially in cultivated areas. It is

very tame, allowing one to approach within two yards of it.

428. Trochilus colubris (?) (Linn.). Ruby-throated Humming-

bird.— On July twentieth, as I was driving along the Middle

Road, not far from the Whitney Institute, a Hummingbird flew

over my head from among some oleanders. As the Ruby-throat

is the only Hummingbird that has been taken in Bermuda (Reid,

'84, p. 210), the bird that I saw was probably of this species.

On July fifteenth another member of our party (Prof. Albert

Mann) saw a Hummingbird in the same neighborhood.

558 THE AMERICAN NATURALIST. [Vor. XXXVIII.

488. Corvus americanus Aud. American Crow.— Messrs.

Bangs and Bradlee (:01, p. 253) say: “.... It is, however,

doubtful if the Bermuda Crow is Corvus americanus. Its notes

are said to be different, more like those of the European Carrion

Crow (Corvus corone Linn.), a hoarse, raven-like croak.

During the season of 1900—1901, crows were seen but seldom

and never more than three together at any one time. They were

shy and their notes were not heard distinctly.”

I have several records, namely: July seventh, several heard

calling on Paynter’s Hill; July tenth, two heard calling in the

direction of Harrington Sound, from the Flatts ; August tenth,

a flock of seven seen at close range (but not heard) on the South

Shore, about opposite the Devil's Hole ; August eleventh, three

together, seen and heard on the southern shore of Harrington

Sound, west of Patton’s Point; and August twenty-first, one

heard from the Biological Laboratory on the side of a neighbor-

ing hil. In neither flight, appearance, nor voice did any of

these crows seem to me to differ from Corvus americanus, and

I heard their cawing distinctly, especially on July seventh and

August eleventh. I wastold that there were three Crows' nests

on Trunk Island, Harrington Sound, and was shown a nest on

Cooper's Island on July twenty-seyenth, said to have contained

young birds earlier in the season.

593. Cardinalis cardinalis (Linn.) Cardinal Grosbeak; Red-

bird.— The * Red-bird” is one of the most abundant of the resi-

dents, and I have an almost uninterrupted series of daily records

from July seventh to August twenty-second, my entire stay, the

single exception being on August first, which was spent on the

" Challenger Bank," twelve miles from shore. At Hungry Bay I

Saw a Cardinal's nest containing three or more young birds, on

July twenty-ninth, a late date, for Reid says (84, p. 201), “Asa

rule, the two clutches of eggs are deposited about April roth,

and May ‘30th, respectively." Moseley (79, p. 25) alludes to

the fact that it is an introduced species, and Ridgway says (: 01,

P. 674), “the Species is said to have been introduced from the

eastern United States”; the date I am unable to determine.

Passer domesticus (Linn.). House Sparrow; “ English” Spar-

Tow.— Reid wrote in 1877 (84, p. 196)! of two importations

‘Reprinted from 1877.

Nos. 451-452] LIST OF BERMUDIAN BIRDS. 559

from New York, one “a few years ago," the second in 1874,

and says, “I have no doubt they will increase and multiply after

their manner, and in time become as much a nuisance as they

are now a curiosity." This prophecy has been fulfilled, and I

am told that the * English ” Sparrows breed in Bermuda at all

seasons. The Sparrows are interesting in one respect, namely,

the fact that the adult birds, and especially the females, are

noticeably lighter and more yellowish than the United States

birds. That this is due to bleaching is proven by a specimen

taken by my friend; Mr. Nichols. This bird is an adult female,

in which the light colored, worn-out tail feathers are being

replaced by a new set of typical dark ones.

‘Carduelis carduelis (Linn. Goldfinch. —Reid (84, pp. 196—

197) says, “I observed a single specimen of the European Gold-

finch, Carduelis elegans, near Harrington Sound, in April, 1875;

it was very wild, and I could not get near it, but I imagine it

must have been an escaped prisoner." This bird is now com-

mon in Bermuda, and I did not find it at all hard to approach, in

spite of the fact that Bangs and Bradlee (: 01, p. 256) state that

it “is exceedingly shy and wary." By far the largest flock that

I saw was at the Flatts, on August tenth, and contained about

thirty birds. On June twenty-ninth Mr. Owen Bryant found a

newly built nest of the Goldfinch on Trunk Island, Harrington

Sound ; it was in a cedar, some twenty-five feet from the ground

and six or seven feet from the trunk of the tree. On July sixth

it contained four eggs.

611. Progne subis ()) (Linn.). Purple Martin.— On August

fourteenth, while between Coney Island and the Ferry Reach,

I saw two Martins flying past the Martello Tower, southward.

I was unable to determine the species, whether Purple or Cuban,

but as the range of the Cuban Martin is, according to Chapman

('95, p. 320), “Southern Florida south to Cuba and probably

Central America," the birds were in alllikelihood Purple Martins.

Reid ('84, p. 190) states that this bird “has only .... appeared

on one occasion, during the ‘entrada’ of September, 1849,

when it was numerous.” I looked this matter up in the ** Nat-

ural History of the Bermudas,” a collection by Miss H. T.

Hurdis of scattered notes by her father, John L. Hurdis. His

560 THE AMERICAN NATURALIST. [Vor. XXXVIII.

only records of specimens fall in September, 1849, but he

records martins seen by other people three times, viz.: August,

1847 (p. 26); September 19th, 1850 (p. 193) ; and April 24th,

I851 (p. 212).

631b. Vireo noveboracensis bermudianus (Bangs and Bradlee).

Bermuda White-eyed Vireo ; “ Chick-of-the-village.’’— This bird

ranks with the Cardinal and the Catbird in point of numbers;

everywhere that one goes, he hears its characteristic song, trans-

lated into *chick-ofthe-village" by the negroes. Bangs and

. Bradlee (: or, p. 253) say, “The iris in V. bermudianus is white

as in V. noveboracensis, and not “brownish, brownish gray, or

gray," as stated by Capt. Reid. This seems to me to be a very

hasty conclusion, as I have never seen a Bermuda Vireo with a

pure white eye. The birds are very tame and familiar, and I

made it something of a point to look at the iris whenever a good

opportunity offered, with the result that I saw just two speci-

mens with “dirty white” irides, and these had every appearance

of being young birds. The irides of all the other birds of which

I took note were dark.

Reid ('84, p. 192) says, “It is on record that the newly fledged

young of this species have been found entangled in the meshes

of the web of the ‘silk’ spider, Æpėira [Epeira] clavipes. These

webs are of great size and strength, extending for many feet

between adjoining cedars, and the number of them among the

woods in summer and autumn is almost incredible. In all my

rambles, however, I never met with an instance of poor little

Vireo having walked into Epéira’s parlor."

On the twenty-ninth of July I saw, at the edge of the man-

grove swamp at Hungry Bay, a Vireo with a large mass of cob-

web on the side of its head, completely covering the right eye.

The left eye only was used, which necessitated the bird's turn-

ing On its perch in order to watch us, instead of simply turning.

its head.

; 704. Galeoscoptes carolinensis (Linn). Catbird; * Black-

bird."— Extremely common throughout the islands. During

my stay in Bermuda I did not once hear the “mew” which

gives the Catbird its name, and the song and the other notes

of the Bermuda Catbirds sounded to me not quite identical with

those of United States birds.

Nos. 451-452] L/ST OF BERMUDIAN BIRDS. 561

766. Szalia sialis (Linn.). Eastern Bluebird.— A very com-

mon bird. Bangs and Bradlee (:ot, p. 255) state that the Blue-

bird is migratory as well as resident, so that ** possibly some of ”

the migrants *remain and breed and thus counteract any tend-

ency to vary that the island birds might develop if wholly cut off

from the main body of the species." Reid (84, p. 175) says of

the nests, *Ihave found them commonly in holes in old quar-

ries or roadside cuttings; alsoin crevices of walls; in rocks,

even when some little distance from the shore ; in holes in trees ;

on the branches of trees ; in stove and water pipes ; in calabashes,

boxes, etc., hung up for them in the verandas of houses, . .

and in several other curious situations."

On July tenth Mr. Owen Bryant found a Bluebird's nest

containing three young birds, built in the capstan of an old

wreck near Coney Island. On the twenty-sixth of July Mr.

John T. Nichols and I found some holes in a bank of sand,

part of one of the cliffs nearly due north from the Devil's Hole,

Harrington Sound; they were some four or more inches in

diameter and a foot and a half to two feet deep, and two of

them contained nests of soggy vegetable materials, one of which

had in it a couple of bad eggs, probably Bluebirds’. This

seemed to us a strange nesting site for the bird which we always

associate with the hollow limb of an old apple tree.

Besides the twenty-one birds listed above, the following were

reported to me:

On July twenty-third Mr. Leon J. Cole saw a Tattler ( Toza-

nus), probably melanoleucus, at St. David's.

During July a large Hawk was seen by various members of

the Bermuda Biological Station. His identification as a Buteo

was all but complete.

On August sixteenth Mr. Louis L. Mowbray told me of a

Snowy Owl (JVyctea nyctea) which had been seen on the after-

noon of the previous day, in the Devonshire marsh. Swallows

(probably Barn Swallows, Hirundo erythrogastra) were reported

common.

562 THE AMERICAN NATURALIST. [Vou. XXXVII.

` INDEX OF SCIENTIFIC NAMES.

PAGE.

Actitis macularia ; : ; ) f : à BS

JEgialitis semipalmata — . : ; ‘ : : ; : l0 E

Ardea herodias i i : : : ; à : : Ee

Arenaria morinella : : ; : ; i : : eee

Buteo . . i 1 à : : ; ; ae

Calidris arenaria : i ; x à s Tu E

Cardinalis cardinalis : : i : f . : ; ieee

Carduelis carduelis : ; : : 1 ; ; ; - A

Colinus virginianus : : E ; j A y

Columbigallina passerina doii i de

Corvus americanus : : : i : à ; j o NE

Galeoscoptes carolinensis i ; ; 1 : 2 e

Hirundo erythrogastra 561

Nyctea nyctea : : : : 561

Oceanites oceanicus : ; 1 : : : : : eo

Passer domesticus 2 à : : ; : i i E os

Phaéthon americanus . : : à : i ; 2 . 4 e

Progne subis 559

Puffinus gravis 555

Síalia sialis conte eM E NM M E 561

Sterna hirundo : : í : : - y . : - Sse

Totanus melanoleucus : : : ; ; A es

Tringa canutus : : E : i : : R : Mir:

Trochilus colubris : i i t ; $ n Lr

Vireo noveboracensis Sed ; : : : : oo

BIBLIOGRAPHY.

BANGs, O., and BRADLEE, T. S.

:01. Resident Land Birds of Bermuda. The Auk, Vol. 18, July.

pp. tn di

CHAPMAN, F. M.

'95. Handbook of Birds of Eastern North America; D. Appleton & ies

New York, xiv, 421 pp. 20 pl.; 115 cuts.

MERRIAM, C. H.

'84. On a Bird New to Bermuda, with Notes upon Several ue of

Rare or Accidental Occurrence. Bull. U. S. Nat. Mus. No. 25-

Part V, pp. 281-284.

Nos. 451-452] Z/S5T OF BERMUDIAN BIRDS. 563

MOoSELEY, H. N.

79. Notes by a Naturalist on the “ Challenger,” etc. Lond., Macmillan

and Co., xvi + 620 pp., 1 map, 2 pl., and cuts.

REID, 5. G;

'84. The Birds of Bermuda. Bull. U. S. Nat. Mus. No. 25, Part IV.

pp. 163-279. [Consists of notes published in 1875 and 1877,

revised.] :

RipGwAY, R. :

‘01. The Birds of North and Middle America. Part I, Family Frin-

gillida. The Finches. Bull U. S. Nat. Mus. No. 5o. xxxi, 715

pp. pl. 1-20.

VERRILL, A. H.1

01. Additions to the Avifauna of the Bermudas with Diagnoses of

two new Subspecies. Amer. Jour. Sci., Ser. 4, Vol. 12, No. 67,

July. pp. 64, 65.

1 Note.— This paper by A. H. Verrill, seems to have been overlooked by the

author.— E. L. Mark.

Neritina virginea, variety minor.

x 2 DIAMETERS.

NERITINA VIRGINEA VARIETY MINOR}

MAYNARD M. METCALF.

WHILE collecting one summer in Jamaica, W. I., the author

visited the “Salt Pond” near Port Henderson. This shallow

pond is separated from the ocean by a narrow strip of sand over

which the waves break during heavy storms, It has no outlet

to the ocean and no fresh water streams flow into it. The water

thus comes to be: considerably more salt than that of the open

ocean. The sample of water from these ponds which the author

examined had a specific gravity of 1.038.

The fauna of this pond is quite rich and includes some very

interesting forms. One is especially attracted to the beautiful

little Neritina which literally covers the beach for rods near the

water’s edge. From a double handful of these animals scooped

up in one spot on a little sand bar near the shore, the author

has selected the shells in the accompanying plate. They are

arranged for use in another publication and are designed to show

a remarkable example of diversity. I would like here to call

attention not only to the diversity but also to the dwarfing of

these shells and its apparent cause.

Neritina virginea is a widely distributed littoral marine spe-

cies. Full grown adults from the ocean are of the size of the

figures on the accompanying plate. They show the same types

of color and color pattern. Other types also are found among

both the open ocean shells and those from the Salt Pond, but I

have not attempted to figure them.

It is interesting to find that the Neritina of the “ Salt Pond”

is a dwarf variety. In almost every regard except size the ani-

mals resemble their ocean relatives, but their size is much less.

The diameter of an average sized shell from the Salt Pond

1 Ry the courtesy of The Macmillan Company the author is permitted to use

for this article the accompanying plate which he has prepared for use in An Out-

line of the Theory of Organic Evolution, now in press. :

565

566 THE AMERICAN NATURALIST. [Vor. XXXVIII.

would be about half that of an average shell from the open

ocean. The figures on the plate are enlarged two diameters.

Apparently the extra salinity of the water in the Salt Pond

has dwarfed the animals. That they are not young individuals

is indicated by the facts (1) that no individuals of larger size

were found, (2) that no smaller ones were found, indicating that

they were not breeding at the time these shells were gathered.

It can hardly be supposed that both the very young and the

adults hide themselves from view, while the half-grown individu-

als crowd upon the shore in numbers that literally cover the .

sand at the water's edge. Yet the author should mention that

he has seen a letter to Professor E. A. Andrews in which refer-

ence was made to these shells, called by the natives Bossu,

saying that they are used for food and are gathered in fresh

water streams at night with lanterns, but that during the day

they hide under stones and can be found only with difficulty.

There seems little doubt that the form from Salt Pond Neritina |

is a dwarf variety, and if So, the most probable cause of the

dwarfing seems to be the density of the water in which they live.

It is still more interesting to find that a similar dwarfed Neri-

tina occurs in fresh water streams in Jamaica. Professor An-

drews has collected them in a small stream at Port Antonio, and

the same letter referred to above mentions their occurrence in

Mabess River, a branch of Spanish River near Buff Bay.

Doubtless they occur in other Jamaica streams.

The author has about five hundred shells collected for him by

Dr. F. S. Conant in “a small stream at Port Antonio,” probably

the same stream in which Professor Andrews saw them. These

Shells are all small, — no larger than those from the Salt Pond.

Among them are some that are very small and apparently im-

mature. Careful search should be made for full sized individu-

als in these streams before we can be positive that none are

Present. This the author has been unable to secure. Such

data as we have, however, especially in comparison with the

Salt Pond conditions, seems to indicate the probability of these

fresh water Neritina shells being a dwarf variety.

If this conclusion is correct we have the very interesting phe-

nomenon of an animal normally living in the ocean able to adapt

Nos. 451-452. NERITINA VIRGINEA. 6

5607

itself to life in fresh water or to water of greater salinity than

that of the ocean, but unable to attain its full size except in the

ocean.

The dwarfed forms may fairly be called a variety, though their

difference from the ocean forms be only one of size probably

directly due to increased or decreased salinity of the water in

which they live.

The great diversity in color and in color pattern found in

Neritina virginea is well known, yet it may be worth while to

call attention to the completeness of the intergrading between

any two types, however divergent.

Color shade. — Note first the differences in the ground color

of the shells: yellow in 54, 19, and 22; white in all the rest.

Observe that the colored lines are black in 1, 3, 5, 54, 6, 6a, 19,

20, 22; purplish in 64 and 11; red in 7a; gray in 23; black

and red in 2; that the major lines are black and the minor lines

red in Ia, 3, 32, 4, 4a, and 5a; that the major lines are black

and the minor lines purple in 42, 17 and 16.

Color patteru. — 1 is marked with a few heavy lines. From

I to 6 these major lines become broken up into small V-shaped

loops. In the shells 2 accessory minor lines are added. In the

shells 6 these are more numerous.

Series 9 to 14 shows diversity in the pattern near the apex

of the coil: 9 has a few slightly larger white dots near the coil,

10 has larger dots here, 11 has them very large; in 12 they

have united to form a continuous white band; in 13 and 14 this

band is wider.

Series 15 to 24 shows diversity in the character of the equato-

rial light band. In r5 and 16 only the minor lines are inter-

rupted or faint along the equator of the shell; in 17 the major

lines also are interrupted; in 18 the band is almost clear white ;

19 and 20 show narrower bands; in 21, 23 and 24 the equato-

rial band is shown by a difference of color in and under the pat-

tern; in 22 the equatorial line is faintly indicated in the pattern

itself, being bordered above and below by large, heavy, black

loops.

The shells figured are but a few selected from the large num-

ber collected. Other types of color and of color pattern were

568 THE AMERICAN NATURALIST. [Vor. XXXVIII.

found. Between any two of the shells collected, no matter how

divergent in type, a completely intergrading series could be

found. 1

It is doubtful if these colors and color patterns are of protec-

tive value. The shell is enclosed by flaps of the mantle most of

the time rendering the coloration invisible. Also all types of

color pattern are seen in one locality. Then, too, the animals

are in such great numbers that they could hardly escape the

observation of their predatory enemies, if such they have, by

means of their coloration.

The shells shown as numbers 52, 84, 19 and 22 are from Port

Antonio, probably from the stream in which Professor Andrews

found Neritina, a small shaded stream with a dark bottom of

vegetable mould. One can readily observe that the shells from

the Port Antonio stream are darker than those on the unshaded,

light-colored sand beach of the Salt Pond. Few of them resem-

ble numbers 1 to 5 or the a shells of the accompanying plate.

Most are heavily marked with dark lines, many of the type

shown in 84 being found. As a whole, the shells from the

stream average much darker than those from the Salt Pond.

Also it is noticeable that the ground color of almost all of these

fresh-water shells is yellow, while that of the Salt Pond shells is

white.

Remembering that it is doubtful if the coloration of these

shells is to any considerable extent protective, it is interesting

to find that the darker-colored shells are found in the midst of

dark-colored surroundings, in a shaded stream, while the lighter-

colored shells are from an unshaded beach of light-colored sand.

Comparison with the collections of Neritina virginea in the

United States National Museum and in the Museum of the

Philadelphia Academy of Sciences shows that the color of the

Jamaica Salt Pond shells averages considerably lighter than that

of shells from the ocean, the ground color being a clearer white,

and shells of the patterns represented in figures 1 to 6 and 14

to 6a being present in larger proportion.

Few, if any, finer examples of diversity in color and in. color

pattern are known. The same types of coloration are found in

the full grown shells from the ocean. In the absence of breed-

Nos. 451-452.] NERITINA VIRGINEA. 569

ing experiments, if it were not for the completely intergrading

forms, systematists would probably have no hesitation in recog-

nizing several species. As it is, all must without doubt be in-

cluded in one species, and it seems hardly worth while to name

as varieties (punctata, lineata, bilineata, trilineata, etc.) even the

most divergent types. The dwarf forms, however, might fairly

be called variety minor.

The writer would suggest that, if the difficulty of rearing

these animals be not too great, it is probable that they would be

favorable subjects for breeding experiments.

THE WoMaN's COLLEGE OF BALTIMORE,

April 25th, 1904.

STUDIES ON THE PLANT CELL— III.

BRADLEY MOORE DAVIS.

SEcTIoN III. HIGHLY SPECIALIZED PLANT CELLS AND THEIR

PECULIARITIES.

VERY much of our knowledge of the structure and behavior

of protoplasm in plants has been derived from the study of cer-

tain cells whose organization has reached an exceptionally ad-

vanced degree of differentiation. The peculiarities of these

cells are obvious and have proved of great interest but we have

as yet scarcely made a beginning in the study which must trace

and relate these characteristics of the most complex products of

cellular evolution in plants to their more simple progenitors.

This section will describe in some detail the structure and

protoplasmic activities of the following six highly specialized

cells: 1, The Zoóspore; 2, The Sperm; 3, The Egg; 4, The

Spore Mother-Cell; 5, The Coenocyte ; 6, The Coenogamete.

I. The Zoóspore.

Zoóspores are interesting not only for their own peculiarities

but also because they are well known to be the progenitors of

the sexual cells or gametes which become later differentiated

into the egg and sperm. Comparative studies upon three cells

so closely related and yet so diverse in their extremes of struc-

ture are sure to yield important results.

The zoóspore is generally an uninucleate cell, colorless in the

Fungi, but containing a chromatophore or plastids in all other

groups of thallophytes. There are usually two or four cilia

| attached to the anterior pointed end which is free from coloring

matter and at this region one may expect to find a red pigment

spot. Some zoóspores are exceptional for special peculiarities,

as those of Vaucheria which are multinucleate, each nucleus

571

572 THE AMERICAN NATURALIST. [Vor. XXXVIII.

being accompanied by a pair of cilia, or those of CEdogonium

whose colorless forward end bears a crown of numerous cilia.

The zoóspore stands among the higher forms for a type of motile

organism that is very close to the bottom of the assemblage of

groups and developmental lines which make up the Alga. The

forms most closely related to the zodspore are in the family

Chlamydomonadez of the Volvocales. But at this general

low level of the plant kingdom there are several groups whose

members pass most of their lives in motile conditions (Volvo-

cales, Flagellates and Peridinales) and the cells of all of these

types resemble zodspores to a greater or less degree in their

structure and habits, so that this condition represents a wide-

spread and well defined stage of evolutionary development.

Therefore when zoóspores are formed in the life history of some

higher plant they represent a return on the part of the organism

for a short time to the structure and mode of life of an ancestry

perhaps related in some way to the groups that still have the

motile habits throughout most of their existence.

For these reasons close comparisons in structure between the

zoospore and motile Algze will be interesting and should help to

explain the peculiarities of these cells. These peculiarities

chiefly concern the organ that forms the cilia (blepharoplast),

which becomes very complex in the sperm, and the pigment

spot.

Unfortunately studies upon these problems have been few and

we are not prepared to make a general statement of the condi-

tions. The most recent investigation on the structure of the

zoöspore is that of Timberlake (: 02), but Strasburger has written

extensively on the subject, especially in the ZZiszo/ogzsche Bet-

trage (92 and :00). The later paper (:00, p. 177-215) reviews

the entire subject of cilia formation. Dangeard has presented

an account of the Chlamydomonadeze, '99, and in :o1 described

especially Polytoma, comparing its structure with that of the

animal spermatozoan. :

Polytoma (see Fig. 94)is a colorless organism but its cell

structure and life history place it unquestionably among the

Chlamydomonadez. The two cilia arise from a small body

(blepharoplast) situated at the extremity of the cell. A delicate

Nos. 451-452.] STUDIES ON THE PLANT CELL. 573

thread-like structure, which Dangeard calls the rhizoplast, extends

from the blepharoplast into the cytoplasm and sometimes ends

at the side of the nucleus in a granule (condyle). The cilia

grow out from the blepharoplast. This apparatus is not known

to bear any relation to centrosomes or to the kinoplasm of

nuclear figures present at the time of spore formation. But it

should be noted that the blepharoplast is situated directly under

if not actually in the outer plasma membrane, which is kino-

plasmic. The filamentous connection between blepharoplast

and nucleus is probably important, especially since it has also

been found in zoó$pores (Timberlake, : 02, for Hydrodictyon) but

we do not even know its developmental history much less its

function. Further study will be necessary to make clear possi-

ble relations to kinoplasm around the nucleus or to centrosomes.

Consequently Dangeard’s comparison of Polytoma to the animal

spermatozoon is not convincing for it seems to be established for

the spermatozoén that portions of the middle piece at least and

the flagellum are derived from a true centrosome. Indeed

from the meager evidence now at hand the blepharoplast of

Polytoma is as likely to be a structure differentiated from the

plasma membrane as to have any relation to the nucleus. But

detailed studies on sporogenesis may discover a history more in

harmony with that of Hydrodictyon.

We have summarized a portion of Timberlake's (: 02) account

of sporogenesis for Hydrodictyon

in the previous section under the

head of “Cleavage by constric-

tion.’ We shall consider now

certain details. Small spherical

bodies are found at the poles of

the spindles during nuclear divi-

sion in the mother-cell. They are

undoubtedly accumulations of kino-

plasm and perhaps stand for centro- |. a1. zosspore. a, Polytoma; 5,

somes. However they have no caftra si ind imesh in Oedo-

gonium. (a, after Dan mis anh o1 ; 6, Tim-

polar radiations nor could they be 14, : 02; c, Strasburger g

followed between mitoses when the

nuclei were in resting conditions. It is not probable therefore

574 THE AMERICAN NATURALIST. [Vor. XXXVIII.

that these structures are permanent in the cell. After nuclear

multiplication is ended segmentation proceeds until the nucleate

masses of protoplasm separate from one another as zoóspores.

Then a body may be found lying in contact with the plasma

membrane and bearing a pair of cilia (Fig. 92). This basal body

(blepharoplast) by its reaction to stains seems to be entirely

distinct from the plasma membrane and is connected with the

nucleus by very delicate threads. There is a time just previous

to the differentiation of the zoóspores when the nuclei lie very

close to the cleavage furrow that finally separates the adjacent

zoospore origins. A granule may sometimes ‘be observed close

to these nuclei and it is possible that this is the first appear-

ance of the basal body (blepharoplast). If this should prove

correct the structure may have a direct relation to the kinoplasm

around the nucleus, a rélation that is afterwards maintained

through the two or three delicate fibers that connect these

Structures. Thus the blepharoplast if not directly derived from

a centrosome may at least have its origin from the same region

of kinoplasm. However these possibilities are mere speculations

and the investigation of these points is very much to be desired

in a number of algal and fungal types.

We are now brought to the views of Strasburger as expressed

in his writings of '92 and :00. His investigations have been

chiefly on Vaucheria, Cladophora and CEdogonium. In all of

these forms the cilia come from a body (blepharoplast) which

he believes to arise from the outer plasma membrane (Haut-

Schicht). The nucleus lies close to the plasma membrane at the

time when the blepharoplast is formed and may determine its

development there as a dynamic center, but the blepharoplast is

not à centrosome according to Strasburger. It is of course

kinoplasmic since it develops from the plasma membrane and

this would accord with its activities as a cilia forming organ.

The blepharoplast is extraordinarily large in CEdogonium (see

Fig. 9c) and develops a ting of numerous cilia on the exterior

while at the same time fibrillar rays grow back into the cyto-

plasm and probably help to give a compact organization to the

zoospore. This structure is very suggestive of the centrosphere

and aster that cuts out the ascospore (see Section II, Free Cell

Nos. 451-452.] STUDIES ON THE PLANT CELL. 575

Formation) and in spite of Strasburger's conclusions that it is

derived entirely from the plasma membrane we are justified in

asking for a fuller description of its development. There is the

possibility of a different origin wherein the nucleus may play an

important part which, in the light of Timberlake's studies on

Hydrodictyon, suggests that Strasburger may not have discov-

ered the earliest beginning of the blepharoplast in CEdogonium.

And the same doubts apply to Cladophora and Vaucheria.

There is thus considerable divergence in the views of the

origin and nature of the blepharoplast in zoóspores, Strasburger

believing that they are developed as a specialized region of the

plasma membrane with no relation to centrosomes, and Timber-

lake holding that the structure in Hydrodictyon is not a part of

the plasma membrane but comes from the interior of the proto-

plasm. The problem is also involved with conditions in the

sperm, where there is likewise a difference of opinion as to the

homologies of the blepharoplast but an undoubted origin at

least in the pteridophytes and gymnosperms from the interior

of the cell. We should naturally expect the blepharoplasts of

zoóspores and sperms to be homologous and consequently the

problem is of great theoretical interest and will be taken up

again in our discussion of the sperm. Its solution demands a

most thorough study of the development of some of the larger

zoóspores as in CEdogonium and certain species of the vise

vales and Volvocales.

The pigment spot is almost universally present in zoóspores

and is also characteristic of the cells of many motile organisms

as in the Volvocales and Flagellates while occasionally found in

other groups. The structure has been called an eye spot from

its fancied resemblance to the simple eyes of certain Crustacea

` (Cyclops, etc.) but this term is unsatisfactory since it is not

established that the pigment spot is primarily a receptive organ

for light or warmth; but even should it prove to be thus sensi-

tive (which is very probable) thereby orienting the cell with

respect to the direction of incoming rays, that is not a function

comparable to sight.

The coloring matter of the igne spot is held as a single

globule or as a collection of numerous small granules in meshes

576 THE AMERICAN NATURALIST. (Vou. XXXVIII.

of the protoplasm. It is frequently associated with a plastid.

The pigment may be readily broken down and dissolved out by

such reagents as alcohol and ether. In chemical composition it

is very close to haematochrome and thus may be related to

- chlorophyll or a derivative of that substance. The cytoplasm

around the pigment spot is undifferentiated and when the color-

ing matter is removed it is very difficult and sometimes impos-

sible to find the situation of the structure. Consequently the

pigment spot can hardly be considered a protoplasmic organ

since it is merely an accumulation of coloring matter at some

point in the cell. Strasburger (:00, p. 193) states that the

pigment spot of certain zoóspores (Cladophora, etc.) is formed

in the plasma membrane but this is not true of many other

motile cells (Flagellata) and there is no doubt that in some cells

(e.g. the gametes of Cutleria) the pigment spot is a portion of

a plastid. The literature upon the structure and function of

pigment spots is reviewed by Zimmermann (Beiträge z. bot.

Centralb. Bd. 4, p. 159, 1894) and since then Wager (99) has

presented a detailed study of Euglena.

2. The Sperm.

The sperm is unquestionably derived from the zoóspore

through primitive types of gametes which were identical with

zoospores in all essentials of morphology. I have described the

origin and evolution of sexual cells of plants in two recent

papers (Popular Science Monthly, Nov. 1901, p. 66 and Feb.

1902, p. 300). We should expect the simplest forms of sperms

to have the characters of zoóspores and this is the fact. The

sperms of the Algæ, as a rule, have the same number of cilia

(usually two) as their ancestral asexual zoóspores. They gener-

ally contain a chromatophore, although sometimes much reduced,

and there is present the pigment spot. The cilia are attached

at the pointed end or at the side, arising from colorless pro-

toplasm that sometimes contains the pigment spot while the

chromatophore, when present, and the nucleus lie at some

distance from this region of the cell. The sperms of bryophytes

and pteridophytes are much attenuated in form and lack the

Nos. 451-452.] STUDIES ON THE PLANT CELL. 577

pigment spot and chromatophore. Those of the bryophytes and

the Lycopodinez are biciliate while other pteridophytes have

multiceliate sperms the cilia being distributed on a band

. (blepharoplast) which lies along one side of the spiral structure.

A large portion of the spiral in these sperms is composed of

nuclear substance and much of the remaining cytoplasm with

granules and vacuolar inclusions may frequently be found in a

vesicle attached to the larger end of the spiral.

The only motile sperm among the Fungi is that of Mono-

blepharis. The male cells of other Fungi are non-motile

bodies (spermatia) generally formed from the ends of delicate

filaments which are found in special organs called spermagonia.

Spermagonia have been described in the Uredinales, the lichens

and in the Laboulbeniacez but their function is only clearly

established for the last two groups. They are very highly

differentiated in the Laboulbeniaceze and comprise several types

of structure. Another type of male cell, found in certain

groups of the Phycomycetes and Ascomycetes, is the coenogamete

(to be described presently) which is however not the homologue

of the sperm but of the mother-cell or antheridium that develops

such structures. Sperms of the red Algze (Rhodophycez) are

likewise non-motile and they are invariably formed singly in

small cells at the ends of filaments. These non motile sperms

of Fungi and red Algz are exceedingly small uninucleate bodies

without further complexity of structure as far as is known.

We shall not attempt to discuss the earlier literature that

treats of the structure and development of the plant sperm. In

1894 Belajeff published a German translation of a paper

written two years before in Russian which presents the views of

previous investigators and to this the reader is referred for such

historical references. At that time various opinions were held

respecting the organization of the sperm, some writers (Campbell,

Guignard and others) believing that it was chiefly or wholly

nuclear in origin, while another group (Zacharias, '87, Belajeff,

Strasburger, '92, etc.) thought that the cytoplasm shared very

largely in its structure. Belajeff (94a) from studies among the

Characez showed with especial clearness that the cytoplasm

was an important constituent of this sperm since the nuclear

578 THE AMERICAN NATURALIST. (Vor. XXXVIII.

material occupied a restricted region in the middle of the spiral

structure. This was the first of a series of investigations which

have given especial attention to cytoplasmic activities during

spermatogenesis and placed the entire subject in a new light.

The year 1897 brought forth almost simultaneously three short

papers by Webber (97a, '97b, '97c) and Belajeff ('97a, '97b,

’97c) respectively. Webber had studied the development of the

motile sperms of Zamia and Ginko, Belajeff certain forms of the

Filicineze and Equisetinez. These were of the nature of pre-

liminary announcements and both authors published later more

detailed descriptions and discussions. The discoveries of motile

sperms in Ginko by Hirase and of Cycas by Ikeno were

announced in several short papers during the years 1896 and

'97 but without descriptions of their development. This litera-

ture together with later papers of Ikeno, Shaw, Belajeff, Hirase,

and Fujii is reviewed in Webber's last contribution (:01) and

also in Strasburger's discussion of * Cilienbildner ” (: 00, p. 177)

to which the reader is referred for the most complete treatments

of spermatogenesis in plants yet published.

The cycads and Ginko are the most favorable subjects known

for studies in spermatogenesis. Detailed accounts of the cycads

are given by Ikeno ('98b) for Cycas and by Webber (: 01) for

Zamia, these forms agreeing with one another in all essentials.

Two sperms are developed from the daughter cells (spermatids)

following the division of the so-called body cell in the pollen

tube. The process really begins in the body cell with the

appearance of the blepharoplasts. Their development has been

followed with especial attention in Zamia. They are formed

de novo in the cytoplasm at some distance from the nucleus and

while the latter is in the resting condition. They appear inde-

pendently of one another, generally on opposite sides of the

nucleus but sometimes much nearer together (Fig. 102). Each

Bis large deeply staining body with numerous radiations

extending into the cytoplasm. The blepharoplasts then increase

in size and, moving farther away from the nucleus, take positions

ay Opposite to one another. The nucleus of the body cell

now divides, its spindle being clearly intranuclear (Fig. 5 7) and

consequently holding no visible relation to the blepharoplasts

Nos.451452] STUDIES ON THE PLANT CELL. | ^" 579

which lie at a considerable distance from the structure (Fig. 10 4).

The latter cannot then be said to occupy the position of centro-

somes in relation to this spindle. Meanwhile important changes,

which are best known for Zamia, take place in the blepharoplast.

In this type the structure forms a hollow sphere which breaks

up into segments and finally into granules as mitosis proceeds:

The radiations disappear without holding any apparent relation

to the spindle. During telophase each of the two blepharoplasts

Dum RSS Nd AW

s TH, f

‘Fic. 10.— Spermatogenesis in Cyc

ae celis i Ż, prothallial cell; a p

ini. t in ; eee wi ith ; a proc viii bs nucleus; d, end of iain roplast attached to the

nucleus at a later stage of a elo , sperm showing section of the flattened spiral

blepharoplast with cilia projecting d the cell. (After ed. 798.)

appears as a mass of granules at some distance from the daugh-

ter nuclei which are to become the sperm nuclei. As a result of

this division the spermatids (sperm mother-cells) are differen-

tiated. At the close of the mitosis the blepharoplast enters

upon its functions of forming in the spermatid a cilia bearing

band which is to lie as a spiral around the sperm. The granules

first extend as a delicate deeply stained line towards the nucleus.

and then in the opposite direction. The nucleus in Cycas puts

forth a papilla (Fig. 10¢) which meets this line of granules and

remains attached to it for some time. The line thickens into a

580 © THE AMERICAN NATURALIST. [Vor. XXXVIII.

band which lengthens and finally takes the form of a spiral of

five or six turns which becomes more or less closely applied to

the plasma membrane (Fig. 10 e, blepharoplast in section). The

cilia develop as protuberances from the outer surface of the

band (Fig. 10c and 7) and grow through the plasma membrane

to the exterior of the cell. The nucleus in the meantime has

increased in size until it occupies the greater part of the top

shaped sperm (Fig. 10 e).

The history of spermatogenesis in Ginko is strikingly parallel

to that of the cycads. The chief features were first described

by Webber ('97c) and in greater detail by Hirase('98). The two

blepharoplasts appear de zovo on opposite sides of the nucleus

in the body cell. They show the same high state of differentia-

tion as those of the cycads, being large and the center of a

number of prominent radiations. Ginko however presents a

peculiarity not reported in the previous group. A large spheri-

cal body lies between each blepharoplast and the nucleus in an

area of granular cytoplasm. This structure stains deeply like

the globules of nucleolar substance which are frequently found

in the cytoplasm after nuclear division. They are probably

accumulations of a somewhat similar material at these points in

the cell to be utilized at later periods of spermatogenesis, since

they decrease in size as the sperms mature. The spindle in the

body cell is formed between the blepharoplasts but its poles lie

at some distance from and are entirely independent of these

structures. During this mitosis the spherical bodies pass to

one side of the spindle so that the daughter nuclei (sperm

nuclei) finally take the position formerly occupied by them.

The blepharoplast becomes granular and begins to lengthen into

a band, one end of which becomes attached to the nucleus that

puts forth a small papilla towards the blepharoplast. The band

elongates and takes the form of a spiral which makes several

turns around one end of the cell just under the plasma mem-

brane. Cilia then develop along this band as in the cycads.

The earlier accounts, describing a short tail on the sperm were

founded upon material that was not altogether normal and have

been corrected by Webber and Fujii. The mature sperms have

essentially the same form as those of Zamia and Cycas.

Nos. 451-452] STUDIES ON THE PLANT CELL. 581

There has been some discussion on the morphology of these

motile sperms of the gymnosperms. The claim has been made

that they are ciliated spermatids (sperm mother-cells) and there-

fore different from the sperms of pteridophytes which are

formed inside of mother-cells that upon their escape are left

behind as empty cysts. However a close analysis of their struc-

ture will show that the sperms in both groups have an identical

protoplasmic organization. There is a nucleus and a greater or

less amount of cytoplasm in which the blepharoplast lies and the

entire structure is surrounded by a plasma membrane. Any

differences in the processes of spermatogenesis can only concern

the greater or less development of a cellulose membrane around

the spermatids. It may be true that this cellulose membrane

is entirely absent in Cycas and Zamia, but if present it would

be merely a shell like envelope around the sperm and cannot

affect its morphological unity and agreement with the sperms

* of pteridophytes. A comparative study of the composition and.

formation of the walls enclosing sperm nuclei in the sperma-

tophytes is much needed to carefully distinguish between plasma

membranes and the cellulose secretions that may be developed

by them.

While the cycads and Ginko have very much the largest

sperms known and are consequently extremely favorable for an

examination of spermatogenesis nevertheless some surprisingly

detailed studies have been made among the Filicineze and Equise-

tineze. Following his preliminary announcements ('97a, '97b,

'97c), Belajeff published in '98 an account of spermatogenesis in

Gymnogramme and Equisetum. These forms present histories

parallel to each other and to the cycads. Two deeply staining

bodies (blepharoplasts) appear on opposite sides of each nucleus

previous to the final mitosis in the antheridium which differen-

tiates the spermatids. Consequently each spermatid receives a

blepharoplast which lies close beside the nucleus. The bleph-

aroplast begins to elongate and is followed by the nucleus so

that both structures form two parallel bands which take a spiral

form. (Illustrated in Fig. 34 of Section I.) The rest of the

cytoplasm remains as a vesicle which comes to lie at the larger

end of the sperm. The cilia of Equisetum could be traced to

582 THE AMERICAN NATURALIST. (VoL. XXXVIII.

definite granules in the band as it develops from the compact

spherical blepharoplast.

There appeared almost simultaneously with the foregoing con-

tribution of Belajeff a paper by Shaw ('98b) on Onoclea and

Marsilia. Shaw investigated the cell divisions preceding the

formation of the spermatids in Marsilia and discovered some

very interesting conditions. The two blepharoplasts which are

found in the mother cell of the spermatid are foreshadowed by

smaller bodies which appear at the poles of the spindle in the

two previous mitoses. The first of these structures was called

a blepharoplastoid. The blepharoplastoid first appears besides

the daughter nucleus after the third mitosis previous to the dif-

ferentiation of the spermatids. There is therefore one for each

nucleus of the grandmother cell of the spermatid. This bleph-

aroplastoid divides but the halves remain close together and

the pair passes to one side of the cell. With the next mitosis

(the second previous to the differentiation of the spermatids)

two new structures are formed at the poles of the spindle and

from these the blepharoplasts arise. They accompany each

daughter nucleus after this mitosis into the mother-cell of the

spermatid. Then each divides and the two blepharoplasts pass.

to opposite sides of the nucleus which prepares for the final

mitosis of the series. This division gives a daughter nucleus to

each blepharoplast and the spermatid is thus organized. The

later history of the spermatid as it changes into the sperm is

identical with Belajeff's results.

Belajeff ('99) followed Shaw’s account of Marsilia with a study

of the same form and came to very different conclusions which .

have to do chiefly with his belief that the blepharoplast is a cen-

trosome, a view that will presently be considered in connection

with the opinions of Strasburger and others. Belajeff found

centrosome like bodies (blepharoplastoids of Shaw) at the poles

of spindles in various mitoses preceding the formation of the

spermatids with their unquestioned blepharoplasts. He is not

wiling to concede that these centrosome like structures pass

into the cytoplasm to disappear there as Shaw states for the

blepharoplastoids. He also found the blepharoplasts at the poles.

of the spindles, which was not observed by Shaw, and holds that.

they have a part in spindle formation.

Nos. 451-4452] STUDIES ON THE PLANT CELL. 583

We are now prepared to take a genéral survey of the proc-

esses of spermatogenesis to harmonize as much as possible the

conflicting opinions respecting the homologies of the blepharo-

plast. Strasburger (:00, pp. 177-215) has critically reviewed

the subject and his conclusions are of great interest. He em-

phasizes the kinoplasmic character of the blepharoplast, whether

it be a differentiated region of the plasma membrane (as he

believes for the zoóspores of Cladophora, CEdogonium, etc.) or ,

a special development in the interior of the cytoplasm (pterido-

phytes and gymnosperms). Strasburger thinks that all kino-

plasmic structures, be they centrospheres, centrosomes or

blepharoplasts, hold a very close physiological relation to the .

substance of the nucleolus and that their appearance and size is

largely the result of nuclear activities. Accordingly the bleph-

aroplast might occupy the position of à centrosome without

being genetically related to that structure, and in fact centro-

somes or centrospheres are to be considered more as products of

the cells’ activities than as self perpetuating permanent organs.

There is abundant evidence that the last possibility is the fact

in many forms both plants and animals. Since centrosomes are

not found at other periods of the life history of gymnosperms

and pteridophytes, Strasburger concludes that the blepharo-

plasts cannot be genetically related (homologous) with such a

structure.

Ikeno and Hirase from their earliest writings have considered

the blepharoplast to be a centrosome. . Ikeno (98a) held that the

blepharoplast corresponded with the middle piece of the animal

spermatozoón. Hirase (94 and '97) although noting for Ginko

that the blepharoplasts did not divide and took no part in spin-

dle formation nevertheless called them attractive spheres. The

conclusions of Shaw (98) and Belajeff ('99) for the same type

(Marsilia) have just been summarized and present very different

points of view. Belajeff believes that the blepharoplast of Mar-

silia holds the same relation to the poles of the spindles as a

centrosome. But Belajeff's conception of the centrosome ('99,

p. 204) is that of a morphological and dynamic center which

may or may not be easily demonstrated according to the amount

of stainable substance present. From these discussions it is

584 THE AMERICAN NATURALIST. [Vor. XXXVIII.

evident that final judgment cannot be passed until certain ques-

tions of fact are established by reinvestigations. Shaw and

Belajeff cannot both be wholly correct in their observations and

interpretations and much depends upon the exactness of future

studies upon Marsilia, other pteridophytes, and in the bryo-

phytes. The problems are also related to the processes of

zoóspore formation among the thallophytes.

With respect to the bryophytes Ikeno (: 03) has recently

published an account of spermatogenesis in Marchantia poly-

morpha. He reports for the mitoses in the antheridium, prelim-

inary to the differentiation of the sperm mother-cells, that a

centrosome appears at the side of each nucleus and divides, the

two daughter bodies passing to opposite sides of the nucleus and

becoming the poles of the spindle. He gives evidence that the

daughter centrosomes sometimes divide again when at the poles

of the spindle in anaphase. The centrosome cannot be found

at the side of the daughter nucleus after the mitosis is com-

pleted but it appears when the nucleus is ready for the next divi-

sion. Ikeno's explanation of the reappearance of the centrosome

is unusual. He believes that the centrosome is formed within

the interior of each nucleus as a deeply staining body among the

linin threads. This body moves to the nuclear membrane and :

is thrust out into the cytoplasm through a protuberance from

the nucleus. It then lies outside of the nucleus and becomes

the functioning centrosome, dividing to form two centrosomes

that separate to preside over the poles of the spindle. After

the final mitoses in the spermatogeneous tissue the centrosomes

remain to become the blepharoplasts of the sperms. Each

blepharoplast passes to the plasma membrane of its sperm cell

and develops two cilia. There is formed at this time another

deeply staining body in the cytoplasm considered by Ikeno equiv-

alent to a “Nebenkérper.” The nucleus begins to elongate

and the * Nebenkórper" takes a position between it and the

blepharoplast and in this manner the much attenuated sperm is

organized from the mother-cell.

Ikeno considers the blepharoplast of Marchantia to be actu-

ally a centrosome as shown by its behavior during mitosis. His

account therefore in the main supports Belajeff 's interpretation

Nos.451452] STUDIES ON THE PLANT CELL. 585

of the blepharoplastoids of Shaw which as just described are

regarded by the latter author as centrosomes.. Both Belajeff and

Ikeno are inclined to use the term centrosome with a looseness

that is unusual since the first accounts of this structure gave to

ita place in the cell which is not strictly followed in these

authors’ descriptions of spermatogenesis. Ikeno’s account of

the intranuclear origin of the centrosome is extraordinary.

Intranuclear centrosomes have been reported in several animal

forms but they do not leave the nucleus in the manner described

by Ikeno.

On the whole the writer is more in sympathy with the views

of Webber (:01, pp. 70 to 81), Strasburger and Shaw than

those of the other authors. Assuming that the observations

upon the cycads and Ginko are correct, Webber is certainly

justified in emphasizing the striking fact that the blepharoplasts

are completely independent of the spindle in the body cell and

that they are formed de novo at a distance from its nucleus.

These are peculiarities which, if established generally through-

out spermatogenesis in plants, will remove the processes entirely

from the activities of centrosomes in certain thallophytes (e. g.

Stypocaulon, Dictyota) and in many animal cells. It is certainly

to be expected that a centrosome when present will always hold

an intimate relation to spindle formation during mitosis. It need

not be a permanent organ in cell genesis and an ever increasing

number of investigations indicate that it frequently is not.

Therefore many authors hold that the centrosome is rather the

morphological expression of a dynamic center than a protoplas-

mic structure with an individuality comparable to the organs of

a cell. But these universal characteristics of centrosomes are

apparently not present in the blepharoplasts of the gymnosperms

nor, according to Shaw, in the pteridophytes (Marsilia). But

then the observations of Belajeff and Ikeno are not in accord

with those of Shaw and it is possible that studies in zoóspore

formation and gametogenesis among the thallophytes may pre-

sent the subject in new lights.

For as shown in our discussion of the zoóspore it is not clear

whether the blepharoplasts in those cells are always derived

in the same manner. We have Strasburger’s view that the

586 THE AMERICAN NATURALIST. [Vor. XXXVIII.

structures are thickenings of the outer plasma membrane

(hautschicht) and opposed to this Timberlake's account for Hydro-

dictyon in which the blepharoplast is considered as a structure

independent of the plasma membrane although lying in contact

with it. It must be apparent that the results of Timberlake

are in essential agreement with the events of spermatogenesis

in the pteridophytes and gymnosperms while those of Stras-

burger introduce new elements in giving to the plasma mem-

brane the functions of forming a blepharoplast. The process of

spore formation in the ascus must also be considered in this con-

nection for in that sporangium a centrosphere associated with

each nucleus develops numerous fibrille that resemble so much

a cluster of cilia as to suggest at once a blepharoplast-like struc-

ture, but this centrosphere of course is an important factor in

spindle formation during the mitoses in the ascus. Indeed we

may well ask for further studies in spermatogenesis and zoóspore

formation before we can expect a solution of the problem of the

blepharoplast.

Comparisons have been made between the sperms of animals

and plants, and some authors (e. g. Wilson :00, p. 175, Belajeff

'97C) consider the two cells in essential agreement as to structure

and development. However these views rest on the assumption

that the blepharoplast is truly the homologue of a centrosome.

It seems to be established that the locomotor apparatus of the

animal spermatozoón is derived chiefly from one or more centro-

somes, generally with the co-operation of archoplasm (idiozome,

Nebenkern) present in some form near the nucleus. It is true

that in plants the locomotor apparatus is derived from kinoplasm

which as we pointed out in Sections I and II corresponds closely

to the archoplasm of Boveri, but this is very far from implying.

that structures formed by the archoplasm and kinoplasm respec-

tively need be homologous. Indeed both archoplasm and kino-

plasm are distinguished by their physiological activities rather

than by their morphological manifestations which are too various

to allow of close genetic relationships. ‘Therefore it seems far -

from established that spermatogenesis in plants is along the

same lines as in animals, especially since the weight of evidence

at present indicates that the blepharoplast is not a centrosome.

Nos. 451-452] STUDIES ON THE PLANT CELL. 587

There are numerous problems connected with the physiology

of the sperm that bear directly upon its protoplasmic structure.

Some of these will be treated in Section IV in connection with

processes of fertilization. But at this time it is well to call

attention to the intimate association that sometimes exists

between the nucleus and blepharoplast. 'These structures come

into actual contact in Cycas and Ginko through a process put

forth from the nucleus. It should also be remembered that

Timberlake and Dangeard found the blepharoplasts in the

zoóspores of Hydrodictyon and in the cells of Polytoma con-

nected with the nucleus by one or two fibers. The nuclear beak

that bears the aster in the ascus suggests a similar relationship.

These conditions indicate that the activities of locomotion may

depend vitally upon the nucleus.

3. Ihe Egg.

The subject of fertilization is reserved for the next section

(Section IV) of this series and the present account will deal

only with the structure of the unfertilized egg. As the sperm

is derived from a motile gamete identical with the zoóspore, so

the egg has had a similar origin. We have traced the steps in

this evolutionary process among the algze in a former paper

(Popular Science Monthly, Feb. 1903, p. 300). The first indi-

cation of a differentiation in the sex of primitive gametes is one

of size. The male gametes tend to become smaller while the

female contains a greatly increased amount of cytoplasm. One

of the important factors determining this differentiation is the

number of nuclear divisions which take place in the cells that

produce respectively eggs or sperms. There are generally a

great many more mitoses in antheridia than in oógonia and con-

sequently a given amount of protoplasm must be very much

divided to provide each nucleus with its quota of cytoplasm.

The tendency of oógenesis on the contrary is to conserve the

protoplasm for relatively few nuclei, provided for several eggs or

fora single nucleus in a solitary egg, with the result that the

egg cell is generally richly supplied with protoplasm. Such proc-

esses result in large cells with a prominent chromatophore or

588 THE AMERICAN NATURALIST. [VoL. XXXVIII.

numerous plastids and not infrequently a considerable amount

of food material. The primitive female gametes were provided

with cilia like the male, but with their increase in size came a

sluggishness of movement which resulted in much shorter peri-

ods of motility on the part of these sexual cells. There are

some algæ (Ectocarpus siliculosus, Cutleria, Aphanochæte)

whose motile female gametes come to rest shortly after their

escape from the oögonia and are fertilized as quiescent cells by

the active sperms. These female gametes at the time of fertili-

zation behave physiologically like eggs although their develop-

ment shows a morphology identical with the sperm. When

such female gametes dispense with cilia entirely they become

eggs.

The absence of cilia does away with very much of the com-

plexity which we have just described for sperms. There is no

trace of the blepharoplast in the egg and no indication of the

activities associated with this structure, so conspicuous in sper-

matogenesis. The large motile female gametes of such Algæ as

Bryopsis, Cutleria, Aphanochæte and certain species of Chlamy-

domonas and Ectocarpus will probably show some interesting

conditions when the details of their cell structure and develop-

ment are known, for some of these types are likely to throw light

on the relation which the blepharoplast bears to other structures

in the cell.

The eggs of all plants (Fungi excepted) are believed to be

richly stocked with plastids in sharp contrast to the sperms which

are entirely destitute of these structures in all groups above the

algæ. The plastids in the eggs of Algæ contain the pigments

characteristic of the respective groups giving these cells a very

rich coloration and sometimes an elaborate internal structure

since these plastids or the single chromatophore generally main-

taina symmetrical relation to the nucleus. Leucoplasts (see

Fig. 1 I2) have been found in the eggs of angiosperms

(Schimper, 85) but detailed studies on the cytoplasm of such

cells in spermatophytes, pteridophytes and bryophytes are

i etermine the history of plastids dur-

ing the development of these germ cells and at later periods

after fertilization.

Nos. 451-452.] STUDIES ON THE PLANT CELL. 589

The distribution of the plastids in the eggs of Alga may be

so general that the entire cell is colored as in Fucus, Volvox and

Spheroplea. Or, the plastids may be largely or wholly with-

drawn from some portion of the egg. It is usual for eggs

retained within the parent cell (oógonium) to present a colorless

area of protoplasm that becomes the point at which the sperm

fuses with the egg. Such a hyaline region is called the recep-

tive spot and is generally situated (see Fig. 114) at the side of

the egg nearest the pore or opening in the oógonium through

which the sperms enter. Excellent illustrations are presented

among the Algae in Vaucheria (Oltmanns, '95), CEdogonium

(Pringsheim, '58, Klebahn, '92) and Coleochatte (Pringsheim,

'60, Oltmanns, '98). It has been suggested that the receptive

spot is related to the clear ciliated end of the ancestral motile

gamete and zoóspore but the structures have not been critically

compared to determine the precise character of their proto-

plasmic structure and development. The receptive spot in

some forms (Vaucheria, CEdogonium, Fig. 114) lies directly

under the opening that is formed in the oógonium and its

protoplasm is probably concerned with the fermentative action

that destroys the wall at that point.

The red Algae (Rhodophycez) do not have eggs although in

their sexual evolution they are at the level of heterogamy. The

female gamete (carpogonium with its trichogyne) is a cell homol-

ogous with an oógonium and its protoplasmic contents corre-

spond to an egg, but the protoplast never withdraws from the cell

wall to lie freely as a naked mass of protoplasm within the

structure. But the general agreement of the carpogonium and .

trichogyne with the oógonium and its neck like extension in

Coleochate seems to determine without doubt the homologies

of the former.

There are very few eggs among the fungi that are strictly

comparable to those of the Algae. Monoblepharis (Thaxter '95a)

however unquestionably furnishes such an example. But the

eggs of the Saprolegniales and Peronosporales are probably in

the author's opinion not directly derived from those of Algz.

They are either a peculiar form of sexual cell called the cceno-

. gamete (Davis :00 and :03) or closely related to this structure

590 THE AMERICAN NATURALIST. [Vor. XXXVIII.

which will be given a separate treatment in this section. The

coenogamete is the homologue of a multinucleate gametangium

but its evolutionary tendencies seem to be towards such a

reduction in the number of nuclei that in the highest expression

of its sexual differentiation the female cell contains a single

nucleus and has the general form of an egg. But this process

of sexual evolution is entirely independent of the well known

lines of development in the Algze (Davis, Popular Science Monthly,

Feb. 1903). The female sexual cell of the Ascomycetes (called

the ascogonium or archicarp) is probably in most forms the

homologue of a gametangium. These subjects will be treated

in our account of the coenogamete.

The egg in the archegonium of bryophytes and pteridophytes

is generally reported to have a clearer region on the side nearest

the neck and this is called the receptive spot. It is reported

by Campbell in his investigations on Pilularia (88), Iscetes (’91),

Osmunda (’92a), Marsilia (’92b), and Marattia ('94), by Shaw in

Onoclea ('98) by Thom in Aspidium and Adiantum (99) and

by Lyon in Selaginella (:01). The receptive spot is generally

believed to be a portion of the egg differentiated to receive the

sperm. It is an open question whether this area is morpholog-

ically the homologue of the receptive spot in the eggs of algz

and the clear area at the ciliated end of motile gametes and

zoospores. The problem demands a detailed study of the finer

protoplasmic structure to determine whether or not it is kino-

plasmic in character. The nucleus is generally situated near the

center of the egg and the portions of the cell farthest away `

from the neck of the archegonium contain coarsely granulate

protoplasm which is evidently trophoplasmic, 7. ¢., much of its

substance is of the nature of food material and the products of

metabolism. The leucoplasts would be supposed to lie in this

region of the cell but we know nothing of their presence and

behavior in the egg of bryophytes and pteridophytes.

The eggs of Symnosperms generally speaking present sharp

contrasts to those of pteridophytes. They are very large, prob-

ably the largest uninucleate cells in the plant kingdom, and

consequently very attractive for cell studies and some of the best

work on the events of the maturation and fertilization of plant

Nos. 451452] STUDIES ON THE PLANT CELL. 591

eggs has been done on this group (to be treated in Section IV).

Passing over earlier investigations that described accurately the

general structure of the egg of gymnosperms we shall consider

the results of a number of comparatively recent papers that

treat especially the pine, spruce (Picea), hemlock (Tsuga), fir

(Abies), cycads, Ginko, Gnetum, Taxodium, etc.

Oogenesis and fertilization in the pine has been the subject of

several extensive studies the chief being papers by Dixon ('94),

Blackman ('98), Chamberlain (99) and Ferguson (:o1ib). The

protoplasm of the egg is at first vacuolate but later takes on a

denser structure which becomes very puzzling because of numer-

receptive spot; c, pine, wi merous proteid vacuoles ; Z, embryo sac of the

lily, gamete nuclei fusing, remains of one Synergid (s) shown. : (a, after Schim-

per, 85; 4, Klebahn, ’92; c, Ferguson, :or.)

Fic. 11, — The Egg, a, Daphne, showing leucoplasts; 4, oedogonium, showing

th nu

ous granular inclusions and masses of amorphous material which

together with fibers present a very complex texture. The

fibers are sometimes collected in fascicles and they may form a

sort of weft at the periphery of the egg or radiate out from the

nucleus which is generally surrounded by a kinoplasmic sheath.

The complexity is greatly increased as the egg grows older by

the development of remarkable structures called proteid vacuoles

(See Fig. 11c) which have been especially described by Blackman

and Ferguson. The number of proteid vacuoles is exceedingly

variable in the egg but they sometimes fill three fourths of the

structure. They are spaces in the cytoplasmic reticulum filled

592 THE AMERICAN NATURALIST. [Vor. XXXVIII.

with granules and irregular masses of a proteid nature some of

which stain like nucleoli. The proteid vacuolee were considered

nuclei by earlier writers (Hofmeister and Goroschankin) and

recently this view has been revived by Arnold (:oob) who

describes the migration of large numbers of nuclei from the cells

of the jacket surrounding the egg into that structure. These

results have not been confirmed by Ferguson who agrees with

the interpretation of other writers that the resemblance of the

proteid vacuoles to nuclei is superficial. Miss Ferguson believes

that the material of the proteid vacuoles is derived in part from

the nucleoli in the cells of the jacket and from those in the egg.

A vacuole is reported (Ferguson) at the end of the egg nearest

the neck of the archegonium and this is regarded as a sort of

receptive spot since the pollen tube discharges its contents into

this cavity. The egg nucleus is very large and its contents are

not arranged with the regularity generally present in resting

nuclei. There are numerous bodies which Chamberlain believes

to be chromatic in composition that look very much like nucleoli

and have been so designated by that writer. But there is gener-

ally one large unquestioned nucleolus and besides this many

smaller nucleoli are reported by Ferguson as held in the linin

reticulum. Then portions of the linin frequently take irregular

forms and stain heavily. There is also present besides the linin,

chromatin and nucleoli much granular material (metaplasm),

especially in the nuclei of younger eggs, which probably holds

some relation to the chromatin although it may readily be dis-

tinguished at certain times from that substance.

Recent accounts of the spruce and fir, by Miyake (: 03a and

: 03b) describe conditions very much as in the pine. The egg of

the spruce (Picea) is apparently not so fibrous in structure but

proteid vacuoles give it a coarse granular structure. He finds

no evidence in support of Arnoldi's (: 00b) peculiar views that

the proteid vacuoles are derived from nuclei that have passed

into the egg from cells of the sheath. They are simply masses

of nutritive material. There is some doubt whether the vacuoles

pu the end of the egg really represent a differentiated

receptive spot. The egg of the fir (Abies) conforms in all

essentials to the structure in the pine and spruce. There are

numerous proteid vacuoles.

Nos. 451-452] STUDIES ON THE PLANT CELL. 593

It is probable that the eggs of other conifers will be found to

present much the same protoplasmic structure and activities as

those of the pine. Thus Murrill (: o0) describes for the hemlock

spruce (Tsuga) a vacuolar receptive spot and figures masses of

food material very much like the proteid vacuoles. The general

features of the egg of Cephalotaxus (Arnoldi, :00a), Thuja

(Land, : 02), Podocarpus (Coker, :02), Taxodium (Coker, ’03)

have been recently described and those of Abies, Larix and

Taxus are familiar from older writers but the pine remains as

the type of conifer in which the events of oógenesis are best

known as regards the details of protoplasmic activities.

Besides the pine we have had some very. complete investiga-

tions on cycads and Ginko (Hirase, '98, Ikeno, '98b and :or,

Webber, :01). In some respects these types and especially the

cycads seem to be the most favorable of all the gymnosperms

for the study of gametes and the processes of fertilization (to

be described in Section IV). The cytoplasm of the egg is com-

paratively homogeneous in structure so that the cell is relieved

from the complicated fibrous structure and proteid vacuoles

present in the pine. Ikeno ('98b) finds that the egg of Cycas

develops a crater like depression just before and at the time of

the fusion of the sperm thus presenting a rather highly special-

ized receptive spot. :

We know almost nothing of the detailed structure of the egg

in the Gnetales. Ephedra (Strasburger, '72) develops arche-

gonia much like those of other gymnosperms and we should not

expect their eggs to be materially different even in details. But

the conditions in Tumboa (Welwitschia) are peculiar and approach

more closely those of angiosperms where the egg nucleus is

scarcely differentiated from neighboring nuclei lying freely in

the protoplasm at one end of the embryo sac. The eggs of

Tumboa (Strasburger, '72) are merely cells of the prothallus that

push out small projections to meet the pollen tubes. Gnetum

presents a further simplification or reduction since the female

nuclei lie freely in the protoplasm at one end of the embryo sac.

In Gnetum gnemon the lower half of the embryo sac is filled

with a tissue (Lotsy '99) but in several other species studied by

Karsten ('92, '93) no cell walls are found in the entire sac until

after fertilization.

594 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The angiosperms present no especial advance over Gnetum in

the organization of the egg except that this structure is generally

reduced to a single female nucleus and the cytoplasm immedi-

ately around it (see Fig. 11 7). This egg nucleus flanked by

two companions (synergids) and the accompanying protoplasm

compose the egg apparatus whose morphology is still a matter

of dispute. It is possible that the synergids may stand for

portions of a reduced archegonium, but the two nuclei bear such

close relations to the egg and polar nucleus that it seems very

probable that they are homologous with these structures which

have clearly defined sexual potentialities.. In spite of the '

numerous studies on embryo sacs in various groups of angio-

sperms we do not yet know precisely how the cytoplasm becomes

gathered around the egg nucleus and the synergids. The

spindles that are formed between these nuclei in some types

(e. g., Lilium) have been supposed to lay down walls by means

of cell plates. But there are other forms in which the proto-

plasm seems to separate along planes of vacuoles without rela-

tion to spindle fibers.

(To be continued.)

NOTES AND LITERATURE.

ZOOLOGY.

Dodge's General Zodlogy.! — The title on the cover and the “book

notice” sent out by the publishers do not suggest that this is other-

wise than an entirely zew book, but in small type on the title page is

the information that this is a “ revision and rearrangement ” of Orton's

Comparative Zoology — a well-known text-book written in 1876 by the

late Professor Orton of Vassar College and revised by Birge in 1883

and by Dodge in 1894. But even the information on the title page

does not prevent the reader from expecting to find that the new

author and new title mean that the book has been recast so com-

pletely that the original author deserves credit for only a minor part

of the book. Careful examination leads to disappointment. Parts I

and II of the original work and its revisions are interchanged, and

there are two new chapters — one on practical work and the other on

origin of animal species. In all other respects, the chapters, sections,

and paragraphs of the greater part of the book stand essentially

unchanged, except in minor verbal changes and revisions of the

classification and morphology in accordance with discoveries made

since the original book and its earlier revisions appeared. Thus

Dodge's General Zodlogy is analyzed into Orton's Comparative Zoology

with probably go per cent of the sentences essentially as Professor

Orton wrote them thirty years ago, and added to this about forty

pages of new material. In the light of these facts any fair-minded

man of science will not hesitate to condemn the change of title and

author's credit and especially the method of advertising the book as

a new one. In all fairness to purchasers and to the principle of credit

for original work, the so-called Dodge's General Zodlogy should here-

after be known to teachers of zoólogy as Orton's Zoó/ogy revised and

extended by Dodge; and as such the chief value of the book is that

of the original work with the changes made necessary by advances in

zoólogical knowledge. These have been great, but in such a general

text-book the majority of the revisions have been possible without

rewriting the original sentences.

M. A. BIGELOW.

! Dodge, C. W. General Zeilogy. New York, American Book Co. 1903.

8vo, pp. 512, 379 figs.

595

596 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Coues’ Key to North American Birds! had nominally four edi-

tions, but the third and fourth were practically reprints of the second

(1884) edition. At his death, Dr. Coues was at work on a new

edition, the MS. of which was fortunately completed by him. It

has been published as the-fifth edition of the Key under the direction

of Mr. J. A. Farley, to whose conscientious care the publishers give

due praise.

The present edition appears in two large volumes containing over

eleven hundred pages. The Historical Preface of former editions is

followed by a short memoir of the author, being the address delivered

by D. G. Elliot at the A. O. U. Congress in 19oo. Part I, on Field

Ornithology is reprinted without change. Part II on the Structure

and Classification of Birds has been somewhat revised. That Dr.

Coues nailed his colors to the mast is evident from a paragraph on

p. 8o, where though he avows his allegiance to the A. O. U. code of

nomenclature, he expressly excepts the canon which requires him to

misspell a name * for no other reason than because it was misspelled

in the beginning."

A less gratifying evidence of his courage in his convictions is given

in the discussion of color-change without moult, or Aptosochro-

matism (p. 92) as Dr. Coues was proud to have termed it. It seems

strange that the pages that follow could have been written after the

publication of Mr. J. A. Allen's paper in the Bull. Amer. Nat. Hist.

1896.

The chief additions in this edition are in Part III, Systematic

Synopsis of North American Birds. Here have been incorporated

the results which the study of systematic ornithology has achieved

within the last twenty years. The descriptions of many species, and

of their nesting habits have been rewritten, particularly in the case

of western birds. Many new species and sub-species have been

accorded recognition. Such difficult groups as the Small Flycatchers

(Empidonax) and the Horned Larks (Otocorys) now appear as they

were left by the more conservative treatment of Brewster and Dwight.

Most of the changes in nomenclature made by the A. O. U. commit-

tee have been accepted, and in some cases some of those not made

at the time of Dr. Coues's death were foreseen and included. The

changes made since his death appear in an appendix. There have

been added to many of the biographies, especially to those of the

less familiar birds, the more important synonyms and bibliographical

! Key to North American Birds. The Fifth Edition. Coues, Elliott, Boston,

Dana Estes and Co. 1903. pp. 1152.

Nos. 451-452.] NOTES AND LITERATURE. 597

references. The accounts of the orders and families have been

largely rewritten, and include references to extra-limital groups so

that the relationships of North American birds to foreign groups is

more clearly brought out. Most of the old illustrations have been

retained, and over two hundred new ones have been prepared for

this edition by Fuertes. Old admirers of the Key will probably

rejoice to see the old familiar cuts, absurdly inadequate or useless as

many of them were, and now cruelly contrasted with Fuertes’ brilliant

work.

The great influence which Coues’ Key exercised on the ornithol-

ogists of the past generation is well known, and it is gratifying to

see his work brought as nearly as possible up to date, and its capa-

city for usefulness thus prolonged. It was always a bulky book, and

now, in its present form, it cannot serve as a manual; it must yield

the field to later excellent “Keys.” But it may still rank as one of

the most valuable works on the reference shelf, especially to those

who grew up with it.

One is curious, however, with regard to this work of Fuertes, to

know whether it represents that artist’s earlier or later style. In

some of it his worst faults in bird portraiture are too prominent. In

his drawing of the Mountain Chickadee (p. 271) in his efforts to show

the plumpness of a vigorous bird, the artist has given us an absurd

little caricature.

à RH

Boulenger on the Classification of Bony Fishes.—The most im-

portant recent contribution to the taxonomy of fishes is “ A Synopsis

of the Suborders and Families of Teleostean Fishes,” by Dr. G. A.

Boulenger of the British Museum, published in the Annals and Mag-

azine of Natural History for March, 1904. It is based on his own

studies of the fish skeletons in the British Museum and on the work

of Woodward, Gill, Jordan, Starks and Regan. The special effort

has been to show the relation of the different members of this great

group of nearly 12,000 species by a classification based on our

knowledge of phylogeny. The result of Dr. Boulenger's work is

not essentially different in fact from that of Dr. Gill, although there

is considerable divergence in nomenclature. Dr. Gill has preferred

to isolate aberrant groups until their true relationship is known. As

a result he has recognized twice as many families and twice as many

orders as Boulenger, many of these families and orders being pro-

visional, The advantage of this method lies in clear definition. The

598 THE AMERICAN NATURALIST. [Vor. XXXVIII.

result of leaving aberrant forms in the group nearest to them is to

destroy all possibility of exact limitation. It isatthe best impossible

to frame exact definitions when transitional groups of all grades

exist. A few orders can be isolated and defined among the bony

fishes, but the group as a whole cannot be divided into orders or

suborders or any other categories which shall have even approxi-

mately equal value.

There is no sharp line separating the Isospondyli or Salmon-

herring series from the Ganoids, from which they are derived. From

the Isospondyli or Malacopterygii, as Dr. Boulenger calls them, all

other bony fishes seem to be descended. But the line between the

Isospondyli and the Haplomi is a narrow one. The Haplomi have

lost the mesocoracoid bone, and as it is abortive in many species

(Synodontidae, Myctophidz) hitherto attached to the Isospondyli,

these are placed by Boulenger among the Haplomi. Among the

Haplomi, on the other hand, are placed the spiny-rayed Percopside

because these have not lost the air duct characteristic of the Iso-

spondyli and their allies.

The eels are divided into two suborders, —the Qstariophysi

remain together as one. The Heteromi are made to include Derce-

tidz, Halosauridz and Fierasferide.

The transitional types with abdominal ventrals, the air duct and

the mesocoracoid being lost, constitute two suborders, which do not

differ from each other in definition although comprising different

categories of families. The first of these is the suborder of Catos-

teomi, comprising the Selenichthyes (Lampris), the Hemibranchii,

the Lophobranchii, and the Hypostomides. The present reviewer

sees no reason for associating Lampris with this group. Its suborder

(Selenichthyes) is worthy of independent recognition. The others

certainly belong together, and almost as certainly are degenerate or

specialized allies of the other transitional group of Percesoces.

In Boulenger's scheme, the Percesoces include not only the Athe-

rinidæ Sphyrznidae-Mugilidze series, but also the Scombresocidz,

the Polynemidz, the Chiasmodontide, the Tetragonuride, the Stro-

mateide, the Ophiocephalidz, and the Anabantida. As thus arranged,

the group is defined only by the imperfect attachment of the ventral

fins to the thoracic arch. Itcannot be exactly defined and its mem-

bers are very divergent. It is fair to say, however, that the fault lies

with the fishes, not with the classifier. The transitional elements be-

tween soft-rayed and spiny-rayed fishes are all still extant, and their

intergradations defy classification. For this reason, Dr. Gill once

Nos. 451-452.] NOTES AND LITERATURE. 599

proposed the name Teleocephali for the order containing the whole

ancestral trunk of the bony fishes, from the primitive semi-ganoid

types to the highly specialized cottoids and labroids, leaving only

the divergent branches to be'recognized as separate orders.

The Anacanthini, relieved of the flounders which have no relation-

ship to the cod-fishes, Dr. Boulenger places near the Persesoces.

The Ammodytidae are wrongly placed by Dr. Boulenger with the

Percesoces; as in Embolichthys, the genus possessing ventral fins,

has these fins at the throat.

The great body of the remaining fishes are placed in the suborder

Acanthopterygii, defined essentially as by Gill, Jordan and Evermann

and Hay. The Beryces are placed among the Perciformes, ‘The

recent discovery by Mr. Starks that all Beryces possess the orbito-

sphenoid bone, absent in other Acanthopterygii and characteristic of

the lower forms, indicates that the Beryciformes (Pempheris and

Aphredoderus excluded) should form a division by themselves.

The group Zeorhombi, containing Zeidz and the flounders, repre-

sents an ingenious guess. Even if the flounders, as is probable, be

descended from ancestors of Zeus, the present differences justify

their separation into a distinct division. Zeorhombi is insusceptible

of definition. The remaining orders, Opisthomi, Pediculati, Plec-

tognathi, are arranged as generally accepted, but the Plectognathi

are only specialized and degraded offshoots from the Chatodontida—

Acanthuridz series with which they form a nearly continuous line of

degeneration.

As Dr. Boulenger extends the range of his work, he will find it

convenient to recognize a greater number of families, while Ameri-

can ichthyologists will mark their progress by the gradual reduction

of the number defined by them. This arises from a different

method of work, a different view as to convenience in regard to

divergent or imperfectly known forms. Thus every year brings

workers on the taxonomy of fishes nearer and nearer together, and

farther and farther from the Cuviernian idea of the perch as a per-

fect fish to be placed first with the others following after.

S.L

Notes on Recent Fish Literature.— In the Buletin U. S. Fish

Commission (1902) Jordan and Evermann describe two new fishes

from Hawaii, 7ropidichthys psegma and Zracundus signifer, the latter

representing a new genus of Scorpznidz, allied to Pontinus.

In the Bulletin of the U.S. Fish Commission for 1902 (1903),

600 THE AMERICAN NATURALIST. [Vor. XXXVIII.

William C. Kendall gives a useful list of the certain river-fishes of

Maine. Leuciscus carletoni,a new species, with a barbel, and there-

fore better referred to Couesius, is described as also Pimephales anuli

(named for Edgar E. Ring) and Coregonus stanleyi.

In the Revista Chilena de Historia Natural (Valparaiso, VII, 1903)

Dr. F. T. Delfin discusses in much detail the fishes known as Cong-

rios in Chile, Ophidiidz of the genus Genypterus.

In the Smithsonian Miscellaneous Collections (XLV, 1903), Dr. C.

H. Eigenmann defines 21 new genera of South American river fishes

of the group of Characinide in advance of his monographic review

of the group. It is to be regretted that a closer adherence to clas-

sical models did not prevent the suggestion of such names as Holo-

shesthes, Stichonodon, Colossoma and Mylossoma.

In the Proceedings of the Zoological Society of London (1903), Mr.

C. T. Regan gives a review of the East Indian fishes of the genus

Triacanthus. Jn this as in many other cases it appears that the

species recognized by Dr. Bleeker are all really valid in spite of the

doubts expressed by subsequent authors with more scanty material.

In the Proc. U. S. Nat. Museum (XXVI). Barton A. Bean de-

scribes a new minnow, Notropis brimleyi (near JV. coccogenis) from

Cane River, North Carolina.

Inthe Atti della Società Italiana di Scienze Naturali (XLII, 1903),

Dr. Cristoforo Bellotti of the Museum of Milan describes an inter-

esting new species of fish, Pteraclis macropus, taken off Yokohama.

In the Proceedings of the U. S. National Museum (XXVII, 1904),

Mr. E. C. Starks discusses the osteology of the berycoid fishes.

This group is one of especial interest because it appears earlier in

geological time than any other of the distinctly spiny-rayed types,

and it is probable that all these are described from Berycoid stock.

Dr. Boulenger unites the berycoids with his perciformes, notwith-

standing the archaic characters retained by most of them. Mr.

Starks shows that all of them retain one hitherto unnoticed archaic

character which is lost in all the other spiny-rayed fishes. This is

the development of the bone called osbitosphenoid.

This bone he finds in Hoplostethus, Beryx, Holocentrus. Myri-

pristis, Polymixia and Monocentris. It is not found in Pempheris or

in Aphredoderus, which genera should be excluded from berycoid

association. The orbitosphenoid is present in the Isospondyli and

in the ganoids. It is wanting in the Haplomi and in Aulopus and

Nos. 451-452.] NOTES AND LITERATURE. 601

Synodus. This fact may indicate an additional character by which

to distinguish the order Haplomi, and that Boulenger is right in

referring Aulopus and Synodus to this order.

Starks finds that the close resemblance of the barbels in Polymixia

to those in Mullus and Upeneus has no phylogenetic significance.

While the organs in the two cases look much alike, they are devel-

oped in quite different fashion.

In Polymixia each barbel is suspended from the hypohyal, where

three short, rudimentary branchiostegals curve around its thickened

base. In Mullus and its relatives, the barbel is the extreme tip of

a slender ray of bone attached to the end of the ceratohyal, and

without relation to the branchiostegals.

In the Bulletin of the U. S. Fish Commission (1902, published

1904), Professor John O. Snyder gives a list of the shore fishes

taken by the “ Albatross” expedition in 1902. Two hundred and

twenty-seven species are recorded, with excellent figures of 25 new

species not taken by the previous expeditions of Dr. O. P. Jenkins

(1889) and Jordan and Evermann (1901). Two new genera are

described, Veternio, a toothless Conger eel, and Collybus, a genus of

Bramide.

In the Bulletin U. S. Fish Commission (for 1902, published in

1904), Dr. Jordan gives a list of species of small fishes from coral

pools, taken at the Tortugas, Florida, by Dr. Joseph C. Thompson,

U. S. N., now surgeon of the “ Albatross,”— Ctenogobius tortuge

Gnatholepis thompsoni, Elacatinus oceanops, and Ericteis iin

Elacatinus is a new genus of gobies, Ericteis a new genus of blennies,

in which group another genus, Acteis (moorei), is detached from

Malacoctenus.

Chaetodon bricei is shown to be the young of Chetodon capistratus,

and the nominal species of Doratonatus are all probably identical

with D. megalepis. The rare Holocentrus siccifer (Cope 1871) is

again recorded. :

Mr. Henry W. Fowler, now of the Academy of Natural Sciences

at Philadelphia, has in the Proceedings of that institution (1903) a

number of papers on fishes largely of the Pacific Ocean. ot

special interest are the following: Atherina sardinella,a new species

from Italy; Atherinomorus, a new sub-genus for the West Indian

Atherina laticeps, Protistius semotilus Cope redescribed. This genus

is allied to Atherinopsis, but has only a single dorsal spine. Gas-

tropterus Cope is the same as the later Pisciregia of Abbott.

602 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Atherinops magdalene Fowler is described from Magdalena Bay,

Lower California. Mugil kelaarti is described from Honolulu. Liza

caldwelli is redescribed from Samoa. A new sub-genus, Aodale-

chilus, is proposed for Mugil labeo. Sphyrena tome is described

from Sambaia, Brazil. Centrobranchus cherocephalus is described

from Hawaii. Centrobranchus is a new genus of Myctophide

allied to Rhinoscopelus, but the gill rakers reduced to bunches of

Spines. |

Numerous species are recorded from Zanzibar. These papers are

illustrated with good plates.

In the Danish Meddelser om Grönland (XIX, 1904), Mr. A. S.

Jensen gives a list of the fishes of Eastern Greenland, based on the

collections of the Amdrup-Hartz expedition of 1900. Thirty-six

species are included, the following being new: Lycodes eudipleuro-

stictus, Lycodes (reticulatus) macrocephalus and L ycenchelys &olthofft.

Excellent faunal notes are given with a very convenient bibliography.

According to Jensen, the American Cottus or Oncocottus hexacornis is

identical with the European guadricornis, as Jordan and Evermann

had suspected. The American Artediellus atlanticus is also shown

to be the same as the European A. uncinatus, from which Jordan and

Evermann thought it apparently distinct. The name Zelus bicornis

is rightly used instead of the later Lcelus hamatus. The Greenland

species known as Liparis tunicata proves to be the same as Liparis

“iparis. Liparis fabricii is different. Paralepis kröyeri is shown to

be different from P. borealis and the Charr of East Greenland is

identified as Samo (better Salvelinus) alpinus. ‘The engravings are

entirely worthy of this admirable piece of faunal study.

Professor Keinosuke Otaki of the Imperial Military Academy of

Tokyo, a graduate of Stanford University, has begun in connection

with Professor T. Fujita and T. Higurashi the publication of a serial

work entitled “ Fishes of Japan." The text is in Japanese and

English. . The species included in part first are Lateolabrax japonicus,

Latilus sinensis, Gymnosarda affinis and Seriola quingueradiata.

These are illustrated by colored plates well executed by Mr. K. Ito.

In the Report on the Sea and Inland Fishes of Ireland for 1901,

Messrs. E. W. L. Holt and L. W. Bryne describe and figure a trans-

lucent symmetrical larval flounder, which they regard as the young of

the “White Sole," Glyptocephalus cynoglossus. The mouth is shown

as much larger than in the adult of this species, which may throw

doubt on the identification.

Nos. 451-452.] MOTES AND LITERATURE. 603

The same gentlemen in a later paper describe and figure all the

British gobies. Their plates will greatly aid in the identification of

these little fishes.

In the Annuaire du Muste Zoologique at St. Petersburg (VIII,

1903), L. S. Berg discusses a collection of Cottoid fishes from Lake

Baical, with observations — unfortunately in Russian — on the clas-

sification adapted by Jordan, Evermann and Starks. Two new

genera, Baicalocottus and Batrachocottus, are defined, with the genus

Procottus of Gratz.

When the present writer was in Japan in 1900, the Imperial Museum

generously placed at his disposal all duplicates of new species of fishes

contained in the Museum. ‘Those represented by single examples

could not be sent away, and these the director of the Department of

Natural History, Dr. Chiyomatsu Ishikawa, has undertaken to de-

scribe for the sake of completing the rich faunal list of Japan.

In the Proceedings of the Imperial Museum, Dr. Ishikawa describes

the following new species, most of them with good figures:

Zesera hilgendorfi, Zacco mitsukurii, Leuciscus dorobaé, Stromateoides

nozawe, Heterognathodon dederleini, Cirrhilabrus lyukyuensis, Amphip-

rion snyderi, Crenilabrus stejnegeri, Tetraroge kagoshimensis, Hemi-

tripterus nipponicus, Podothecus tokubire.

Of these the Crenilabrus from the Riu Kiu Islands is the most inter-

esting, as that genus, largely represented in the Mediterranean, had

never before been recorded in other waters. The Tetraroge should

rather be referred to the genus Erisphex.

In the Records of the Australian Museum (V, 1904) Mr. Edgar R.

Waite gives an account of his explorations of Lord Howe Island.

Seventeen known species of fishes were added to the fauna of this

interesting island. Ten new species were discovered, most of these

with a number of other interesting forms being figured by Mr. Waite.

The new genera are Xenogramma (Gempylida), Allogobius, Lim-

nichthys (Trichonotida) and Lepadichthys (Gobiesocidz). he

fauna as a whole, although distinctly tropical, is very different from

that of Hawaii or Samoa.

In the same Records (V, 1904) Mr. Waite gives a list of the fishes

known from Lord Howe Island, 18o in number, with a bibliography.

The list is especially useful for comparison with faunal lists of the

South Seas, the genera being largely identical, the species different.

In this list Mr. Waite has taken special care to verify the dates of

publications.

604 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In the Transactions of the American Philosophical Society (XX,

1901) Dr. O. P. Hay gives a valuable table showing the chronologi.

cal distribution of the genera of elasmobranchs. He calls attention

to one remarkable fact. While many genera and species (Clado-

selachidz, Acanthoésside, Psammodontide, Pleuracanthidz, Clado-

dontidz, etc.) occur in the Devonian and Carboniferous ages, only

three genera remain in the Permian and but four in the Triassic.

All these are Heterodontidz, the successors or descendants of the

Orodontidz of the Carboniferous. In the Jurassic, numerous fami-

lies with many genera appear, and through these from Heterodontid

stock the modern sharks seem to have sprung. Apparently the Pale-

ozoic families, except the Orodontida, have no modern descendants.

In the Bulletin of the Museum of Comparative Zoólogy (XLI, 1904)

Samuel Garman gives an elaborate and excellent account with beau-

tiful plates of the anatomy of the long-nosed Chimzra of Japan, RAz-

nochimera pacifica (Mitsukuri). In connection with the study of this

species, he has given a general account of the living chimzroid

fishes, with figures of some of them. On grounds of priority, Mr.

Garman substitutes the name Chismopnea Rafinesque, 1815, for the

later name Holocephala of Muller (1834).

In the Geological Magazine (IX, 1902) Dr. Charles R. Eastman

founds a new family of fossil sharks, Peripristidae, on the genus

Peripristis (Pristodus). i

In the American Geologist (XXX, 1902) Dr. Bashford Dean gives

illustration of a startlingly perfect preservation of the muscular fibre

of a shark, C/adose/ache fyleri, in the Cleveland shales. This was an

estuary deposit originally of fine mud, in which the fish remains

became phosphatized.

In the Proceedings of the Academy of Natural Sciences of Philadel-

Phia (1903) Dr. Carl H. Eigenmann and Clarence H. Kennedy dis-

cuss a large collection of river fishes, made in Paraguay by Prof. J.

D. Anisits of the Paraguay National University. One hundred and

nine species are noted, many of them new, and the paper is followed

by a useful biography. A feature of special convenience is a synop-

sis of the genera of Cichlidz, 2 5 in number, Biotodoma and Biotce-

cus being new names,

In the Annotationes Zoologice Japonenses of the Imperial University

of Tokyo Dr. Bashford Dean gives a study of the development of

the egg of the Japanese Shark, Heterodontus japonicus. He finds

good evidence of the existence of holoblastic cleavage.

Nos. 451-452] | NOTES AND LITERATURE. 605

In the Christiana Videnskabs-Selskabs Vorkandlinger for 1902 and

1903 (the last volume being published from the Fridtjof Nansen

fund) Dr. Robert Collett gives additional records of additions to the

fish fauna of Norway. These admirable notes are of great value to

the student of the fishes of Northern Europe, as well as of Northern

regions generally.

Jordan and Starks continue in the Proceedings of the U. S. National

Museum (XXVII, 1904) their monographic accounts of the fishes of

Japan.

In the review of the Scorpznidz 52 species are described as

Japanese, 14 of these being new. These are arranged in 23 genera,

the following being new: Thysanichthys, Sebastiscus, Lythrichthys,

Ebosia, Decterias, Inimicus, Ocosia, Erisphex. The new species and

most of the new genera are figured.

The Cottidz of Japan are also described in detail by the same

authors. In these fishes as in the Scorpznidz, of which the Cottidze

are the Arctic descendants, the waters are particularly rich. Fifty-

seven species are described, 19 being new. These are arranged in 40

genera, the new genera being Stlengis, Schmidtia, later called

Schmidtina, the name Schmidtia being preoccupied. Daruma,

Ricuzenius, Rheopresbe, Ainocottus, Crossias, Elaphichthys, Alcich-

thys, Furcina, Ocynectes, Bero, Vellitor.

Similar reviews cover the less numerous species of the families of

Agonide and Hexagrammidz.

The two species of white Chimera (Chimera phantasma and

Chimera mitsukurii) found in Japanese waters are described and

figured by Jordan and Snyder.

Dew.

PALZEONTOLOGY.

Eastman's Translation of Zittel, Vol. II.'—— Since the publica-

tion of the first volume oi Dr. Eastman's translation of Zittel's

Grundzüge der Paleontologie, English speaking students have awaited

with keen interest the appearance of the second volume, and it may

lvon Zittel, Karl A. Text-Book of Paleontology, Vol. II. Translated and

edited by Charles R. Eastman. London and New York, Macmillan, 1902. Svo,

283 pp., 373 figs.

606 THE AMERICAN NATURALIST. [Vor. XXXVIII.

be said that the latter fully maintains the high standard set in the

earlier portion of the work. The present volume treats of the Ver-

tebrates, with the exception of the mammalia, to the consideration of

which class the third and last volume will be devoted.

The general method of treatment which has made Zittel’s works

so valuable to the student happily has been closely adhered to in

the translation; we refer especially to the introductory sections on

the various classes, and to the brief diagnosis in italics with which

the consideration of each group,— class, sub-class, order and family,

— isintroduced. It is largely to this latter feature that these works

owe their superiority as books for ready reference.

In comparing the English edition with the original we note first

very considerable amplification and revision, representing the ad-

vance in Palaozoólogy during the seven years since the publication

of the earlier work, and in this enlargement and revision Dr. East-

man has had the collaboration not only of the author but of a num-

ber of competent English and American specialists. As regards

Taxonomy the changes from the original have been very slight, far

less indeed than we had hoped to see, and it is to be regretted that

the editor and his collaborators so far subordinated their own views

on classification to those of the author, since some rearrangement,

especially of the fishes and reptiles, was quite desirable.

The entire section on fishes has been translated and revised by

Dr. Arthur Smith Woodward of the British Museum. Dr. Wood-

ward has been rather conservative in the matter of revision and has

kept his own opinions on taxonomic points very much in the back-

ground. Of this section the part on ostracoderms has undergone

the most extensive revision, chiefly based on the recent researches of

Traquair. Thus we find Traquair's order Anaspida included, and in

all nine families of ostracoderms are recognized. The Arthrodira,

which in the original were given ordinal rank among ostracoderms,

are now very properly placed as a sub-class, but the Macropetalich-

thyds —the Anarthrodira of Dean — are ranked incerte sedis. It is

rather to be regretted that the old division of the bony fishes iuto

Ganoidei and Teleostei has been adhered to, instead of the better

morphological differentiation of Crossopterygia and Actinopterygia.

The old system has, however, the merit of familiarity and conven-

lence. The section on ganoids has been amplified about one half

ue s Noise revised. Little is said regarding the sys-

à € Dipnoi, but we can scarcely approve of their

place in the book, in immediate sequence to the Arthrodires; it sug-

gests tacitly the former supposed relationship of the two groups.

Nos. 451-452] NOTES AND LITERATURE. 607

The section on Amphibia, revised by Dr. E. C. Case, differs but

little from the original, except in the added descriptions of several

genera. The taxonomic system is unaltered and, on the whole, is

quite satisfactory, but it must be conceded that the Phyllospondyli of

Zittel (the Branchiosauria) do not form a group coórdinate with

Lepospondyli, and there is much to be said in favor of comprising

the Branchiosauria, Microsauria, and Aistopoda as sub-orders of the

order Lepospondyli.

The treatment of the reptiles, in which the editor had the collab-

oration of H. F. Osborn, S. W. Williston, J. B. Hatcher and O. P.

Hay, occupies nearly one half of the book, and is admirable in details,

such as the descriptions of smaller groups, but the taxonomic

arrangement shows nothing of the relationships of the various orders

which compose the class. It is regrettable that the revision of this

section was completed before the appearance of recent papers by R.

Broom, H. F. Osborn and others, which show in a convincing man-

ner that the reptiles have had a diphyletic evolution. Some of these

papers have considerably amplified our knowledge of the Anomo--

dontia and have indicated that the Parasuchia constitute an order

quite distinct from the Crocodilia.

To Mr. F. A. Lucas is due the credit for the enlargement and

thorough revision of the section on birds. This part has been almost

entirely re-written, and a different taxonomic scheme, based chiefly

on Stejneger's classification, has been adopted, to the marked

improvement of the book, although, as Mr. Lucas explains, it is not

possible in the case of birds to do more than group related forms,

and quite out of the question to arrange the groups so as to indicate

phyletic relationships.

It will be noted that the points here criticised are all points of

classification, points on which the editor and his collaborators were

not entirely free to express their personal views. The translation

and revision have been excellently done, and we may say that Dr.

Eastman has given us in the present volume the most useful text-

book published on the Paleontology of the verebrates below mam-

mals. We trust that the volume on mammals may be as good.

J. H. McG.

608 THE AMERICAN NATURALIST. [Vor. XXXVIII.

BOTANY.

A New Book on Ferns.' — For all who study or wish to study our

native ferns, Dr. Waters has prepared a book which is sure to prove

both helpful and inspiring. The numerous photographic illustrations

include enlarged views of the fructification in which the generic char-

acteristics are often surprisingly well brought out. Specific and vari

etal differences which frequently prove perplexing to beginners are

clearly shown in photographs of fronds or entire plants. Sometimes

these are grouped instructively in series to exhibit the range of varia-

tion. Especially charming and significant are the views showing

typical habits and habitats.

The text besides pointing out the features by which forms are dis-

criminated, directs attention most happily to the out-door aspects of

ferns — their adaptations and preferences — in a way to encourage

the best sort of field work. As a help to determining specimens not

in fruit the key based upon characters drawn from the stipes will

doubtless prove welcome to all students of the group. While excep-

tion might be taken to such unnecessary departures from botanical

accuracy as the use of “stem” for stalk or stipe, the substitution of

untechnical for technical expressions has been, on the whole, skillfully

done. Dr. Waters writes as a lover of ferns and his contagious

enthusiasm suffuses the whole book.

F l9

Porter's Flora of Pennsylvania.? — For sixty-five years Professor

Porter was an assiduous and intelligent collector and student of the

flora of the State in which he lived. A contemporary of Darlington,

Torrey and Gray, he shared their knowledge and views; but he

lived to see a school of natural grouping of orders, species segrega-

tion, and nomenclatorial reform, quite different from theirs, come to

the front, and the editor of this posthumous Flora tells us that being

thoroughly acquainted with the author's ideas he can confidently say

! Waters, Campbell E., Ph.D. Ferns: A manual for the Northeastern States,

with Analytical Keys based on the Stalks and on the Fructification, and over 200

Illustrations from Original ee and Photographs. New York, Henry

1903. 4to, xii+ 3

? Porter, J. N. Flora of A eRe Edited, with the addition of analytical

keys, by J. K. Small. Boston, Ginn & Co., 1905. Svo, pp. xv + 362, with out-

line map.

Nos. 451-452.] NOTES AND LITERATURE. 609

that he would have heartily subscribed to everything in it as published.

The book is essentially a list of the spermatophytes or flowering

plants of the State, arranged in the phylogenetic sequence of Engler

and Prantl, with keys to the orders, families, genera and species,

and copious indexes to Latin and vernacular names. The nomen-

clature adopted is apparently based on the Rochester code, a com-

mon name is given for each species, with a reference to its descrip-

tion in Britton’s Manual, and its figure, if one is published, in

Britton and Brown's /Yora,—to supplement the terse characters

embodied in the keys; and each is accompanied by a brief indica-

tion of habitat, general geographic distribution, and an enumeration

of the counties in which it occurs in Pennsylvania. As a determina-:

tion hand-book, it is one of the best local floras thus far published.

Its defect — if it be one lies in the omission of all synonymy,

leaving a very large proportion of the plants under names different

from those by which they were known to the author during the

greater part of his life and by which his contemporaries referred to

them in their classic publications, and without intimation to the

novice that this is the case or direct indication of any means by

which the two may be correlated, — the Manual referred to being of

no assistance in this matter.

The Journals.— Botanical Gazette, February: — Darwin, “On a

Self-recording Method applied to the Movements of Stomata ” ;

Holferty, “The Archegonium of Mnium es ag "i Lewis,

“Studies of some Anomalous Dicotyledonous Plants”; Shriner and

Copeland, “Deforestation and Creek Flora about ie Wis."

Snow, “The Effects of External Agents on the Production of Root

Hairs”; Duvel, “Preservation of Seeds Buried in the Soil”;

MacMillan, * Cumaphytism in Alaria.”

Botanical Gazette, March : — Garber, “The Life History of Riccio-

carpus natans? ; Merriman, “ Vegetative Cell Division in Allium”

American Republics — XXV”; Thom, “ Craterellus taxophilus a

New Species of Thelephoracex ”. Greenman, “Notes on South-

western and Mexican Plants."

Botanical Gazette, April: — Bennett, “Are Roots Aerobic?” ;

Nelson, “Contributions from the Rocky Mountain Herbarium —

V”; Lyon, “The Evolution of the Sex Organs of Plants";

610 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Robertson, * The Structure of the Flowers and the Mode of Pol-

lination of the Primitive Angiosperms”; Coville, *Arcterica, the

Rarest Genus of Heathers”; Ganong, “ New Precision-Appliances

for Use in Plant Physiology.”

The Bryologist, March: — Fink, “ Further Notes on Cladonias —

Cladonia fimbriata” : Holzinger, “The Genus Anacolia in North

America”; Cardot, “Notes on Some North American Mosses” ;

Collins, “Some Erroneous References”; Grout, “A New Brachy-

thecium — B. rivulare tenue”; Towle and Gilbert; “The Fruiting

Season of the Hair-cap Moss.”

Bulletin of the Torrey Botanical Club, March : — Arthur, * Taxono-

mic Importance of the Spermogonium” ; Harshberger, “ A Phyto-

Geographic Sketch of Extreme Southeastern Pennsylvania”; Cush-

man, “Desmids from Southwestern Colorado”; Britton, “ Four new

North American Birches”; and Richards and MacDougal, “The

Influence of Carbon Monoxide and other Gases upon Plants, — a

Correction."

The Fern Bulletin, January : — Parish, “The Fern Flora of Cali-

fornia”; Clute, * Measurement of Variation in Equisetum "; Hill,

“Remarks on some Fernworts of Western New York" ; Clute, ^A

New Species of Equisetum [E. Ferrissi]"; Eaton, “The Genus

Equisetum in North America — XVI”; Klugh, “Treasure Trove” ;

Fetherolf, * Astlenium ebenoides ”; and a biographic sketch, with

portrait, of Sarah Frances Price.

Journal of the New York Botanical Garden, April: — Anderson,

“Protection of our Native Plants” ; Rusby, * Beverages of Vegetable

Origin"; and Howe, * The Pike Collection of Alge.”

Ohio Naturalist, February : — Kellerman, “Index to Uredineous

Culture Experiments, with List of Species and Hosts for North

America"; Schaffner, *Some Morphological Peculiarities of the

Nymphaeacez and Heliobize.

Ohio Naturalist, March : — Schaffner, “Ohio Plants with Extra-

Floral Nectaries and other Glands."

Ohio Naturalist, April: — Cook,

“Galls and Insects producing

them, with Appendix."

The Plant World, February : — Safford, “ Extracts from the Note-

Book of a Naturalist on the Island of Guam — XV ”; Price, “ Ken-

tucky Oaks"; Safford, “ Henry Elwood Baum (with portrait) " ;

Nos. 451-452.] NOTES AND LITERATURE. 611

Squires, * Wild Flowers of Prairie and Canyon in Northern Idaho" ;

Beal, *What is a Bud and how long does it retain its Identity ” ;

Pepoon, “ Destruction of a Farm Flora.”

The Plant World, March :— Safford, “ Extracts from the Note-

Book of a Naturalist on the Island of Guam — XVI”; Waters, “ Plant

Wounds and Natural Pruning”; Blodgett, “ Frost-Weeds and other

Winter Notes”; Bailey, * Undraped Trees”; Britton, “ Relations of

Plants to Birds and Insects.”

Rhodora, February :— Davenport, “Recollections of Charles

Christopher Frost (with portrait) ”. Ames, “Spiranthes neglecta” ;

Davenport, * Miscellaneous Notes on New England Ferns — VI” ;.

Fernald, “Preliminary Lists of New England Plants — XIII.

Juncacez"; Collins, “The Black Spruce in Rhode Island”;

Weatherby, “ Panicum Commonsianum in Connecticut”; Andrews,

“Some Interesting Mosses from a Southern Vermont Peat-Bog”;

and Ames, “ Spiranthes Grayi.”

Rhodora, March : — Leavitt, * Partial Reversion in Leaves of the

Fern-leaved Beech”; Graves, “Noteworthy Plants of Southeastern

Connecticut" ; Robinson, “Identity of Anychia dichotoma” ; Rehder,

Preliminary Lists of New England Plants — XIV."

Rhodora, April: —Chase, “North American Allies of Scirpus

lacustris”; Andrews, “ Bryophytes of the Mt. Greylock Region —

III”; Slosson, “A New Hybrid Fern from Vermont”; Sanford,

“The Range of Saururus cernuus extended into R. I.”; Harger,

“Some Introduced Plants of Ct.”; Nye, “Bulblets of Microstylis

ophioglossoides” ; Knight, “Cleome serrulata in Me.”; Graves,

* Helenium nudiflorum in Groton, Ct.”; and Osmun, “ Further Sta-

tions for Botrychium matricariafolium in Ct.”

Torreya, February : — Stone, “ Physiological Appliances — II” ;

Arthur, “An interesting unpublished Work on Fungi ”; Britton,

Juncacez of the West Indies"; Britton, “Agdestis Clematidea ”;

Harper, “New Station for Arabis Georgiana ". Burnham, * Notes

on Zpigea repens”; and Atkinson, “A New Lemanea from New-

foundland. |

Torreya, March:— Osmun, “A Summer in Salisbury, Ct.";

Sumstine, “The Slime-Mounds of Pennsylvania"; Eggleston, " The

Crategi of Ft. Frederick, Crown Point, N. Y.”; Britton, “ Crategus

Porteri.”

W. T.

(No. 450 was mailed August 24, 1904.)

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THE

AMERICAN. NATURALIST.

VoL. XXXVIII. September, 1904. No. 453.

SOME OBSERVATIONS ON RORQUALS OFF

|J SOUTHERN NEWFOUNDLAND.

GLOVER M. ALLEN.

Unt very recently it has been the usage, in books on nat-

ural history, to picture Cetacea, when in their native element, as

floating lightly on the surface of the water and sending forth

from the blow-holes great columns of spray which break and fall

in showers over the back. In the works of the older writers,

as Bonnaterre and Lacépéde, the spouts of whales are repre-

sented as solid columns of water, of nearly uniform diameter

` throughout, which after reaching their maximum height, curve

over, either to the front or to the rear, and, breaking slightly,

vanish away. Such representations, however, were recognized

as entirely inadequate, being merely the conventional vagaries of

the artists. K. E. von Baer (’64) seems to have been among the

first to attempt an accurate delineation of the whale’s spout. He

figures a Finback whale in the act of “blowing,” the column

being a vertical one, expanding very slightly until the maximum

height is reached, when it bushes out and gradually becomes

dispersed. Henking (:01) also represents in a very diagram-

matic way his impression of the form of a Finback whale's

613

614 THE AMERICAN NATURALIST. [Vor. XXXVIII.

spout. The outline he makes retort-shaped; and the whole is.

directed slightly backward. Both these authors add that their

observations were made in calm weather with a smooth sea.

Not until 1903 have there been published any actual photo-

graphs of the larger whales alive and free in the open ocean.

The first published photographs of this nature appear in the

report on the Cetacea of the Antarctic expedition of the

* Belgica." These represent the Humpback whale (Megaptera

nodosa) and the Sulphur-bottom (Ba/enoptera musculus) in the

various positions assumed during their appearance at the surface

of the ocean, and were taken by Dr. E. G. Racovitza and Dr.

F. A. Cook, in 1898. Only one view is shown of the spout,

and this is so indistinct as to be rather unsatisfactory. Later

in the year 1903, Dr. F. W. True (:03*) published some very

excellent photographs of Finback whales (Ba/enoptera physalus}

taken from the bow of a whaling steamer off the east coast of

Newfoundland. These views show very well the appearance of

this whale in its various postures following the spouting, until

its final plunge. No photograph of the spout itself was ob-

tained, however, so that it seems worth while to publish a few

views of spouting whales obtained by the present writer a few

months ago.

Through the courtesy of Mr. Alexander McDougall, manager

of the Newfoundland Steam Whaling Company, I had the privi-

lege of visiting the whaling station at Rose-au-Rue, in Placentia

Bay, Newfoundland, during the second week of September,

1903. A number of interesting observations were made at this

time and a valuable series of photographs was secured, some of

which are reproduced here.

Four species of rorquals are taken on the Newfoundland

coast: the Humpback (Megaptera nodosa), the Sulphur-bottom

(Balenoptera musculus), the common Finback (B. physalus), and

the Pollack whale (B. borealis) or, as the Norwegians call it, the

“Sejhval.” True (: O3) was the first to record the presence of

the last named species on this side of the Atlantic, on the basis

of four specimens taken at the Rose-au-Rue station during the

season of 1902. The steam whaling industry at Newfoundland

1s one of recent origin, having been established in 1898. Accord-

No. 453] OBSERVATIONS ON RORQUALS. 615

ing to the Morning Chronicle, of Halifax, N. S., the amount of

whale oil produced in Newfoundland for the fiscal year ending

June 30, 1902, was valued at $125,287. In addition to the oil

which is tried out from the blubber and carcass, an excellent

“ guano” is prepared from the refuse flesh and the bones are

ground up into lime..

The fishery itself is carried ‘on by means of small and

staunchly built iron steamers of something over one hundred

tons. A cannondike gun is mounted on a pivot at the bow,

and discharges a five-foot harpoon of over roo pounds weight,

which at short range is nearly buried in the body of the whale.

A hollow, iron cap filled with blasting powder is screwed to the

tip of the harpoon, forming its point. A timed fuse discharges

this bomb inside the body of the whale. The harpoon carries

a stout cable which is handled by a powerful 5-sheet winch on

the steamer's deck.

On September 9th the writer accompanied the whaling steamer

* Puma," Captain Christaffersen, on the daily hunt in the lower

part of Placentia Bay, and obtained several successful photo-

graphs of living whales at close range. A few of these are here

reproduced, and illustrate particularly the spout of the Sulphur-

bottom whale (Balenoptera musculus), no photograph of which

has hitherto been published, with the exception of the one by

Racovitza.

Both Finback and Sulphur-bottom whales observed on this

occasion seemed to go through a regular series of evolutions,

and were doubtless feeding. They rose to spout about once in

every 12 to 15 seconds with great regularity for perhaps twelve

times, after which they dove for a much longer stay of several

minutes. The precise length of the longer periods was not

accurately determined, but could hardly have been more than

5 or 10 minutes. On rising, the first part of the animal to

reach the surface is the top of the head; at the same time it

spouts, and a portion of the long back comes into view. The

head is then lowered, the body arches slightly and the descent

begins. The back comes curving out of the water and down

again, till finally the dorsal fin appears. By the time the fin has

reached the surface again in its forward and downward move-

616 THE AMERICAN NATURALIST. (VoL. XXXVIII.

ment the entire body has disappeared. The flukes were not

thrown out of water by either of the two species seen alive, as

has been noted by True and others.

The whale, in diving, leaves a long “slick” on the water at

the spot where it went down, and comes up again in regular

course several times its length farther on when making a series

Fic. 1.— A sulphur-bottom whale spouting.

of “spouts” or breathings. The distance between successive

spouts seemed to be nearly two or three times the length of the

whale, z. e., 150 to 200 feet.

When a whale is sighted the steamer is put about to overtake

it, but the endeavor seems to be not so much to head it off as

to cut in behind so as not to unduly frighten the animal. On

overtaking the quarry, the steamer is manceuvered so as to come

No. 453-] OBSERVATIONS ON RORQUALS. 617

to a stop at about the spot where the whale is expected to rise

for the next spout. On one occasion a Sulphur-bottom was thus

followed for a considerable distance till finally the vessel came

to a standstill at about the place where the next appearance of

the animal was expected. The distance had been well judged,

and the writer, standing with camera ready, was able shortly to

perceive the shadowy form rising obliquely under the port bow.

As the whale broke water and shot forth a column of vapor, the

click of the camera and the crash of the harpoon gun sounded

almost at the same instant. The photograph obtained (Fig. 1)

shows the Sulphur-bottom with the region of the blow-holes just

$4 S ln}

Fic. 2. Head p

out of water. The spout itself was a very short one and is seen

to have somewhat the form of a narrow, inverted flask. The

wind, blowing from left to right of the picture, carries the upper

portion of the vaporous stream away to leeward. The most

interesting feature of this view is that the area at each side of

the blow-holes is clearly seen to be elevated above the apertures

themselves as the breath escapes. The elevation of these

ridges is well shown in side view among the photographs

obtained by True (:03*) and in the drawings by Racovitza

(:03), but neither of these observers was able to determine

satisfactorily whether it was the blow-holes themselves, or only

618 THE AMERICAN NATURALIST. (Vou. XXXVIII.

the adjacent parts, that were thus raised. In the view here

shown, which was taken from directly behind the animal’s head,

there can be no doubt that the portion elevated in spouting

is the region lying along the external side of each nasal aper-

ture. The broad, shallow groove or depression extending down-

ward from each side of the blow-holes may possibly be due to

the muscular contraction incident to the raising of the two

ridges. The same feature in side view is possibly shown in one

of the photographs by True (:03, Pl. 25, Fig. 2). No such

groove was seen in the dead specimens. The column of vapor

itself is clearly single, even though arising from two apertures,

for the latter are situated so close together that the two jets

Fic. 3 Spout f Iphur-b

of vapor must unite at once. The photographs do not, there-

fore, bear out Packard's (66, p. 272) statement, on the testi-

mony of'another, that the Sulphur-bottom blows in a ** double

stream which is directed backward toward the tail." The blow-

holes of a large whale of this species are represented in Figure

2. The animal lies on its left side with the upper surface of the

head toward the observer. The mouth is partly open, and from

it projects the fringe of baleen. The two slit-like nasal open-

ings are seen near the lower right hand of the figure and appear

to be situated between the arms of a V-shaped prominence

whose point is directed forward, and is continued as a slight

No. 453.] OBSERVATIONS ON RORQUALS. 619

median ridge toward the tip of the snout. In the dead animal,

however, there is hardly more than this slight suggestion of the

nasal ridges which are so prominent in life.

The form of the spout, in both the Sulphur-bottom and the

Finback whale, unless distorted by the wind, is that of a simple

column, narrow at the base and gradually increasing in diameter

with the height, like a jet of steam forced through a small open-

ing. Such a spout is shown fairly well in Figure r, Plate r, of

Racovitza's (:03) paper. The views obtained by the present

writer all show the effect of the light wind blowing at the time,

in that the vapor is carried off to leeward to a greater or less

Fic. 4.— An irregular spout of a sulphur-bottom.

extent. Figure 3 shows the spout of a Sulphur-bottom which

is fast by aline to the whaling steamer. This view shows the

general outline of the column, with a slightly rounded top.

Figure 4 shows a spout of an irregular outline from the same

whale at closer range. The two harpoon lines by which it is

fast to the vessel are seen at the lower right hand. The top

of the column is of thin vapor and is being wafted away by the

breeze. The lower part of the column is much denser and some-

what in the form of an inverted cone. Possibly the irregular

Shape may be in part due to a slight wave breaking over the

animal's head as it commenced to spout.

The height to which the larger rorquals spout varies consider-

620 THE AMERICAN NATURALIST, (VoL. XXXVIII.

ably according to circumstances. The same individuals are seen

at times to make a low spout and again, one twice or perhaps

thrice as high. Estimates of the height of the column by seem-

ingly reliable persons run from ten feet up to fifty. It is some-

times stated (cf. Beddard, :00, p. 153) that the Sulphur-bottom

whale may be recognized by the great height of its spout as

compared with that of other large species of rorquals. The

writer was unable, however, to distinguish between the spouts

of the Finback (Balenoptera physalus) and the Sulphur-bottom

whale (Balenoptera musculus), nor did the men on the whaling

FIG. 5.— Lancing a spouting sulphur-bottom.

vessel believe that the height of the spout afforded any criterion

for such a distinction. One of the photographs obtained by the

writer, however, affords an opportunity for the direct comparison

of the relative heights of a man and of the spout of a whale.

Figure 5 shows the captain in the act of lancing a 77-foot Sulphur-

bottom which two harpoons had failed to despatch. He stands

in the bottom of the boat, alongside the exhausted animal, and

the spout, extending up to the skyline in the photograph, is one

of average height. The standing height of the man is about

No. 453] OBSERVATIONS ON RORQUALS. 621

5 feet 8 inches, and the height of the spout is 21 times as great,

or about 14 feet. A maximum spout would probably be close

to 20 feet high, which is the estimate I find in my notes taken

at the time.

I had no means of accurately estimating the speed at which

these whales travel through the water, but it not infrequently

happens that the whaler, steaming at ten knots an hour, is un-

able to overtake a free whale even after a considerable chase.

Beddard's statement that the maximum speed of a Sulphur-bot-

tom whale is in the neighborhood of twelve miles an hour is

Fic. 6.— Throat of a finback whale.

therefore probably not far from the truth. The ordinary rate of

swimming, however, is apparently a little less than this.

During my short stay at the Rose-au-Rue station six whales

were taken and all were males. The men told me that at that

season (September) the females seek the shallow and more quiet

waters of the bays to bring forth their young, and their shyness

at this time renders it difficult to approach them. Shortly

before my arrival, at about September 4th, a female Finback

whale (Balenoptera physalus) was killed which contained two

calves nearly ready to be brought forth. They were said to

have been male and female, about twelve feet long, and were

622 THE AMERICAN NATURALIST. [VoL. XXXVIII.

lying side by side in the uterus with the head of one by the tail

of the other. This was the first time that a whale containing

more than a single foetus had been taken by the Company's

steamers.

In addition to the photographs of spouting whales it seems

worth while to introduce one showing the throat folds. These

are usually represented in drawings as simple longitudinal plicze.

Figure 6 represents the ventral side of the throat in a Finback

whale (Balenoptera physalus). The folds are seen to start from

the border of the lips as single plications, but as the expanse of

the throat increases posteriorly they fork dichotomously in a

fairly definite manner, so that the number of folds at a given

part of the center of the throat is greater than that at either

end of the corrugated area. Posteriorly the folds run together

in reverse order, so that a reduction is effected similar to that

found at the anterior region of the throat. Curiously, however,

forking may take place in either direction, so that the two new

branches may point either anteriorly or posteriorly, but the lat-

ter mode of branching was not noticed in the posterior part of

the area covered by the folds. Sometimes, also, two folds run-

ning parallel to each other may be connected by a short cross-

fold, which aids in binding all together.

Up to the time of my visit the whaling steamer Puma, oper-

ating at Chaleur Bay and at Placentia Bay, had taken in 1903

107 Sulphur-bottoms, 66 Finbacks, 14 Humpbacks, and 1 Pol-

lack whale (B. borealis}. The last named was captured in Pla-

centia Bay, as were the four taken in 1902.

No. 453.] OBSERVATIONS ON RORQUALS. 623

BIBLIOGRAPH Y.

VON BAER, K. E.

'64. Noch ein wort über das blasen der Cetaceen, mit bildlichen darstel-

lungen. Bull. de l'Acad. imp. des sci. de St. Pétersbourg, vol. 7,

col. 333—341, 3 text figs.

BEDDARD, F. E.

A Book of Whales. New York. $8vo, illus.

HENKING, H.

:01. Ueber das blasen der wale. Zool. anzeiger, vol. 24, p. 103-111,

text fig. 1-2.

PACKARD, A. S.

'66. List of Vertebrates observed at Okak, Labrador, by Rev. Samuel

Weiz, with Annotations. Proc. Boston, Soc. Nat. Hist, vol. 10,

p. 264-277.

RacovitTza, E. G.

:03. Cétacés. Résultats du voyage du S. Y. Belgica, Zoologie., 142 pp.,

4 pls. Anvers : 4to.

TRUE, F. W.

:08. First Record of the Pollack Whale (Ba/enoptera borealis) in the

Western North Atlantic. Science, n. s., vol. 17, no. 421, p. 150.

TRUE. F. W.

:03a. On some Photographs of Living Finback Whales from Newfound-

land. Smithsonian Miscell. Collections, vol. 45, p. 92-94, pl. 24-

26.

PHYSICAL IMITATIONS OF THE ACTIVITIES OF

AMCEBA.

T H. S. JENNINGS.

PHYSICAL imitations of the activities of lower organisms, such

asare given us by Bütschli and Rhumbler, have always taken

a place among the “startling achievements " of science. They

arouse a lively interest in the popular mind as well as in the

minds of men who are seriously studying the problems which

such activities present. Anything which promises a bridge —

from the inorganic to the organic, from the physical to the vital,

demands attention. Almost all men have définite convictions

as to the relation of these two fields, — convictions which are

foundational for the whole superstructure of intellectual or

religious life; anything which touches these convictions must

awaken interest.

How far have the physical imitations of vital activities gone?

What do they really show as to relation of physical and vital ?

In the present paper such physical imitations as relate to one

of the lowest organisms, Amceba, will be examined with these

questions in view. The greater number of the physical experi-

ments relate directly to Amoeba, attempting to imitate its

behavior. The writer has recently made a thorough re-examina-

tion of the behavior of this animal, the results of which have

been published elsewhere (Jennings, : 04), $0 that opportunity

is presented for a comparison between the imitations and the

reality. By determining to what extent the physical imitations

throw light on the behavior of Amoeba, we shall perhaps have

a fair measure of what has been accomplished in this way, and

of the promise for the future.

What is the real purpose of the physical imitations of vital

activities? Clearly, the final purpose is to show what physical

factors are at work in these activities. But this end may be

followed in many ways; what is the special purpose of the

imitations ?

625

626 THE AMERICAN NATURALIST. [Vou. XXXVIII.

In the best cases the physical imitations arise as follows:

There is first a study of certain vital activities. This is fol-

lowed by construction of a hypothesis as to the nature of the

factors at work, — an explanation of the activities in terms of

phenomena already known. The third step is to determine by

experiment whether the supposed known factors can produce

such activities ; these factors are combined in appropriate ways

and the results observed. If they bring about activities similar

to those shown in the vital phenomena, then the explanation

gains much in probability, and we have an “imitation” of the

vital activities. What the imitation shows is then, as Rhumbler

(98, p. 108) has well said, that the factors assumed to be at

work really can produce such activities as we attribute to them,

— and this is a long step in advance. There still remains the

question whether the factors in our imitation actually ave those

at work in the vital phenomena.

To enable us to. judge intelligently on this final question we

need an accurate knowledge of the phenomena to be explained

and of the forces at work in the imitation, that they may be

closely compared; imitations founded on external resemblance

are likely to be misleading. We have indeed three factors to be

compared, — the explanation as it exists in the mind of the

investigator, the physical experiment, and the vital activity. In

the best cases these three must agree; the explanation fits the

experiment, and the experiment is essentially similar to the vital

phenomenon, so that the explanation fits the latter also. But the

explanation given may fit the physical experiment and not the

vital activity, or it may not even fit the experiment; we shall

find examples of both these cases.

In the commoner case, where the explanation given does fit

the physical experiment, how are we to judge whether the vital

activity is to be similarly explained ? Evidently an explanation

based on an imitation can at best fit the vital activity only in so

far as the latter agrees with the imitation. Points in which it

does not agree must be attributed to other factors, and if these

points are essential ones for the explanation given, then we must

conclude that the vital activity is not explicable in the way pro-

posed. Further, we must determine whether certain conditions,

No. 453.] ACTIVITIES OF AMQEBA. 627

preceding or following, which the explanation requires are actu-

ally fulfilled in the vital phenomena.

Imitations of the movements and of the variations in form

have been oftenest attempted. Almost without exception the

imitations are based on the hypothesis that these phenomena in

Amoeba are due to local changes in the surface tension of a fluid

mass. Among the earliest experiments of this sort were those

of Gad (78). Gad placed drops of rancid oils (oils containing

fatty acids) in weak solutions of alkali; for example, cod liver oil

in 0.2 to 0.5 % sodium carbonate. As. a result of the reaction

between the fatty acid and the alkali soap is produced. This

lowers the surface tension of the drop of oil here and there; as

a result the drop changes form, sending out projections having

an external resemblance to the pseudopodia of Amoeba. A

number of figures showing the forms taken by oil drops under

these conditions are given in Verworn’s General Physiology.

Gad pointed out the resemblance of these forms to those shown

by Amoeba, but did not carry the matter farther.

Quincke ('79, '88) pursued further the study of move-

ments caused in the manner just described, and put forth dis-

tinctly the view that the movements of Amoeba (as well as of

other protoplasmic masses) are due to similar causes. Quincke

found that egg albumen might take the place of the sodium

carbonate in the experiments above described ; soap is then

formed and movements occur as when the alkali is used. He

held that Amceba is covered externally by a thin lamella of oil ;.

that albuminous soaps are formed on the inner surface of this,

thus decreasing the surface tension, and that the movements

and changes of form are due to these changes in surface tension.

Most celebrated of all imitations of amoeboid movements are

those of Bütschli ('9o, '92). Bütschli mixed slightly damp,

powdered potassium carbonate with old olive oil, of a certain

degree of rancidity, and brought drops of the mixture into water

on a slide. (Directions in Bütschli, '9o.) After standing

twenty-four hours the drops are washed and new water or glyc-

erine supplied. The drops now show streaming movements,

send forth projections (see Fig. 1, 4), and move about. The

external resemblance to the phenomena shown in Amoeba is.

628 THE AMERICAN NATURALIST. [Vor. XXXVIII.

very striking. The movements are caused as follows: The

potassium carbonate is dissolved by the water and acts on the

oil, forming soap. Thus after a time the oil drop is permeated

throughout by minute globules of soapy water, forming a foam-

like emulsion. At times one of these globules of soap bursts

on the outside of the drop of oil; the soap then spreads over

the surface of the oil, lowering its surface tension in the region

affected. At once a projection is formed here, currents flow

from within the drop toward the region of lowered tension, and

the entire drop may move in that direction.

Bütschli held that the movements of Ameeba take place in a

similar manner. He considers that protoplasm has an emulsion

structure similar in a general way to that of the oil drops, —

though of course the constituents are not the same. At times

the meshwork enclosing the globules breaks at the outer surface

of the Amoeba, allowing some of the enclosed fluid to spread

over the surface. This lowers the surface tension, causing

Amoeba to move in the same manner as the drop of oil.

Bütschli is inclined to attach much significance to the fact

that the oil drops which move in the way described have a foam-

like emulsion structure, and to consider this as a support to his

view that the similarly moving protoplasm is similarly consti-

tuted. But such movements are by no means specially charac- .

teristic of fluids having a foam-like or emulsion structure; many

drops having this structure do not show the movements, while

other drops which have not this structure show the movements

equally well, as we shall see. The movements require only that

there shall be some method of producing local changes in sur-

face tension; this may be easily brought about without the

emulsion structure.

Bernstein (:00) produced similar movements in drops of

mercury. Sufficient mercury to make a drop or disk five to ten

millimetres in diameter is placed in a flat-bottomed watch-glass.

Over it is poured some 20% nitric acid, and thereto is added a

quantity of a strong solution of potassium bichromate. The

mixture acts chemically on the mercury, lowering its surface

tension. The intensity of the action varies locally, so that the

surface tension is decreased now here, now there. As a result

No. 453.] ACTIVITIES OF AMGBA. 629

the mercury moves and changes form in a striking manner,

sending out projections or becoming wholly irregular, at the

same time moving from place to place.!

The present author (:02) has given another method of

observing such movements. A mixture of three parts glycerine

and one part 95 % alcohol is placed on a slide and covered with

a large cover-glass, supported near its ends by glass rods. Be-

neath the cover-glass a drop of clove oil is introduced by means

of a medicine dropper drawn to a fine point. The alcohol acts

locally on the surface of the clove oil, decreasing its surface ten-

sion here and there. As a result the clove oil drop changes

form, sends out projections and moves from place in a strik-

ing manner. The phenomena shown are similar to those in

Biitschli’s drops of oil emulsion. The experiments are much

easier to perform than those of Biitschli; by varying slightly

the amount of alcohol in the mixture one can always be certain

of getting marked results. But the movements do not continue

so long as in Biitschli’s experiments.

In all these experiments the movements are due to local

changes in surface tension. When such a local change is pro-

duced on the surface of a fluid drop a characteristic set of cur-

rents results. From the region of least tension surface cur-

1 The attempts of Herrera to imitate protoplasmic movements read almost

ike a traves sty of those of the authors above mentioned. Herrera made a

showed to exist in the npa of € one of the vnde pit

one well co usion currents. Herrera considers these as a

: diaithfal reproduction of the internal movements

is Tieghem." In a later contribution Herrera ('98a) gi

mæboid motion based on the theory that Ameeba is iive by the bubbles ng

pi dioxide which it gives off in its respiration. Mix bicarbonate of

printer’s ink so that a product is obtained having a sirupy consistency. Place

surface wet with a weak solution of tartaric acid. Bubbles of carbon dioxide

to change form and move ; “ the illu-

.” It is only just to say that Herrera later

gave up the idea that the movements of Amoeba are caused in this manner

630 THE AMERICAN NATURALIST. [Vor. XXXVIII.

rents pass in all directions, while an interior current passes

toward the region of least tension. The reason for these cur-

rents may be seen by imagining that the drop is covered with a

stretched India rubber membrane in place of the surface film.

If this stretched membrane is weakened or cut at a certain

point the remainder of the membrane will pull away from this

point, simulating the surface current. At the same time fluid

from within will be pressed out at the weakened point, — thus

simulating the central current toward the point of least tension.

IG. 1 «c.

gus 1.— Currents produced by local decrease of surface tension, after Bütschli. a, Currents

in an oil drop when the surface tension is decreased at one end by contact with a soap

solution (s); surface currents away from the point of lowered tension; a central current

TA this point. 2, One of the drops of oil emulsion, showing the irregular form and

ec ti i t th E £ } : :

projection.

The characteristic currents may be seen in Bütschli's experi-

ments or in those with the drops of clove oil, if some soot or

India ink has been mixed with the oil. Such currents are repre-

sented in Fig. 1, taken from Bütschli. If the axial current

carries forward more fluid than the superficial currents carry

backward, the drop may elongate in the direction of the axial

current and move as a whole in the same direction. This often

occurs.

Such currents as are shown in Figure 1 are an invariable

feature of movements of fluids due to local decrease in surface

tension. Indeed, these currents are the characteristic phenom-

ena ; they may be the only movements that occur.

No. 453.] ACTIVITIES OF AMGBA., 631

If, then, the movements of Amoeba are really produced as

they are in the imitations, by means of local changes in surface

tension, we must expect to find in Amoeba these characteristic

currents. In an Amoeba moving in a certain direction there

should be a central current forward and superficial currents

backward. In an extending pseudopodium the central current

should be toward the point, the superficial currents away from

it. Do such currents exist ?

There is evidently a central current forward. But are the

superficial currents backward, as the theory requires? In

studying the movements from

above, without the aid of experi-

ment, it is difficult to determine

this point. But there are certain 4— g

appearances on the lower surface

and at the lateral margins which

give the impression that such back-

ward currents may exist. In fact

Biitschli, Rhumbler and others be-

came convinced of the existence b

of such currents. The movements

of Amceba were thus brought into

full agreement with those of the

drops moving as a result of local

decrease in surface tension. This

is brought out clearly by an exam- ing Ameba, according to Bütschli and

ination of the figures of the Cur- ,, seen from above, ns haute’

rents in Amceba given by Bütschli sdb endo i pepo sana eet

and -Rhumbler, copied in Figure 2. rents in an advancing peeudopodiam,

It was then almost inevitable to — *'^ Korm

conclude that the same causes are at work in the two cases;

that the movements of Amoeba are due to local changes in sur-

face tension.

In the extended experimental study of the activities of

Amaeba recently made by the present writer (:04), it was

shown that the supposed backward currents of the surface do

not exist. On the contrary, all parts of the surface which are

not attached to the substratum are typically moving forward, in

632 THE AMERICAN NATURALIST. [Vor. XXXVIII.

the same direction as the central current, while the attached

parts of the surface are at rest. The movement of Amoeba is

thus of a rolling character; the upper surface continually passes

around the anterior end to form the lower surface; this then

remains quiet until it is taken up by the posterior end as the lat-

ter moves forward. The movements in an advancing Amoeba

are indicated in Figure 3. -In a projecting pseudopodium the

movements are of the same character as those at the anterior

end (Fig. 3), save when the pseudopodium projects freely into

Fic. 3.— Diagra

show the direction of the currents; the longer arrows indicating more rapid movement.

he large arrow above shows the a of locomotion. The anterior end (A) is thin

and attached to the substratum as

m of the movements of an Amceba in locomotion, side view. The arrows

back ower surface from a to x is at

The posterior end mld is high a rounded, and free from the substratum a, 5,c

successive positions occupied by the anterior end. The broken outline shows the position

occupied by the Amceba a e later.

the water, being nowhere in contact with a solid. In the latter

case the entire surface moves outward, in the same direction as

the tip.

Details of the observations and experiments which demon-

strate the movements to be of the character just set forth are

given in an extensive paper published elsewhere (Jennings, : 04).

The movements were determined chiefly by observing the motion

of objects attached to the outer surface of Amoeba, of objects

partly imbedded in the outer layer, and of particles within the

body. The movements as thus studied are clear, and exclude

the possibility of the typical existence of backward currents on

the surface.

It appears then that Amoeba does not move in the same man-

ner as do the imitations based on local changes in the surface

tension of a fluid mass. The currents which form the charac-

teristic features in the latter case are not present in Ameeba.

No. 453.] ACTIVITIES OF AMGBA. 633

Neither theoretically nor practically does there appear to be any

evidence that movements due to changes in surface tension can

take place without these characteristic currents. We cannot then

consider the movements of Amæœba to be due to a decrease in

surface tension at the anterior end, as in the “imitations.” In

precisely the feature which led tothe supposition that the move-

ments in the two cases were of the same character we find that

there is actually an absolute contrast. In Ameeba the surface

currents are in the direction of movement of the mass, and in the

same direction as the central current; in the imitations they

are in the opposite direction.

Clearly the surface tension theory will not account for the

phenomena as they actually exist. This becomes still more evi-

dent when we consider the formation of pseudopodia not in

contact. In these there is not only no backward current, but

also no resting surface; axis and surface move outward in the

same direction as the tip. Such movements are not producible

by local changes in surface tension. The * imitations " are imi-

tations only to the extent that they are fluids and that they move ;

they are not imitations so far as the nature of the movements

and their cause is concerned.

A much more nearly accurate imitation of the movements of

Amoeba may be produced with gravity as the active agent in

place of surface tension. A drop of water moving down hill on

a surface to which it does not cling strongly shows the same

rolling movement that we find in Amoeba. The lower surface

(in contact with the substratum) is at rest, while the upper sur-

face moves forward and passes continually around the anterior

end to the lower surface. But we know that gravity is not the

active agent in the movement of Ameeba.

An imitation of the usual locomotion of Amoeba that is accu-

rate even to minute details is described by the present author in

the paper on the behavior of Ameeba already cited (Jennings

:04). A drop of fluid resting on a substratum is caused to

adhere to the substratum more strongly at one edge than at the

other. Thereupon the drop moves toward the more adherent

edge, and in so doing it shows exactly the form and movements

of an Amceba in locomotion. The experiments may best be

634 THE AMERICAN NATURALIST. [Vor. XXXVIII.

performed as follows: A piece of smooth cardboard, such as

the Bristol board used for drawing, is placed in the bottom of a

flat dish and on a certain spot on the cardboard is placed a drop

of water. The whole is then covered with bone oil. This soaks

into the cardboard, except where the latter is protected by the

drop of water. After the board is well soaked in oil the drop

of wateris removed, leaving the whole surface covered with oil

some millimeters deep. Now a drop of water or glycerine, to

which has been added some fine soot, is placed on the cardboard

under the oil. This drop is allowed to come in contact by one

edge with the area which had been protected from the oil. To

this area it adheres, the edge in contact spreads out as a thin

sheet, and the rest of the drop is pulled over to. the area. Its

movement is then exactly that typical for a flowing Amoeba, so

that Figure 3 would do equally well for a diagram of the move-

ments of such a drop as for those of Amoeba. The resemblance

extends to minute details ; many of these are set forth in the

author's paper above cited (:04). Among other things, the

formation of pseudopodia in contact with the substratum may

be imitated by making the area to which the drop adheres at one

edge very small; thena projection is formed merely of the width

of this area.

But this imitation, like the others, fails when we take into

consideration the formation of pseudopodia which are nowhere

in contact with a solid. Projections corresponding to these can-

not be formed in the physical experiments just described, for in

these adherence to a solid is the essential point. Since the

entire anterior end of the Amceba can be pushed out into the

free water, we find that Amceba can perform all the active

operations concerned in locomotion without adherence to a

solid. This effectually blocks any attempt to explain the move-

ments ot Amoeba as due, like those of the drops in the exper-

iments just described, to one-sided adherence to the substratum.

Thus none of the physical imitations gives us a clue to the

physical agent actually at work in the movements of Amoeba.

VR a last described are perhaps useful in giving us

; e direction of action of the forces at work in pro-

ducing locomotion. Not even so much as this can be said of

No. 453.] ACTIVITIES OF AMGBA. 635

the surface tension experiments; the direction of action of

the forces in these is evidently different from that in Amoeba.

We may then turn to imitations of other activities of Amoeba.

Many attempts have been made to imitate certain of the reac-

tions to stimuli — particularly the positive reaction to chemicals.

Such imitations depend on the fact that a local decrease in the

surface tension of a drop of fluid mày be caused by contact

with a chemical; the drop then moves in the direction of low-

ered tension. Some of the experiments based on this are the

following :

Rhumbler (99, p. 585) placed a small drop (60 to 90 yw in

diameter) of castor oil in alcohól, and brought close to it the

open end of a capillary tube containing clove oil, chloroform, or

5 % potassium hydroxide. The substance within the tube dif-

fused out against the dropof castor oil and decreased its surface

tension in the region of contact. Thereupon the usual currents

were produced (Fig. 1), and the drop moved in the direction of

lowered tension, finally entering the tube.

Bernstein (: 00) placed a drop of mercury in twenty per cent.

nitric acid, then brought near it a crystal of potassium bichro-

mate. By the chemical action the surface tension on the side of

the drop next to the crystal is decreased. Thereupon the drop

moves rapidly over to the crystal, and may push it about from

place to place.

In the drops of clove oil in a mixture of glycerine and alcohol,

described above (p. 8), similar movements may be caused (Jen-

nings :02). With a capillary pipette a little alcohol is brought

near one side of the drop. This decreases the surface tension

of the part affected; thereupon a projection is sent out toward

the alcohol, and the drop as a whole moves toward it. If the

drop is heated at one edge, by touching the cover glass near it

with a hot wire, the clove oil moves toward the heated side, and

may be induced to follow the wire for some distance.

In all these experiments the movement is due to local altera-

tions in surface tension; the drop moves toward the region of

lowest tension ; there is a central current in the direction of loco-

motion, and surface currents in the opposite direction. In

Amoeba, on the other hand, as we have seen, the movements

636 THE AMERICAN NATURALIST. [Vor. XXXVIII.

cannot be considered due to local decrease in surface tension.

There are no superficial currents away from the region toward

which the animal moves, but all parts that are in motion move

toward the object causing the reaction. (For details, see Jen-

nings, :04.) The experiments do not imitate the essential

features of the action of Amoeba, and do not show us the

causes at work in its behavior. The reactions of Amoeba are

not simple direct results of the physical action of the agents

producing them, but are indirect, like those of higher ani-

mals.

Many imitations have been devised for the taking of food by

Amoeba. Rhumbler ('98) holds that the ingestion of food by

Ameeba is due to physical adhesion between the liquid proto-

plasm and the solid food. He shows that drops of all sorts of

fluids take in certain solids in this manner. A drop of water

placed at the edge of a plate of glass draws to itself and envel-

opes splinters of wood and various other solids which come in

contact with it. Glycerine, oils, white of egg, gum arabic, mas-

tax varnish, etc., are shown to do the same. A convenient way

of showing this is to fill a capillary glass tube with the fluid,

then to bring a small piece of the solid in contact with the free

surface of the liquid at the end of the tube. The pulling of

the solid into the liquid is due to the adhesion of the two, in

connection with the surface tension of the liquid.

These experiments of Rhumbler show that food might be

taken in this manner, not that it is so taken. Careful study

shows that there is in most species of Amoeba no adhesion

between the protoplasm and the food body. Food is taken by

actively enclosing it along with a small quantity of water; the

fact that no adhesion exists between it and the protoplasm is.

strikingly evident, and occasions much difficulty in the ingestion

of food. (For details, see Jennings, : 04, and compare the simi-

lar account of food-taking by Le Dantec, ’94.) Thus the

hi quietam do not really imitate the essential features of the

behavior in Amoaba. Only in Amoeba verrucosa and its close

relatives is there evidence of adhesion between the animal and

its food. But even here there is adhesion equally to bodies

which do not serve as food and are not ingested, so that for the.

ingestion itself an additional factor is necessary.

No. 453.] ACTIVITIES OF AMGBA. 637

One of Rhumbler’s most striking experiments is an imitation

of the method by which Amoeba takes as food a long filament

of Oscillaria, coiling it up and enclosing it. The Amoeba set-

tles down somewhere along the filament, lengthens out upon it,

and bends it over, forming a loop. This process is repeated

until the long filament forms a close coil within the Amoeba

(figures in Rhumbler, 1898, p. 211, Lang, :01, p. 39; a simi-

lar account with figure in Leidy, '79, p. 86). Rhumbler con-

siders this remarkable process to be brought about as follows :

The Amceba adheres to the filament. It lengthens out along it,

just as a drop of water lengthens out along a filament to which

it adheres. Owing to the surface tension of the fluid proto-

plasm, impelling it to take the spherical form, it pulls on the

two halves of the filament, producing a thrust inward from both

directions. Gradually the enclosed parts of the filament are

softened in the digestive processes of the Amoeba. The soft-

ened portion then yields to the thrust from both directions

and bends, so that more of the filament can be pulled into the

Ameeba by the tension of its surface film. The Amoeba then

lengthens out farther, owing to adhesion ; more of the filament

is softened and yields farther, so that more is pulled in by sur-

face tension. This process continues until the filament is com-

pletely coiled up and enclosed.

On the basis of this explanation Rhumbler devised an imita-

tion of the process. A chloroform drop is placed in the bottom

of a watch-glass of water. A long fine thread of shellac,

` obtained by heating two pieces of shellac in contact over a flame

and rapidly pulling them apart, is brought in contact with the

drop. The latter envelopes the filament in some portion of its

length, then proceeds to coil it up, as Amoeba does with the

Oscillaria filament ; after a time the shellac thread is completely

enclosed within the chloroform drop. The mechanism of the

process is conceived to be the same as that above given for

Ameeba and the Alga filament.

This experiment is an interesting example of one of the

numerous difficulties which beset the worker along such lines,

— of the fact, namely, that even the imitation may not agree

with the explanation given. The coiling up of the shellac

638 ., THE AMERICAN NATURALIST. (VoL. XXXVIII.

thread by the chloroform is not explicable in the manner sup-

posed by Rhumbler ; the surface tension of the drop has really

nothing to do with it. This is shown by the fact that such

a thread of shellac is coiled up in exactly the same manner if

submerged in a large vessel of chloroform, so that it is nowhere

in contact with the surface film. The coiling up is apparently

due to strains within the shellac filament, produced when it was

pulled out, and to the adhesiveness of its surface when wet with

chloroform. There are no corresponding factors in the Oscil-

laria thread ; this will indeed, as Rhumbler has shown, straighten

out again when released by the Amceba. The process by which

Amoeba coils up the Oscillaria filament must thus be of an

essentially different character from that occurring in the experi-

ment. The explanation given by Rhumbler may of course still

be correct for the process in Amceba, though it is not correct

for his imitation of the process.

Ameeba does not ingest every small object with which it

comes in contact, but exercises an evident choice as to the

substances which it takes as food. Physical explanations and

imitations of such choice have been given. We may notice

especially those set forth by the present author (: 02) in exten-

sion of certain experiments of Rhumbler. A drop of chloroform

is placed in the bottom of a watch-glass of water, and with fine

tweezers pieces of various substances are brought in contact

with its surface. Some are at once taken in; others are not,

or are thrown out if forced into the drop. Glass, sand, dirt,

wood, gum Arabic, and chlorate of potash are rejected; shellac,

paraffin, styrax, and hard Canada balsam are accepted. The

selection or rejection depends upon the relative amount of adhe-

sion between the solid object on the one hand and the chloro-

form and water on the other. Those which adhere more strongly

to the chloroform than to the water are taken in; others are

rejected.

These experiments show how choice might occur in an organ-

ism; they do not show how it actually occurs in Amoeba.

Food-taking is usually, as we have seen, not accompanied by

adhesion between Amceba and the food, so that choice of food

cannot be explained as due to the fact that some substances

adhere while others do not.

No. 453.] ACTIVITIES OF AMGBA. 639

Rhumbler ('98) has given a physical imitation of the taking

in of a food body and of later giving off the undigested residue

(defecation). A rod: of glass covered with a thin layer of shellac

is taken in by a drop of chloroform (as a result of adhesion).

The shellac is dissolved off by the chloroform and the glass rod

is then thrown out, since the chloroform does not adhere to it.

This imitation, like the others, loses much of its force in view

of the fact that food-taking is not usually due to adhesion and

that substances which do not adhere are taken as food ; defeca-

tion cannot then be explained as due simply to lack of adhesion.

In all the imitations thus far we find that the physical factors

at work cannot be considered the same as those that are acting

in Amoeba. The imitations are such only in purely external

features. There exist certain imitations, however, in which this

has not been proved to be the case. Thus, Rhumbler (’98)

found that when chloroform drops are placed in water, the water

gradually passes into the chloroform, collecting in minute glob-

ules, which later gather in a larger drop. This larger drop is

finally given off to the outside. This process Rhumbler con-

siders analogous to the formation and discharge of the contractile

vacuole in Amoeba. The present author (:04) has described

imitations of certain movements of the pseudopodia in Amoeeba,

produced in liquids partly covered with a solid layer; these are

hardly of sufficient general interest to be detailed here. The

most striking experiments which can still be considered with

some degree of probability to indicate the factors really at work

in certain processes occurring in the Rhizopoda are undoubtedly

Rhumbler’s imitations of the production of Difflugia shells.

Since these deal with an organism closely related to Amoeba,

they may be described here.

The experiments may be performed as follows: Chloroform

is rubbed up with fragments of glass in a mortar until the glass

is reduced to the finest dust. Then with a pipette drawn out to

a small point drops of this mixture of chloroform and glass are

injected into water. At once the grains of glass come to the

surface of the drops so formed and arrange themselves in a

single layer, without chinks or crevices, exactly as in the shell

of Difflugia. The chloroform drop is thus covered with a shell

640 THE AMERICAN NATURALIST. [VorL. XXXVIII.

having a striking resemblance to that of Difflugia. In place of

chloroform, linseed oil or other oils may be used. The drops

must then be injected into 70 % alcohol, since the oil would float

on water.

The factors at work in the formation of the “artificial shells”

are diffusion currents within the chloroform, the adhesion of the

bits of glass to its surface, and the action of surface tension in

arranging and fitting together the bits of glass. Studies of the

process by which the shell of Difflugia is formed at the time of

division of the animal seem to indicate that the same factors

may be at work in the living organism. (See Rhumbler, '98,

p. 289.)

Reviewing our results, we find that few of the experimental

imitations of the activities of Amoeba stand before a critical

comparison with what actually takes-place in the animal. Such

comparison shows in almost every case that the factors at work

in the imitations are essentially different from those acting in

Amoeba. In particular, almost all the imitations based on local

changes in surface tension break down completely.

What are we to conclude from this fact as to the part played

by surface tension in vital phenomena? The tendency has been

of late to attribute more and more of a róle among life processes

to surface tension. Amoeba has been the chief place where the

important part played by surface tension seemed really demon-

strable; the movements, the reactions to stimuli, the taking of

food, and the choice of food, were all attributed to this and

closely related factors; With the demonstration of the complete

failure of surface tension to account for the phenomena that

were chiefly relied on to prove its importance, the supposition

that it plays an immensely important róle in life processes loses

much of its weight. Surface tension may of course, in a more

refined way than was supposed for Amoeba, still play the large

róle in vital phenomena that some attribute to it. In the

meshes of Bütschli's protoplasmic meshwork, or in the muscle

fibrillae (Bernstein), it may perhaps do what is demanded of it.

Possibly the study of surface tension is still the most promising

field for detection of the physical factors underlying life proc-

esses. But the surface tension theory must come to us shorn

No. 453.] ACTIVITIES OF AMQEBA. 641

of the trophies of its prowess, — its supposed full explanation of

most of the activities of Amoeba, — and bearing instead the

record of a complete defeat.

What positive results of value have the physical imitations of

vital activities in Amoeba to show? As we have seen, there are

still two or three of these that may really give us a clue to the

factors at work in the vital processes; at least this has not yet

been disproved. Beyond this the positive results are of a very

general character. The imitations show that a drop of fluid

might, through physical factors, show locomotion, move toward

certain agents and away from others, and exhibit choice in the

taking in of certain substances and the rejection of others. But

they do not show specifically through what physical factors the

activities are as a matter of fact brought about in Amoeba or any

other particular organism.

The chief value of most of the attempted physical imitations

is that of a trial. The method of trial and error is a method of

progress in science as elsewhere. In these imitations a definite

explanation of the phenomena is put on trial. The “trial” con-

sists in a more careful study of the phenomena in question ; it

is as an inspiration to such study that the imitations are of great

value. If as a result the explanation given is recognized as

*error, that is in itself an advance; this particular trial will

not need to be made again. Continued application of this

method of trial and error must result finally either in the dis-

covery of the real factors at work, or in the recognition that we

are dealing with a new class of factors not found in physics.

UNIVERSITY OF PENNSYLVANIA.

BIBLIOGRAPHY.

BERNSTEIN, J. NE

1900. Chemotropische Bewegungen eines Quicksilbertropfens. Arch.

f. d. ges. Physiol., Bd. 80, pp. 628—637.

BUTscHLI, O. :

'00. Experimental Imitations of Protoplasmic Movement. Quart.

Journ. Micr. Sci., vol. 31, pp. 99-103.

642 THE AMERICAN NATURALIST. [Vor. XXXVIII.

'92. Untersuchungen über mikroskopische Schäume und das Proto.

plasma. Leipzig. 234 pp., 6 plates.

GAD, JOHANNES.

"78. Zur Lehre von der Fettresorption. DuBois Reymond's Arch. f.

d. Physiol., Jhrg. 1878. pp. 181-205. Taf. 2.

HERRERA, A. L.

Sur le protoplasma synthétique et la force vitale. Bull Soc.

Zool. France, vol. 23, pp. 118—120.

'98a. Sur la manière de produire certains mouvements amiboides par

un dégagement d'acide carbonique. Bull. Soc. Zool. F rance, vol.

23, pp. 128-130.

JENNINGS, H. S.

Artificial Imitations of Protoplasmic. Activities and Methods of

demonstrating them. Journ. Appl. Micr. a. Lab. Methods, vol. 5,

Pp. 1597-1602.

:04. The Movements and Reactions of Amceba. Contributions to

the Study of the Behavior of the Lower Organisms, No. 6

Carnegie Institution of Washington, D. C., Publication No. 16.

LANG, A.

:01. Lehrbuch der vergleichenden Anatomie der wirbellosen Thiere.

Zweite Auflage, zweite Lieferung: Protozoa. Jena. 311 pp.

LE DANTEC, F.

'94. Études biologiques comparatives sur les Rhizopodes lobés et

réticulés d'eau douce. Bull. Scient. de la France et de la Bel-

gique, T. 26, PP. 56-99.

LEIDY, J.

"79. Fresh-water Rhizopods of North America. Rep. U. S. Geol

Survey of the Territories, vol. 12, 234 pp., 48 pl.

QUINCKE, G

Ueber Emulsionsbildung und den Einfluss der Galle bei der

Verdauung. Ar

'80. Ueber periodisch

RHUMBLER, L;

'98. Physikalische Anal

Bewegung,

und Geháu

yse von Bebenserscheinungen der Zelle. I.

Nahrungsaufnahme, Defákation, Vacuolen-Pulsation

sebau bei lobosen Rhizopoden. Arch. f. Entw.-Mech.,

Bd. 7, pp. 103-350.

99. Allgemeine Zellmechanik. Merkel und Bonnet’s Ergebnisse der

Anatomie und Entwicklungsgeschichte, Bd. 8, pp. 543-625.

THE INFLUENCE OF THE MUTATIONS OF THE

PLEISTOCENE LAKES UPON THE PRESENT

DISTRIBUTION OF CICINDELA.

H. F. WICKHAM.

Wrrz the propagation of the late theories of life-zones founded

upon temperature conditions, has come about a neglect of those

considerations, of a more obscure though not less important

nature, which we must recognize if we are to attain anything

more than a superficial understanding of geographical distribu-

tion. The zonal theory as expounded by some of its more stead- .

fast adherents has the apparent advantage of simplicity and

moreover appeals directly to the ‘ practical man ' since it is with-

out doubt correctly assumed that most of our cultivated plants

as well as numerous wild ones are limited to certain belts which

are more or less closely coincident with the isotherms. The

theory works well with agriculture in general, but it only

imperfectly expresses the truth if we apply it to the natural

distribution of Coleoptera on this continent. By natural, I mean

original, in the sense of not being modified through the agency

of civilized man ; for cultivation and settlement have been potent

factors in changing the range of numerous species, often

resulting in the rapid extermination of the most characteristic

types of a district and their replacement by others, Barriers

which a few years ago were amply sufficient to prevent the inter-

mingling of life of two nearby regions become no longer effective

and the Faunz and the Flore become contaminated by the

encroachment of strangers. Irrigation changes the nature of

entire counties, lakes are drained, mountains are denuded of their

forests and can no longer support the life which has been theirs

for thousands of quiet years.

While it has been known for many years that the geological

history of a given region has exercised a profound influence upon

the present Fauna and Flora; while the effects of the Glacial

643

644 THE AMERICAN NATURALIST. (Vou. XXXVIII.

Epoch upon the life of this and other countries. have long been

recognized and carefully studied so that numerous distributional

phenomena due thereto have thus been referred to their proper

cause ; and although modern lines of insect migration are being

mapped and their relative importance determined, there is never-

theless a vast amount of detail to be worked out in order that

certain local problems may be understood. It is with one of

these local problems that we have now to do.

Some years ago, as the result of several seasons' collecting in

the Great Basin, I became convinced that the distribution of

certain species of Coleoptera: therein occurring could not be

accounted for by the theory of zonal arrangement of life, but

apparently had as ultimate cause some condition or combination

of conditions which belonged to the geological rather than the

present history of the area under discussion. With the aim of

testing this belief, further collections were made and all available

data bearing upon the matter were collated, in order that it

might be seen whether or not the above conclusion rested upon

tangible grounds.

Since none of the rivers rising within the basin empty into

bodies of water outside the limits, one of the most common

methods of introduction of extra-limital species — by migration

along the line of a water-course — is eliminated, and the fauna

1S not exposed to this source of modification. The two great

"western rims have

The southern boundary is less

» SO that those Species which are suited to desert

that quarter, while the northern

hills that mark the divide between

No. 453] DISTRIBUTION OF CICINDELA. 645

We have here, then, a great enclosed region which, though :

comparatively open to migrations from the north and south, is

nearly closed against encroachment from the east and west

except in those cases where man may be conceded to have been

a factor. We should expect the more characteristic forms to

show extensive north and south distribution or that they may be

confined to the basin and the more accessible adjoining areas.

Some species no doubt originated, as such, within the limits of

the basin proper, and I believe that, in some cases at least, we

can determine which these are. It is with certain forms of this

nature — that is to say with true indigenes — that we have now

to deal.

Two types of littoral beetles may be said to be very character-

istic of the Great Basin and to be dependent upon the peculiar

conditions that occur there in the way of saline and alkaline flats

in connection with springs and lakes. These are the Cicindelz

of the echo type and the species or subspecies of Tanarthrus

which belong with T. salicola. Neither of these genera is con-

fined to the basin, Cicindela being of wide distribution and

evidently of southern origin, while Tanarthrus is not known out-

side of the southwestern United States. Besides the forms of

this latter genus described from the lake shores of the Great

Basin, a few species of somewhat different aspect are known to

occur in saline spots in California and Arizona. The species of

Bembidium of the %enshawi type have also, in my opinion,

attained their present specific structures within the limits of the

Great Basin and are not to be considered migrants from the

outside. For the present, I prefer to leave out of the discussion

all of those Coleoptera not directly connected with the existence

of alkaline and saline lakes, since the problem of their dispersal

or distribution is different, in some respects, from that con-

cerning the littoral forms and needs a separate body.

I am prepared to go farther than the simple statement that

we can correctly indicate certain species as having arisen, as

such, within the limits of the Great Basin. I believe it is also

possible to show that some of these have been inhabitants of the

region for long periods of time, and that in the course of their

existence they have been played upon by conditions which arose

646 THE AMERICAN NATURALIST. [Vor. XXXVIII.

as the consequence of geological changes ; that we can point out

the modifications which have taken place in the species and, to a

considerable extent, we can trace the geological phenomena

which are the fundamental or underlying causes of the modifica-

tions. Of course not all species will be equally affected by

changes in their environment, nor will the modifications neces-

sarily be parallel. Minute differences in the organic structures

of insects indicate a probable diversity of physiological charac-

ters, and varying physiological activities may well modify such

details as patterns of coloration or even the colors themselves.

As an example of an insect which has, in all probability, had

its range determined and its specific characters modified by a

series of geological changes, the history of which is not too

remote and therefore fairly well known, we may take that aggre-

gate of forms of Cicindela described under the names C. echo

Casey and C. pseudosenilis W. Horn. These are without doubt

modifications of one type; in fact they are so closely related as

to be separable only in series. They are also very close to e

willistoni, and are ranked as races thereof by Dr. Walther Horn.

However, the exact status of these names, specific or subspecific,

does not concern us at present; the fact remains that these

forms are closely related and may well have come from a com-

mon stock. Cicindela fulgida, to which willistoni was formerly

referred as a variety or race, is readily separable by the thickly

pn front. It may perhaps be an older offspring of the same

The Ways in which the beetles differ from one another may

be briefly outlined, in order that the reader may understand the

relationships and the better appreciate the account of the varia-

ker - each. It must be borne in mind that the descriptions

-— especially to series of specimens than to individuals,

unless definitely stated to the contrary.

a e a Easily separated from the allied spe-

CONEY Ted, di ye t ickly hairy. The color is usually more

' 8^ specimens occur at Lincoln, Nebraska, in

bem us sround color is nearly black. Markings moderate,

ow as a rule, the mi

margin (or € middle band not expanded along the

but very slightly so), the humeral lunule very oblique

No. 453.] DISTRIBUTION OF CICINDELA. 647

behind. This species belongs especially to the broad strip of

plains lying to the eastward of the Rocky Mountains. It is

known from Wyoming, Colorado, Kansas, Nebraska and New

Mexico, but has not been reported from Central America nor

from Old Mexico.!

Cicindela willistoni Leconte. A little stouter than C. fulgida,

the color much less metallic. The front is sparsely haired ; these

hairs are readily lost, but the group of large punctures from

which they spring may always be made out with little difficulty.

Markings broader, the humeral lunule and median band always

united along the margin ; the apical lunule, also, usually, but not

always, connected on the outer edge with the expanded marginal

portion of the middle band. The tip of the humeral lunule (in

those specimens in which it is free) is much less oblique than in

C. fulgida. This insect is known only from the beaches of

small lakes near Medicine Bow, Wyoming, where it was taken

first by Dr. Williston, next by Mr. Warren Knaus and lastly by

myself. I am not certain which of the little lakes was visited

by Dr. Williston. He gives the locality Como Lake, but the

usage of the village is not uniform and it may have been any

one of three small lakes which lie within about twenty miles of

one another. Mr. Knaus and I visited the one near the old sta-

tion of Aurora, now several miles from the track. He calls it

Como Lake,? while I have followed the prevailing custom of the

villagers in speaking of it as Aurora Lake.

Cicindela echo Casey. Form of body nearly that of C. wil-

listoni. Color brownish with an obscure purplish or coppery

overcast. Front very sparsely hairy, the hairs being so readily

removed that most specimens show only the punctures whence

they originate. Markings usually rather narrow, the humeral

lunule not or but slightly oblique at tip, middle band attaining

the margin, usually but slightly dilated thereon but occasionally

reaching the humeral lunule though not quite attaining the

apical one which is complete. My series from Great Salt Lake

(the original locality), most of which I collected at Saltair, in

1 It is not included in Dr. Walther Horn's List of the Cicindelidæ of Mexico,

Jour. N. Y. Ento. Soc., Vol. 11, p. 213.

? Entomological News, Vol. 13, p. 147-

648 THE AMERICAN NATURALIST. [Vor. XXXVIII.

June, shows quite a good deal of variation in the markings, the

changes chiefly affecting the middle band. The marginal por-

tion of this band is variously developed so that it may reach

quite to the humeral lunule and almost to the apical one, or it

may not be expanded at all. The discal part may be quite rec-

tangularly or very obliquely bent near the middle and the termi-

nal knob-like appendix may be abruptly formed and angularly

bent from the stem or it may appear simply as an enlargement,

without being noticeably deflected at all. A set from Humboldt

Lake, Nevada, taken by myself in June, while not identical with

the Saltair forms, shows a range of variations almost exactly

corresponding thereto, all of the important features of the one

being duplicated in the other.

On the shores of Honey Lake, near Amedee, California, I

took a fine series of Cicindela which I refer, for convenience, to

C. echo, though they are not typical. The form is nearly the

same and the ground colors are about alike, but the Amedee

specimens almost entirely lack metallic gloss and the surface

sculpture of the elytra is notably shallower. The Amedee

beetles also differ, as a species, from the Saltair specimens in

the broader markings, the greater obliquity of the median band

(the terminal knob less deflected), and in the expansion of this

band along the margin so as to connect broadly with the humeral

lunule. The apical lunule is free in all of my specimens.

One individual is entirely blackish, except that each elytron

bears two small spots, one of which represents the anterior por-

tion of the humeral lunule, the other the posterior part of the

apical.

Cicindela pseudosenilis Walther Horn. Green, shining, a few

varying to brownish or reddish. Form of body as in C. echo.

Front of head sparsely hairy. The elytral markings are very

close to those of the Saltair echo ; the middle band does not

show a distinct tendency to spread along the margin in any of

"y specimens, so that it is not connected with nor closely

approximated by the apical and humeral lunules. The chief

oi. those exhibited in the descending portion of the

a vhs: te though this band is usually rectangularly bent

nob shows numerous modifications. In some

No. 453.] DISTRIBUTION OF CICINDELA. 649

specimens this knob is largely developed though not hooklike, in

others it disappears entirely, so that the descending part of the

band is of nearly uniform size to the tip. None of the speci-

mens in my series show a strong tendency towards the willistoni

type of marking, the lead to that form going through C. echo.

But some of the 5seudosenz/zs approximate the Saltair C. echo so

closely that if they were mixed they could scarcely be separated

again. The shores of Owens Lake, in southeastern California,

are the only definitely known haunts of the true C. pseudosentiis,

though Dr. Howard writes me that in the National Museum is

a single specimen said to have been taken by Mr. Coquillet in

Los Angeles County — not very far distant. None of the other

Californian collectors have found it in this latter locality, how-

ever. I found it in great abundance about the overflow of a

spring on the upper beaches of Owens Lake, but none occurred

in the immediate vicinity of the bitter waters of the lake itself.

Now that the variations of the insects have been described,

we must turn again to the geological records and see what can

be said of the early conditions of the country they inhabit.

The geology of the Great Basin has been worked out by Dr. G.

K. Gilbert) and Dr. Israel C. Russell? the results of their

labors appearing in two fine volumes from which the main geo-

logical items used in this discussion are compiled. We find

that in the early Pleistocene the basin held two great fresh-

water lakes: Bonneville, covering the greater part of western

Utah and a small portion of eastern Nevada and southern Idaho,

and Lahontan, occupying an extensive area in western Nevada

and eastern California. Between them lay a broad plateau or

divide, forming a watershed, the hydrographic basins of the lakes

being contiguous. Both of these lakes were of irregular shape,

Lahontan being especially so, with numerous arms and bays

extending up narrow, flooded valleys. Each lake had two great

periods of high water, which had been preceded by times of

drought and desiccation, the second stage of flood being higher

than the first. These times of plenty correspond to the two

lake Bonneville. Monographs of the U. S. Geological Survey, I, 1890

2Geological History of Lake Lahontan. Monographs of the U. S. Geological

Survey, XI, 188s.

650 THE AMERICAN NATURALIST. (VoL. XXXVIII.

glacial epochs of the Sierra Nevadas, though the climate is sup-

posed to have been only moderately humid and rather cold. The

increase in the size of the lakes is not ascribed to the melting of

the glaciers, though this must have added considerably to their

volume, as the ice cap was not large enough to furnish so much

water. While Lake Lahontan seems not to have overflowed,

Lake Bonneville broke through the barrier to the north and

found an outlet to the sea by way of the Columbia River basin.!

The desiccation of the lakes during the dry times is thought to

have been more complete than at present.

The relation of our modern lakes to those of Pleistocene

times is principally that of occupying the same area ; that is to

say, the bodies of water now existent have, in the main, been

formed since the old lakes dried up, and are not to be consid-

ered remnants left by incomplete evaporation, since in the latter

case the waters must have been much salter than they are. An

exception may perhaps be made of Great Salt Lake, the evi-

dence being inconclusive; and I have no data of this nature

concerning Owens Lake and Mono Lake, which never formed

part of these two larger bodies, but were separate even during

the high-water periods of the Pleistocene. The littoral Fauna

may easily have been preserved, even through times of great

drought, by clustering about the edges of springs. In this way,

even to-day, we know that some are carried over ; for example,

this very Cicindela echo flocks on the damp ground in the vicin-

ity of the little springs about the edge of the now dried-up

Humboldt Lake. Many springs are of a far more permanent

nature than the shallow lakes into which they discharge, and I

think that we may safely assume that they lasted, in many

instances, through the times of most complete desiccation.

. Now let us consider the relation of existing lakes (which have

been examined for material for this paper), and those of ancient

times, in order to see what opportunity the beetles have had for

differentiation through isolation. I have visited Great Salt

Lake, Utah Lake, Humboldt Lake, Honey Lake, Walker Lake,

1 ; : ;

This may possibly account for the present occurrence in the Columbia dis-

trict of such widely distributed Great Basin species as Cicindela hemorrhagica

and Saprinus estriatus.

No. 453.] DISTRIBUTION OF CICINDELA. 651

Owens Lake and Mono Lake, and found Cicindela echo or C.

Bseudosenilis at all but three — namely, Utah, Mono and

Walker. I cannot definitely assert that the species, in some of

its forms, is absent from all of these three, but we have no evi-

dence to show that it occurs there. Mono Lake may perhaps

lie at too great an altitude for the insect to flourish, or it may

never have been introduced into the district. It is possible that

a more extended search may yet disclose some form of it there,

my visit having been a hurried one and productive of no Cicin-

dela whatever, though C. pseudosenilis and C. hemorrhagica

must have been flying in abundance at Owens Lake, distant

about a hundred miles. "Walker Lake has been twice visited by

me with the special object of looking for some form of C. echo,

but I took only C. hemorrhagica and a variety of C. oregona,

both in plenty. I hope that if any entomologist has the oppor-

tunity, he will visit the flats at the upper end of the lake, in

the vicinity of the mouth of the river, since I have noticed that

the colonies are sometimes confined to a limited area and may

easily’ be missed. It will be well also to examine the neighbor-

hood of the large spring said to exist near Wabuska ; it dates

back to considerable antiquity and may yield something inter-

esting. Utah Lake has been visited by several entomologists at

different times, and it seems that if C. echo occurs there it

should have been met with before this.

These three lakes which have failed to yield material. after

exploration may be left out of the rest of the discussion. Ot

those which have been productive, Owens Lake has been dis-

tinct fora vast period, having, as we have seen, been separate

even during the greatest extension of the others. Great Salt

` Lake was a part of Bonneville, while Humboldt Lake and

Honey Lake lie in the ancient bed of Lahontan. The small

bodies of water in Wyoming, which harbor C. willistoni were of

course not included in either and had no communication there-

wth

Looking again at the beetles, we find :

(a) The Owens Lake form, which has probably been isolated

since (at least) early Pleistocene times, is sufficiently well differ-

entiated to have been separated by systematists as a distinct

species, though this view is no longer held.

652 THE AMERICAN NATURALIST. [Vor. XXXVIII.

(b) The form from Honey Lake, on the extreme western

border of the old Lake Lahontan, is also well differentiated,

though not in the same way. This lake is extremely shallow

and evanescent, having dried up at least twice within the mem-

ory of the present generation, and has no doubt been separated

from the main body of the old lake since a comparatively early

period in the second great decline in size of the latter. The

beetles live about the seepage on the lake beach that comes

from the hot springs a few hundred yards distant. While it is

impossible at present to point out the details which lead to vari-

ation in any one direction, it is evident that local conditions,

aided by long isolation, have so modified this form that it is now

readily separable, in series, from those taken elsewhere.

are more of the type of typical ecZo, differing but little from the

series taken at Great Salt Lake, Utah, whence the species was

originally described. While the Humboldt colony must have

been separate from the latter fora great length of time, it has

been comparatively little differentiated therefrom. The reason

for this cannot now be assigned. It may be that the local con-

ditions requisite to incite variation were not present, or perhaps

the two colonies have simply varied along parallel lines. Though

isolation is unquestionably favorable to differentiation, it does

not follow that every isolated colony must differ from every

other. It is well known that with certain species of Cicindela

some colonies produce mostly specimens of one type, others

will produce those of another, while a third will be composed of

a mixture of both, with all the intergrades.

(d) The form from Great Salt Lake, to which the name C.

echo is properly applied, is not like that from Owens Lake nor `

that from Honey Lake, though closely approached by the speci-

mens from Humboldt Lake. :

(e) On the shores of the small lakes in Wyoming, we have

C. willistoni, a closely related form occurring (as far as we

know) nowhere else, which according to geological evidence can

scarcely have had any communication with the Great Basin col-

onies during Pleistocene times, Iam inclined to look to a still

more remote date for the cause of this phenomenon, and con-

No. 453.] DISTRIBUTION OF CICINDELA 653

sider C. willistoni a survivor of an early stock which, during the

Tertiary period, inhabited the present Rocky Mountain region

and in all probability the adjacent districts to the east and west

at the time of the great extension of the Tertiary lakes. C.

willistoni is much better differentiated from the races of C. echo

than those races are among themselves.

My conclusions are these: That in C. echo (with its several

races, including C. pseudosenilis) and C. willistoni, we have two

branches of a stem which were probably separated by some of

the orographic movements which gave rise to the upheaval of

the Wasatch and Rocky Mountains. That this stem form was

a littoral species and the branch remaining in the Great Basin

was carried over the interval between the dissolution of the Ter-

tiary lakes and the appearance of the great Pleistocene lakes by

clinging to the borders of springs and other bodies of water.

That with the growth of Lake Bonneville and Lake Lahontan

and the contemporaneous filling by water of the smaller basins,

it spread over the whole habitable area between the Wasatch

and the Sierras. The subsequent desiccation of the greater

part of the Lahontan and Bonneville basins left a series of

smaller lakes of varying permanence and more or less complete

isolation. Local conditions, acting on the members of colonies

of the beetles thus separated from their neighbors, have given

rise to variations of different kinds and certain of these varia-

tions have been preserved and accentuated through this isolation.

As a consequence, we have the phenomenon of local races,

strongly or slightly marked according to the strength of the

conditions exciting variability and to the comparative degree of

isolation of the colony after a certain character had made its

appearance. Owens Lake and Honey Lake, having been long

since separate from the larger bodies and presumably subjected

to different conditions, climatic and otherwise, because of their

proximity to the Sierras, have at length produced upon their

shores racial types which are easily distinguished from each

other and from the forms found farther east. j

If the species of Cicindela used in illustration were alone in

presenting the general phenomena noticed, one might well doubt

the sufficiency of the evidence adduced in support of the foregoing

654 THE AMERICAN NATURALIST. [Vor. XXXVIII.

conclusions : and as the matter stands, I am well aware of the

circumstantial nature of much of it and of the many possibili-

ties of wrongly interpreting the facts. However, I have been

collecting and studying other species of littoral Coleoptera in

the Great Basin, with a view to further prosecution of the prob-

lem, and find much in corroboration. It is evident that the

same general law is followed in the variations of Tanarthrus,

and I hope also to demonstrate a parallelism in certain Carabidze

and Histeridze which are associated with it.

STATE UNIVERSITY OF Iowa,

lowa City, March 19, 1904.

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF

THE MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD

COLLEGE. E. L. MARK, DiRECTOR.— No. 154.

A SIMPLE APPARATUS FOR AERATING LIQUID

SOLUTIONS.

S. O. MAST.

IN studying the effect of dilute and concentrated solutions of

sea water on marine algae it became necessary to aérate the solu-

tions in order to keep them fresh. For this purpose a very

simple piece of apparatus was devised. The apparatus was used

slowly to concentrate or dilute the solutions as well as to aérate

them. It has now been in continuous operation for over three

months and has proved to be so effective, to require so little

care, and to be so easily made, that a brief description of it may.

be useful to others.

The general structure of the apparatus will be readily under-

stood by referring to the accompanying figures. It will only be

necessary then to discuss a few details of its structure and the

principles on which its operation depends.

The aérating tubes used in the experiments mentioned above

are 9o cm. long and 4.5 mm. in diameter (inside measure), which

is constricted to 0.5 mm. at a point 7 cm. from the upper end.

The capillary siphon is 0.75 mm. in diameter; the solution in

the culture jar is 27 cm. deep. It will readily be seen that,

since the aérating tube is much larger than the capillary siphon,

if both are to be constantly full the solution must flow much -

more slowly in the tube than in the siphon.

Owing to the action of gravitation, however, the motion of the

solution tends to accelerate as it falls, and therefore tends to

flow faster in the aérating tube than in the siphon ; so that the

solution in the tube as it flows through the constriction is

broken into very short columns separated by small bubbles of

655

656 THE AMERICAN NAFURALIST. [Vor. XXXVIII.

i ; hese

air, which incidentally aërates the solution. pida: : ies

Minos unite, forming larger ones as they flow git He.

boe as do the bubbles also. The relative lengths o

tube,

F1c. 2.

Fic. 1.

e jar.

Fic. 1. — An aérating apparatus in which the liquid used to a&rate flows puedes e :

r k ,

IG. 2. milar 4 apparatus in Wapi the “qma does not tlow into the cu ii = oven

clr ja Jar, 2. e., a jar o be a&ra aem B,a ply jar;

j c Ne

: D, a capillary ien: Ea an skating die FLA nAi siphon ;

accessories to the aërating tube.

ee fac-

columns of tite and spaces of air depend upon M ie

tors: (1) The difference between the diameter of the

No. 453.] AERATING LIQUID SOLUTIONS. 657

lary siphon and that of the aérating tube, (2) the length of the

aérating tube, and (3) the depth of the solution in the culture

jar. All that is necessary in order to increase the amount of

air, as compared with that of the solution in the tube, is to

increase either the diameter or length of the aérating tube or to-

decrease the depth of the solution in the culture jar.

If the diameter of the tube is increased and the flow kept

constant it is clear that more air must be taken in to fill the

increased space due to the enlargement. But as the diameter

increases the adhesive force between the walls of the tube and

the solution in it decreases, as compared with the cohesive force

of the solution, so that when a certain diameter is reached the

bubbles of air break through the columns of solution and thus

can no longer be forced to the bottom of the tube in the culture

jar. As a matter of fact, the maximum limit of diameter in the

case of water is practically about 4.5 mm. Further, if the rate

of flow in the aérating tube be in any way increased while that

in the siphon remains constant, more air will be required to keep

the tube full, just as in case of an increase in diameter. The

rate of flow in the tube, however, varies directly with the length

of the tube; and since there is no theoretical limit to the possi-

ble length of the tube, the amount of air compared with that of

solution can be theoretically increased indefinitely. The depth

of the culture solution is really a function of the length of the

aérating tube. In order to overcome the pressure of the culture

liquid and force the air out at the bottom of the aérating tube,

it is necessary to have a column of solution in the tube above

the level of the solution in the culture jar slightly greater than

the air space in the tube de/ow that level. Consequently the

portion of the tube which is below the level of the solution in

the culture jar and enough of it above that level to contain as

much solution as there is air below, have nothing to do with

increase in acceleration due to gravitation, for they simply bal-

ance each other, and therefore these parts are not effective in

increasing the amount of air introduced. This being true, it

will readily be seen that increasing the depth of the culture solu

tion has the same effect as decreasing the length of the aerating

tube, and that therefore the deeper this solution, other things

being equal, the less will be the air forced through.

658 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The purpose of the constriction in the aérating tube is merely

to break up the column of solution in the tube. If this were not

thus broken up it would accumulate until it became slightly

greater than the air spaces in the tube below the level of the

culture fluid and then would suddenly flow down, forcing the air

out with a rush, after which it would again accumulate, flow

down, etc. This is just what happens if the constriction is not

small enough in comparison with the size of the capillary siphon.

A few experimental trials, however, will be sufficient to enable

one to decide on the proper relative sizes of the two. No defi-

nite instruction can be given with reference to this since the

diameter of the siphon must vary with the diameter and length

of the aérating tube and with the depth, specific gravity and

viscosity of the culture solution. In general the constriction

should be somewhat smaller than the siphon.

In some forms of apparatus! the lower end of the aérating

tube is bent on itself so as to project upwards in the culture

solution and a small piece of sponge is then pressed rather

tightly into this bent free end, or the tube may be drawn out

into a capillary ending. It is claimed that the sponge, espe-

cially, breaks up the air into very small bubbles. This is un-

doubtedly true, but it has been found that in the apparatus

represented in Figures r and 2 the pressure required to force

the air through the sponge is so great that the value of this

adjunct is questionable. |

The most efficient and simplest method discovered to break

up the air bubbles is as follows: Cut off the lower end of the

acrating tubé square, grind it quite flat, and let it rest on a firm

piece of rubber, leather, ór wood on the surface of which a con-

siderable number, of radiating grooves has been made. The

piece of rubber rests on the bottom of the culture jar and the

tube is placed over the point whence the grooves radiate. The

Srooves In any event must be narrow. Their depth should depend

Sron the weight of the aërating tube. If the tube is heavy t

will sink into the surface upon which it rests, and therefore

under such conditions the grooves must be deeper than if the

ds he H. Lenz. Verbesserung an den Durchlüftungsapparaten der Seewasser

quanen. Zool. Anzeiger, 1879, Jahrg. 2, pp. 20, 21. *

No. 453.] AERATING LIQUID SOLUTIONS. 659.

tube is light. Although by this device the bubbles are not

broken up as much as they can be by using a piece of sponge

forced into the end of the tube, this method has the decided

advantage of requiring much less pressure, and there is also

much less danger of clogging the tube than if a sponge is used.

In making the aérating constriction the tube should be heated,

without being drawn out, until the walls fall in, as this causes.

them to become much thicker and consequently stronger than

they otherwise would be.

It is much better to use a capillary siphon than a larger tube

drawn out to a capillary ending ; first, because a capillary siphon

never needs to be filled, for when the supply jar is filled to within

about 1 cm. of the top capillary attraction will fill the siphon

and start it running ; secondly, because a large tube drawn out

at one end is much more readily clogged than a small one of

equal diameter throughout.

The constant-level siphon (F) should be unquestionably large

enough to drain off the solution as fast as it comes into the cul-

ture jar, but not much larger; neither should its outer end be

more than a few centimeters lower than its highest point,

for if it is the decrease in pressure at the highest point causes

sufficient increase in vapor pressure of dissolved gases to cause

them to be thrown out of solution, to collect at this point, and

thus to clog the siphon.

The principles of operation in the second apparatus, that,

represented in Figure 2, are in all respects similar to those

discussed with reference to the apparatus represented in Figure

I. The upper end of the aérating tube, the capillary siphon,

and the supply jar are the same as represented in Figure r.

The chamber G, which serves to separate the air from the

liquid, can be readily made by cutting off a test tube. It should

be about 1.5 cm. in diameter and 4 or 5 cm. long. It must of

course be air-tight. Ordinary corks soaked in paraffin will serve

very well to close the ends.

The tube Æ should be at least 5 mm. in diameter, slightly

larger than the aérating tube proper, 7. e., large enough to pre-

vent air being forced through it with the solution. Its length

must be such that its outer end is somewhat farther above its.

660 THE AMERICAN NATURALIST. [Vor. XXXVIII.

lower bend than the outer end of the tube / is below the surface

of the solution in the culture jar; for if it is not the air will pass

out through it instead of through the culture solution.

The tube / serves to convey the air from the chamber G into

the culture solution. It may be of any size, length or form.

The lumen through the constriction in it is very small; though

not absolutely necessary, it serves to cause a more steady flow

of air. A rubber tube with an adjustable pinch-cock might serve

this purpose still better.

All that is necessary to operate either apparatus described, is

to pour the solution from the overflow jar into the supply jar

from time to time. The frequency required in this operation

depends of course upon the size of the jars and the diameter of

the capillary siphon. If, as in the experiment mentioned above,

4liter jars and a 0.75 mm. siphon are used, a transfer once in

three days will be quite sufficient.

In closing I wish to thank Dr. E. L. Mark for furnishing

material which made the experiments referred to above possible,

and for valuable criticism of the manuscript of this article.

SPIRE VARIATION IN PYRAMIDULA

ALTERNATA.

FRANK COLLINS BAKER.

I. INTRODUCTION.

Tue object of the present investigation has been to ascertain

by quantitative means the amount of variation in the same spe-

cies of a pulmonate mollusk from several localities. For this

purpose the shell of the common species, Pyramidu/a alternata,

has been selected, as that species shows a large amount of varia-

tion in the form of the shell, particularly in the height of the

spire.

Among the mollusks little biometric work was done previous

to the year 1898. Since that time this group of animals has

been receiving marked attention and several exhaustive papers

have been published, notably those by C. C. Adams on Io and

C. B. Davenport on Pecten. These papers, however, have dealt

with marine or fresh water forms ; in this investigation the bio-

metric study has been applied to an air breathing land mollusk.

II. MATERIAL.

The material used in this investigation, Pyramidula alternata

Say, was secured from the following localities :

1. Rochester, New York, on the steep hillside of the “ Pinna-

cle” and the banks of the lower Genesee River. This locality

is heavily wooded and there is a large accumulation of fallen logs

and the ground is covered by a rich, black loam. Collected by

the writer.

2. Auburn, New York, in damp, low, flat woodlands. Col-

lected by Dr. Howard N. Lyon.

3. Bowmanville, near Chicago, Illinois, in flat woodlands.

The timber is large and heavy, the ground is strewn with fallen

661

662 THE AMERICAN NATURALIST. [Vor. XXXVIII.

logs and the soil is a rich, black loam. Collected by the writer.

The shells studied are not of uniform size, although they are

nearly all adult measuring from twelve to twenty-four millimeters

in diameter. The young shells of this species always have a flat

or nearly flat spire, and as they would materially affect the results

they were excluded.

In Pyramidula alternata we find a good example of variation

caused by individual environment. The species lives for the

most part under started bark, in crevices and under flat-lying

tree trunks; hence its shell varies with its abode. For example,

a specimen living between the “started” bark of a tree and the

tree trunk, the space being very narrow, measured 23 x 11 mill,

the height being about 48 per cent. of the width, while another

specimen living under a fallen tree trunk measured 15 x 13 mill,

the height being about 87 per cent. of the width, or 39 per cent.

more than the first example. These specimens were from the

same locality and from adjacent trees. Their habit of crowding

into narrow crevices and between the bark and the tree trunk

has caused this species to become one of the most variable of

land shells as regards the form of the shell.!

On account of the extreme variability it was thought that a

quantitative study of material from several rather widely sepa-

rated localities would produce results of some interest. This

species has a wide geographic range, being found throughout

the eastern and central parts of the United States and Canada.

Its western limit is said to be Minnesota.

III. METHOD or OBTAINING QUANTITATIVE DATA.

To obtain a variation index the diameter and altitude of the

shell was measured in millimeters and the altitude divided by

the di d altitude. f : :

diameter ; 7. e., diameter Lhe per cents. obtained in this way

provide the shell index. In the diagrams the individuals or

classes having the same per cent. are indicated on the horizontal

line and the number of specimens in these classes (the frequen-

cles) are indicated on the vertical line.

1 See y . ,

bae de autilus, Vol, 10, p. 63, fora good article on this subject by C. C. Orms-

' Influence of Environment upon the Form and Color of Helix alternata.”

No. 453.] PYRAMIDULA ALTERNATA. 663

The number of specimens available for this study has not

been as large as is desirable in investigations of this kind; but

as the three lots are of about the same number the results will

not be materially affected.

In the tables of data the per cents. are indicated by a numer-

ator and the individuals or frequencies having the same per cent.

is noted as a denominator.

IN. DiscUssIoN oF DATA.

The Rochester Shells.

Figure I.

The shells from Rochester show a rather wide range of varia-

tion, as is seen in Fig. 1, which is strongly trimodal. The most

H ti

10 | 10

g | 9

f Li 8

7 | Lil 7

$ i eH Lii 6

5 x ILI "

4 N 4

3 3

2 2

/ — LI

0 o

Fic. r.— Rochester shells. Variation curve of 67 specimens.

peculiar aspect of this curve is the broad mode from 47 to 52

per cent., the frequencies being 9. The two sharp modes are

$5 with a frequency of 8 and 59 with a frequency of 12. The

variation is from 44 to 65 per cent. The data for Fig. 1 1s

` Shown in Table A.

Table A.

44 46 47 $0 52 53 54 59

ga 58 59 60 61 65

I $5 9 9 9 » * — ? .

The number of shells examined was 67.

664 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The Auburn Shells.

Figure 2.

The Auburn shells are not as variable as those from Roches-

ter, a fact shown by the greater regularity of the curve, which

| Wi 23

| "

Ms wu, S» SOUS Re m m m

~ ho w a ^ 6 90 5G

i / 7 j 70 JI 7. F 6

Fic. 2. — Auburn shells. Variation curve of 104 specimens.

is practically bimodal, the minor mode being at 54 per cent. with

a frequency of 21 and the major mode at 57 with a frequency

of 24. The data for this curve is shown in Table B.

Table B.

ao 3° 33 54 55 56 57 58 59 60 61 02 "EN

ED o6 3 3 6 24 160 5 7 4 UM

04 65 66 68 7o

A P Y 1. rr

The number of shells examined was 104. The range of vari-

ation is from 45 to 79 per cent.

No. 453.] PYRAMIDULA ALTERNATA. 665

The Bowmanville Shells.

Figure 3.

The Bowmanville shells are the most variable of the three

lots, the curve showing a marked multimodal tendency, which

i | | | n

I0 * | 10

9 | 9

$ I | 8

/ TZN ;

6 i 6

: i Y ;

4 4

4d \ 3

i \ y 2

/ I

o " 0

61 69.63 64 65 66 67 68 6970 7) 72 13 74 75 “

Fic. 3. — Bowmanville shells. Variati f 85 specimens.

always stands for extreme variability. The range of variation

is from 46 to 87 per cent. There are four modes of promi-

nence, at 50—53, 57, 59 and 65, with frequencies of 5, 10, 9 and

5, respectively. It will be noted that the curve for the Bow-

manville shells is very similar to that of the curve for the Roch-

ester shells, the peculiar broad mode being present in each.

The data for this curve is shown in Table C.

Table C.

46 50.33 54 4i 35 90 Ee i Ed

ow priri ie ee o v 3 *

65 67 68 jo Jt PR uron

s 4 Y 4'q 9 9 17 47 4*4 «^ '

The number of specimens examined was 85.

V. COMPARISON OF THE THREE LOCALITIES.

Figure 4.

In comparing the three localities it will be noted that the

Bowmanville and Rochester shells resemble each other very

666 THE AMERICAN NATURALIST. [Vor. XXXVIII.

closely in the form of their curves, which is quite different from

" n

r 2

1 t 29

H i 21

» 1 i 20

7 i H

1 i 18

J 4

; 16

! Hi 15

^ T "L4

: 13

: 12

: //

, E 10

r 9

Ctr 8

a ` D

LITE ji Ay A

/ E

ZLT Bet " r | py $

{CACHAN ;

L = = 3

E AAC \ 2

1 ^ Y-F -la l

0 m i 9

758 37 60 6] [; 70 71 7% 73 14 25 76 17 78 29 80 8| 8485 8

Fic. 4. — Comparison of the three localities. r. Bowmanville. 2. Rochester. 3. Auburn.

the Auburn curve, which is very symmetrical, showing a smaller

amount of variation. The major mode for the three localities is

about the same, standing as follows :

Bowmanville .57 %. 1

Rochester 59%.

Auburn 57 fb.

The mean per cent. for the three lecalities is as follows:

Bowmanville 65.1 %.

Auburn 59.7 %.

Rochester — $4.1 %.

This

highest

The wi

the ext

shows that the Bowmanville shells have relatively the

spires, while the Rochester shells have the lowest spires.

dest variation is found in the Bowmanville shells, where

remes are 46 and 87 per cent.

1 100% would mean that the d

iameter and height were the same; hence the

per cent. shows the relation of height to diameter

No. 453.] PYRAMIDULA ALTERNATA. 667

Geographically the western shells show a wider range of vari-

ability than do the eastern shells. With one exception (Auburn

79%) the eastern shells have the per cent. of spire elevation

between 44 and 68, while those from the west range between 46

and 87, the per cents. between 68 and 81 being numerous.

One of the most noticeable features in the curves as plotted

in this paper is their tendency to assume a multimodal form.

This is clearly shown in the individual diagrams, but stands out

prominently in the comparison of the localities (Fig. 4). This

is, of course, indicative of great variability.

From the study of these three lots of shells we may conclude

that the western a/ternata has a higher shell, on the average,

than does the eastern form, and that it shows a much larger

amount of variation in spire elevation. It would be very inter-

esting to have curves plotted from other localities, east and west,

to ascertain whether their results would coincide with the con-

clusions of the present paper.

BIBLIOGRAPHY.

BAKER, FRANK C.

Rib Variation in Cardium. Amer. Nat. XXXVII, 481-488, 1903.

BAKER, FRANK C. :

The Digitations of the Mantle in Physa. Bull Chi. Acad. Sci., II,

No. 4, 1901.

BATESON, W.

On Some Variations of Cardium edule apparently correlated to the Con-

ditions of Life. Phil. Trans. Roy. Soc. London, 180, B, 297,

1889. ,

BicELow, R. P. AND RATHBURN, ELEANOR P. mE

On the Shell of Littorina litorea as Material for the Study of Variation.

Amer. Nat., XXVII, 171, 1903.

BuwPvs, H. C. :

The Variations and Mutations of the Introduced Littorina. Zoological

Bulletin, I, 247, 1898.

! This bibliography contains some of the more important mp oS :

the variation of the Mollusca, from a biometric standpoint; the list is no

exhaustive.

668 THE AMERICAN NATURALIST. [Vor. XXXVIII.

DAVENPORT, C. B.

On the Variation of the Shell of Pecten irradians Lamarck from Long

Island. Amer. Nat., XXXIV, 863, 1900.

DAVENPORT, C. B.

Comparison of Some Pectens from the East and West Coasts of the

United States. Mark Anniversary Volume, article VI, 121, 1903.

DAVENPORT, C. B.

Comparison of Pecten opercularis from Three Localities of the British

Isles. Proc. Amer. Acad. Arts and Sci., XXXIX, 123, 1903.

Dimon, ABIGAIL C.

Quantitative Study of the Effect of Environment upon the Forms ot

Nassa obsoleta and Nassa trivitata from Cold Spring Harbor,

Long Island. Biometrica, II, 24, 1902.

DOWNING, ELLIOTT R.

Variation in the Position of the Adductor Muscles of Anodonta grandis.

Say. Amer. Nat., XXXVI, 395, 1902.

MAYER, A. G.

Some Species of Partula from Tahiti. A Study in Evolution. Mem.

Mus. Comp. Zool., XXVI, 117, 1902.

THE CHICAGO ACADEMY OF SCIENCES.

une 4, 1904.

FOSSIL PLUMAGE.

C. R. EASTMAN.

PRESERVATION of avian remains in the fossil state is neces-

sarily of very rare occurrence, and extraordinarily so, if fossiliza-

tion takes place in marine sediments. Cases of the latter

description presuppose the creature either to have perished at

sea, or to have been swept out at a distance from the land as a

floating carcass without having been destroyed by predaceous

animals. The body must have sunk to the bottom before decom-

position had advanced far enough to disrupt the skeleton and

scatter the plumage. Burial by fine sediments must have fol-

lowed almost immediately, in order that the body be preserved

intact. And mineral replacement of the organic tissues must

have proceeded in a wonderfully subtle manner, transforming

the most delicate particles into stone without obliterating their

microscopic structure.

Supposing a dead bird to have reached the bottom in a toler-

ably complete condition, the feathers naturally become loosened

and scattered with decomposition of the skin, and the least cur-

rent is liable to sweep them away except they become entangled

and covered by the sediment at once. If the material happens

to be a fine calcareous ooze, the feathers may leave in it an

exceedingly delicate impression, or, in the rarest cases of all,

their structure may become replaced molecule for molecule by

mineral matter, generally calcareous or carbonaceous.

Chance, controlling thus absolutely the fate of this class of

remains, goes away after sealing them up in the rocks to remain

hidden for ages; but may peradventure come back again, and

disclosing them to the light of day, permit them to fall into

the domain of scientific investigation. Great as is the miracle,

it has actually happened a few times, as witness the two

complete individuals of Archzeopteryx that are known, and one

of Hesperornis, with their plumage preserved. Scarcely less.

669

670 THE AMERICAN NATURALIST. [Vor. XXXVIII.

interesting and remarkable are the complete examples found in

lacustrine and fluviatile deposits, the most perfect in our own

country being Palæospiza from the insect-bearing shales of

Florissant, Colorado, and Gallinuloides from the Green River

Eocene of Wyoming. In the former the plumage is pre-

served, in the latter it is wanting.! Skeletons nearly as per-

fect as these have also been described from fresh-water deposits

of the south of France and elsewhere in Europe.

But it is fossil plumage from marine deposits with which this

article is especially concerned. The feathers of Archzeopteryx

are too well known to require more than a mere mention of their

occurrence, and those of Hesperornis, recently made known from

the Kansas Cretaceous, are still fresh in the minds of students?

There is yet another marine horizon from which feathers have

been obtained, these constituting, however, the only indication

of avian life which exists in the formation. This is the Upper

Eocene limestone of Monte Bolca in the Veronese, famous for

nearly four centuries on account ef its remarkable fish and plant

remains. But as for the occurrence of detached feathers to the

exclusion of other remains, it should be remembered that ma-

rine and shore-birds are constantly shedding them, and hence, if

circumstances favoring preservation are equal, they are likely to

be numerically more abundant than bones.

Owing to their excessive rarity and wonderful preservation,

it is not surprising that the few feathers thus far obtained from

Monte Bolca should be objects of considerable interest. In the

early days of palzontology, no little rivalry was created over the

acquisition of the first specimens brought to light, one having

been found in 1777, and another some twenty years later, both

issant locality in Colorado; the Lower Miocene of Ronzon, near Puy-en-Veday ;

a Upper Miocene lignite of Rott, near Bonn ; and from the Upper Miocene

lacustrine deposits of Oeningen, Switzerland. Good examples from the last-

named locality, and of Ratite feathers from the Quaternary of New Zealand, are

preserved in the Museum of

Eora

Williston, S, W., Kansas Univ. Quar. Vol. 5, p. 53, 1896. —Marsh, O. C^

No. 453] FOSSIL PLUMAGE. 671

being in counterpart. These two specimens, which are now pre-

served in the Paris Museum of Natural History, were described

and figured in the early part of the last century by Faujas-St.-

Fond,' but not without misgivings lest his readers be inclined to

doubt their avian nature. The names are given of four profes-

sors at the Museum who agree with the author in his conclusions,

and it is observed that with reference to one of the feathers, that

“on ne sauroit la confondre avec certains fucus qui ont quelques

rapports apparens avec des plumes, parce que celle-ci a ses

barbes garnies d'autres petites barbes." This is the only state-

ment which is given in regard to the finer structure, and the

latter is not illustrated in the figures. Both of these feathers,

it may be added, are of the pennaceous, and not of the plumu-

laceous variety.

Except for a casual mention by Milne Edwards? of his having

seen one or two fossil feathers

in Verona, where they are still

on exhibition in the Public

Museum, no other references

occur in literature to this sort

of remains from Monte Bolca.

It may therefore be of interest

to examine the figure which

is given herewith of a speci-

men recently acquired by the

Museum of Comparative Zoöl-

ogy at Cambridge, along with

a fine suite of fish-remains

from a famous old Veronese

collection. This is a small

contour feather, only 1.5 cm. our Beda, dl

long, but remarkable for its dreams wr) RA anie

perfect preservation of details.

It is possible to distinguish each separate barb of the symmetri-

cal vane, and even the barbules along either side of the branches.

*Faujas-Saint-Fond, B. Mémoire sur quelques fossiles rares de Vestena Nova

dans le Véronais. Ann. Mus. d' Hist. Nat. vol. 3, pp- 18-24, 1804.

2 Milne Edwards, A. Oiseaux Fossiles de la France, vol. IL., p. 544, 1871.

672 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The barbules appear relatively shorter, coarser, and less closely

spaced than those in the body-feathers of recent Carinates, and

the apex of the vexillum is more pointed. The shaft not being ,

prolonged at the base, it is probable that the specimen is com-

plete in itself, and not the tip of a larger feather. No inferences

are warranted regarding even the remote affinities of the form it

belonged to, beyond that chances favor its having been a shore

bird of small or moderate size. Although the beds were un-

doubtedly laid down under deep-water conditions, the presence

in them of crocodilians, chelonians, and plant remains indicates

that the Bolca locality was not far removed from land at the

time these strata were deposited. It deserves to be stated

that, according to Walther, the Solnhofen lithographic stone

in which Archaopteryx occurs was deposited within a coral

island lagoon.

NOTES AND LITERATURE,

" ZOOLOGY.

“An Introduction to Vertebrate Embryology” by Dr A. M.

Reese! is *the result of a need that the author has felt, for some

years, for a concise text-book of embryology that described the devel-

opment of both the chick and the frog” (Preface, p. v). It is ques-

tionable if such a need has been generally felt in the presence of

existing treatises on embryology, though they may be more * cumber-

some and expensive" books than Dr. Reese's. There is something

to be said in favor of students becoming acquainted, at first hand, with

standard works such as the text-books of Balfour, Hertwig, Marshall,

and Minot.

To give a satisfactory account of the development of both the frog

and chick within the space limits of a book like Dr. Reese's, the

author must be a master of the subject and, equally important, he

must be a master of the art of expression in written language. It is

unfortunate that Dr. Reese's book does not do justice to its author's

knowledge. ‘The fundamental defect of the book — a defect which

overshadows good qualities — lies in poor presentation of the subject,

and, in a book which does not offer new material, form is the all-

important thing, so long as facts are not violated. The account of

the development of the optic nerve may be cited as an example of

the quality of description. “It is sometimes stated that the optic

nerve is formed by the hollow stalk of the optic cup; but it is proba-

ble that it is formed by an outgrowth of cells from the retina, this

outgrowth extending along the optic stalk to the brain, and forming

the fibres of the optic nerve. The growth of these fibres may have,

as has been mentioned, something to do with the formation of the

choroid fissure” (p. 181). What, in the light of this paragraph, is

the manner of development of optic nerve fibres? The account of

the development of the chick begins with the sentence, “ The egg of

the chick (Fig. 33) is of large size, ovoid in shape and usually some-

*Reese, A. M. An Introduction to Vertebrate Embryology Based on the Study

of the Frog and the Chick. New York, G. P. Putnam’s Sons, 1904. XVII +

291 pp., 84 figures.

673

674 THE AMERICAN NATURALIST. [Vor. XXXVIII.

what larger at oné end than at the other” (p. 90). (If the egg of

the “chick” were not ovoid, what would its form be ?) Then follows

what apparently is a description of the egg at the time of laying. In

the course of this description it is stated that the yolk exhibits on

one side “a small, whitish circle, the d/astoderm or cicatricula” (p. 91).

The next paragraph tells us that “ Although of so large a size, the

yolk of the hen’s egg is a single cell, its great size being chiefly due

to the large number of yolk granules which it contains "(p. 92).

Fortunately, the last paragraph of the section on “ The Egg " explains,

as if by an afterthought, that “The preceding is a description of the

egg at the time of its laying” and that “The statement that the yolk

is a single cell is really true only from the time it leaves the ovary

until it is fertilized, or until a short time after fertilization, when seg-

mentation begins" (p. 93). Even then we are left in doubt as to the

precise time when the egg ceases to be a single cell. It does not

tend toward conciseness to make an incorrect statement and then

add a paragraph to explain what is “really true."

In the accounts of the visceral apparatus of both frog and chick,

an indiscriminate use of the terms 277, branchial, and visceral, as

applied to the several arches and clefts, leads to hopeless confusion.

It is doubtless by an oversight that the author states that the ecto-

dermal auditory invagination of the frog gives rise to the lining of

the middle ear (p.43). Eia surprise to find the terms somato-

leure and Splanchnopleure applied to the parietal and visceral layers

of mesoderm respectively. The use of the incorrect plurals “ diver-

ticulae " and “lumena,” an apparent failure to appreciate that mares

IS a plural, and reference to nascent organs as rudiments are com- .-

paratively unimportant matters. The figures, mainly from Marshall,

Duval, and Minot, are well reproduced, but their arrangement in rela-

tion to the text could be much more convenient.

A book such as Dr. Reese has proposed should lay down in a few

firm bold Strokes the main outlines of the subject. Unimportant

details should be omitted, important details must be adequately

treated. The account must proceed in an orderly constructive way,

always complete so far as it has progressed, like the development it

portrays. There must be such unfailing accuracy and such clearness

of Statement that misconception is impossible. If Dr. Reese’s book

I5 not an unqualified Success, it is because the author undertook a

very difficult task.

H. W. R.

No. 453] NOTES AND LITERATURE. 675

A Hermit’s Wild Friends..— The first impulse of a reviewer on

reading this book is to rage and to utter vain things. There is

something peculiarly irritating in the cock-sureness and the conde-

scension of ignorance; there mingles, no doubt, with our zeal for the

defense of truth a little of the wrath of the orthodox prophet against

him who would lead the people after false gods and therefore after a

false prophet. If it were only a question of the scientific world, it

would be hardly necessary to pay any attention to the book, but

the reviews quoted by the publishers show how easily the general

press are mislead.. The Hermit is hailed as a second Thoreau, or

put before Thoreau, since he “spares us Thoreau’s philosophy."

One reviewer admits that there is much “ out-of-the-way information,"

but is convinced that it all bears *the stamp of truth." Another

reviewer assures us gravely that the book never goes “beyond the

observed facts." It is rather against an unscrupulous publisher and

irresponsible reviewers that our wrath should be directed ; the Her-

mit's sins are those of ignorance and vanity; the publisher's are

those of greed.

The book purports to give true and detailed records of intimacies

with wild animals of the Gloucester woods, where the author has

lived for years. He has numbered among his intimates, song

sparrows, chewinks and chickadees, squirrels, mice and crows. It

will only be necessary to give an extract taken almost at random to

show the character of the book. The “ Hermit” has found (p. 187)

a crippled sparrow and has been feeding him in his camp in the

woods. “The fourth day, while I was feeding him, an old chewink

hopped to the loaf of bread [always put out for the birds], and

called him. The sparrow did not respond at first, but after awhile

hopped over to see what the chewink wanted. He seemed surprised

to find the bread, and began at once to help himself, The chewink

called him into the bushes. Z suppose he intended to give him an

introduction to his family. The next day the sparrow came into

the dooryard alone. He made for the bread and did not look at

me. I tried to catch him, but he hopped into the bushes, apparently

filled with terror, I think that old chewink had told the sparrow

that I was a very bad man. The old fellow might have been jealous,

: and had frightened the young sparrow, so that he would fly from me

in wild alarm, The next time the sparrow visited the yard the

1 Walton, Mason A. A Hermit’s Wild Friends. Boston, Dana Estes & Co.

[1903]. pp. 1-304.

676 THE AMERICAN NATURALIST. (Vou. XXXVIII.

chewink was with him. They departed together, and three days

later I saw the sparrow near the old barn..... Jt was evident that the

chewink had piloted him three-fourths of a mile to his friends.....

How did the chewink know where to take the sparrow?” How

indeed? The italics are the reviewer’s and this passage is com-

mended to the critic who could say that the book never went

“beyond assured facts.” When we add that the illustrations are

in keeping with the text we have done all that is possible to put the

public on their guard against this book.

Ry HS

The Sino-Australian Continent.—The existence of this continent,

first assumed by Neumayr for the Jurassic period, and which was

accepted by various subsequent writers for the Cretaceous, and

upward in the geological scale to the beginning of the Tertiary,

apparently needs restriction with regard to its duration. It now is

rendered more or less probable that it was not present at all in the

Jurassic period. Lately G. Boehm! has demonstrated that, in the

region of the Moluccas, Mesozoic marine deposits of European type

are largely developed, and are chiefly represented by various horizons

of the Jurassic series. Boehm draws the conclusion, “it becomes

apparent that a Sino-Australian Jurassic continent, as conceived by

Neumayr, did not exist.”

On the other side, deposits of Cretaceous age are absent or scarce

in this region, so that this old continent might have existed at least

during a part of the Cretaceous period. Boehm does not discuss

this question, but we must bear in mind that zodgeographical facts

positively demand a connection of Australia with eastern Asia, and

all evidence tends to show (see H. von Ihering, C. Hadley. H. A.

Pilsbry, A. E. Ortmann, M. Weber) that this connection was a broad

and important one in pre-Tertiary times, while, during the Tertiary,

it became more irregular, and was subject to many changes which

amounted frequently to complete interruption, which latter condition

prevails at present. The restriction of the Sino-Australian continent

to a certain part of the Cretaceous times consequently would meet

the postulates both of geology and zoógeography.

A. E. O.

1 . " Li

Geologische Ergebnisse einer Reise in den Molukken, in: Compt. Rend. 9.

Congr. Geol., Wien., 1904.

No. 453.] NOTES AND LITERATURE. 677

Origin of the Large Mammals of North America.—M. Grant!

discusses the old continental connections of North America with the

Old World and with South America. Most important is his idea

about the old “ Beringian connection” between. northeastern Asia

and northwestern America, which, according to him, was not a con-

tinuous one in time, as generally accepted (from the upper Creta-

.ceous to the lower Pleistocene). Grant believes that it existed in

the lower Eocene, lower Oligocene, middle Miocene, upper Pliocene

and lower Pleistocene, but that it was interrupted chiefly in the mid-

dle and upper Eocene, upper Oligocene and lower Miocene. The

evidence supporting this assumption is not very convincing, since in

part it may be founded only upon a deficiency in our knowledge of

the fossil Mammals both of the Old and the New World. Moreover,

the geographical distribution of maririe animals does not support this

view, at least as far as it refers to the older Tertiary. There is

hardly any trace of an exchange of faunas between the northern

Pacific and the northern Atlantic by way of the Arctic basin during

earlier Tertiary times, the similarities in the faunas of these oceans

generally being clearly indicative of a very recent connection of

them. There is either no resemblance at all, or very close affinity

of forms generally amounting to specific identity, the latter cases

being found among forms that are apparently circumpolar cold-water

types of recent origin.

It shall not be denied that there are a few cases of allied or even

identical species in both oceans belonging to more temperate cli-

matic conditions (for instance, resemblances of Japanese and Medi-

terranean forms) which might possibly indicate a former interruption

of the Beringian bridge in the beginning of the later half of the Ter-

tiary, but this point needs further elucidation.

A. E. O.

PAL/EONTOLOGY.

Walther's Solnhofen Fauna.?— One of the most philosophical

! discussions of extinct Faunz is this essay by Dr. Walther, which

forms part of the Haeckel Anniversary Volume. The geological

8th Ann. Rep. New York Zool. Soc., 1

g W 904.

alther, J. Die Fauna der Soluhofener Plattenkalke, bionomisch betrachtet.

1904.

ena,

678 THE AMERICAN NATURALIST. [Vor. XXXVIII.

evidence is interpreted as demonstrating the extension of coral reefs

throughout the shallow sea covering Central Europe in late Jurassic

time, which was a period of oscillation. The fine calcareous sedi-

ment interbedded with lithographic stone, for which the district

about Solnhofen is famous, represents the infilling of a lagoon, out-

- side of which the coral limestone carries a totally distinct fauna.

The peculiar mixture of land and marine organisms, the occurrence

of large trunks of trees and seaweed with roots, the interbedding of

apparently wind-blown material, and the tracks of undoubted air-

breathing animals, are among the facts which indicate that the

bottom of the lagoon was barely below tidewater, and probably was.

even exposed at times. Creatures straying into the lagoon and

becoming entrapped there, or volant forms like insects, Pterodactyls.

and Archzopteryx, which met their death in the paste-like, rapidly

accumulating sediment, were covered before any injury had been

done to their bodies through decomposition or other causes, the

most delicate structures being perfectly preserved. Although the

variety of forms is large, yet Solnhofen fossils are surprisingly rare

as compared with the majority of horizons, and a great many species.

are known only by one or two individuals; several important groups

are not represented at all, and on the other hand, a large percentage

of species is restricted to this locality. There appears to be no room

for doubting that the assemblage is an accidental one, and this vast

cemetery gives us a unique but by no means typical reconstruction

of the late Jurassic fauna.

Karl Alfred von Zittel.— Of the numerous biographical sketches

which have appeared of the great master of paleontology this recent

memorial of Pompeckj, pupil, associate and intimate friend of the

late Geheimrath, is the most complete, and most satisfactory. This.

is not a eulogy of von Zittel, but a plain and sufficient account of

his career, with his achievements mentioned in such a way that they

speak for themselves, and with the light so distributed upon his

personal traits, his ambition, energy, concentration — and above all

upon his aptitude as a teacher, helpful, inspiring and commanding of

respect,— that his character is revealed naturally before us without

addition or subtraction, as must be acknowledged by anyone who

had the good fortune to know him well Dr. Pompeckj has told us:

1 ; ;

Pompeckj, J. F. Karl Alfred von Zittel: Ein Nachruf. Paleontogr aphitdy.

vol. L., 1904.

No. 453.] NOTES AND LITERATURE. 679

in measured and dignified language much that is good to know and

to remember in connection with the life-work of one of the torch-

bearers of science, but there is one respect in which we would like

to have been told more. Zittel as a teacher, text-book writer,

ardent collector and museum administrator, Zittel as an investigator

thirsting for scientific discovery — in all these capacities he is pre-

sented to us; but enough has not yet been said in regard to him as

a philosopher, as a theorizer upon the vast store of empirical knowl-

edge of which he was the possessor. He was an excellent systema-

tist, and the faculty of coórdination was developed in him to a

remarkable degree. Though he discovered no new laws of natural

history, yet he had faith in the discovery of others, and he believed

in certain principles and methods of drawing philosophical con-

clusions, as sincerely as he disbelieved in certain others, nor did he

always insist upon his own personal judgment, often deferring to the

opinions of colleagues in whom he had confidence. On such matters

as these we should eagerly welcome more light.

BOTANY.

Maple Sap Flow..— This paper, by Messrs. Jones, Edson, and

Moore, and edited by J. H. Hills, Director of the Agricultural Exper-

iment Station of Vermont, is unusual in two ways. It is a very

good paper, giving the carefully considered results of experiment and

observation sufficiently extended to justify general conclusions. In

the second place the paper is unusual for it is the first on this sub-

ject since Clark's papers in 1873 and 18742 As I have said else-

where; it is surprising that American botanists at the Agricultural

Experiment Stations in the states where maple-syrup and maple-

sugar making is an important industry have not carefully studied

the phenomena, at least from an economic standpoint. The present

paper is written both from the economic and from the physiological

standpoint, and the plant physiologist will find iu it data which he

' Bulletin Vermont Agric. Exp. Station, No. 103, Dec., 1903. ;

* Report Mass. Agric. Coll., 1873-4; Report Mass. State Bd. Agric, No. 22,

1874.

? Text Book of Plant Physiology, 1903.

680 THE AMERICAN NATURALIST. [Vor. XXXVIII.

has long wished to have. The paper is of such length (184 pages,

8vo) that even an abstract may easily be too long for this journal.

After an introduction, stating the importance of the maple-sugar

industry, the process of sugar-making, and former investigations, the

authors proceed to describe their plan of work and the scope of their

investigations. A few pages are given to the structure and general

physiology of the inaple. In sixty pages the authors discuss (1) the

water and gas contents of the maple at different seasons, comparing

it with other trees; (2) pressure, positive and negative, at different

seasons, in different parts of the tree, and the direction of pressure

and sap movement; (3) temperatures, comparing internal with exter-

nal. Itis to be hoped that this study of external temperatures will

be supplemented by further ones. The authors studied only air

temperatures, but it is obvious that the temperatures of the soil must

have at least an equal, if not a greater, bearing on the phenomena

of sap flow in the spring than the air temperatures. As others have

shown, the roots begin to be active much earlier than the aérial parts

of trees and shrubs growing in the temperate zone. This early re-

sumption of active life in the roots, and the energetic absorption of

water from the soil, depend more directly on soil temperatures

than on air temperatures. Hence, if we are thoroughly to under-

stand the process of periodic sap flow, we must know all the condi-

tions, not merely those in the air and in the aérial parts of a tree.

It may not be too much to suggest also that studies of the water-

content of soil and air before, during, and after sugaring time would

be a valuable addition. This is intimated by the authors, though the

subject is not pursued further.

The remaining seventy pages are occupied by a discussion of eco-

nomic problems and by tabular reports of weather conditions, etc.

G. Je E

The Journals.— 77e Botanical Gazette, May :— Sargant, "ss.

Evolution of Monocotyledons”; Smith, “The Nutrition of the Egg

in Zamia”; Opperman, “A Contribution to the Life History of

Aster”; Cardot and Thériot, “New or Unrecorded Mosses of North

sucia ”; Livingston, “ Physical Properties of Bog Water "i ME

William M. Canby”; Ramaley, “ Anatomy of Cotyledons."

The Bryologist, May :—Grout, “The Peristome — VI "; Holzinger,

Rhacomitrium Fletti, n. sp.” ; Holzinger, “ A Bryologist’s Glimpse

into Geological History"; Harris, “Lichens —Collema and Lept

No. 453.] NOTES AND LITERATURE. 681

gium"; E. G. Britton, “Notes on Nomenclature — III" ; Grout,

“The Specific (?) Value of the Position of the Reproductive Organs

in Bryum " ; and a number of short notes by various persons.

The Fern Bulletin, April : — Curtiss, “The Fern Flora of Florida”;

Eaton, “The Genus Equisetum in North America—XVII" ; Clute,

“New or Rare Ferns from the Southwest”; Eaton, “ Prelims

list of Pteridophyta Collected in Dade Co., Florida, during Novem-

ber and December, 1903”; Kalbfleisch, “ Polystichum acrostichoides

and some Insects that infest it"; Burnham, * Ferns of Ann Arbor,

Mich." ; Clute, * Raynal Dodge” (with portrait) ; and short notes by

various persons.

Journal of Mycology, March: — Morgan, “ A New Melogramma ”

Cockerell, “Some Fungi Collected in New Mexico”; Dudley and

Thompson, * Notes on Californian Uredinez and Descriptions of

New Species "; Kellerman, “ Ohio Fungi, Fascicle IX ?; Kellerman,

“Minor Mycological Notes — III”; Kellerman, “ Index to Uredine-

ous Culture Experiments, with List of Species and Hosts for North

America —I” (concluded); Kellerman, “Notes from Mycological

Literature—IX ”; and Kellerman, “ Elementary Mycology.”

Journal of the New York Botanical Garden, May :—MacDougal,

“ Botanical Explorations in the Southwest "; Broadhurst, “The Pro-

tection of Our Native Plants.”

The Ohio Naturalist, May : —Claasen, “List of the Mosses of

Cuyahoga and Other Counties of Northern Ohio”; and Schaffner,

“Deciduous Leaves.”

The Plant World, April: — Safford, “Extracts from the Note

Book of a Naturalist on the Island of Guam — XVII”; Broadhurst,

“Nature Study as a Training for Life" ; Nehrling, ^ The Beginning

of Spring in Florida — I."

The Plant World, May :— Morris, “The Bush Morning Glory”

Safford, * Extracts from the Note Book of a Naturalist on the Island

of Guam — XVIII"; Nehrling, “The Beginning of Spring in

Florida—II." ;

Rhodora, May : — Parlin, “Some Casual Elements in the Flora of

Western Maine"; Pease, “Preliminary Lists of New England

Plants—XV "; Sanford, “Occurrence of Verbena stricta and Heti-

anthus mollis in Mass.” ; Hill, “Note on the Polygamy of Chionan-

thus”; Woodward, “Two Noteworthy Plants of New Haven, Ct.” ;

682 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Robinson, “ Ste//aria glauca established in the Province of Quebec”;

Knight, “Some Plants New to the Flora of Maine”; and Eaton,

“Note on Equisetum pratense.”

Proceedings of the Society for the Promotion of Agricultural Sci-

ence, Vol. 25: — Hansen, “ Possibilities of the Western Sand Cherry”;

Pammel, “Some Unusual Fungus Diseases in Iowa during the Sum-

mer of 1903”; King, “ Promising Methods for the Investigation of

Problems of Soil and Plant Physiology, and Some Lines of Investi-

gation to which they are Adapted.”

Bulletin of the Torrey Botanical Club, April: — Peck, “New

Species of Fungi”; Evans, “Hepatic of Puerto Rico — IV ";

Morgan, “ Polarity and Regeneration in Plants.”

Bulletin of the Torrey Botanical Club, May:— Nelson, “New

Plants from Wyoming — XV " ; * Code of Botanical Nomenclature y

(in three languages); Watterson, “The Effect of Chemical Irrita-

tion on the Respiration of Fungi.”

Torreya, April :—Underwood, “ Early Writers on Ferns and their

Collections— II ”; Rusby, * William Marriott Canby ” ; MacKenzie,

“ Notes on Evening Primroses ” ; Cockerell, “ Mutations and Ferns E

Sumstine, “ A New Hydnum.”

Torreya, May: — Eggleston, “A Canoe Trip on the St. Francis

River, Northern Maine ” ; Kobbé, * Notes on the Local Flora" ; and

Britton, * Viburnum molle Michx."

Zoe, April: — Brandegee, ^A Collection of Mexican Plants"; .

Greenman, “ New Species of Mexican Plants”; T. S. Brandegee,

"Palms of Baja California” ; and Katharine Brandegee, “ Notes

on Cactez."

Notes. — A “Flora of Los Angeles and Vicinity," by Abrams

(Stanford University Press, April s, 1904) forms an octavo volume

of 474 pages, and contains analytical keys and full descriptions of

the Spermatophyta of the coast slope of Los Angeles and Orange

Counties, California. The Orders are arranged in the Engler and

Prantl sequence, and the Neo-American nomenclature is adopted —

with synonymic citation where the generic name is unfamiliar to the

ordinary reader.

A Catalogue of the Bryophyta and Pteridophyta of Pennsy Ivania,

by the late Professor Porter, edited by Dr. Small (Boston, Ginn &

No. 453.] NOTES AND LITERATURE. 683

Co., 1904) forms an octavo of 66 pages. Each entry is followed by

habitat and distribution data by counties.

A check-list of the higher plants of Hamilton County, Ohio, and a

list of medicinal plants growing in the vicinity of Cincinnati, both by

Aiken, form no. 4 of Vol. 20 of the Journal of the Cincinnati Society

of Natural History.

A few separates of the several chapters of Vol. 5 of the publica-

tions of the Harriman Alaska Expedition, dealing with the crypto-

gams, have been distributed by the authors.

A dictionary of plant names of the Philippine Islands, by Merrill,

forms a bulletin from the Philippine Bureau of Government Labora-

tories.

A paper on the flora of St. Andrews, New Brunswick, by Fowler,

is published in “Contributions to Canadian Biology," — a supple-

ment to the 32d Annual Report of the Department of Marine and

Fisheries, Fisheries Branch, of Canada.

Fascicle 2 of’ Millspaugh's *Plantz Yucatanz," forming Vol. 3,

no. 2, of the botanical series of Publications of the Field Columbian

Museum, deals with Composite, by C. F. Millspaugh and Agnes

Chase, and is admirably illustrated.

The first fascicle of Vol. 3 of Halacsy’s " Conspectus Flore

Grecex,” recently issued, covers Lentibulariacez to part of Cyper-

acez.

A general comparison of the Alpine floras of Australia and Eu-

rope is given by Weindorfer in ZZe Victorian Naturalist of Septem-

ber las

Forbes and Hemsley's enumeration of the plants of China, etc.,

forming Vol. 36 of the Journal of the Linnean Society (Botany), has

reached the 18th part, dealing with parts of Cyperacez and Gram-

inez.

Warburg and de Wildeman have begun the publication of an

account of Ficus as represented in the Congo district, in the Annales

du Musée du Congo, the first fascicle being issued in January, 1994.

The newly established Records of the Albany Museum, of Grahams-

town, is in part devoted to South African botany.

Part 10 of Hough’s American Woods, comprising nos. 226 to 250,

represents chiefly western and southwestern species, — perhaps the

684 THE AMERICAN NATURALIST. [Vor. XXXVIII.

most interesting being Cereus giganteus. Like earlier fascicles, this

is accompanied by leaf and fruit keys and indexes for the entire

issue, and a systematic account of the species now distributed.

An excellent winter key to the genera of woody plants, wild or

cultivated, in New York State has been issued by Wiegand and Fox-

worthy, of Cornell University.

A polyglot Code of Botanical Nomenclature, by a committee of

the Botanical Club of the American Association for the Advance-

ment of Science, has been separately printed from the May Bulletin

of the Torrey Botanical Club.

The sixth fascicle of Dalla Torre and Harms’ “Genera Siphono-

gamarum” includes Gentianacez (in part) to Acanthacez (in part),

— genera 6492—7927.

The roses of Pecos, N. M., are discussed by Cockerell in Vol. 56,

part 1, of the Proceedings of the Academy of Natural Sciences of Phil-

adelphia.

Figures of Jamesia Americana are published by Späth in Garten-

fora of May 1.

À paper on Echeveria, by Berger, is published in Gartenflora of

April 15. :

Professor Greene contributes a paper on Some Canadian Anten-

nares to The Ottawa Naturalist tor May.

Gaylussacia resinosa is figured in Vol. 4, fascicle 7, of Zcones Select

Horti Thenensis.

Abronia is recorded for the Tertiary flora of Europe by Laurent

the Comptes Rendus ot the French Academy, of April 18.

Kreenzlin’s

“Orchidacearum Genera et Species,” which had

reached the end of Part I of Volume 2, has been broken off.

i Several tropical American orchids are added to the known flora of

Florida by Ames in a leaflet of Vol. 17 of the Proceedings of the

Biological Society of Washington, issued May 19th.

An interesting article by Miller on cultivated orchids, well illus-

trated, is contained in Country Life in America for June.

A preliminary synopsis of the Southern California Cyperaceæ, by

Parish, is in course of publication in the Bulletin of the Southern

California Academ Ly Of Sciences,

No. 453.] NOTES AND LITERATURE. 685

The leaf characters of Pinus are discussed by Masters in No. 248

of The Fournal of the Linnean Society, Botany.

An important paper on ferns of central China, by Christ, is pub-

lished in No. 173 of the Bulletin de Académie Internationale de

Géographie Botanique.

Anaccount of the liverworts found about Chapel Hill, N. C., is

given by Coker in the January Journal of the Elisha Mitchell Scien-

tific Society.

Part 2 of Grout's “ Mosses with Hand Lens and Microscope” was

issued in May, and extends from Ceratodon to Tortula.

A list of the lichens and mosses of Montana, by W. P. and C. W.

Harris, forms Bulletin No. 7, Biological Series, of the University of

Montana.

An account of the diatoms of the Gulf of Naples, by Balsamo, is

being published in the Bollettino della Società di Naturalisti in

Napoli.

A study of the plankton of Lake Winnebago and Green Lake, by

Marsh, forms Bulletin r2 of the Wisconsin Geological and Natural

History Survey.

A monographic account of 77iphragmium, by Milesi and Traverso,

is published in Annales Mycologici, of March.

An illustrated account of Séictis Panizzei, and its disease of the

olive, is given by Brizi in Z'Z/a/ia Agricola of April 15.

An account of Colletotrichum gleosporioides and its diseases of

Citrus is published by Rolfs as Bulletin 52 of the Bureau of Plant

Industry of the U.S. Department of Agriculture.

The report of the botanist of the Connecticut Agricultural Experi-

ment Station for 1903 is devoted to diseases of plants cultivated in

that State, and the inducing fungi.

Fungous diseases of fruits in Michigan are discussed by Longyear

in Special Bulletin No. 25 of the Experiment Station of that State.

The fungous parasites of Hevea form the subject of No. 34 of the

Notizblatt des k. botanischen Gartens und Museums zu Berlin, by

Hennings.

An exhaustive paper on the yeasts which ferment milk-sugar, by

Heinze and Cohn, is published in the Zeitschrift für Hygiene und |

Lnfectionskrankheiten of March 31.

686 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Popular resumées of the plants used for various economic pur-

poses are being published by Lagermann in current numbers of the

Fournal of the Columbus Horticultural Society.

'The use of plants in controlling and reclaiming sand-dunes is con-

sidered by Hitchcock in ZuZ/etin 57 of the Bureau of Plant Industry

of the U. S. Department of Agriculture.

A paper by Sherman on gutta percha and rubber of the Philip-

pine Islands is published as a bulletin from the chemical laboratory

of the Bureau of Government Laboratories of the Philippines.

An account of Senecio jacobwa as the causative agent of hepatic

cirrhosis of horses and cattle, is contained in the Annual Report of

the Department of Agriculture of New Zealand, for 1903.

An economic account of annual flowering plants, by Corbett, is

published as Farmers’ Bulletin No. 195 of the U. S. Department of

Agriculture.

An economic account of the date palm, by Swingle, forms Buletin

53 of the Bureau of plant industry of the United States Department

of Agriculture.

Noriega contrasts true and false Jalap in Vol. 6, No. 3, of the

Anales del Instituto Médico Nacional of Mexico.

An account of poison ivy is contributed by Brownell to Country

Life in America for June.

A readable account of English herbals, by Agnes Robertson, is

contained in the Popular Science Monthly, for May.

| One of the most suggestive of recent horticultural books is “ The

Tree Doctor," by John Davey (Akron, Ohio).

An important paper on root trichomes of Pteridophytes and Angio-

sperms, by Leavitt, constitutes Vol. 31, No. 7, of the Proceedings of

the Boston Society of Natural Histor Y.

A PEN by Hus, on spindle formation in the pollen-mother-cells of

Cassia tomentosa, forms the concluding number of Vol. 2 of the third

series of Proceedings of the California Academy of Sciences, Botany.

Deformation of plants through external influences is discussed by

. Reinke in the Botanische Zeitung, Abtheilung 1, of May 1.

The connection of leaf structure and environment, in Z teris

aquilina, is considered by Boodle in No. 248 of The F ournal of the

Linnean Society, Botany. :

No. 4553] NOTES AND LITERATURE. 687

Popular botanizing has received an admirable aid in Miss Niles's

* Bog-trotting for Orchids" (New York, G. P. Putnam's Sons, 1904),

a book for lovers of the country, written by a nature lover, in which

is included a compiled synopsis of the orchids of New England.

Plain and tricolor half-tones in large number are interleaved with the

text.

An illustrated account of the botanical garden of Carlsaue (1580),

the first of its class in Germany, is published by Jungein Gartenflora

of April 1.

A biographic sketch of Askenasy, with portrait, appears in the

Berichte der deutschen botanischen Gesellschaft of April 11.

A portrait of Professor Rothrock is contained in Forestry and

Irrigation for May.

A biozraphic sketch of Karl Schumann, with portrait, is contained

in the April Monatsschrift für Kakteenkunde.

D fr

Aes

VEU MA

CORRESPONDENCE.

To the Editor of the American Naturalist :

SIR : — Apropos of Dr. Theodore Gill’s letter in the March num-

ber relative to early collections of the vernacular names of animals,

it is interesting to recall that Thomas Gray, “the English poet who

has written less and pleased more than any other,” was an accom-

plished naturalist for his time, and busied himself during the last ten

years of his life in compiling, amongst other notes, a voluminous

catalogue of the familiar names of plants and animals. Not only

the common English synonyms are given of Linné’s species, but

also their equivalents in more than a score of languages, some of

the citations being from remote and little-known tongues.

These lists were written down by Gray in his interleaved copy of

the tenth edition of the Systema Nature, and portions of them, some

25 pages in all, were published in the second volume of Mr. T. J.

Mathias’s edition of Gray’s Works, which appeared in 1814. Other

selections from the same source, with facsimiles of some of his

drawings, have recently been published by Mr. Charles Eliot Norton,

who now possesses Gray’s original copy. No one can take up this

little booklet ! without feeling grateful to Professor Norton for having

placed this “monument of Gray’s learning and industry " within

general reach. It is stated by the editor that these annotations, if

printed, would form a volume at least equal in size to one of Linné's,

and that the light they throw on the poet's occupations and interests

during his latter years helps us to a "more just appreciation of his

character and his acquisitions."

Those interested in the derivation of the common names of

animals may find it worth while to consult a paper by J. W. Gibbs

on the “Origin of the Names of Beasts, Birds, and Insects,” pub-

lished in vol. xli, of the American Journal of Science (pp. 32-39;

1841).

C. R. EASTMAN.

1 The Poet Gray as a Naturalist with Selections from his Notes on the

Systema Nature of Linneus etc. Boston, Charles E. Goodspeed. 1903. PP- 67..

Nos. 451-452 were mailed September 30, 1904.

689

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VoL. XXXVIII, No. 454. OCTOBER, 1904

THE

AMERICAN

NATURALIST ^

DEVOTED TO THE NATURAL SCIENCES a

IN THEIR WIDEST SENSE |

CONTENTS

~ L The Anatomy of the Coniferales (concluded) PROF. D. P.

The American Naturalist.

ASSOCIATE EDITORS:

j. A. ALLEN, PH.D., American Museum of Natural xs a York.

E. A. ANDREWS, PH.D. » Johns Hopkins University, Bal:

J WILLIAM S. BAYLEY, FH.D., Coig University, geste

g DOUGLAS H. L C aetna PH. D., Stanford eee:

e RH B., Cornell University, ft

— WILLIAM M. DAVIS, ME, Harvard = Cambridge.

M ALES HRDLICKA, M.D., U.S. National Museum, Washington

D.S. JORDAN, LL.D., Stanford University.

— CHARLES A. KOFOID, Pu.D., University of California, Berkeley.

= ARNOLD E. ORTMANN, PH.D., Carnegie Museum, Pittsburg.

eal.

2j A

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

mo LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington.

W ed : |, S.D., Missouri Botanical Garden, St. Louis

as HENRY B. WARD, Pap, University of Nebraska, Ti

x WILLIAM M. WHEELER, Pu. D., American Museum of Natural History

i New York.

: Tue AMERICAN Natura ist is an illustrated monthly magazine

of Natural History, and will aim to present to its readers the leading _

facts a discoveries in Anthropology, General Biology, Zoology,

‘Botany, Ute Geology and Physical Geography, and Miner-

alogy : and graphy. The contents each month will consist of

: est | | cientific questions of the .

cri ritical. reviews of recent literature, bad a quarterly record. oo

appointments, retirements, and deaths. aa

st: who have anything interestne to say are invi: ee es ee

mh s a es ll endeavor to select ee a

THE

AMERICAN NATURALIST.

VoL. XXXVIII. October, 1904. No. 454.

THE ANATOMY OF THE CONIFERALES.

D. P. PENHALLOW.

(Continued from page 554)

RESIN PASSAGES.

Distribution and Taxonomic Value.

PRANTL (44, p. 37) States that resin passages occur in the

wood of “most Abietinez, namely, Pseudotsuga, Picea, Larix,

Pinus and Abies firma.” This statement requires some modifi-

cation in detail, especially with respect to the last named genus,

and in order to make the results of the present studies clear it

will be expedient to discuss d the distribution of the

resin cysts and the resin passages

.. The first species to which our attention may be directed is

Tsuga mertensiana. This is the only species | of the genus in

_ which definite resin cysts are to be found. - Such structures are

never numerous, and they take the form of short rows of con-

tiguous cysts in the initial layer of the summer wood of distant

&rowth rings. Longitudi y the c mits,

692 THE AMERICAN NATURALIST. [Vor. XXXVIII.

through the entire longitudinal growth of the season, at least.

There is no obvious alteration either in the position or volume

of the resinous contents of the isolated resin cells which lie on

the outer face of the summer wood. The constancy with which

these structures occur gives to them a definite value for the

recognition of the species, and permits us to differentiate it from

T. caroliniana on the one hand and from the remaining three

species on the other.

In the genus Abies only four species out of eleven show resin

cysts. These are A. bracteata, A. nobilis, A. concolor and A.

firma. Referring again to Prantl's observation (44, p. 37), it

must be pointed out that his statement with respect to the

occurrence of resin passages in A. firma requires modification

in detail, in so far as these structures are not passages but cysts ;

while he also appears to have overlooked the occurrence of simi-

lar structures in the three other species mentioned. In all of

these cases the cysts are contiguous and disposed in tangential

rows of considerable length, either in the summer wood (A. con-

color and A. nobilis), in the outer spring wood (A. firma), or in

both the spring and summer wood (A. bracteata). Such varia-

tions appear to be of no specific value, conforming as they do to

similar variations in the zonate distribution of the resin cells.

It is found, however, that in only one case (A. concolor) are these

cysts associated with isolated resin cells. In the three other

cases the resin cells are entirely wanting, a relation which is.

strongly suggestive of their replacement by the cysts.

Sequoia sempervirens is the only species of that genus which.

develops resin cysts in the secondary wood, though Jeffrey (24)

has shown that such structures are normal to the primary wood

zone of S. gigantea, and not elsewhere. As already shown, such

cysts are much more highly organized than those of either Tsuga

or Abies, though they are similarly contiguous and even coales-

cent, and form extensive tangential rows in the initial layer of

the spring wood of distant growth rings. They form a much

more prominent feature than in any of the preceding species,

because of their generally larger size and the greater extent of

the series in which they lie. Unlike Abies, however, there

appears to be no diminution either in the number or the extent

No.454] ANATOMY OF THE CONIFERALES. 693

of the prominent resin cells which are often intimately associ-

ated with the cysts.

The normal course of development for such cysts as thus

described is subject to special alteration under conditions which

involve an unusual stimulus to growth, and under such circum-

stances they may become definitely associated with, or may even

be regarded as indicative of, pathological conditions. Thus

Anderson (1, p. 28—29) has shown that such cysts are definitely

developed in association with the formation of Witches' brooms

in Adies firma. Under such circumstances the cysts become

much larger, more distant and more numerous than in normal

growth, but they form well defined tangential rows in the earlier

spring wood of successive growth rings. In the development

of such secondary features the cysts manifestly exhibit a distinct

approach to that higher type of structure and distribution which

is exhibited in Picea. In the following year Anderson (2, p. 336)

further showed that while resin cysts are absent from the nor-

mal wood of A. balsamea they do arise under the influence of

the special stimulus connected with the formation of tumors

produced by the action of Æcidium elatinum. He furthermore

points out that such cysts attain their greatest development and

largest number in the region of greatest stimulation, 7. ¢., in the

middle of the tumor, decreasing above and below until they

eventually become pointed and finally disappear between four

tracheids * which, in their meristematic condition, probably

function as epithelial cells." It is unfortunate that the histo-

logical details of these cysts and their endings are not given,

since such facts would serve to throw much light upon the rela-

fion of the cysts to similar structures in normal tissues, but

there is no reason to suppose that they differ in their essential

structure from those which occur in the normal tissues of the

same or other species. The tracheids above referred to are un-

doubtedly parenchyma tracheids, and it is probable that further

examination would show that they ultimately replace the resin

cells remaining over after the disappearance of the cyst proper.

More recently Jeffrey (24) has contributed an important

paper on the anatomy of Sequoia in which he brings out sev-

eral facts of considerable value. He shows that resin cysts may

694 THE AMERICAN NATURALIST. [Vor. XXXVIII.

arise in the roots of Aes balsamea which have been injured,

while they may also be produced experimentally by injury, thus

confirming the observations of Anderson that they may be trau-

matie in their origin. The most significant facts, however,

relate to the normal occurrence of such cysts in Sequoia. He

shows in the first instance that they are absent from the wood

of the first year's growth in Seguota sempervirens, while they

are present for the same period of growth in S. gigantea, though

absent from the growth of later years. In both species they

arise in the earlier spring wood. Jeffrey concludes that the

tangentially disposed resin cysts of Seguota sempervirens repre-

sent the result of injury, and he would apply this rule to all

similar cases in the various species of Abies and Tsuga. :

Some years since, De Bary (9, pp. 490 and 495) formulated

the law that “resin canals . . . . occur in the ligneous bundles of

the same Abietineze which possess horizontal canals in the med-

ullary rays." This is a law of very great constancy, and, as

recently shown by Penhallow (39), it is applicable without

exception to all living species. But as the same author (41,

P. 42) has shown more recently, Segauoza burgessii from the

Eocene of the Northwest Territories offers a remarkable excep-

tion to this law, since it shows well defined resin passages in the

medullary rays, but without corresponding structures in the vas-

cular bundles.

As presented by existing species, Pseudotsuga, Larix, Picea

and Pinus, without exception, show resin passages in both the

radial and longitudinal positions. In transverse section they are

scattered throughout, sometimes appearing chiefly in the sum-

mer wood, sometimes chiefly in the spring wood, or again about

equally in the two regions, and they rarely conform to the pre-

cise law stated by De Bary (9, p. 495), that “they lie scattering

in a ring in the external region of every annular layer." he

constancy of their occurrence in the four genera mentioned in-

volves very few features which call for special comment. In

Pseudotsuga and Larix the resin passages are scattering.

Sometimes they not infrequently unite in pairs so as to form |

Short, tangential series and they thus approach the type of

Tsuga or Abies, while yet again they may become definitely Be o

No. 454] ANATOMY OF THE CONIFERALES. 695

lated and scattering, thereby approaching the distribution of

Picea and Pinus. In Larix occidentalis the tendency to a primi-

tive form of distribution is expressed in the formation of a tan-

gential zone essentially similar to that of Tsuga mertensiana.

In both Pseudotsuga and Larix there is an obliteration of resin

cells from all parts of the structure except the extreme outer face

of the summer wood. In Picea, however, without exception,

there is a complete obliteration of all resin cells except such as

enter into the structure of the resin passages, and this is directly

correlated with a higher type of structure in such passages.

In the genus Pinus, as already shown, the resin passage

reaches the highest degree of organization in all respects. It

shows little if any tendency to those primitive associations which

are expressed in the formation of tangential series, while it has

entirely replaced the isolated resin cells which are never to be

found in that genus.

If, then, we ask what value such structures have for taxo-

nomic purposes we find them to be of well defined importance.

It has already appeared that in Tsuga the occurrence of resin

cysts is of well defined value for specific differentiation, and the

same rule is also applicable to Sequoia sempervirens and to four

species of Abies. In the higher Abietineze, inclusive of Pseudo-

tsuga, Larix, Picea and Pinus, the invariable association of resin

passages in the wood and in the medullary rays not only serves

to separate these genera from all those in which resin cysts only

may occur, but it differentiates them absolutely from all the

remaining genera. Such association therefore constitutes a

feature of great value. More particularly, the thin-walled epi-

thelium of Pinus at once separates that genus from the other

three, which are invariably characterized by thick-walled epithe-

lium, Such generic differentiations are greatly emphasized by

the occurrence of thyloses. These are typically developed in

Pinus, where they are always thin-walled and almost invariably

present. They are therefore of definite value as supplementing

other features previously described. In the other genera, how-

ever, their presence in either the cyst or the resin passage,

where they are generally thick-walled, is of so sporadic a nature

as to give them no definite value, and we therefore find that for

specific diagnoses such structures may be neglected.

696 THE AMERICAN NATURALIST. [Vor. XXXVIII.

PHYLOGENETIC.

We are now in a position to present an answer to the question,

* How are the resin passages related to the phylogeny of the

Coniferales?” In order to present an intelligent answer to this

question it will be necessary to recall the facts already discussed

in connection with the resin cells and bring them into relation

with our discussion of the resin passages.

In the genus Sequoia it has been shown that the general

course of development of the resin cells is essentially the same

as in Cupressus, etc., while it has also been shown that the

genus presents in other respects a somewhat remarkable devia-

tion. Of the two existing species, both show the distribution

of the resin cells to be of the typically primitive form, x

scattering. Nevertheless there are also in Seguota sempervirens

definitely organized resin cysts but without exhibiting the tran-

sitional form of a zonate disposition. Among fossil representa-

tives Penhallow (41, p. 41) has shown precisely the same fea-

ture to be present in S. langsdorfii. This is the less remark-

able, however, because that species is undoubtedly the ancestral

form of, and practically identical with, S. sempervirens. The

fact made clear by Jeffrey (24, p. 457), that resin cysts occur in

the first annual ring of vigorous branches of adult trees, as well

as in the roots of S. gigantea, also tends to make it apparent

that the genus presents a very striking advance upon even the -

type presented by Juniperus, since the aggregation of resin cells

and the formation of cysts from them has arisen abruptly and

without the transitional forms presented by Juniperus and Taxo-

dium. While, therefore, Sequoia is obviously related to Thuya

and Cupressus on the one hand, it is on the other hand related

to such types as Abies. In this sense it may be regarded as

the terminal member of a developmental series embracing the

Taxodiinz, Cupressineze, Taxoideze as follows : —

Taxus and Torreya.

"Thuyopsis.

Cryptomeria.

Podocarpus.

Aua o oc

No. 454] ANATOMY OF THE CONIFERALES. 697

Cupressus.

Thuya.

Libocedrus.

Taxodium.

Juniperus.

Sequoia.

9 0 MN AN

—

In the Abietinez a new series is presented. This is not in

any sense strictly coterminous with the first, but the two appear

to make a fault, as it were, whereby there is a lateral displace-

ment, but of such a nature that Sequoia still serves as the con-

necting link. Within the eleven species of Abies investigated

three important phases are presented: (1) Resin cells scattering

on the outer face of the summer wood, (2) resin cells grouped

and forming cysts, and (3) resin cells entirely wanting. View-

ing these phases in the order given, it is to be observed that in

those four species which develop cysts only one shows isolated

resin cells, and it is probably correct to interpret the variations

noted as expressions of developmental phases in such a way that

the occurrence of cysts represents the highest position. The

genus Tsuga is closely related to Abies in the occurrence of iso-

lated resin cells on the outer face of the summer wood, as also

in the formation of resin cysts, but it obviously occupies an infe-

rior position because (1) of the greater abundance of resin in the

individual cells, and (2) the occurrence of definite aggregates of

resin cells without the formation of cysts. This series is directly

extended by those genera in which definite resin passages replace

the simple cysts, since the latter are convertible into the former

by easy and natural transitions. Both Pseudotsuga and Larix

occupy equivalent positions because they not only present resin

passages of an equal degree of development, but they show a

survival of the isolated resin cells on the outer face of the sum-

mer wood. Their affinities are therefore directly with Abies

and Tsuga on the lower side, but on the upper side their alli-

ance is with Picea, which presents a very similar though some-

what higher organization of the resin passage and a complete

obliteration of the isolated resin cell. Yet again, the structure

of the resin passage in Picea at once connects that genus with

698 THE AMERICAN NATURALIST. (Vor. XXXVIII.

Pinus, in which the most complete development is attained, and

it therefore terminates the series upwardly.

Having special reference to the particular forms of the secre-

tory reservoirs, and leaving out of account all other considera-

tions than their particular evolution, it is possible to indicate the

general sequence of the genera and, to a more limited extent, of

their species, as follows : —

I. Tsuga caroliniana.

* — mertensiana.

Abies bracteata.

" prina.

* — nobilis.

* — concolor.

Sequoia.

. Pseudotsuga and Larix.

Picea.

Pinus.

From this it is manifest that Sequoia is superior to Tsuga and

Abies but inferior to Pseudotsuga, Larix, etc. But if we now

view the general phylogeny with reference to the entire course

of development of the resin cells and the resin passages, the

relations just explained must be modified with reference to the

particular position of Sequoia, and the sequence would then

become : —

. I. Thuyopsis and Cryptomeria.

2. Podocarpus.

3. Cupressus

4. Thuya

5. Libocedrus.

6. Taxodium.

7. Juniperus.

8. Sequoia.

9. Tsuga.

IO. Abies.

II. Pseudotsuga and Larix.

I2. Picea.

I3. . Pinus.

No. 454] ANATOMY OF THE CONIFERALES. 699

But it may assist in the general argument to view this question

from another standpoint. Regarding the resin cells and the

secretory reservoirs as falling within a definite series, we may

apply to the various forms of distribution and to the various

grades of resin reservoirs arbitrary values of such a nature as to

represent our conception of their relative positions in the scale

of development as expressed by percentages, thus: —

Resin cells scattering. ; ; ; ; 25.0%

A "* -zonate : i ; : i rv. e

xe Qporodpeg : ; ; j : 50.0 “

a « on the outer face of the inner wood,

Pseudotsuga and Larix. ‘ e

k * on the outer face of the summer

wood, as in Abies (partial only), o"

« * wholly wanting . ; go”

Resin cysts, as in Tsuga, Abies and Sequoia . 70.0 *

Resin passages with constrictions, as in Pseudo-

tsuga, Larix and Picea i : ; i 80.0 *'

Resin passages without constrictions and of the

highest type of organization, as mmus . 100.0 “

We obviously have two subordinate series here, which for con-

venience may be regarded as coterminous, but which as already

shown are “faulted” in such a way that the grouped resin cells (*)

and the resin cysts (T) jointly represent the point of divergence for

two separate courses of development, the latter continuing up-

ward, while the former descends and thereby represents degra-

dation. These features are best exhibited graphically, and the

accompanying curves clearly show how, on the one hand, resin

cysts and resin passages directly result from special modification

of cell aggregates, while on the other hand, from the same start-

ing point, there arises a course of degradation which finally

results in the complete obliteration of the. resin cell as an inde-

pendent structure.

The facts thus far set forth have thrown important light upon

the general course of development of certain anatomical features,

and they also show the general course of development for genera

700 THE AMERICAN NATURALIST. [Vor. XXXVI

and species with reference to particular structures. They do

not, however, convey any information with respect to the origin

bo rd

SERRES, aio

nai ~—

P ojo

a SIE

DOLES 1 n mlt.

O 49 E

E ZI / E:

[o ©

ANRE Coreen ae Maney A

bo jm

Dr ISI QUEUE rz

; p a

Curve for Resin PaSsages

50]———— — — —— — |

Edi i

bpopup eegne E YT ee

p poH

Epio sls sf 4

25 Curve for Resin |Cells

Een 0 oor] \

nn | s. *

atone x.

BOREAM L

ems P

ju. t

Curve ch t at

d 1

EH blitera-

tion of resin cells.

of the phylum as a whole, or the relations of the particular genera.

and species from the standpoint of collective data, and such a-

No. 454.] ANATOMY OF THE CONIFERALES. JOI

discussion will be more appropriately reserved for the general

summary. There is, however, one feature arising out of recent

investigations which calls for consideration at this point, since

certain of the conclusions reached are not in harmony with our

own, the divergence of opinion indicated being the result of

different methods of interpretation.

Jeffrey states (24, pp. 447 and 457) that a// such resin cysts as

occur in Sequoia sempervirens and Abies are of a traumatic nature,

and therefore pathological. To this category he would also doubt-

less assign the corresponding structures of Tsuga. This opinion

appears to be shared by Anderson (1 and 2), and it is also appar-

ently supported by Pierce (42). Both Jeffrey and Anderson

show that the development of such cysts is sometimes definitely

associated with the production of tumors through the operation

of parasites, and that they may also be induced by wounds experi-

mentally produced. The facts they cite show conclusively that

resin cysts may, and often do, arise traumatically, but in such

cases they lie outside the usual course of development.

The occurrence of resin passages in the fundamental tissue

of the Coniferales is a well known fact, as pointed out by De

Bary (9, p. 441) many years since, when he summarized the

general facts in the statement that *all investigated species of

Coniferze, with the single exception of Taxus, have resin pas-

sages or resin reservoirs which vary in distribution and number

according to the species." This statement would include the

leaves and bark and sometimes even the pith of species which

produce neither isolated resin cells nor resin reservoirs of any

kind in the xylem tissue of the stem. It directs attention some-

what forcibly to the fact that while the occurrence of resin

reservoirs in the fundamental tissue is a legitimate inheritance

of the mucilage canals of the Eusporangiate ferns and the

Cycadofilices, as also later of the resin cells of Cordaitales, the

xylem structure is the very last to receive the impress of such

a course of development ; and it is therefore in nowise surpris-

ing that the resin passages do not appear there until a very late

period of development and that their organization can even then

be brought about only through a somewhat prolonged series of

changes which are initiated by the occurrence of isolated resin

702 THE AMERICAN NATURALIST. (Vor. XXXVIII.

cells, much as the formation of mucilage canals may be traced

back to specialized cells which separately have the same function

in the Eusporangiate ferns.

The local occurrence of resin passages in the xylem of the

floral axis in no way invalidates the obvious conclusions to be

drawn from these statements, since it may be readily accounted

for in other ways. Ina structure so unresponsive to influences

which would induce profound alterations as the xylem, it is to be

expected that important structural changes could be effected

only after a prolonged interval during which the fixation of any

particular character would be preceded by a period of sporadic

development within which such character would be liable to

recur under special conditions ; and as such conditions are obvi-

ously of fundamental importance we may inquire somewhat more

fully into their nature and results.

The statement of Prantl (44, p. 35) that ‘‘those genera

which are devoid of resin passages in the wood of young and

vigorous growth later produce single parenchyma elements in

the wood, which contain resin" requires some modification in

view of what Jeffrey has shown in the case of Sequoia and

Abies, as well as what has been shown in the course of the

present studies, and in its more comprehensive and exact form

it should read, *those genera which are usually devoid of resin

passages in the wood, but some species of which may neverthe-

less contain resin cysts in the young and vigorous growth, later

produce single parenchyma elements in the wood, which contain

resin."

Taken by itself this statement as applied to Sequoia and

Abies might be held to indicate that the growth of the first

year represents the most stable structural region of the entire

stem, in the sense that it embodies characters which are most

fully established, and that it will therefore embrace elements

which may be eliminated from the older parts, or which may be

replaced there by degenerate forms only. From this point of

view 1t would be necessary to regard the complex resin passage

as the primitive form of structure from which the cysts, groups

of cells and isolated resin cells have been derived by a process

of progressive degradation. . This view appears to have been

No. 454.) ANATOMY OF THE CONIFERALES. 793

adopted by Jeffrey (24, p. 454), who supports his position by

citing the occurrence of resin passages in the vascular structure

of the peduncle of certain fossil Cycads, interpreting this to

mean that such structures represent a survival of features which

have been obliterated from the structure of the stem. Such a

view does not seem to be in harmony with the facts which our

own studies, have: brought out, to the effect that resin passages

of the type found in the xylem structure are in no sense primi-

tive or vestigial, since they are wholly wanting in the primitive

gymnosperms, and their organization does not arise until a very

late period in the evolution of the highest forms. If our inter-

pretation of observed facts is correct as applied to the origin of

the resin passages, it shows as clearly as one could well expect

a progressive development from the isolated resin cell through

various phases of aggregation to the highest form of structure

as found in Pinus. That there is such a series cannot be

doubted and we must interpret it in one of two ways: either as

progressive evolution or as progressive degeneration. Tous the

arguments all seem to be very emphatic with respect to lending

support to the former view, most especially as all anatomical

data confirm the relative positions of the genera as determined

by the development of the resin passage. But assuming for the

moment that the latter view is the correct one, let us see where

it would lead us. It would first of all necessitate a direct

reversal of the structural sequence, and this in turn would

impose the necessity of placing the genus Pinus at the bottom

of the scale, while those genera like Taxus, Torreya, Agathis,

etc., which have no resin cells even, would be at the top. I

venture to suggest that such a proposition would meet with

instant opposition even from the advocates of the idea that the

resin passage has preceded the other forms of resin bearing

structures. The whole question appears to turn upon our rec-

ognition of what constitutes the most impressionable portions

of the stem structure, and therefore the regions within which

structural changes are initiated. In this connection the evi-

dence of both palzeobotany and recent botany brings out certain

facts with great force and throws them into strong relief. They

are as follows : | |

704 THE AMERICAN NATURALIST. (Vor. XXXVIII.

1. The mucilage canals of the Eusporangiate ferns may be

regarded as the ancestral forms of the resin passages among the

higher plants, but they are obviously the successors of, as they

are derived from aggregates of, mucilage sacs as simple, paren-

chyma cells.

2. Resin passages are wholly unknown in the wood of the

stem of ferns, the Cycadofilices, the Cycads, Cordaites or

Araucarioxylon.

3. Resin cells are known and are abundant in the pith and

bark of Cordaites, but they are absent from its wood.

4. Resin passages are known in the bark and in the pith of

the Cycadacez, of Agathis, Araucaria and of the Coniferales in

general. They also occur in the wood of the peduncles of

Sequoia and Cycas, and in the xylem of the first year's growth

of vigorous shoots in Sequoia and Abies. They likewise occur

in the leaves generally.

5. In Sequoia burgessii from the Eocene, resin passages

occur in the medullary rays, but they do not traverse the wood

longitudinally, though isolated resin cells do occur there. `

From this it would seem that the fundamental tissue is the

most impressionable with respect to the development of these

structures, and that after it we have in the same order the

peduncle of the inflorescences and the wood of the young

shoots, to which latter category would also belong the develop-

ment of resin passages in fasciated stems, and such a sequence

is precisely what we should expect from our knowledge of the

relation which the fundamental tissue bears to other structures.

According to this conception the resin passages may appear in

any. part of the woody structure where growth is sufficiently

VIGOTOMS, but such appearance would be temporary and indica-

tive only of a future course of development which has not as yet

become sufficiently well impressed upon the organism to form à.

permanent feature of it. In other words, the tissue exhibits

what in other cases would be termed *sports." Such structural

forecasts are well known and of frequent occurrence and as

applied to the development of tissues no better example is

afforded than that shown by the central strand of mosses, which

Is generally accepted as prophetic of the future vascular system.

No. 454.] ANATOMY OF THE CONIFERALES. 795

in the sporophyte. They serve to suggest that the law of

mutation as proposed by De Vries finds expression in the evolu-

tion of internal structures as well as in the development of exter-

nal forms. Such cases as Sequoia gigantea, which shows resin

cysts in the wood of the first year and nowhere else, being

replaced later by resin cells, appear to us to show that young

and vigorous growth in general, and therefore the growth ring

of the first year, constitutes a transitional zone within which

many changes of structure wholly apart from the strictly normal

may arise; and such a law would similarly be applicable to the

wood of peduncles. This feature is manifested in the structure

of the medullary ray, the character of the tracheids as exhibited

in transverse section, the genesis of the bordered pits from

spiral tracheids, and in all probability also in the formation of

resin passages in Sequoia and Abies as noted by Jeffrey.

Changes of this nature are to be regarded as tendencies in

development in the direction of higher types of structure

whereby potentialities assume a more Or less definite form.

From this it may be assumed that the primary growth ring is

a zone within which sporadic characters are common, but it is

only in the later rings that the various anatomical characters

become permanently developed and properly express the normal

features of structure and development.

This view is justified not only by observed facts but also by

analogy which shows that as plants ascend in the scale they

exhibit sporadic characters or “ sports ” as tendencies toward the

development of otherwise potential characters. As plants gain

in complexity such tendencies become manifested not simply in

alterations of external form but with respect to particular details

of structure and development. We therefore find ourselves

compelled to conclude that the development of resin cysts and

resin passages from resin cells, and the occurrence of the latter

in the Coniferales, shows that all of these structures are features

in the development of higher types of plants, and it is difficult

for us to accept the statement of Jeffrey that such resin pas-

Sages represent primitive structures and are of the nature of

survivals.

Returning to the question of the traumatic nature of the

706 THE AMERICAN NATURALLIST. [Vor. XXXVIII.

resin cysts in Sequoia, Abies and Tsuga, which Jeffrey appears

disposed to formulate as a general law, it is not clear to us how

this can be made to harmonize with facts coming under our own

observation. It has been shown that such resin cysts occur in

one species of Sequoia, four species of Abies and one species of

Tsuga, yet another species of which also shows them in a poten-

tial form. The same elements appear in each case, vzz. :—

I. The cysts assume a definite form in distribution.

2. They always occupy a definite place in the scale of struc-

tural organization.

3. They are constant features of the same species.

4. They occur at frequent intervals in the same transverse

section, showing them to be repeated at intervals of from one to

several years. .

It is exceedingly difficult to conceive how injuries could be

inflicted upon particular species with such constancy, and in

such a way as to produce uniform results in the production of

resin cysts which occupy a definite place in the structural scale.

It is a well known fact in the physiology of plants, that con-

ditions which induce a premature development of parts also

bring about the conversion of potentialities into actualities, and

under such circumstances the latter become evidences of a

pathological condition. The swamp maple normally develops a

brilliant foliage in the autumn, but it is not uncommon to find

individual branches which have been injured, or even entire

trees, which exhibit the characteristic autumnal foliage in mid-

summer, a condition which is correctly interpreted as pathologi-

cal. Special conditions of nutrition, e. g, an excess of mineral

food elements, may similarly induce a premature development

of the reproductive process. It has been shown by Richards

that in cases of injury the rate of respiration is greatly in-

creased, an alteration in functional activity which he rightly

interprets as due to efforts directed toward the repair of injured

parts. But this implies a local increase of nutritive materials

and their application to a more intensive process of nutrition.

Such features are well known in the case of all hypertrophies,

and they must be similarly applicable to all forms of wounds, no

matter what their origin, differing only in degree. We cannot

No.454] ANATOMY OF THE CONIFERALES. 707

very well conceive of such profound functional disturbances

without assuming a corresponding alteration in or development

of those structures upon which the activities are dependent.

The structural alterations may thus become characteristic, /oca/

features, and they may even represent the tangible expression

of potentialities which are not manifested elsewhere in similar

regions of the plant body. Furthermore, normal resin canals

are invariable features of Larix, Pseudotsuga, Picea and Pinus.

But Anderson (1, p. 29, etc.) has shown that in Picea axcelsa,

Larix japonica and Pinus strobus there is an enormous increase

in the number of resin canals, arising through the unusual stim-

ulus afforded by the operations of the mycelium of Agaricus

melleus. With respect to Adbzes firma he also clearly shows that

the general effect of the stimulus afforded by the parasite Æcid-

zum elatinum (which gives rise to hexenbesen) is to produce a

more perfectly organized form of the secretory reservoir than is

present under normal conditions. To us, therefore, these facts

offer a reasonable explanation of the appearance of resin cysts

under conditions of injury, when they assume a pathological róle,

while they also serve to harmonize their occurrence under such

circumstances with the general course of their evolution as

already set forth.!

The results to which we are now brought are based entirely

upon developmental phases in anatomical elements of the vascu-

lar cylinder. While our studies lead us to certain definite con-

clusions, we do not in any sense regard the latter as final, but

only as affording one step in the solution of a question which

must be viewed not only from the broader standpoint of more

extended anatomical data, but from that of Physiology as well,

although we feel disposed to insist that the final answer will be

found to rest chiefly upon an anatomical basis. That there may

be room for a different interpretation of the facts here recorded,

is quite possible, since Dr. Jeffrey has recently permitted me to

1 Since the above was written, Dr. Jeffrey has very kindly shown me several

specimens which appear to afford strong evidence in support of his pounos, and

in view of such facts the conclusions here stated are made with reserve until fur-

ther evidence is at hand through the publication by him of studies now in prog-

ress . :

708 THE AMERICAN NATURALIST. [Vor. XXXVITII.

examine the manuscript of an important contribution to our

knowledge of the Abeitineze, in which he brings out very signifi-

cant facts suggestive of the idea that this group is of a much

more primitive character than has hitherto been supposed, or than

is indicated by our own studies. It is therefore of importance

that final judgment should be suspended until the results of

these various studies, as well as those of Coulter and Chamber-

lain, all directed to the same end but prosecuted along entirely

distinct lines, can be brought together and co-ordinated. It is in

this sense, therefore, that we offer the following.

SUMMARY AND GENERAL CONCLUSIONS

In discussing the phylogeny of the higher Gymnosperms,

three subordinate phyla must be taken into consideration in the

following order :— (1) Cordaitales, (2) Ginkgoales, (3) Conife-

rales.

Regarding the Cordaitales as the most primitive gymnosper-

mous stock of which we have present knowledge, it is possible

to trace its origin to the Cycadofilices. The genera Lyginoden-

dron, Heterangium, Calamopitys and Pityoxylon present many

structural features which are common to all, and which not only

establish their relation to the Cycadean line of descent, but they

offer many suggestions of that course of development which is

realized in the higher Coniferales. They, therefore constitute

the real starting point for two lines of descent, the first of which

embraces the Cycadales. With this we have little or nothing to

do at the present moment, beyond establishing its probable rela-

tion to the other gymnosperms. The second line emerges in a

type of plants having characteristics distinctly allied to those of

the Coniferz, and it is this line of descent with which we are

now chiefly concerned. It is now possible to define the origin

of this phylum somewhat more exactly than Coulter has done

(7 & 8), since there is good reason to believe that it emerges

from the Cycadofilices through Poroxylon. Scott (52, P- 398)

has already pointed out the relations of this genus to the Cyca-

dofilices and the Cycadacez on the one hand, and to Cordaites

on the other, so clearly as to remove the necessity for detailec

No. 454.] ANATOMY OF THE CONIFERALES. 199

discussion at this time, beyond giving emphasis to one or two

important structural relations. It has been noted that in Ca/a-

mopitys saturni, the most primitive distribution of the bordered

pits upon both the radial and tangential walls, is represented in

the protoxylem structure. Such distribution, however, under-

goes rapid modification whereby it is wholly limited to the radial

walls in the secondary wood. A similar limitation appears in

other, somewhat closely related genera, and it is fully expressed

in Poroxylon where the multiseriate disposition and hexagonal

form are typically preserved, though there is, at the same time, a

tendency to segregation to such an extent that the pits some-

times become round. In this it is possible to notice the first

indication of a character which, while infrequent, is nevertheless

occasionally expressed among the Cordaitales, though it is gen-

erally characteristic of the related phyla Ginkgoal s and Conife-

rales.

Among the Cordaitales there is but one genus (Cordaites)

"which we have heretofore been accustomed to associate with

that phylum, and, so far as our present knowledge goes, it

undoubtedly stands in the closest relations to Poroxylon. It is,

however, improbable that the two were in any sense coterminous,

and it is altogether probable that there may have been some one

or more intermediate forms of which we have no present knowl-

edge. Our present studies on the other hand, show clearly, that

we must bring into this phylum two other genera of an obviously

higher degree of development, but which have commonly been

ranked with the Abietineze and which, according to Eichler (11),

occupy the highest position in the scale. This position is unten-

able upon anatomical grounds which give us reason to believe

that Agathis and Araucaria (including, of course, Araucarioxy-

lon) are not only inferior to the Coniferales as a whole, but that

they are distinctly Cordaitean. Accepting this view, and the

fact that Agathis is the inferior genus, the sequence would place

Cordaites at the base and Araucaria at the top, with Walchia =

the immediately ancestral form of the latter. This relation is

not only natural, but it is justified on anatomical gr E um 4

The tendency to segregation of the bordered pits as en ite

by Poroxylon suggests the relation of this genus to others in

710 THE AMERICAN NATURALIST. [Vor. XXXVIII.

which such a feature is fully expressed, and it thereby forms the

basal member of another series. From the opposite point of

view, it has been shown that the occurrence of two-seriate pits

in Pinus and others of the Coniferales, as well as in Ginkgo,

points to a common origin for such genera in a type with multi-

seriate, hexagonal pits, and that both Agathis and Araucaria

must likewise center in the same generalized form. This grad-

ual convergence is justified on other grounds, and the genus

Poroxylon among known forms most nearly fulfills the require-

ments of the case. We may therefore look upon it as lying

between the Cycadofilices and all the higher gymnosperms, giv-

ing rise to two lines of descent, the first of which embraces the

Cordaitales as already described, while the second shortly divides

once more. This secondary division gives rise on the one side

tothe Ginkgoales, and on the other to the Coniferales. The

anatomical data already discussed when viewed collectively show

that the general sequence within the latter would be (1) the

Taxoideæ, (2) the Taxodiinz, (3) the Cupressinez, (4) Abies,

(5) Tsuga, (6) Pseudotsuga, (7) Larix, (8) Picea and (9) Pinus,

of which one division (II) represents the highest type of devel-

opment. The sequence of species for each genus cannot always

be determined with a full measure of satisfaction, and these

difficulties may possibly be made clear by reference to à particu-

lar case. The succession of the two species of Sequoia is diffi-

cult to determine on purely anatomical grounds, but the general

tendency of the facts already recited is to give to S. semper

virens the more primitive position, a view which is sustained by

its palæontological history.

The relations brought out in the foregoing studies, and the

conclusions reached, may be made more obvious without the tedi-

ous method of a detailed discussion by reference to the accom-

panying table of anatomical data, which substantially summarizes

all the results derived from the study of particular structures.

No. 454] ANATOMY OF THE CONIFERALES.

711

Data for Table of Anatomical Characters, in identical series.

M mU me uw Mo»

ub wone oo MW.

Spiral tracheids.

Bordered pits in 1—3 rows.

Bordered pits in 1—2 rows.

Bordered pits in one row.

Pits on the tangential walls of the summer wood.

Lateral walls of the ray cells with bordered pits.

Uniseriate rays.

Terminal walls of the ray cells thin and entire.

Resin cells. |

Terminal walls of the ray cells locally thickened.

Terminal walls of the ray cells strongly pitted.

Ray tracheids.

Resin passages.

Fusiform rays.

Thyloses in the resin passages.

Lateral walls of the ray cells with simple pits.

Ray cells of two kinds.

Resin cells scattering.

Resin cells zonate.

Resin cells grouped.

Resin cells on the outer face of the summer wood.

Ray tracheids marginal.

Ray tracheids interspersed.

Ray tracheids dentate.

Number of species.

Percentage value of genus.

THE AMERICAN NATURALIST. (VoL. XXXVIII

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716 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In preparing this table, the various anatomical features have

been chosen with reference to (1) the constancy of their occur-

rence, (2) their structural prominence and (3) their obvious

relation to diagnostic purposes. In their horizontal extension,

an attempt has been made to arrange them in accordance with

the law of frequency, as well as with reference to their relation

to development, in such wise that while the spiral tracheid is

assumed to be the most primitive type of the vascular structure,

the presence of two kinds of cells in the medullary ray may be

held to express the highest form of development. To the mem-

bers of the series so constituted we may then assign arbitrary

values in arithmetical sequence from one to seventeen ; while those

subordinate characters which are represented by different forms

of distribution may be regarded as forming a second series simi-

larly valued. Any primitive or other character which has

become obliterated through development may be held to retain

its original value with respect to the general course of such

development, and it is always indicated by —. Vestigial struc-

tures occurring sporadically are designated by (1), and to them

one half the value of the fully developed character is assigned.

All normal features are designated by x, which becomes x +

when they show development toward the next higher form, or

by x — when they show a definite tendency to degeneration.

Sporadic characters which are obviously in the line of develop-

ment are indicated by (o), but they are assigned only half values.

On this basis it is possible to arrange a sequence of genera and

species in such a manner as to exhibit a progressive develop-

ment from the simple Agathis with a minimum of characteristics

to the complex Pinus in which the greatest number of anatomi-

cal features are involved. Furthermore, through such a series

it is possible to determine the relative position of the various

genera by percentage values, and this gives the most valuable

insight into the approximate relations of the various members

within the general line of descent. Such relations are deter-

mined not only for each anatomical character, but for the collec-

tive characters. Reducing these facts to a graphic form, the

accompanying curves will assist in making the relations more

clear, especially in emphasizing the general course of develop-

sip ou») 3

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718 THE AMERICAN NATURALIST. [Vor. XXXVIII.

ment which in its final form is best expressed by a biological

tree. A figure of this sort is difficuit to construct, and there is

no agreement among investigators as to the particular form it

should take. While the figures in common use indicate a cer-

tain relationship in descent, they completely fail to convey any

impression of the way in which the succession arises, and they

furnish no indication of possible gaps. They therefore consti-

tute a very poor working basis.

I have long been accustomed in teaching to compare the vari-

ous lines of descent among plants with the branchings of a deli-

quescent tree, since it has always seemed reasonable to suppose

that the laws which govern the branching of a limb, which give

rise to all the varying forms of arrested development, and which

thereby determine a particular modification of the figure which

would otherwise result from unmodified growth, must be equally

applicable to the general evolution of the higher forms of plants

from a common ancestral type. In endeavoring to secure a

natural growth which would best express all the conditions |

involved, a sympodium was first of all suggested, inasmuch as it

conveys the idea of succession through lateral members in such

a way as to indicate the direct line of descent. But sympodia

as we usually know them fail to adequately express the idea of

arrested development and suppression in their various forms.

In the branching of Acer platanoides all the conditions appear |

to be met in a very satisfactory manner. The branch of the

Norway maple, when of vigorous growth, is a monopodium,

and it is obvious that such would not answer the object in view,

since its most prominent feature would suggest the idea of a

continuous series of coterminous members from which lateral

members would arise at intervals. There is no evidence that

any phylum represents stich a series; on the contrary there is

every reason to believe that such relations do not exist among

the various groups of plants.

But in those branches of the Norway maple which exhibit

slow growth various forms of arrested development are mani-

fested. These take the form of atrophied buds, or of branches

in all stages of development, and there thus arises a modified

monopodium which eventually becomes, in many cases at least. |

720 THE AMERICAN NATURALIST. (VoL. XXXVI

a true sympodium. In comparing this with the monopodial

branch of vigorous growth, it appears that the alterations in-

volve more than mere suppression. In the monopodium the

average angle of divergence for the lateral members is 45.3°,

while for the derived form it is 34.1°. The latter will be seen

to completely fulfill all conditions with respect to the develop-

ment of a phylum, even to indicating the position of missing

members. Selecting from this such portion as may serve the

requirements of the present case, we obtain the following figure

which may be held to embody our final conclusions as to the

general succession of the different gymnosperms, and from it

we may gather that the highest representative — Pinus — is the

terminal member in the main line of descent from the Cycado-

filices through Poroxylon, while from such a central line both

the Cordaitales and Ginkgoales have been given off as side lines.

In the construction of this figure an attempt has been made to

show all normally developed buds (0) and their relative dimen-

sions ; atrophied buds (0), the position of which is recognizable ;

and atrophied branches (/ —) which are still visible, but it is

obvious that the figure does not show many members, all evi-

dence of the former existence of which has completely dis-

appeared.

The general results of these investigations serve to confirm

in a very striking manner the probable monophyletic origin of

the Gymnosperms as already expressed by Coulter (7), while

they also show that the real transition ground, at least for all

but the Cycadaceze, was probably represented by Poroxylon as

indicated by Scott (52).

No. 454.] ANATOMY OF THE CONIFERALES. 721

LITERATURE.

ANDERSON, A. P.

(1. Ueber abnorme Bildung von Harzbehaltern. München, 1896.

(2) Comparative Anatomy of the Normal and Diseased Organs of

bies balsamea affected with Aecidium elatinum. Bot. Gaz.,

XXIV, p. 309-344. (2 pl.) 1897.

BENTHAM & HOOKER

3.) Genera Plantarum.

BRITTON, N. L., & BROWN, A.

(4) An strated Flora of the Northern United States and Canada.

Y., 1896. Vol. I, 49-61.

CHAPMAN, A. W.

(5. Flora of the Southern United States. 1884.

CooPER, J. G

(6. Smithsonian Report, 1858.

COULTER, J. M.

(7) The Gymnosperms and the Seed Habit. Bot. Gaz., XXVI, 153-

COULTER, J. M., AND CHAMBERLAIN, C. J.

(8.) Morphology of the Spermatophytes. New York, 1901.

De Bary, A.

(9. Comparative Anatomy of the Phanerogams and Ferns. Oxford,

ENGELMANN, GEORGE.

(1o. Botanical Works. Cambridge, 1887.

EICHLER, A. E.

(rt) Naturlichen Pflanzenfamilien. II, 6 et seq.

ENGLER, A., AND PRANTL, K.

(12.) Naturlichen Pflanzenfamilien. I1, 7-127.

ENGLER, A.

(13. Naturlichen Pflanzenfamilien. II, 24 et seq.

ESSNER, B.

(14. Diagnosticher Werth der Anzahl und Hohe der Markstrahlen bei

den Coniferen. Bot. Centralbl., XII, 407, 1882.

FiscHER, H.

(15) Ein Beitrag zur vergleichenden Anatomie des Markstrahlgewebes

d der Jahrkichen Zuwachszonen im Holzkorper von Stamm,

reir und Aesten bei Pinus abies, L. Flora, vol. 68, p. 263 et

seq. 1885.

GOEBEL, K. .

(16) Outlines of Classification and Special Morphology of Plants.

rd. 1887.

GOEPPERT, H. R.

(17.) Fossilen Coniferen. eden 1850.

722 THE AMERICAN NATURALIST. [Vor. XXXVIII.

GRAY, ASA.

(18) Manual of Botany of the Northern United States. 189o.

HARTIG, GEORG LUDWIG.

(19.) Lehrbuch für Forster. Stuttgart, 1877.

HARTIG, R.

(20.) Das Holz der deutschen Nadelwaldbäume. Berlin, 1885.

HARTIG, THEODORE.

(21.) Anatomie und Physiologie der Holzpflanzen. Berlin, 1878.

HOOKER, SIR W. J.

(22.) Flora Boreali Americana. II, 165.

INDEX KEWENSIS.

23.)

JEFFREY, E. C.

(24. The Comparative Anatomy and Phylogeny of the Coniferales.

Part I, The Genus Sequoia. Mem. Boston Soc. Nat. Hist. 1903.

KALM, P.

(25.) Travels in North America. II, 646.

KNOWLTON, F. H.

(26.) A Revision of the Genus Araucarioxylon of Kraus, with compiled

descriptions and partial synonymy of the Species. Proc. U. S.

Nat. Mus., XII, No. 784, 601-617. 189o.

MAcoun, Jonn.

(27) Catalogue of Canadian Plants. Exogens. Geol. Surv. of Can.

1883.

(28.) Rept. Geol. Surv. Can., 1875-76. 211.

MASTERS, MAXWELL T.

(29.) Notes on the Genera of Taxaceae and Coniferae. Jn’l Linn Soc.,

XXX, I.

(30.) The Cypresses of Monterey. Garden & Forest, VII, 298.

Mayr, H.

(31) Waldungen von Nordamerika. München, 189o.

MICHAUX, A.

(32. Flora Boreali Americana. II, 1820.

MICHAUX, F. A.

(33) The North American Sylva. III.

MULLER, N. J. C.

(34) Atlas der Hólzstructur. Halle, 1888.

(35-) Botanische Untersuchungen. Heidelberg, 1872.

Murray, A.

(36.) Edinburgh New Phil. Jn. n.ser. I.

NORDLINGER, H.

(37.) Die Technischen Eigenschaften der Hölzer. Stuttgart, 1860.

NUTTALL, T.

(38.) The North American Sylva. III.

No. 454.] ANATOMY OF THE CONIFERALES. 723

PENHALLOW, D. P.

(39.) Taxaceae and Coniferae. Trans. R. Soc. Can, II, iv, 33-57.

(40.) North American Species of Dadoxylon. Trans. R. Soc. Can, VI,

iv, 51—7 1900

(41. Noteson Tatary Plants. Trans. R. Soc. Can., IX, iv, 33-95. 1903.

PIERCE. LE

(42.) Studies on the Coast Redwood. Cal. Acad. Sc., II, 83-106, 1901.

PoroNiE, H.

(43-) Lehrbuch der Pflanzenpalaontologie. Berlin, 1899.

(44-) Nafurlicben Pflanzenfamilien. II, 33-40.

PursH, F.

(45.) Flora Americana. 1816.

RorH, FILIBERT.

(46.) Timber Pines of the Southern United States. U. S. Dept. of

gric., Div. of For., Bull. 13, 133-143.

Russow, E.

(47-) pui Kenntniss des Hólzes, insonderheit des Coniferenhólzes. Bot.

ntralbl XIII, 29. 1883.

E m S.

(48. Forest Trees of North America. Tenth Census of the U. S., IX.

(49.) Tia "dba of North America. X, XI, XII. Boston, 1897.

SCHROEDER, J.

($0. Holz der Coniferen. Dresden, 1872.

Scorr, D. H.

(51) The Anatomical Characters presented by the Peduncle of Cyca-

daceae. Ann. Bot., XI, 399-419. 1887.

(52.) Studies in Fossil Botany. London, 1900.

SEWARD, A. C. AND GOWAN, J.

(53) The Maiden Hair Tree. (Ginkgo biloba, L.) Ann. Bot, XIV,

109-15

WATSON, SERENO. :

(54) Botany of California. II. 1880.

Weiss, G. A.

(55. Anatomie der Pflanzen. Wien, 1878.

WILLIAMSON, W. C.

(56.) n the Structure and Affinities of some Esogmóus Stems from

e Coal Measures. M. Mic. Jn3, II, 66-72.

Wido o.c.

( 57.) pie Constituents of Plants, trans. by Baron F. von Mueller.

— 1878.

ZEILLE

8) " Eléments de Palzobotanique Paris, —

STUDIES ON THE PLANT CELL.-—IV.

BRADLEY MOORE DAVIS.

Section III. HiGury SPECIALIZED PLANT CELLS AND

THEIR PECULIARITIES (Continued).

4. The Spore Mother-cell.

THE spore mother-cell and its homologues the pollen mother-

cell and certain embryo-sacs have furnished some of the most

interesting subjects for cell studies in the plant kingdom.

Sporogenesis in all plants above the thallophytes seems to be

a period when nuclear structures are especially clearly differ-

entiated and when the mechanism of mitosis reaches the highest

degree of complexity. These intricate conditions are only

equalled by processes in the development of the female game-

tophyte of some angiosperms, and during endosperm formation,

also in the events of spermatogenesis and with the segmentation

of the egg nucleus of certain gymnosperms.

Sporogenesis is one of the critical periods in the life history

of a higher plant since it is the time when the asexual genera-

tion (sporophyte) passes over to the sexual (gametophyte). This

provides certain important features such as the reduction phe-

nomena concerned with chromosomes and greatly adds to the

interest in these cells. These matters will receive special atten-

tion in Section V, but they must be borne in mind to appreciate

fully the significance of many events of spore formation.

The general history of the spore mother-cell may be described

as follows: It is the product of the last mitosis in the repro-

ductive tissue called the archesporium. This mitosis always

presents the number of chromosomes characteristic of the

sporophyte which is double the number found in the game-

tophyte. Therefore the nucleus that passes into the spore

mother-cell has the sporophyte number of chromosomes. Two

725

726 THE AMERICAN NATURALIST. [Vor. XXXVIII.

mitoses occur successively in the spore mother-cell in all forms.

The first mitosis presents half the number of chromosomes

found in the last nuclear division in the archesporium and is

consequently the reduced or gametophyte number. The reduc-

tion of the chromosomes then takes place during the period of

rest between the last mitosis in the archesporium and the first

in the spore mother-cell. There are two mitoses in the spore

mother-cell. In some forms these are exactly alike and present

essentially the same characters as the usual typical mitoses of

plants. But among the spermatophytes there are likely to be

peculiarities in the arrangement and distribution of the chromo-

somes. In consequence the first mitosis may be heterotypic

and the second homotypic in contrast to the normal typical con-

ditions. The description and explanation of these characters.

wil be reserved for the groups that illustrate them the best.

They have nothing to do with qualitative reduction phenomena

as was formerly supposed.

There is sometimes a well defined period of rest after the first

mitosis with the formation of a wall between the two daughter

nuclei, but frequently the second mitosis follows immediately

after the first so that the spore mother-cell comes to contain

four daughter nuclei. Cell walls may then be formed between

these nuclei simultaneously so that the resultant spores are dis-

posed in a radially symmetrical arrangement that is termed

tripartite. These cell divisions are almost universally present in

the spore mother-cell, the only exceptions being certain sperma-

tophytes whose megaspore mother-cells develop directly into

embryo Sacs, the two mitoses (heterotypic and homotypic) being

included within these structures and forming a part of the game-

tophyte history. Why the number of spores should generally

be four is unexplained. There does not seem to be any physio-

logical significance in the number or other reasons why it sho

not be more or less. Indeed it is somewhat variable in the

spermatophytes for microspore or pollen mother-cells form two

and three pollen grains in certain types and five, six and seven

have been found in others, while much larger numbers have

been occasionally reported. In no case is the microspore

mother-cell known to develop directly into a pollen grain, al-

No. 454.] STUDIES ON THE PLANT CELL. 7?7

though the megaspore mother-cell regularly becomes an embryo

sac in some forms (e. g., Lilium). But an increasing number of

observations indicate that the megaspore mother-cell generally

develops two, three or four potential megaspores although nor-

mally only one of these becomes an embryo sac.

The interest in the protoplasmic activities of sporogenesis lie

chiefly in the elaborate methods of spindle formation and mech-

anism of mitosis, in the organization and distribution -of the

chromosomes, in the functions and activities of the nucleolus,

and in the organization of the cell plate and development of the

cell wall. There is a very extensive literature on the spore

mother-cell some of which, however, merely treats the broad

features noted in studies of a general morphological character on

the development of sporophylls or floral structures. We shall

only attempt to consider the most important contributions, and

for convenience will begin our treatment with the Hepaticz and

conclude with the spermatophytes where the conditions are the

most complex.

The Hepatice or liverworts furnish some remarkable spore

mother-cells, and are now the subject of considerable interest

and some discussion. They were first brought conspicuously to

the attention of botanists by a paper of Farmer ('94) on Pat

lavicinia decipiens. Farmer described a remarkable series of

events in this type. The nucleus of the spore mother-cell

became surrounded before division by dense protoplasm that

extended into the four lobes of the cell in the form of a four-

rayed star which he called a “quadripolar spindle." After its

development four chromatic droplets appeared in the nucleus to

indicate its approaching division. . These chromatic droplets

became four chromosomes which by division were doubled in

number. The eight rod shaped chromosomes moved in pairs

towards the four lobes of the spore mother-cell. There was a

further division of each chromosome, making sixteen in all, and

the four groups of four each passed simultaneously to the poles

of the *quadripolar spindle " which persisted to the end. It

should be noted that the striking peculiarities of Farmer's account

lie in the division of the four primary chromosomes into sixteen,

and in their simultaneous distribution through a “ quadripolar

728 THE AMERICAN NATURALIST. [Vor. XXXVIII.

spindle” to form at once four daughter nuclei. These events

are unparalleled, as far as the writer is aware, in the plant or

animal kingdom, and consequently the account deserves especial

attention. A four-rayed figure around the nucleus is not surpris-

ing because the spore mother-cell of the Jungermanniales is four

lobed, and its centrally placed nucleus lies in a restricted area.

But the simultaneous distribution of quadrupled chromosomes

to form four daughter nuclei is a process whose establishment

would be of fundamental significance. Farmer also described a

centrosome at each pole of the ** quadripolar spindle."

Farmer (95a, 4, and c) followed his paper on Pallavicinia with

studies on other liverworts. He reported the “ quadripolar

spindle " in the early stages of mitosis in several of the Junger-

manniales, but did not find the quadrupling and simultaneous

distribution of the chromosomes as in Pallavicinia. The * quad-

ripolar spindle" when present was a temporary structure replaced

later by the bipolar spindles of two successive mitoses with a

longer or shorter interval between. Farmer considers the

* quadripolar spindle" of these forms as transitional between

that of Pallavicinia and the normal bipolar spindle. The Ric-

ciales, Marchantiales and Anthocerotales present two successive

mitoses after the usual manner in the spore mother-cell.

The writer has described the events of sporogenesis in Pellia

(one of the Junger iales) in a paper covering the nuclear

activities at several periods in its life history (Davis, :01), and

confirmed much of Farmer's account of the mitoses in this spore

mother-cell. These are two in number and successive, with a

very well defined resting period between the first and the second.

There is a four-rayed figure present during the prophase of the

first mitosis, and this seems to correspond to Farmer's “ quad-

ripolar spindle." The nucleus lying in the center of the four

lobed spore mother-cell becomes invested by a kinoplasmic

Sheath which develops a fibrillar structure. Many of these

fibrillae extend into the lobes of the spore mother-cell because

the nucleus is confined to a narrow space in the constricted cen-

tral region of the cell and the lobes offer the only possible relief.

for the crowded conditions. However, the four-rayed structure"

is not present when the chromosomes are ready for distribution,

No. 454.] STUDIES ON THE PLANT CELL. à 729

but there is found instead one large, broad poled spindle. (See

Fig. 5 e) A cell wall is formed between the two daughter

nuclei (Fig. 8 d) which divide again after a very short period of

rest, the two spindles lying at right angles to one another. The

poles of the spindles are rather blunt, and there are no centro-

somes or centrospheres in either mitosis. The four-rayed struc-

ture of prophase must be regarded as preliminary to spindle

formation because the chromosomes are not ready for distribu-

tion, and when that period arrives the structure has been re- i

placed by the true spindle of the first mitosis. These facts led

me to question Farmer's account of mitotic phenomena in Palla-

vicinia and his conception of the *' quadripolar spindle," and I

suggested that this structure might prove to be a phenomenon of

prophase, a view to which Farmer (:01) has taken exception in

a criticism of my results.

Recent investigations of Moore (: 03) on Pallavicinia are flatly

contradictory to the conclusions of Farmer for Pa//avicinia

decipiens and support my suggestions. Moore finds that there

are two mitoses in the spore mother-cell of Pallavicinia lyellit,

the second (Fig. 12 c, d) following immediately upon the first

(Fig. 12 6), each with bipolar spindles and without centrosomes.

The chromosomes, eight in number, appear in the usual way

with each mitosis (Fig. 12 c, d). There is no *'quadripolar

spindle" in Farmer's sense, no quadrupling and simultaneous

distribution of the chromosomes. The prophases preceding the

first mitosis present a tetrahedral form as is shown in Fig. 12 a.

This is accentuated by the fibrillae which gather at the points

to make a four-rayed structure extending into the lobes of the

spore mother-cell. This condition is identical with similar stages

in Pellia and in other leafy liverworts, and is a feature to be

expected from the fact that the spindle fibers develop chiefly or

wholly externally to the nuclear membrane in a rather crowded

region of the cell. - The nucleus at this time is unquestionably

in prophase as shown by the undifferentiated chromosomes and

because this stage passes immediately into a bipolar spindle of

the normal type (Fig. 12 b). It seems very probable that

Farmer was mistaken in his conclusions for Pa//avicinia decipi-

ens, and that the mitoses jn the spore mother-cell of this form

730 THE AMERICAN NATURALIST. [Vor. XXXVIII.

are not different in any essentials from those of other plants.

the four lobes of th ther-cell; the nuclear membrane has not yet broken down;

similar stages of p rophase were probably considered by Farmer as poesi olar spindles.

4, metaphase of is first mitosis; the spindle in all respects a normal bipolar Seger

g, metaphase o i A

ing in the spindle between two nuclei. wo nuclei at the side of their respective

detis res and the be pee prea, after the second mitosis ; a third chromato-

phore shown with strands g it with me regions of thecell. (a,

ó e, P after Moore, :03; hs $; afte Van Hook,

The *quadripolar spindle" proves to be nothing more than a

condition of prophase. |

Besides Pellia and Pallavicinia, which are the most thoroughly

studied of the lower liverworts, we know the processes of sporo-

No. 454.] STUDIES ON THE PLANT CELL. 731

genesis in the highest type, Anthoceros (Davis, '99). This form

is exceedingly attractive for such investigations because the

spore mother-cells may be found in all conditions upon the same

sporophyte. However, the small size of the nuclei and spindles

is a disadvantage. Just previous to the first mitosis the nucleus

becomes surrounded by a mesh of delicate fibrillae (kinoplasmic).

Later the nucleus takes an angular form, and the fibrillz are

found conspicuously at the prominent poles (Fig. 12 ¢). The

nuclear membrane breaks down and the fibers become arranged

to form a bipolar spindle (Fig. 12 f) without centrosomes or

centrospheres. There is a short period of rest after the first

mitosis, but no wall is formed between the two daughter nuclei.

The small spindles of the second mitosis (Fig. 12 g) are like-

wise bipolar. They lie at right angles to one another and the

cell plates that are laid down determine, in part, the position of

the walls that are formed between the four granddaughter

nuclei and which divide the spore in a tripartite manner. These

cell plates are very small (Fig. 12 4 and 7), but they have been

observed in a favorable species of Anthoceros by Van Hook

(:00). It is not clear how these plates become extended to

the wall of the spore mother-cell unless (as suggested in Sec. II)

their edges make use of planes of vacuoles when the protoplasm

separates to develop the cleft between the four daughter cells.

The poles of the spindles in Anthoceros are-flattened and entire: y

free from structures that might be considered centrosomes.

Other interesting events of sporogenesis in Anthoceros are

the division of the chromatophores and the nuclear condition

termed synapsis. The young spore mother-cell contains a single

large chromatophore. This increases greatly in size and becomes

filled with starch grains. The chromatophore divides succes-

sively into two and then four portions which arrange themselves

symmetrically in the cell with the nucleus in tbe center. The

mitoses then follow and the four daughter nuclei are distributed,

one for each chromatophore in the cell. This provision of four

chromatophores long before the mitoses in the cell seems very

remarkable (Davis, '99, p. 94 and 95). Synapsis is a condition

very common in the nucleus of spore mother-cells before divi-

sion. The chromatic material becomes gathered into a compact

732 THE AMERICAN NATURALIST. [Vor. XXXVIII.

mass besides the nucleolus. The significance of synapsis is not

clear, but the subject will be discussed in Section VI. How-

ever, there is good evidence from Anthoceros that the phenome-

non is a normal event and not an artefact, because synapsis is

always found at a certain period of sporogenesis, and nuclei in

neighboring spore mother-cells a little older or younger present

their chromatic material with the usual arrangement (Davis, '99,

p. 96 and 97).

To summarize the conditions in the spore mother-cells of the

Hepatice, all conclusions, in the author's opinion, indicate: (1)

That the spindles develop from a surrounding weft of fibrillae

without the assistance of centrosomes. (2) That the mitoses

are always two in number and successive with the same number

of chromosomes for each division. (3) That the cell walls mày

be formed successively as in Pellia and some other of the Jun-

germanniales or simultaneously, to give tetrahedral spores, as in

Anthoceros, types of the Marchantiales and Ricciales, Pallavi-

cinia and some companion forms in the Jungermanniales. It

will be interesting to note the essential agreement in these

matters between the Hepaticæ and the higher plants.

Nothing is known of the nuclear activities during sporogenesis

in the other great division of the bryophytes, the mosses

(Musci). The spore mother-cells in this group are always small

and unattractive for cell studies but the Sphagnales appear to be

rather the most promising for such investigations, which are

greatly to be desired.

The pteridophytes have furnished some important contribu-

tions to our knowledge of the spore mother-cell. There is first

the paper of Osterhout ('97) on spindle formation in Equisetum,

which was one of a group of three contributions (Mottier, '97,

Juel, '97) that did much to dispose of a then prevalent belief

that the development of the spindle in higher plants was con-

trolled by centrosomes. This investigation was followed by

a study of Smith (:00) on spindle formation in Osmunda.

Calkins ('97) and W. C. Stevens (98a) considered especially

the formation and reduction of chromosomes in several of the

ferns, and arrived at contradictory conclusions. Strasburger

(:00, p. 76 to 79) has reviewed these results in relation to

Studies of his own on Osmunda.

No. 454] STUDIES ON THE PLANT CELL. 733

Osterhout's (’97) account of spindle formation in Equisetum

is noteworthy. He found that the nucleus of the spore mother-

cell became surrounded by a web of delicate fibrillae, which,

extending radially into the surrounding cytoplasm (Fig. 13 2),

were later (Fig. 13 4) gathered into numerous pointed bundles

or cones. After the dissolution of the nuclear membrane these

c ^

Kn —-

SAY

2 | li BN o,

Fic. 13.— Spore mother-cells o

first mitosis; the radially disposed fibrilla are gathering. together into cones. 4, prophase,

older than a; the nuclear membrane has broken down and the fibrilla have entered the

c ; the cones lie in two ps opposite one another. ju ore' meta-

phase; the fibrillar cones are nearer together and the chromosomes have gathered to form

the nuciear plate ef 85 regalis. d, very early prophase of the mito-

e, prophase, somewhat older than d; fibrillar kinoplasm showing polarity. /, still older ;

chromosomes formed; one pole of spindle developed. g, metaphase; a tri-polar spindle.

(a, 6, c, after Osterhout, '97; d, e, £,» Smith, : 00.)

cones arranged themselves side by side in two sets to form the

spindle of metaphase (Fig. 13¢). The spindle is then from the

outset multipolar, and even though some of the cones unite when

they become grouped around a common axis, nevertheless the

poles of the spindle at metaphase show their composite nature in

the absence of a common focal point for the fibrilla. There are

no centrosomes at the poles and no reason for their presence at

any stage in the process of spindle formation.

Smith’s (:00) study of Osmunda presents an important con-

firmation of Osterhout's conclusions that the spindle in pterido-

734 THE AMERICAN NATURALIST. [Vor. XXXVII.

phytes developed without centrosomes, while illustrating a proc-

ess of spindle formation along somewhat different lines. Smith

distinguished a zone of kinoplasm around the nucleus previous

to spindle formation. This zone became granular, and then the

granules arranged themselves in rows to form fibrillze (Fig. 13 £),

which, however, did not extend into the cytoplasm radially, but

lay generally parallel to one another, so that the spindle appeared

bipolar from the beginning (Fig. 13e). One pole of the spindle

was generally formed considerably in advance of the other (Fig.

13/). The fibers did not meet at a common point but over a

broad area, and there were no centrosomes. There is, then, nor-

mally no multipolar stage in Osmunda, although tripolar spindles

(Fig. 13g) were occasionally found. During anaphase secondary

fibers were put forth from the vicinity of the daughter nuclei and

these met in the equatorial region of the cell. The spindle of

the second mitosis was formed exactly as in the first. After this

division the four granddaughter nuclei lay connected with one

another by six spindles (two primary and four secondary). Cell

plates were laid down in the equatorial regions of these spindles

so that the protoplasm became divided simultaneously and sym-

metrically into tetrahedral spores.

The studies of Calkins (97) and Stevens ('98a) were chiefly

upon the division and distribution of the chromosomes in con-

nection with reduction phenomena. Calkins believed that the

processes of sporogenesis followed the same course as the matu-

ration of sexual cells in animals, with a transverse division to

give a qualitative reduction in Weismann's sense. Stevens dis-

agreed with Calkins in several particulars, holding that the

reduction was merely quantitative. Reduction phenomena in

plants is now much better understood than at the time of these

papers which dealt with plants much more difficult to study than

some other forms (e. g., types of the Liliaceze). We shall con-

sider the subject in Section V, but may state now that Calkins'

conclusions have not been sustained.

Strasburger (:00) gives considerable attention to spindle

formation in his well known review and critique of cytological

literature. He proposes the following classification of spindles

in higher plants which lack centrosomes. Those that pass

No. 454.] STUDIES ON THE PLANT. CELL. 735

through multipolar stages and later become bipolar are called

multipolar polyarch spindles. When the spindle has a well

defined axis from the beginning, as is generally true of the cells

in vegetative tissues of higher plants, it is termed multipolar

diarch. Strasburger has shown that these types, while easily

separated in the extremes, grade into one another so that the

classification is not founded on distinctions of a very funda-

mental character. The spindle of Osmunda, for example,

resembles a multipolar diarch, but its method of development is

more closely related to that of other spindles in spore mother-

cells (multipolar polyarchs) than to those of vegetative tissues.

The gymnosperms offer in Larix an excellent subject for

studies on the formation of pollen, and this type has been

treated in several important papers, notably by Belajeff (94%),

Strasburger ('95) and Allen (:03). Belajeff's contribution is

important as the first investigation that considered the multi-

polar spindle as a preliminary stage in the development of the

bipolar structure. Other authors, at this time and previous to

his publication, had noted multipolar and tripolar spindles

(Strasburger (80) and (’88) in several forms), but the lily had

received the greatest attention in this connection (Farmer ('93)

and ('954), Strasburger ('95), Sargent ('97) and Mottier (’97) ).

Mottier's investigation presented the first detailed account of

spindle formation in this angiosperm and will be discussed

presently.

Allen's (:03) paper on Larix includes one of the best discus-

sions of the literature bearing on the subject of spindle forma-

tion that has yet appeared. He finds that the cytoplasm around

the nucleus just previous to mitosis comes to contain a loose net-

work of fibrille. Some of the fibers may be followed through

the nuclear membrane and may be seen attached to chromatin

bodies in the interior (Fig. 144). Later the cytoplasmic fibrillae

become arranged radially and extend from the nucleus even to

the outer plasma membrane at the periphery of the pollen

mother-cell. The radiating fibers are connected with one another

by branches which indicate that the structure is in part an

expanded condition of the original network, but the fibers also

grow. The fibers now fold over sothat they tend to lie parallel to

736 THE AMERICAN NATURALIST. (Vor. XXXVIII.

the surface of the nucleus and thus form a dense felt around

the nuclear membrane. Presently the nuclear membrane which

was before a definite film becomes wavy in outline and often

granular in appearance. The nucleolus shows signs of dissolu-

tion and there is a marked increase in the number of intranuclear

fibers, which are chiefly or wholly of nuclear origin. After the

M ACES

wi à

WIN

" S

bj Ox

i»

eit!

bi,

AM if

QÈ

E MN, " "M

LAD Ws ROS

ANS

Fic. 14.— Pollen or microspore mother-cells of spermatophytes. a, 6,c, Larix europea. @,

prophase of first mitosis; kinoplasmic fibrillze forming a felt around the nucleus. 4, late

prophase}; the nuclear |i has broken d 2a d ior space has ! fied

with fibrille which have g 1 to form a multipolar spindle. c, metaphase; a completed

spindle with polar radiations. d, e, Lilium candidum. d, prophase of first mitosis ; the

ch would have become poles of the spindle. ¢, late prophase; the nuclear mem-

disappearance of the nuclear membrane some of the peripheral

fibers push outward to form several cones of a multipolar figure

(Fig. 144). The fibers attached to the chromosomes gather

into bundles that extend towards these poles. Finally the bun-

dles of fibers become more regular and straighten out so that

they come to lie approximately parallel to one another, and thus.

No. 454.] STUDIES QN THE PLANT: CELE, - 737

the multipolar structure, developing a distinct axis (Fig. 14¢),

becomes bipolar (multipolar polyarch). There is no central

body at the poles and no place fora centrosome in this develop-

mental history. ;

The first detailed study of spindle formation in Angiosperms

was, as before stated, that of Mottier (97) which treated especi-

ally of Lilium, Podophyllum and Helleborus. This paper with

one by Juel (97) on Hemerocallis and Osterhout ('97) on Equi-

setum effectually disposed of previous views very generally held

(Guignard, '91, followed by other authors), that spindle forma-

‘tion and mitotic phenomena in higher plants was involved with

the activities of centrosomes or other kinoplasmic centers.

Mottier found that the nucleus in the pollen mother-cell of the

lily became invested just before mitosis with radiating fibers that

shortly after increased in quantity to form a felted web around

the structure. Some of the fibers gathered into cones (Fig.

14 d) which pointed towards the periphery of the cell so that

there resulted, with the disappearance of the nuclear membrane

and the entrance of the fibers into the nuclear cavity, a multi-

polar spindle (Fig. 14 e). The poles gradually came to lie

parallel to one another in a common axis, some. of them disap-

pearing, so that the spindle generally became distinctly bipolar

at metaphase (multipolar polyarch). Essentially the same his-

tory was repeated during the second mitosis in the lily.

From this time on there have been a succession of papers

verifying the general conclusions of Mottier and Juel and extend-

ing these results to many other forms until now it seems to be

well established that centrosomes are never present in the pollen

mother-cell and that multipolar spindles, developed from felted

stages and changing to bipolar spindles, may be expected in

most if not all forms. Guignard (97 and '98) described multi-

polar spindles in several types (Nymphaea, Nuphar, Limoden-

dron, etc.), and while he believed that these poles were occupied

by granules that sometimes fused to form typical centrosomes,

nevertheless he admitted that the multipolar spindle might be

formed independently of centrosomes.

The most important papers on spindle formation in Angio-

sperms following those of Mottier ('97 and '98) and Juel (797).

738 THE AMERICAN NATURALIST. (VoL. XXXVIII.

have been Lawson ('98) on Cobea, W. C. Stevens ('982) on

Asclepias, Atkinson ('99) on Ariseema and Trillium, Duggar

(99) on Bignonia, Wiegand ('99) on Convallaria and Potamoge-

. ton, Gregoire (99) on Lilium and Fritillaria, Guignard (99) on

Naias, Williams (99) on Passiflora, Duggar (:00) on Symplo-

carpus and Peltandra, Lawson (: 00) on Gladiolus, Byxbee (: 00)

on Lavatera, Andrews (:01) on Magnolia and Liriodendron,

Schniewind-Thies (:01) on Galtonia and Osterhout (: 02) on

Agave.

Of the papers listed above several demand especial attention

for the completeness of the studies on the early stages of spindle '

formation in the pollen mother-cell. Lawson ('98 and :00)

found that the nuclei of Cobea and Gladiolus previous to mitosis

were surrounded by a zone of granular kinoplasm which he

named perikaryoplasm. This zone developed a felted envelope

of fibrillæ from which projections extended to form the cones of

a multipolar figure. The cones by fusing in two groups devel-

oped the bipolar spindles. The spindle fibers of Gladiolus are

formed entirely from the perikaryoplasm, the nucleolus and linin

apparently taking no part in the development of the spindle.

The nucleolus remains intact until after the dissolution of the

nuclear membrane when the spindle is practically completely

organized. Miss Williams (99) found for Passiflora that the

nuclear cavity became filled with a network developed from the

linin. The nuclear wall became also transformed into a mesh

which connected the network from the linin with the surround-:

ing cytoplasmic reticulum, thus forming a continuous system

throughout the cell The central region of this network,

enclosed by a granular zone, developed a multipolar figure

whose poles finally fused to form a bipolar spindle. The con-

trast between this type of spindle in which so much of the

fibrous structure is derived from the linin and that of Gladiolus

just described is very marked. A granular region outside of

the fibrous network around the nucleus is much more conspic-

uous in Lavatera, described by Byxbee (: OO), than in Passiflora.

It forms in Lavatera a dense zone that suggests a gathering of

nutritive material (deutoplasm). The fibrillæ are developed as a

felt around the nuclear membrane and enter the nuclear cavity

No. 454.] STUDIES ON THE PLANT CELL. 739

with the breaking down of this structure. The fibers gather

into projecting cones presenting a multipolar structure, and two

of these, becoming more prominent, absorb the others and thus

form a bipolar spindle.

One of the most recent studies on spindle formation is that of

Osterhout (:02) on Agave. This investigation is of especial

interest for the extensive experimentation in the technique of

fixation. The author proposes a new terminology for the stages

of mitosis that need not be presented here. Agave offers a

striking peculiarity in the presence of a special membrane

around the early stages of the spindle. The fibrillae form inside

of this membrane and finally push through it radially into the

exterior cytoplasm where they gather into cones (Fig. 14 /).

The cones separate into two opposite groups with a general

parallel arrangement of the fibers and in this manner a bipolar

spindle is formed.

It is becoming possible to make some general statements

respecting the methods of spindle formation in the spore

mother-cell. Just previous to prophase it is almost always

possible to differentiate a region of kinoplasm around the

nucleus. This zone has been found to be either granular, e. g.,

Pellia, Anthoceros (Davis, '99 and : o1), Osmunda (Smith, :00),

Cobea and Gladiolus (Lawson, '98 and :00), or it presents the

appearance of a fibrous reticulum, e. g., Equisetum (Osterhout,

'97), Larix (Allen, :03), Lilium (Mottier, '97 and '98), etc.

The latter condition probably develops from the former by the

arrangement of granules into fibers and the gradual expansion of

a very close network thus formed into a coarser structure. The

fibers in this reticulum sometimes surround the nucleus as with

a heavy web. They later extend radially into the cytoplasm,

partly by the expansion of the network and partly by their own

growth and frequently take a radial arrangement. In some

instances the spindle fibers are developed very largely within

the nucleus from the linin (Passiflora, Williams, '99). They

then become gathered into bundles or groups forming the cones

which collectively constitute a multipolar figure that is often

called a multipolar spindle. By the rearrangement of these

cones somewhat parallel to one another, together with more or

740 THE AMERICAN NATURALIST, (VoL. XXXVIII.

less fusion, the multipolar structure becomes a bipolar spindle

(multipolar polyarch) generally just previous to the period of

metaphase. The formation of cell plates and the disappearance

of the spindle fibers have been discussed in Section II under the

title ** Cleavage by cell plates."

Mention should be made of some irregularities in the division

and distribution of the chromosomes that are conspicuous in cer-

tain spore mother-cells and which have been the cause of much

discussion. The subject has especial reference to certain older

views of the reduction phenomena in plants. Chromosomes split

once longitudinally in all typical mitoses and the halves are drawn

apart in a symmetrical manner which is very easily understood.

This division is really determined by the longitudinal fission of

the spirem thread. But appearances during the first nuclear

division in the spore mother-cell of many forms have puzzled

investigators for many years and have given rise to a number

of interpretations. It seems to be pretty clearly established

now that in these types there is a double longitudinal splitting

of the chromosomes at the time of this mitosis. The first divi-

sion takes place during prophase and the second follows closely

after the first and is generally clearly seen at metaphase or dur-

ing anaphase. Therefore the chromatic bodies which appear at

the nuclear plate during the first mitosis are in reality divided or

about to be divided into quarters and they separate after this

mitosis as pairs of granddaughter chromosomes instead of simple

daughter elements. These pairs are either firmly united at one

end into a V or irregularly drawn out so that the bodies have

very unusual and sometimes bizarre forms. Nuclear figures of

this irregular appearance were originally described by Flemming

for the first mitosis in the spermatocyte of Salamandra and named

by him “heterotypic.” These in the spore mother-cell of plants

are of similar character and the designation * heterotypic" has

been adopted by botanists for this condition. The pairs of chro-

mosomes that enter the daughter nuclei after the first mitosis

fuse end to end to form a spirem thread which breaks up again

during the second mitosis, without longitudinal fission, into pairs

of chromosomes which are believed to be identical with those

that entered the nucleus after the first mitosis. Since there is

' No. 454.] STODIES ON THE FLANT CELL. 741

no longitudinal splitting of the spirem thread before the second

division this mitosis differs from that of the “typical” mitoses of

cells and is called **homotypic " to distinguish it on the one hand

from the former and on the other from “ heterotypic " divisions.

Several illustrations of heterotypic and homotypic mitoses to

be described presently are presented in Fig. 15, showing the

peculiar V-shaped pairs of granddaughter chromosomes, charac-

teristic of the first group. It is important to note that whatever

the significance of this premature fission of the chromosomes

before the second mitosis it is not of the nature of a qualitative

reduction division in Weisman’s sense. The details and signifi-

cance of reduction phenomena will be considered in other con-

nections (Section V). The topics discussed above have been

recently studied and reviewed by Mottier (: 03).

We have as yet said nothing of the megaspore mother-cell in

Spermatophytes. An increasing number of investigations have

clearly established the fact that the embryo-sac in many forms

is one of a group of two, three or four cells, each of which is a

potential megaspore because its nucleus contains the reduced

number of chromosomes. We are accustomed to think of the

well known conditions in the lily, where the megaspore mother-

cell develops directly into the embryo-sac. But this type with

some others (e. g., Fritillaria, Tulipa, Erythronium, etc.) are the

exceptions and present a very highly differentiated condition in

which the usual developmental history is shortened in a very

interesting manner, which will be described presently.

The embryo-sac arose undoubtedly as one of four megaspores

developed after essentially the same manner as microspores or

pollen grains, excepting that their arrangement was generally in

a row, which is even true of somè pollen grains (e. g., Asclepias,

Zostera). As stated above, an increasing number of investiga-

tions have established the row of four potential megaspores in a

large number of forms in various groups. They may not always

be distinguished by the form of the group, but their homologies

are established by the mitoses that lead to their differentiation.

Two mitoses are of course required to establish the group of

four cells and both are identified by the reduced number of

chromosomes. Some detailed studies on these mitoses have

742 THE AMERICAN NATURALIST. [Vor. XXXVIII.

established the fact for certain forms that the first is heterotypic

and the second homotypic, exactly as in divisions of the micro-

A EE

S ES

api a adir

CIF

GZ EoD

SRS IY

wes

at, first mitosis in megas

ac or megaspore mother-ceil of spermatophytes. a, 4, Galtonia candicans.

re mother-cell. 42, second mitosis. 23, group of four mega-

em. c2, after the first mitosis. c3, after the second mitosis, the lower cell of the pair to

th bryo sac. c4, after the second mitosis, the upper cell of the pair to become

the embryo sac. di, anaphase of the first mitosis (heterotypic). 42, anaphase of the sec-

ond mitosis (homotypic). e, Lili tagon; portion of embryo sac mother-cell, nucleus

surrounded by a felt of fibrille. /, Lilium candidum; embryo sac mother-cell, nucleus

surrounded by radiating fibrille. g, 4, 7, Lilium martagon. g, late prophase of first

mitosis in embryo sac mother-cell, a multipolar spindle. 4, anaphase of first mitosis

(heterotypic). i, anaphase of second mitosis (homotypic). (a, 4, c, d, after Schniewind-

Thies : o1 ; e, f, g, A, i, Mottier '97.)

spore or pollen mother-cells. Schniewind-Thies (:01) figures

very completely the mitoses in Galtonia. The first mitosis in

the megaspore mother-cell (Fig. 154) is heterotypic because the

No. 454] STUDIES ON THE PLANT CELL. 743

chromosomes (Fig. 154, 1, 2) show clearly the V-shaped forms

characteristic of this division. The second mitosis (Fig. 15 4, 3)

is homotypic. The lowest cell of the group of four (Fig. 15 4,

3) becomes the embryo-sac and the mitoses that take place

within it as the female gametophyte develops are all typical.

This account illustrates a simple history in megaspore mother-

cell development and is considered the first of three types ina

classification proposed by Schniewind-Thies (: 01).

The second type of development is one in which two mega-

spores are generally developed from a mother-cell and one of

these becomes the functional embryo-sac. Schniewind-Thies

presents an excellent illustration of this type in Scilla. The

. first mitosis in the megaspore mother-cell (Fig. 15 c) is hetero-

typic (Fig. 15d, 1) and results in two cells (Fig. 15 c). The

second mitosis in both cells is homotypic (Fig. 154,2). Either

the lower (Fig. 15 ¢, 3) or the upper (Fig. 15 ¢, 4) of the pair

may become the embryo-sac. The embryo-sac then includes the

homotypic or second mitosis within its development, making it

the first nuclear division of the gametophyte history. The typi-

cal mitoses of the gametophyte begin with the second nuclear

division in the embryo-sac. Three megaspores may be formed

in such a. group when the cell of the pair that does not become

the embryo-sac divides again.

The third type of development is illustrated by several forms,

of which the best known are Lilium (Mottier, '98 and :03) and

Tulipa (Schniewind-Thies : or). The lily has been much studied,

but Mottier presents the most detailed account of spindle for-

mation and the behavior of the chromosomes. He supports the

observations of. Schniewind-Thies, based upon the tulip, and her

explanation of this type of development. The megaspore

mother-cell of the lily and tulip develops directly into the

embryo-sac. The first mitosis in this cell (Fig. 15 7) is hetero-

typic and the second (Fig. 15 i) homotypic. These divisions

give the four-nucleate embryo-sac and one more mitosis presents

the mature structure. This last is a typical mitosis, the only one

found in the embryo-sac before the development of the endo-

sperm and sporophyte embryo. Thus the two mitoses charac-

teristic of the spore mother-cell are here included within the

744 THE AMERICAN NATURALIST. [Vor. XXXVIII.

embryo-sac and appropriated as a part of the gametophyte

history.

We can see in these three types of embryo-sac development

an evolutionary process of which the third stage is plainly

derived from the simpler second and first, and is consequently

a highly developed and very complex condition, far removed

from primitive gametophyte structures among the angiosperms.

The embryo-sacs of these forms (Lilium, Tulipa, Fritillaria,

Erythronium, etc.) are probably the most complex spore mother-

cells that we know. The studies of Schniewind-Thies and

Mottier have been supported by other investigations, and more

especially by the results of Ernst (:02) on Paris quadrifolia

and Trillium grandiflorum, who followed the history of the

heterotypic and homotypic mitoses in these forms in detail.

They illustrate the second type of embryo-sac development in

the classification of Schniewind-Thies.

Spindle formation in the embryo-sac mother-cell has not

received as much attention as in the pollen mother-cell, probably

because material of the latter structures may be obtained much

more readily than the former. There have been numerous

descriptions and figures of the spindles but few accounts in full

of their development. 'Of the latter the investigation of

Mottier (98) on Lilium is the most complete. This paper was

written at the time when the centrosome question was under

discussion and served, with other papers on the spore mother- -

cell (Osterhont, '97, Juel, '97, Mottier, '97) to discredit the pres-

ence of these bodies in this structure. Mottier found that the

nucleus of the embryo-sac became invested with a close network

of fibrille (Fig. 15 ¢) from which fibers developed into the cyto-

plasm radiating from the nucleus in all directions (Fig. 15 f).

With the dissolution of the nuclear membrane the fibrilloe

entered the nuclear cavity, filling it with masses of fibers which

gathered into cones to form a complicated multipolar spindle

(Fig. 15g). These cones later come together into two poles,

but even in the mature spindle the fibrilla are frequently in sev-

eral groups atthe poles. Essentially the same history is repeated

in the second mitosis. A large number of later papers have

described and figured multipolar spindles in embryo-sacs, con-

No. 454.] STUDIES ON THE PLANT CELL. 745

firming the conclusions of Mottier that these structures are

developed here after the same methods as in the pollen mother-

cell, from surrounding investments of fibrillæ and without cen-

trosomes. Indeed the embryo-sac is remarkable for the quan-

tity of the cytoplasmic fibrillæ present during its mitoses.

In concluding this account attention should be called to

some forms whose microspore mother-cells were formerly sup-

posed to omit the mitoses of sporogenesis and develop directly

into pollen grains. These conditions were reported in Zostera,

the Cyperaceæ, and the Asclepiadaceæ. However, Juel (:00)

finds the two mitoses present in Carex acuta, although three of

the nuclei break down and the cytoplasm is appropriated for the

fourth to form a single pollen grain whose wall is developed

from that of the mother cell. The history is very similar to

the development of the megaspore in certain heterosporous pteri-

dophytes (e.g. Marsilia, Selaginella) and to the embryo-sac,

which functions while its companion polential megaspores degen-

erate. The development of the pollen in the Asclepiadaceæ has

been shown to be normal in the nuclear activities by several

investigators (Frye, : or, Strasburger, : o1, and Gagner, :02), the

tetrad consisting of four pollen grains in a row, instead of the

usual arrangement. In Zostera (Rosenberg, : O1) there are lon-

gitudinal divisions of the very much elongated pollen mother-cell

to give four extraordinary filiform pollen grains.

5. The Coenocyte.

This remarkable type of cell has reached an extraordinarily

high state of development in certain plants, notably among the

Siphonales and the filamentous Phycomycetes (Mucorales, Sap-

rolegniales and Peronosporales). Ceenocytes are multinucleate

cells. The simplest types are developed by the limited division

or fragmentation of a nucleus accompanied by an increase in the

size of the cell but without extended growth. Excellent illus-

trations are found in the older cells of the red alga, the inter-

nodal cells of the Characeze and in old parenchyma cells of many

higher plants. :

A higher type of coenocyte is presented when the multinucle-

146 THE AMERICAN NATURALIST. [VOL XXXVIII.

ate cells show some definite activity resulting in extensive growth

or peculiarity of form. Thus some laticiferous coenocytes are

branching tubes that grow for considerable distances among the

cells of the tissues in which they are contained. The embryo-

sac and the female gametophytes of Selaginella and Iscetes in

the early stages of their development are interesting coenocytes.

Among the lower algæ there are numbers of Coenocytic forms.

(e. g., Hydrodictyon, Cladophora) whose cells present very little

change with age except an increase in size. Yet some of these

conditions, especially those illustrated in the Cladophoracez, are

probably related to the higher types of coenocytes.

The best differentiated coenocytes are found in the Siphon-

ales, Mucorales, Saprolegniales and to a lesser extent among the

, Peronosporales and are especially well illustrated in a few aquatic

forms, such as Monoblepharis and Myrioblepharis. The pecul-

larities of these forms lie in elaborate structures which result

from the ability of the coenocyte to respond to several directive

stimuli in its growth. The most complicated responses and con-

sequently the most highly differentiated morphology is shown

among the Siphonales, where some very elaborate forms are

found. In many types the plant body is clearly composed of

root and shoot regions and in the highest expressions (e. g.,

some species of Caulerpa) there are rhizoids, shoots and leaf-

like structures presenting a remarkable degree of specialization.

The behavior of the protoplasm in these most highly differenti-

ated types of the Siphonales is known to us chiefly through

studies of Noll and Klemm. .

There is a very conspicuous layer of clear protoplasm next to

the cell wall which constitutes an outer plasma membrane (haut-

schicht). This outer plasma membrane is stationary while the

granular protoplasm within changes its position readily and fre-

quently in different portions of the plant streaming in various

directions. The nuclei are all situated in the granular cytoplasm.

so that they must shift their positions with its movements. Noll

No. 454.] STUDIES ON THE PLANT CELL. 747

method of growth a process of eruption in contrast to Nageli’s

conception of growth by intussusception. Increase in thickness

comes with the laying down of successive lamelle inside the

older wall and is consequently growth by apposition, Caulerpa

is very favorable for such investigations and Noll's results greatly

strengthen the theory that a cellulose wall results from the

direct transformation of a plasma membrane in which carbo-

hydrate molecules gradually replace those of albuminous mate-

rial. Accordingly the cellulose wall is not strictly a secretion

and its growth is not by the intercalation of new molecules

among the old (intussusception) in'a non-living membrane.

The wide space in the interior of the filaments of Caulerpa

and some other members of the Siphonales is frequently crossed

by cellulose bars at various angles. These are at first strands

of protoplasm. which become. gradually filled with a carbohy-

drate material and finally solidified. Noll (884), while recog-

nizing that these structures may have value in strengthening

the filament, believes that they are also the paths of metabolic

exchange between the interior regions of the protoplasm and

the Water outside the plant. They are surrounded by the

plasma membrane which in consequence presents a much greater

extent of surface to the water permeating the cell wall.

It is plain that because of the constant movement of the

granular cytoplasm carrying with it the nuclei which change

their position in the cell, the outer plasma membrane is the only

portion of the protoplasm that can receive fixed stimuli for an

extended period. Consequently Noll regards this membrane as

the responsive or irritable region of the cell that reacts to the

stimuli which largely or wholly direct growth. Some of these

stimuli are well established. Thus it is light which directs the

formation of leaves and shoots. The behavior of Caulerpa in

relation to prominent stimuli (light, darkness, gravity, etc.) has

been studied by Noll (887) and Klemm ('93). The latter author

believes that the response is due to the presence of foods or

other substances at certain points which make them especially

sensitive to the external stimuli. Injuries to a filament of the

Siphonales brings about an immediate flow of protoplasm to the

wounded part (Klemm,’ 94), after which the plasma membrane

is quickly repaired and new portions of the wall laid down.

149 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Mitotic phenomena in the Siphonales is known to us only

through the investigations of Fairchild ('94) on Valonia. He

found that nuclei in the same individual may divide directly or

indirectly. The first process is one of simple fission, the latter

takes place with the formation of an intranuclear spindle.

Studies in sporogenesis and gametogenesis are very much to

be desired in the Siphonales that we may understand the

behavior of the nuclei at these periods. The author's recent

studies of oógenesis in Vaucheria (Davis, : 044) have shown an

interesting process of nuclear degeneration similar to that in the

Saprolegniales and Peronosporales, and suggests some very inter- |

esting lines of investigation. :

The protoplasmic structure in the hyphz of the larger fila-

mentous Phycomycetes, especially the Saprolegniales and Perono-

sporales, is undoubtedly much the same as in the Siphonales.

But the absence of chlorophyll and the greater delicacy of the

filaments makes it more difficult to recognize the different

regions of the protoplasm. There is an outer plasma membrane

inside of which the granular material slowly moves in proto-

plasmic currents that may sometimes be observed in rapidly

growing tips. Delicate strands which are the paths of stream-

ing currents are beautifully shown in developing sporangia of

the molds and the oógonia of the Saprolegniales and Perono-

sporales. The nuclei are undoubtedly carried by the protoplas-

mic movements, sometimes collecting in cnnsiderable numbers

in growing regions of the filaments which always contain much

dense protoplasm.

Another type of ccenocyte, and in some respects the most

remarkable, is the plasmódium of the Myxomycete. These

structures are too well known to need description here. We

shall only refer to them as they help to break down an old

theory that the coenocyte is a compound structure composed of

many energids, represented by the nuclei, which coóperate to

make up the whole. The plasmodium and the protoplasmic

mass inside the cellulose tubes of the Siphonales and Phycomy-

cetes agree in all essentials of structure and mode of growth.

PAs: growth of the plasmodium, as is also true of the

» begins with the prolongation of the outer plasma mem-

No. 454] STUDIES ON THE PLANT CELL. 749

brane (hautschicht, ectoplasm) into a process (pseudopodium)

which advances and is followed immediately by an inflow of the

granular cytoplasm. And the growth of the filaments of the

Siphonales and higher Phycomycetes is a pushing forward of

the outer plasma membrane followed by the granular proto-

plasm, but this growth is slow because the plasma membrane is

at all times under the restraint of a cellulose envelope.

Mention should be made of the remarkable coenocytic zoóspores

well known in Vaucheria-and also described by Thaxter (954),

for the Phycomycete Myrioblepharis. In Vaucheria the entire

contents of the sporangium becomes transformed into an im-

mense multinucleate zoóspore, the cilia being distributed in pairs

above the nuclei. In Myrioblepharis the contents of a spor-

angium usually forms four large multiciliate zoóspores.

These zoóspores of Vaucheria have often been called com-

pound zoóspores, and the idea has been expressed that they

stand for the coóperative union of many hundreds of zoóspores

(energids) represented by the nuclei and their respective pairs of

cilia. And this explanation of the zoospore of Vaucheria is a

part. of a broad view, formerly very largely held, that the

coenocyte is an assemblage of energids (uninucleate masses of

protoplasm) coóperating in a fused structure.

The theory of the coüperative association of energids in a

coenocyte (Sachs) has been very much modified. While the

nucleus and some other organs of the cell, such as groups of

cilia, plastids, etc., are homologous with the same structures in

uninucleate cells nevertheless the behavior of the coenocyte is

not the same as a group of coóperating protoplasmic units. The

coenocyte reacts to the usual stimuli in precisely the same man-

ner as a uninucleate cell, and must be regarded as physiologi-

cally presenting no peculiarities over the latter structure except-

ing those of an increased bulk of protoplasm demanding a greater

number of nuclei for its metabolic processes. The most impor-

tant contribution presented by the coenocyte to our knowledge

of the physiology of the cell is the establishment of the plasma

membrane as the region of the protoplasm responsive to the

stimuli that determine the form assumed in growth. The con-

stant shifting of the nuclei and plastids in the movement of the

750 THE AMERICAN NATURALIST. [Vor. XXXVIII.

granular protoplasm eliminates them as structures immediately

concerned with the form of a cell or organ thus limiting their

functions more especially to metabolism.

6. The Coenogamete.

The conogamete is a multinucleate sexual cell.. The name

was first applied by the author (Davis, :00, p. 307) to the

remarkable multinucleate eggs of A/bugo bliti, and the concep-

tion has been considerably extended since, as explained in his later

writings on Saprolegnia (Davis, :03, p. 320-331) and on ** The

relationships of sexual organs in plants” (Davis, : 04%). Ste-

vens ('99) discovery of the multinucleate eggs of Albugo bliti

opened a field of research that has been greatly extended in the

past four years and which is likely to yield very important con-

clusions on the relationships and evolution of the Phycomycetes

and Ascomycetes. Conditions similar to Albugo bti were

reported the following year by Harper, : 004, for Pyronema, and

several later papers have described, with greater or less fullness,

the structure and behavior of coenogametes in some other Asco-

mycetes, types of the Peronosporales (species of Albugo) and in

the Mucorales.

We shall not discuss the details of these investigations with

their bearings upon the problems of phylogeny as this has

become a very complicated subject and is treated elsewhere

(Davis, : 04 a—6), but merely describe the structure and behavior

of coenogametes so far as they are known to us.

Stevens and Harper both found that the multinucleate female

cell of Albugo bliti and Pyronema was fertilized by the introduc-

tion of a large number of nuclei from the antheridium. These

sexual nuclei paired off and fused, a male with a female, in the

common mass of cytoplasm so that the fertilized cell finally con-

tained a large number of fusion nuclei. A similar history was

reported later by Stevens (:01), in Albugo portulacae and Albugo

tragopogonis. These events have been so thoroughly studied

that we know the processes of fertilization in the above forms as

well perhaps as for any plant type.

The structure and especially the nuclear history of other

No. 454.] STUDIES ON THE PLANT CELL. 751

coenogametes is less perfectly understood. The multinucleate

character of the fusing gametes is well known, but the later dis-

tribution and fate of the sexual nuclei has not been followed, and

it is by inference that we believe these coenogametes to behave

in essentially the same manner as those of Albugo and Pyronema.

Coenogametes fall into two classes according as they involve

all of the protoplasm contained within the mother-cell or only a

portion of such protoplasm. The first group probably represents

the simplest and most primitive conditions.

Ceenogametes of the first class are found in the Mucorales

(Gruber : o1) and in the Gymnoascez (Dale, :03). In these

types the entire contents of the terminally formed sexual cells

unite to produce the zygospore in the former group and the

fertilized ascogonium in the latter, from which arises the system

of ascogenous hyphe.

Coenogametes of the second class contain only a portion of

the protoplasm in the mother-cell which is usually a terminal

structure. The protoplasm that is not involved in the coeno-

gamete proper generally bears some important relation to the

sexual element. Thus the periplasm of the Peronosporales

assists in the formation of the wall of the oóspore and the con-

jugation tube of Pyronema becomes the path through which the

contents of the antheridium enters the ascogonium. But in

some forms the superfluous protoplasm is merely cut off from

the coenogamete as a sterile cell (Monascus). In Albugo and

Pyronema the sterile and fertile portions of the protoplasm are

so closely associated that the mother-cell really acts as a whole,

very much as the simplest types of coenogametes which shows

the close relationships between the two. Moreover the anthe-

ridia of these forms are types of coenogametes almost as simple

as those of the molds or the Gymnoascee.

The coenogamete is a type of sexual cell unknown in the

animal kingdom and among plants is probably restricted to the

Phycomycetes and Ascomycetes. The problems of its homol-

ogies and origin are very interesting.

The simplest types of coenogametes (Mucorales and Gymno-

ascez) are cells situated at the ends of filaments in the same

position as the sexual organs of the Siphonales. The mother-

752 THE AMERICAN NATURALIST. [Vor. XXXVIII.

cells of the more complicated coenogametes (oógonia and anthe-

ridia) are also terminal cells. All of these sexual organs are

multinucleate. In the Siphonales (Vaucheria excepted) all of

the nuclei are functional gamete nuclei. This is also true of

simplest types of Coenogametes, but in the more complicated

forms (Albugo, Pyronema, etc. large numbers of the nuclei

degenerate or fail to function sexually in sterile accessory re-

gions of the protoplasm. The same conditions of sexual degen-

eration are also found in the oógonia of Vaucheria (Davis, :04)

and Saprolegnia (Davis, :03). The agreement of all of the

structures mentioned above in structure and protoplasmic behav-

ior seems to establish beyond question their common homology.

The problems of the origin of the coenogametes are very

difficult with the meager evidence at hand. The author believes

that the simplest types have probably been derived from struc-

tures like the sexual organs of the isogamous Siphonales, which

Structures gave up the habits of forming uninucleate gametes

and acting as ccnocytic units became multinucleate sexual ele-

ments. A physiological development very similar to such a

change must have taken place in Peronospora and some species

of Pythium when their conidia ceased forming zoóspores and

took the habit of germinating directly by a tube. This view

regards the ccenogamete as a coenocyte derived from a proto-

plasmic structure that at óne time produced a large number of

independent sexual elements, represented in the coenocyte by the

numerous nuclei,

(Albugo, Pyronema, etc.) have developed directly from the sim-

evels of the heterogamous algae, such as are

not be treated here, T

have been considered in recent papers of the author (Davis, : 03,

:0944, 046). Ccenogametes are proving to be among the most

interesting types of sexua] cells in plants and research in this

field is likely to prove very fruitful of results.

No. 454.] STUDIES ON THE PLANT CELL. 753

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758 THE AMERICAN NATURALIST. [Vor. XXXVIII.

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No. 454.] STUDIES ON THE PLANT. CELL. 159

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760 THE AMERICAN NATURALIST. [Vor. XXXVIII.

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ena.

(To be continued.)

THE AFFINITIES OF THE OPHIOGLOSSACE/E

AND MARSILIACEZE.

DOUGLAS H. CAMPBELL.

Tne last ten years have been notable for many important

contributions to our knowledge of the pteridophytes, due in part

to a more exact investigation of many forms already studied, but

still more to a critical study of tropical types, which hitherto

have been studied more or less superficially, owing to the diffi-

culty of procuring suitable materials for complete investigations.

The fossil forms have also yielded much important material for

a better understanding of the affinities of some of the living

ones.

The work has covered a wide field and comprises very valu-

able additions to our knowledge of the anatomy and develop-

ment of many interesting types which were but imperfectly

known before. These investigations have added materially to

the data available for a critical comparison of the different

groups, and we are in a much. better position than ever before

to understand the affinities of some of the more puzzling types

of pteridophytes. It must be admitted, however, that the con-

clusions of botanists in regard to the relationships of certain

groups are by no means entirely in accord.

There are two orders of ferns about whose relationships there

has been a good deal of controversy, and it is the bearing of

some of the recent investigations upon these relationships to

which the writer would direct attention. The two orders referred

to are the Ophioglossaceze and the Hydropteridinez, especially

the Marsiliacez.

THE OPHIOGLOSSACEJE.

The Ophioglossacez constitute a very natural family of fern-

like plants, evidently closely related among themselves, but

761

762 THE AMERICAN NATURALIST. [Vor. XXXVIII.

whose affinities to the other pteridophytes have been much dis-

cussed. They are generally considered to belong to the Fili-

cinez, or ferns with which they agree in their general structure.

The writer has expressed the opinion that the sporangium of

Ophioglossum represents the lowest type found in the fern

series. This conclusion is based upon the structure of the living

forms, as the Ophioglossacez are almost unknown in a fossil

condition, perhaps owing to the soft tissues which characterize

them. These would hardly be preserved as fossils except under

the most favorable conditions.

As is well known, the Ophioglossacez differ from the typical

ferns in the position of the sporangia, which are borne upon a

peculiar structure (sporangiophore) arising from the inner sur-

face of the leaf, either directly from the lamina or from the

petiole. The sporangiophore may be a simple spike with two

rows of large sporangia scarcely projecting at all, or it may

be much branched and the numerous sporangia quite distinct

and even stalked. (Campbell, Mosses and Ferns, p. 296.) The

former structure occurs in Ophioglossum, the latter in Botrychium

and Helminthostachys. The simpler species of Botrychium, such

as Botrychium simplex and Botrychium lunaria, form a transition

between the type of Ophioglossum and the larger species of

Botrychium and Helminthostachys, in which both the sterile and

fertile leaf segments are much branched, and the free sporangia

bear a certain resemblance to those of Osmunda or Angiopteris.

It is possible that the latter types, or at any rate Osmunda, may

have arisen from forms like Botrychium or Helminthostachys,

through which they would be connected with the more primitive

type of sporangium found in Ophioglossum.

Much the most important recent work upon the sporangia of

these puzzling forms is that of Bower (* Studies in the Morphol-

ogy of Spore-Producing Members"; Phil. 7; rans., 1894—1903,

185-196), whose views as to the relationship of the Ophio-

glossacez differ radically from those of most students of the

pteridophytes. He considers the whole spike of Ophioglossum

as the equivalent of a single sporangium of Lycopodium, pro-

duced by growth and septation, from an originally unilocular

structure. He is not inclined to admit any direct relationship

No.454] OPHIOGLOSSACE 4 AND MARSILIACE4 763

between the Ophioglossacez and the true ferns, and thinks they

are probably more nearly related — although very remotely — to

the lycopods. In his most recent work (doc. cit. No. V, p.253),

however, he recognizes the importance of the decidedly fern-like

character of the gametophyte and ‘recedes somewhat from the

extreme views expressed in the earlier paper. He seems still

unwilling, however, to give up the homology between the sporo-

phyll in Ophioglossum and Lycopodium. It is held that all the

pteridophytes are reducible to a common strobiloid type, which

is most clearly seen in the lycopods and Equisetacez, but which

is believed to be recognizable in the ferns as well. With this

view the writer has been unable to agree.

Bower thinks it inconceivable that a large-leaved type could

originate directly from any conceivable form of bryophyte sporo-

gonium, and therefore concludes that all the primitive pterido-

phytes must have been microphyllous. To the writer there

seems to be no greater difficulty in assuming that a single leaf-

like organ may have been developed at the base of the sporogo-

nium (Mosses and Ferns, p. 515) than that a series of small

leaves should be developed as eruptions from its surface. Why

both forms may not have been developed quite independently is

not clear, either from the evidence of comparative morphology,

or from the fossil record. That a special assimilative structure,

remotely comparable to a leaf, may originate from the base of

the sporogonium is shown in many mosses, where the apophysis

is of this nature. In certain species of Splachnum the apophysis

assumes a relatively very large size, and is expanded into a disc

which might almost be compared to a perfoliate leaf. Of course

there is no question of a direct homology between the apophysis

of a moss and the leaf of a vascular plant, but the fact is patent

that the sporogonium of a moss may give rise to a special assim-

ilating organ, and it is therefore quite conceivable that the pro-

genitor of the large-leaved ferns may have arisen from some

bryophytic type by an analogous formation of a special organ

which became a’ true leaf subtending a sporogenous structure

much as is actually seen in Ophioglossum. i

That the Ophioglossacez represent an ascending series, as

Professor Bower (Bower, /oc. cit. No. 5, P- 233) believes, is

764 THE AMERICAN NATURALIST. [Vor. XXXVIII.

extremely probable; but that the lowest members of the series

are to be connected with forms in any way like the Lycopodiaceze

appears to the writer exceedingly doubtful. The direct origin

of a type like Ophioglossum from an Anthoceros-like prototype

has been considered at length elsewhere (Campbell, — /oc. cit.,

p. 515) and will not be discussed further here.!

The numerous structural resemblances in both gametophyte

and sporophyte between the Ophioglossaceze and Marattiaceze

are too great to be passed over, and indicate a genuine relation-

ship between the two. It seems to the writer that the impor-

tance of these obvious resemblances has been rather minimized

by some recent authors, and it may be just as well to call atten-

tion afresh to some of the most striking ones.

Our knowledge of the gametophyte of the Ophioglossaceze is

now pretty complete, owing to the important researches of Jef-

frey (The Gametophyte of Botrychium virginianum, Proc. Canad.

Inst. V, 1898) and Lang (On the prothalli of Ophioglossum pen-

dulum and Helminthostachys zeylanica, Ann. Bot. 16, 1902).

The former has described in detail the prothallium of Botry-

chium virginianum, while Lang was fortunate enough to pro-

cure fairly complete material of Ophioglossum pendulum and

Helminthostachys seylanica, so that all three genera are repre-

sented. As might be expected, there is a good deal of similar-

ity, this being most marked between Botrychium and Helmin-

thostachys. All agree in having a subterranean prothallium

which is destitute of chlorophyll, thus resembling the prothallia

of certain species of Lycopodium. It is highly improbable that

this condition is a primitive one, however, and too much stress

cannot be laid upon it in making a comparison with the green

prothallia of the Marattiacez, with which otherwise there is a

close agreement, especially in the case of Ophioglossum. As

some species of Lycopodium have a green prothallium, and chlo-

rophyll has been found under certain circumstances in the pro-

! Since this paper was written there has been published by Bower (Ann. of Bot.

Apr. 1904) an account of a remarkable species of Ophioglossum from Sumatra, in

which the leaf consists simply of a long-stalked sporangiophore with scarcely a

trace of the sterile segment. This interesting plant almost realizes the hypothet-

ical form assumed by the writer as the ancestor of the ferns. |

No. 454] OPHIOGLOSSACEÆ AND MARSILIACE&. | 265

thallium of Ophioglossum (Mettenius — /2/ices Hort. Bot. Lips,

1856), it is extremely likely that the absence of chlorophyll is a

secondary condition, and that so far as the prothallium is con-

cerned, the Marattiacez approach more nearly the ancestral

type than do the Ophioglossacez.

Both Jeffrey and Lang are agreed as to the essentially fern-

nium; 2, young antheridium of O7 so. C, D, after Lang.

Fic. 1, — A, archegonium ; 2, young antheridium of Marattia douglasii, X 350. C, archego-

' hioglossum pendulum, X 2

. like character of the prothallium. This is shown in its apical

growth, strongly dorsiventral form, and especially in the repro-

ductive organs, which in Ophioglossum (Fig. 1, 4, 4) resemble

to an extraordinary degree those of Marattia. The large multi-

ciliate spermatozoids are also entirely fern-like in form. When

to the close resemblance in the sexual organs is added the

marked correspondence in the development of the embryo,

which is very similar to that of the Marattiaceze and not in the

least like that of Lycopodium, the case seems a very strong one.

There next remains to be considered the structure of the

adult sporophyte. In most species of both Ophioglossum and

Botrychium the stem is a short upright rhizome about which the

leaves are spirally arranged. In Ophioglossum pendulum, how-

ever, and in Helminthostachys the plant is dorsiventral.

Among the Marattiacez both types also occur. Marattia and

Angiopteris are radial, Danza is dorsiventral. Bower (oc. cit,

766 THE AMERICAN NATURALIST. (Vou. XXXVIII.

p. 219) is inclined to consider the radial type as older than the

dorsiventral, but as both types occur in both families under con-

sideration, this has little bearing upon the question of their rela-

tionships.

In the structure of the apical meristems the Ophioglossacez

are even more fern-like than the Marattiacez, as they regularly

show a single tetrahedral initial in both stem and root. In this

respect they resemble the leptosporangiate ferns more than they

do the Marattiacee. In the very young sporophyte of the lat-

ter, however, the root shows a single apical cell which is replaced

by a group of initials in the more massive roots of the older

sporophyte.

The very complicated vascular system of the older sporophyte

in the Marattiacez is undoubtedly associated with the massive

stem. A study of the young plant shows that in the earlier

stages there is a single cylindrical stele much like that which

occurs in the adult rhizome of Helminthostachys and such lep-

tosporangiate ferns as Gleichenia. This is considered by Jeffrey

to be a primitive condition from which the more complicated

arrangement found in the adult stem of the Marattiaceze has been

derived. The arrangement of the vascular bundles in the stem

of Ophioglossum is much like that in the typical ferns, while

Botrychium closely resembles Osmunda. (Jeffrey, “Structure

of the Stem in the Pteridophyta and Gymnosperms,” PL

Trans. 195, 1902.)

The leaves of the Ophioglossacez, which are in some respects

different from those of the typical ferns, nevertheless are much

more like these than like either the lycopods or Equisetacez.

The venation is not essentially different from that of the Marat-

tiaceze, and although there is never found the circinate form of

the young leaf, so characteristic: of the ferns, still in the larger

species of Botrychium there is an approach to this which sug-

gests that the coiling of the young leaf is connected with the

very great development of the lamina of the leaf which charac-

terize most ferns.

While the reticulate venation found in Ophioglossum is differ-

ent from that in most ferns, still there are many ferns which

show a very similar venation. Thus among the Marattiacez

No. 454] OPHIOGLOSSACE 4! AND MARSILIACE 767

there may be instanced Kaulfussia, whose venation is not very

different from that of Ophioglossum, and possibly a further

comparison might be made in the form of the leaf with the

palmately lobed leaf of Ophioglossum palmatum. On the other

hand, the venation of Agiopteris or Danza is almost identical

with that of Helminthostachys. (See Fig. 2.)

Farmer (Farmer & Freeman, “The Structure and Affinities

of Helminthostachys," Ann. Bot. 1899) has referred to the stip-

ules of the latter genus and thinks they are not directly compa-

Fic. 2.— A, rhizome of Helminthostachys zeylanica, after Farmer ; B, part of a leaflet E the

same plant, showing the venation; after Hooker & Baker: C, Rhizome of Danea alata,

(reduced); D, base of leaflet showing venation ; st., stipules.

rable to those of the Ophioglossacez, although it is not quite

clear wherein the difference consists. There certainly seems to

be no essential difference between the stipular structure at the

leaf base in Botrychium (Campbell, Mosses and Ferns, p. 242)

and those in Marattia or Danza, although in the latter genera

the stipules are more distinct. The development of stipular

structures is especially marked in both Ophioglossacez and

Marattiacez, while among the Leptosporangiate they are gen-

erally absent except in the Osmundacee, which are admittedly

the nearest relatives among the Leptosporangiate to the euspo-

rangiate ferns.

The markedly dorsiventral rhizome of Danza is strongly sug-

168 THE AMERICAN NATURALIST. [Vor. XXXVIII.

gestive of that of Helminthostachys (Fig. 2, 4, C), and the

position of the leaves and roots is much the same. The cylin-

drical stelar tube with relatively small foliar lacunze in Helmin-

thostachys may perhaps be compared to the stele in the young

rhizome of Danza, which, according to Jeffrey, is of much the

same type. (Jeffrey, /oc. cit., pp. 120, 121.)

The most marked biscolriràl difference between the Marat-

tiaceæ and the Ophioglossacez is perhaps the occurrence of

conspicuous mucilage ducts in the former. These are, however,

also wanting in the other ferns. The circinate form of the

young leaves is probably associated with the great development

of the lamina of the leaf, but is certainly a difference of some

importance.

To the labors of Bower we owe by far the most important

contributions to our knowledge of the development of the sporan-

gium that have been made for many years. These enable us to

make a detailed comparison of all the developmental stages in

each group, and a careful study of his descriptions and figures

of the two groups in question have led me to a somewhat dif-

ferent result from that reached by him.

The progression from the sporangial spike of Ophioglossum

to the much branched sporangiophore of Botrychium or Helmin-

thostachys, with its numerous distinct sporangia, is comprehen-

sible enough; but the relation of the sporangiophore of the

Ophioglossacez to the sporangia of the other pteridophytes is

not so obvious. Bower believes (/oc. cit., p. 250) that all pteri-

dophytes may be reduced to the strobiloid type, this being

reduced to a single sporophyll in Ophioglossum. As we have

stated elsewhere, this theory seems hardly satisfactory when

applied to the ferns, however probable it may appear for the

other pteridophytes. The comparison of the sporangial spike

of Ophioglossum to the single sporangium of Lycopodium

appears rather improbable, and it seems more in accordance

with the facts to consider it an entirely distinct development,

derived directly from the whole upper part of the sporogonium

of the ancestral form.

Egi Pom gin account of the sporangia in Ophio-

arattiaceze suggests a possible point of con-

No. 454.] OPHIJOGLOSSACE 4A AND MARSILIACEZ 769

tact between the two groups. In the very peculiar Ophzoglossum

palmatum, there may be several nearly sessile sporangiophores

borne near, or actually upon, the margin of the large palmately

lobed sterile segment of the leaf. Among the Marattiacee

Esh

Fic. 3.— A, leaf of Ophioglossum palmatum, showing the positi — oia

phores, s., — after Bower; Z, part of a sporophyll of Danæa alata, T r

synangia, sf.; X 2.

Danza shows a marked resemblance to Ophioglossum inthe

arrangement of the loculi in the greatly elongated synangia,

which almost completely cover the lower surface of the con-

tracted sporophylls. A study of the development of the sporangia

in the two, to judge from Bower’s account and figures, shows a

close resemblance in many particulars.

Bower thinks that the elongated synangium of Danza has

been secondarily derived from a circular one, like that of Kaul-.

770 THE AMERICAN NATURALIST. (VoL. XXXVIII.

fussia, but there is no certain evidence, either morphological or

geological, that the type found in Danæa may not be the older.

While a direct comparison of the sporangiophore of the existing

species of Ophioglossum with the synangium of Danæa would

be ħardly justifiable, still it is conceivable that synangia of the

type of Danæa might have originated from the complete adhe-

sion to the surface of the sterile leaf segment of a series of

small sporangiophores not so very different from those actually

found in Ophioglossum palmatum (Fig. 3). If the primitive

character of the synagium of Danæa is admitted, it is easy to

trace the origin of the marattiaceous types by the expansion of

the leaf lamina. Bower suggests that the primitive type of

Marattiaceæ probably had an undivided leaf like that of Daneæa

simplicifolia, with scattered round sori similar to those of Kaul-

fussia. We agree with this except as to the character of the

sori, which we believe were quite as probably like those of the

actual Damea simplicifolia.

In comparing the two families, Ophioglossaceze and Marat-

tiacez, it is clear that they closely resemble each other in the

character of the reproductive

organs and embryo, and also in,

the apical growth and dorsiven-

tral form of the gametophyte.

The position of the archegonia

and the absence of chlorophyll

are undoubtedly associated with

the subterranean life of the

gametophyte.

While in the sporophyte such

e differences as the coiled verna-

iubes = wh pore of ite et — = tion of the leaf, and presence of

Xa B, cross-section of the same; C,cross. Mucilage ducts in the Marat-

trace de ment pocos T" tiaceze must be taken into ac-

count, the two orders neverthe-

less agree in the general morphology of both stem and leaf,

including the venation of the leaves, while in regard to the

apical growth of the stem and root, the Ophioglossaceze are

more like the higher ferns than are the Marattiaceze. Finally,

No. 4:34] OPHIOGLOSSACE 4 AND MARSILIACEK. 77i

whether or not the possibility of a direct connection between the

two is admitted, there is no question of the strong resemblance

between the sporangia of Ophioglossum and Danza. In view

of all these facts, we can see no valid reason for removing the

Ophioglossaceze from their association with the ferns, and we

believe that their association with the Marattiacez among the

eusporangiate ferns is entirely justified.

THE MARSILIACEXE.

That the Marsiliacez are directly allied to some group of

homosporous leptosporangiate ferns is apparent; but there has

been little attempt to connect them directly to any of the existing

ferns. The writer (/oc. cit., p. 421) has suggested a comparison

| 20 6 | |

SIN oS eleg -

Sail

mets!

Ses

29.

Fic. 5.— A, cross-section of a fertile leaf-segment of Schizæa pennula, highly magnified; s.,

sporangium mother-cell — after Prantl; Z, cross-section of very young sporocarp Py lar-

silia quadrifolia, X goo, after Johnson. The marginal cell, sf., gives rise to a row of

sporangia. |

with the peculiar Ceratopteris, and has also compared thé anthe-

ridium with that of the Polypodiacee. !

The more recent work of Belajeff (“Die mannlichen Proth-

alien der Wasserfarne," Bot. leit, 1898) on the antheridium,

and of Johnson (* On the leaf and sporocarp of Marsilia," Ann.

Bot. XII, 1898) (** On the leaf and sporocarp of Pilularia,” Bot.

Gaz. XXVI, 1898) and Goebel (Organographie der Planzen,

772 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Part II, 1900) upon the sporocarp, have suggested a somewhat

different affinity, and it was thought worth while to look into the

evidence carefully, supplementing it by a direct comparison of

such material as was available.

Goebel has called attention to the similarity between the young

sporocarp of Marsilia and the fertile leaf-segment of Schizza.

Johnson's studies on the sporocarp of both Marsilia and Pilularia

bear out this, and a comparison with Prantl’s figures of the

Fic. Ed » Sporophyll of Z4 xeimia hirsuta, showing the very conspicuous fertile pinnze, sf. ;

B, Plant of Mearsilia salvatrix, with the long-stalked sporocarps, s., after Sachs.

young sporophytes of Schizea emphasizes the marked resem-

blance between the two (Fig. 5). Johnson does not admit the

foliar nature of the sporocarp ; that is, he does not believe the

capsule is homologous with the leaf lamina, a view which has

been expressed by the writer, and which is supported by most

students of the subject. A comparison of the developing fertile

segment of the leaf of Schizæa, however, shows that at the

time the first rudiments of the marginal sporangia appear, there

IS no more trace of a lamina than ina corresponding stage of

the sporocarp in Marsilia. In short, the structure of the very

No. 454.) OPHIOGLOSSACE4 AND MARSILIACEX. | 773

young sporocarp of Marsilia and the fertile leaf segment of

Schizea is remarkably similar; moreover, the origin of the spo-

rangia from originally marginal cells is noteworthy.

The origin of the sporocarps in the Marsiliacez, however,

o6)

Frc. 7.— A, cross-section of the rhizome of Marsilia vestita, X about 15; B, similar section

‘Aneimia hirsuta. The outer cortical tissue is composed in both of

he tubular stel loses a medulla al 1 of sclerenchyma ;

he tubular steie Į , 1

odermis, outer (o. 7.) and inner (7. £)

the stele has an outer (o. e.) and

of the rhizome of

tb. 1 1

e stel n

pericycle; outer (a. #4.) and inner (7. ph.) phloem; and a central xylem (x).

is different from that of the fertile leaf-segment in Schizza, but

finds a close counterpart in the allied genus Aneimia (Fig. 6)

where the remarkably developed fertile pinnze bear much the

i mia hii x8; both

— A, leaflet of Nearsilia vestita, X 10; B, pinnule of Anemia hirsuta, ;

tio:

Fic. 8

a strictly dichotomous venation.

hes

hat the sporocarp does in

porocarp in Marsilia and

same relation to the rest of the leaf, t

parable to what obtains

Marsilia. The pinnate form of the s

the dichotomous sterile segment, are com

774 THE AMERICAN NATURALIST. [Vor. XXXVIII.

in some species of Schizzea, where the leaf is dichotomously

divided, but the fertile segments are pinnate.

A comparison of the vegetative organs of the Marsiliaceze

and Schizaaceze shows equally marked resemblances. The

creeping rhizome of Marsilia and Pilularia is remarkably similar

to that of Schizzea and certain species of Aneimia (Fig. 7), as

may be clearly seen from the figure. The leaves also show

interesting resemblances. Thus the filiform leaf of Pilularia

finds its exact counterpart in the sterile leaves of Schiz@a pusilla,

while the four-lobed leaf of Marsilia is comparable to such spe-

cies as Schizea pennula, with dichotomously branched lamina.

The venation in Schizza and Aneimia is characteristically dichot-

omous, closely resembling that of Marsilia. Except in Lygo-

.dium, the pinnate venation

found in the majority of the

ferns does not occur among

the Schizzeacez.

So far as I am aware, no

direct comparison of the spo-

rangia in the two families has

been made, although the early

stages of the sporangium have

been carefully studied. The

Schizeeaceze differ from all

the other ferns in the very

_an peculiar form of the sporan-

gium which is strongly oblique

and has a terminal annulus.

X The sporangia of Pilularia

s ote, ad americana and Marsilia ves-

FIG: 9.4, sporangium of Schisea pennula, show- fita were examined, and it

ing the apical annulus, a». x 45, (after Prantl), eos

koe parue d Pilularia Americana; the Was found, especially in the

C, apical region, more mr bh. se ug case of the former, that the

sporangia resembled strongly

those of the Schizzeacez. In Marsilia the closely packed micro-

spore rendered it difficult to make out the form of the parietal

cells, but in Pilularia the spores are widely separated and the

form of the parietal cells is easily seen when the sporangia,

KEN

£N

—

No.454] OPNIOGLOSSACEÆ AND MARSILIACEÆ 775

. after placing them in water for a short time, are treated with

alcohol to check the swelling of the mucilaginous cell walls

which otherwise are soon completely destroyed. The oblique

form of the sporangium is especially conspicuous in Pilularia

(Fig. 8, B), and the arrangement of the cells near the apex is

very suggestive of the annulus of Schizza, although of course

in these aquatic plants, no hardening of the annulus cells takes

place.

This remarkable correspondence in the structure of the spo-

rangium, taken together.with the other close structural resem-

blances, justifies the assumption of a not very remote relation-

ship between the Schizaeacez and Marsiliacez.

It may be added that this view would tend to confirm the

contention of Belajeff, that in the male gametophyte there are

two antheridia, and not a single one as the present writer has

held. The antheridium of the Schizzeacez 1? is relatively sim-

ple in structure, and might very well be compared to each of

the groups of sperm-cells in Marsilia, with its two peripheral

cells.

1 Bauke — Bertrüge zur keimungsgeschichte der Schizeacee Prings. Jahrb.

XI, 1878.

2 Britton, E. G. & Taylor,—A life history of Schizza pusilla, Bull. Torrey Bot.

Club, XXVIII, 1901. ;

(Wo. 453 was issued Oct. 15, 1904.)

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AMERICAN NATURALIST.

Vor. XXXVIII. November—December, 1904. Nos. 455-456.

THE EMBRYOLOGICAL DEVELOPMENT OF THE

SKELETON OF THE HEAD OF BLATTA!

WILLIAM A. RILEY.

Our knowledge of the embryological development of the

sclerites of the head of insects is very meager. With but few

exceptions the subject has been treated in the most incidental

manner and the observations are few and scattered. This is due

not only to the fact that attention has been directed to the more

general features of insect embryology, but is due also to a lack

of systematizing of our knowledge of the structure of the head

of the adult insect and to the difficulty of determining the

limits of the sclerites before they become firmly chitinized.

Of all the students of insect embryology, Heymons (’95 and

'97), Janet ('99), and Comstock and Kochi, (:02), are the ones

who have devoted the most attention to this subject and these

workers differ radically in their conclusions.

It was in hopes of determining the conditions in one of the

more generalized of the pterygote insects that I undertook the

study of the embryological development of the skeleton of the

1Contribution from the Entomological Laboratory of Cornell University.

777

778 THE AMERICAN NATURALIST. [Vor. XXXVIII.

head of Blatta germanica. This was proposed by Prof. Com-

stock and to him I am indebted for many suggestions.

In thë choice of an object for this study I was guided by the

desire to, select one of the more generalized forms and one which

at the same time would afford an abundance of material. Both

of these conditions seemed to be met by Blatia germania.

Though undoubtedly highly specialized in some respects, the

cockroach is regarded by many students as the most primitive

of ourpterygote insects. I believe that in this respect one of

the Plecoptera would have been more valuable, but as regards

abundance of available material it would be difficult to select a

‘form more satisfactory than is Blatta.

On thé other hand, as objects of study the embryo of Blatta

present certain difficulties which are accentuated in the course

of the work. Of these there are **die ausserordentlichen tech-

nischen Schwierigkeiten" which Cholodkowsky has so empha-

sized. These are due primarily to the great mass of yolk, which

becomes so brittle as to make it almost impossible to secure

complete series of sections by ordinary methods. A more

serious difficulty in the work under consideration was the small

size of the cells and the indistinctness of the invaginations of

thé body wall. Of the latter, Wheeler ('89, p. 349) says: “In

Blatta the formation of the nervous system in its earlier stages

cannot be clearly seen from the exterior. The same holds true

of the small tracheal invaginations, though several pairs, especially

those of the thorax and basal abdominal rings, may be seen on

the plurz in good preparations before revolution. Still they are

so much less distinct than in Doryphora that I have given them

little attention."

THE PRIMITIVE SEGMENTS OF THE HEAD.

The question of the number of segments in the body of the

perfect insect early attracted the attention of entomotomists.

As the history of the development of our knowledge of the

number entering into the structure of the head has been very

fully discussed by Comstock and Kochi (:02), I have but little to

add to their account.

- Nos. 455—-456.] | HEAD OF BLATTA. 779

As early as 1839, Newport discussed this question in consid-

erable detail. He believed in the “correctness of the opinions

advanced by Savigny and others, that the organs of manduction

are the properly articulated members of distinct segments and

are perfectly analogous to the proper organs of locomotion.” He

attributed the conflict of views to a “ too exclusive examination

of the head in perfect insects, without reference to the corre-

sponding parts in the larva.” ;

It was thus early realized that it is to the ontogenetic record

that we must appeal for a settlement of the question. Unfor-

tunately Newport chose as the basis of his study ‘the larva of

Musca vomitoria, a form so highly specialized as to be worthless

for this purpose. His evidence regarding the possible presence

of a fifth segment must therefore be rejected. The value of

his conclusion that there were at least four segments was due to

his acceptance of the criterion proposed by Savigny.

Zaddach ('54), believed that six segments entered into the

composition of the head. Of these the first was the ocular,

while the third was the second antennal segment. His evidence

regarding the presence of the latter, however, was based upon

the erroneous conclusion that in the phryganids the larval

antennz represented the second pair of the Crustacea, and that

they were later replaced by the definitive antennz of the adult,

which corresponded to the antennules of the Crustacea.

In 1858 Huxley also discussed the mooted question from the

view point of embryology. With characteristic thoroughness

and penetration he not only treated of the embryology of an

insect, Aphis, but compared its development with that of Crus-

tacea and Arachnida. Granting that the presence of paired

appendages was indicative of an equal number of segments, he

argues that the absence of the appendages does not necessarily

imply the absence of the segment. “No one will pretend that

it is so in the abdominal and thoracic regions." In conclusion he

proposes the hypothesis * that in the Articulata the head is

normally composed of six somites, which are all fully developed

only in Podopthalmia, Stomapoda, and some Branchiopoda,

while in other Crustacea some one or more of the preoral somites

is more or less abortive, and in Arachnida and Insecta the

780 THE AMERICAN NATURALIST. [Vor. XXXVIII.

appendages of the first somite are sessile and those of the sec-

ond or third undeveloped.”

I shall have occasion to revert to this paper of Huxley's, for,

as we shall see, he devoted considerable attention to a consider-

ation of the development of the head, and brought out some

facts of prime importance.

. Packard (66), believed that the insectean head was composed

of seven segments. In 1870, in the second edition of the Guzde

to the Study of Insects, he gives a table of these segments and

their appendages. He believed the segments to be a first ocel-

lary, second ocellary, opthalmic, antennary, mandibular, first

maxillary, and a second maxillary or labial. In addition to these

seven he includes a hypothetical first segment whose tergal parts

give rise to the labrum, epipharynx, and clypeus.! Thus at this

time he regarded the insectean head as composed of eight prim-

ary segments.

In 1871, however, Packard presents evidence to show that

the ocelli and the compound eyes do not represent appendages.

* Accordingly," he says, “we seem forced to the belief that the

head of the hexapodous insect consists of but four segments, 7. £.,

the second maxillary, the first maxillary, and mandibular seg-

ments behind the mouth opening arid the antennary, or first and

preoral segment situated in front of the mouth." This view he

reiterates in 1883. In 1898 he tabulates six segments.

One of the most recent workers to approach this subject is

Janet ('99). Basing his conclusions largely on a study of the

musculature of the adult Vespa, this worker states that the

so-called primary head segment — the protocerebral segment of

authors — is in reality composed of four segments and that there-

fore the head of an insect is made up of not less than zzze seg-

ments, all of which were primitively postoral. According to

Janet these segments are :

1° Le somite du gésier

o ou somite proto-stomodæal

2' Le somite cesophagien

o ou somite deuto-stomodzal

3 Le somite cly péo-pharyngien ^ ou somite trito-stomodzal

1% The tergal parts (z. e., the labrum,

of the ocelli,

posed by the

epipharynx, and clypeus) situated in front

are left out in enumerating the seven segments as they are not sup-

author to belong to either of those segments." JZ. c. p. 20.

Nos. 455—-456.] HEAD OF BLATTA. 781

4 Le somite du labre ou somite proto-cérébral

5° Le somite antennaire ou somite deuto-cérébral

6° Le somite post-antennaire ou somite trito-cérébral

7 Le somite mandibulaire ou somite proto-gnathal

8° Le somite maxillaire ou somite deuto-gnathal

9° Le somite labial ou somite trito-gnathal

From the embryological view point Janet sees support for

these conclusions in the development of the sympathetic nervous

system. He considers “ the nervous system of the stomodaum

(or the unpaired sympathetic system) as being simply the mor-

phological prolongation on the ectodermic stomodzal invagina-

tion, of the chain of the central system." The untenability

of this view has been pointed out by Heymons, (:00), who has

devoted especial attention to the study of the development of

the sympathetic system, and whose argument, therefore, carries

especial weight.

Of the work of Patten, Wheeler, Viallanes and Folsom, the

excellent résumé by Comstock and Kochi leaves little to be said.

These last writers accept Folsom’s conclusion as to the presence

of seven primitive segments, and agree with him that the append-

ages of the superlingual segment may be represented in the

Orthoptera by a pair of small sclerites lying behind the lingua,

within the mouth cavity. Neither Heymons nor Wheeler!

accept this view. As regards the condition in the cockroach, I

have only the negative evidence that there is wholly lacking a

trace of a ganglion belonging to this segment. The position of

the above mentioned sclerites is such as to make them readily

susceptible of explanation on purely mechanical grounds, as

secondary structures. Though Folsom's work was done on

Anurida, and his corroborative evidence was drawn from a study

of other Collembola, it is not to be expected that we should find

such a fundamental difference between the Apterygota and the

Pterygota.

!In a forthcoming paper, which Dr. Wheeler has kindly allowed me to antici

pate by the above statement, he very fully and conclusively presents the arguments

against such a view.

782 THE AMERICAN NATURALIST. [Vor. XXXVIII.

THE GENERAL FEATURES OF THE FORMATION OF THE HEAD

OF BLATTA.

The changes during the first six days! of the embryonic life

of Blatta lead to the formation of the ventral plate, the blasto-

dermic thickening which represents the first rudiment of the

germ band. This extends along about two-thirds of the ventral

surface of the egg, beginning at about one-fourth of the length

of the egg from its cephalic end.

The newly formed germ band is not an evenly developed struc-

ture, but there are to be seen, on each side, groups of rapidly

proliferating cells. These active cells give rise to an indistinct

segmentation in the earlier stages of the germ band. This was

noted by Cholodkowsky (89), who says, (p. 91): ‘ However,

when I say that the young germ band of Blatta gives no indica-

tion of cross divisions I do not mean by that that there are no

traces of metamerism. Very early when the germ band is still

very imperfectly marked off from the surrounding undifferen-

tiated blastoderm, one may observe a noteworthy grouping of its

cells around certain points which are none other than the centers

for the formation of the future appendages.”

Heymons (95), has correctly interpreted this description of

Cholodkowsky's as referring to-certain formative centers in the

undifferentiated blastoderm, which by their extension and fusion

give rise to the germ band. A similar method of formation of

the germ band of Astacus had already been pointed out by

Reichenbach (86). That Cholodkowsky was not clear as to the

nature of these centers is evident not only from his reference to

them as the precursors of the appendages, but also from his

Figure 2, which shows an embryo of about ten days, in which

Mc pen and thoracic appendages are already prominent,

Pte 1S to be seen “die beginnende Gruppirung der Zellen

agen des ersten Paares von Abdominalfiissen.”

1 Cholodkowsky C91)

the age of the embryos

this, I have found that t

of a single capsule. However, the few observations I

confirmed Wheeler's approximations, and I have

etermining the ages of the embryos studied.

Nos. 455-456.] HEAD OF BLATTA. 783

An interesting fact is that, as pointed out by Heymons (95b),

the cephalic lobes arise from three pairs of these centers of pro-

liferation, which very early fuse. Wheeler found that two such

centers entered into the formation of the lobes of Xiphidium.

The presence of the three pairs would preclude Cholodkowsky's

idea that these centers represent future appendages. On first

thought it might seem that their presence confirms Janet's view

as to the compound nature of the preoral region. Janet believes,

however, that there are three segments lying cephalad of the

cephalic lobes, and that these do not fuse with the lobes, but in

the later stages are invaginated with the stomodzeum. It, is evi-

dent that the formative centers of the cephalic lobes lend no

support to that view. It has been suggested that they might

represent the three segments whose ganglia enter into the for-

mation of the brain. That this is not the case is shown from

the fact that these areas precede the definite formation of the

germ band, and that their fusion and consequent obliteration

has taken place long before the deuto- and tritocerebral seg-

ments have united with the protocerebral segment.

It is during the seventh day, after the appearan

blastoporic thickening of the caudal end, that there.

becomes prominent a pair of thickenings laterally

placed at the cephalic end of the germ band (Fig.

BOLJ These thickenings represent the cephalic

réle in the formation of the head capsule. From

the first they are connected with the germ band.

Of an independent development, such as described

by Will (88) for Aphis and by Heider (89) for

Hydrophilus, I find no indication in Blatta. as

During the eighth day of embryonic life the 5, ; rne germ

most striking change is the appearance and very a Ths ceshe

rapid development of the amnio-serosal folds, — tic lobes(c. /.)are

phenomena which have been described in aye

Wheeler. The cephalic lobes develop rapidly, and during the

first half of the ninth day they become prominently demarcated

from the strap-like trunk region. The embryo in this stage

covers about half of the ventral face of tHe egg, being somewhat

ce of the

aoe: y

D rtu

782 THE AMERICAN NATURALIST. (Vor. XXXVIII.

THE GENERAL FEATURES OF THE FORMATION OF THE HEAD

OF BLATTA.

The changes during the first six days} of the embryonic life

of Blatta lead to the formation of the ventral plate, the blasto-

dermic thickening which represents the first rudiment of the

germ band. This extends along about two-thirds of the ventral

surface of the egg, beginning at about one-fourth of the length

of the egg from its cephalic end.

The newly formed germ band is not an evenly developed struc-

ture, but there are to be seen, on each side, groups of rapidly

proliferating cells. These active cells give rise to an indistinct

segmentation in the earlier stages of the germ band. This was

noted by Cholodkowsky (’89), who says, (p. 91): “ However,

when I say that the young germ band of Blatta gives no indica-

tion of cross divisions I do not mean by that that there are no

traces of metamerism. Very early when the germ band is still

very imperfectly marked off from the surrounding undifferen-

tiated blastoderm, one may observe a noteworthy grouping of its

cells around certain points which are none other than the centers

for the formation of the future appendages.”

Heymons (95), has correctly interpreted this description of

Cholodkowsky's as referring to-certain formative centers in the

undifferentiated blastoderm, which by their extension and fusion

give rise to the germ band. A similar method of formation of

the germ band of Astacus had already been pointed out by

Reichenbach (86). That Cholodkowsky was not clear as to the

nature of these centers is evident not only from his reference to

them as the precursors of the appendages, but also from his

Figure 2, which shows an embryo of about ten days, in which

the mouth parts and thoracic appendages are already prominent,

MK iere is to be seen ** die beginnende Gruppirung der Zellen

ür die Anlagen des ersten Paares von Abdominalfüssen."

states that on account of the peculiarity of ovoposition,

of Blatta is not easily determined. As an illustration of

have made have, i A

lodi aa ed in the main, confirmed Wheeler’s approximations, and I have

Nos.455-556] __ HEAD OF BLATTA. 785

logues à celui que nous décrivons maintenant qui a décidé la

plupart des embryologistes à considérer, à tort croyons-nous,

l'antenne comme un membre post-buccal." (p. 289.)

Viallanes was influenced in his interpretation by his studies

of the nervous system of the adult, in which the deutocerebrum,

the ganglion of the antennal segment, is clearly pre-oral, as is

also its commissure.

More recently, Comstock and Kochi (: 02) have challenged the

commonly accepted view. Believing that the clypeus *''repre-

sents the median field of the sternite of the antennal segment,"

they argue that a study of the figures given by authors describ-

ing a post-oral position of these appendages “shows that while

a line connecting the two antennze would pass in some cases

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

of the rudiment of the antennal sclerite

does not abut against the procephalon. In

fact, the very figures given to support the

view that the antennze are post-oral in the

early embryo support the opposite view."

(p. 31.)

A study of Blatta has served to confirm

the view of Zaddach. At the time of the

first appearance of the antennz the stomo-

dzeal invagination has not yet manifested

itself. Its position is marked, however, by

an area of rapidly proliferating cells (Fig.

2, m.). The antennal rudiments are clearly

caudad of this area, and are thus at their

earliest appearance post-oral (Fig. 2, ant.).

They increase rapidly in size, and at a stage

but slightly later than that represented in

Fig. 2, they lie caudo-laterad of the newly

formed stomodaum. It is at this stage

that they were studied by Viallanes. Very Juge ds

soon they are pushed more markedly post-orally, and it is in this

stage that they have usually been figured (Fig. 3 ant.).

From the view point of comparative anatomy we have in Apus

strong evidence of the original post-oral position of the antennae.

G. 2. — The ventral aspect of

by a shght thickening of the

cells. X

786 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Lankester thought that in this form the antenna of the adult

were innervated from subcesophageal ganglia. Though this has

proven to be incorrect, Pelseneer ('85) has

shown that the antennal ganglia of the adult,

though associated with, are histologically per-

fectly distinct from those of the archicere-

brum, and that they are perfectly homologous

with the ganglia of the ventral cord.

By the end of the ninth day there are to be

seen, in addition to the antennal rudiments,

those of the thoracic appendages, which are

distinctly outlined, while those of the mouth-

parts are merely indicated. This is the stage

figured by Wheeler (’89, Fig. 44). Wheeler

found that in many embryos, as in the one

which he figures, “one or, more rarely, both

antennary lobes are temporarily bilobed."

This he thought might be due to a tem-

porary reversion, ‘tending to show that the

antennary lobes originally gave rise to two

pairs of appendages which were perhaps homologous with the

TWO pairs of antennæ in the Crustacea.’ In embryos but

slightly more advanced I, also, have seen indications of this

bilobulation. Believing that the antennæ of insects are not to

be homologized with those of the annelids but, rather, are

homodynamous with the other appendages, I should regard

this temporarily bilobed condition as a possible reversion toward

à biramous condition of the appendage.

The next notable change is the appearance, during the tenth

day, of the mouth and of the rudiment of the procephalon. The

mouth is not invaginated from within the latter, but, as is shown

by Figure 3, lies caudad of it.

Same has been much discussion concerning the origin of

Is rudiment of the procephalon which, by many, is spoken of

as merely that of the labrum. A number of observers have

reported that in certain forms it originates as a paired structure

(Hydrophilus, Kowalevsky, 71, Heider, '89, Graber, '9o; Bombyx

mort, Tichomiroff, '82; Pieris, Gastropacha, Zygaena, Graber,

second antenna (2d

nt.. X 80.

Nos. 455-456] HEAD OF BLATTA. 787

'9o ; Hyalotoma, Graber, '90; Chalicodoma, Carriére, '9o, Bürger,

'97). Kowalevsky, Patten, Carriére, and others maintain that

not only does this structure arise from paired rudiments, but

that it represents a fused pair of appendages. Bürger in his

edition of Carriére’s notes, does not commit himself to the latter

VIEW.

On the other hand a greater number of species have been

noted in which the procephalon is, from the first, unpaired.

Among these are included a number of Apterygota (see Folsom,

: 06, pp. 93—96), while of the Pterygota may be mentioned Aphis

(Huxley, '58) ; Apis (Grassi, '84); CEcanthus (Ayers, '84) ; For-

ficula (Heymons, '95); Mantis (Viallanes, '91); Blatta (Cholod-

kowsky, '90); e¢a/. Heymons has emphasized the fact that the

place of origin of the labrum, between the two halves of the

nervous system rather than laterad of them, fundamentally distin-

guishes this structure from the true appendages.

Especial attention was directed to this point in Blatta on

account of finding an embryo in which the procephalon was

quite clearly in two distinct halves. A study of a large number

of other preparations of the same stage failed to duplicate this,

though in some there was to be seen a slight mesal constriction.

As the embryo first mentioned was otherwise distorted I am

inclined to regard the appearance of the procephalon as abnor-

mal, and to agree with Cholodkowsky that a

in Blatta its rudiment is unpaired.

By the end of the tenth day the embryo

: pr-

has increased considerably in length, and o, 777"

the mouth part appendages are all distinct. Ei E mad

In an embryo of 1.5 mm. length, the ocular 2p

and antennal segments, which at this time nz

i eas- is

constitute the head of the embryo, m zamr

ured 275 u, while the mouth part region ge

was 350 p in length, their relative extent Fic. i are ge de

being thus as 11:14. The ist and 2nd sonal S bo

maxilla: are subequal and have relatively

the same situation upon the germ band. The mandibles are

considerably smaller, but are equally distinct (Fig. 4)-

A striking feature of embryos of this period is the relation

788 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of the mandibles to the antennz. They are separated by a

space much greater than that which separates the mandibles

from the rst maxilla. This appearance, which in Hydrophilus

proved so puzzling to Heider, is due to the presence of the

so-called **intercalary," * Vorkiefer," ** premandibular," or: “ tri-

tocerebral" segment. As the evidence all tends to show that

this segment is the homologue of the second antennal segment

of the Crustacea, it should be designated either so or as the

tritocerebral segment rather than by the indefinite and non-com-

mital term ** premandibular.”’

Wheeler (93) was the first to demonstrate in Anurida a pair

of vestigial appendages, which bear to its ganglion the same

relation as do the other appendages to the ganglia of their

respective segments. Wheeler’s work has been abundantly con-

firmed by workers on the embryology of the Apterygota, Uzel

(98) and Folsom (99) having determined that in certain forms

these appendages persist even to the adult stage.

. Though these results have been generally accepted as applying

to the Apterygota, embryologists have been unwilling to accept

the scant evidence of the presence of such vestigial appendages

in the Pterygota. As long ago as 1870 Bütschli observed in the

embryo of Apis a paired rudiment just behind the mouth, which,

as he said, “sich fast wie ein Paar innerer Antennen ausnimmt."

Bütschli himself did not regard this structure as homologous

with the other appendages. Grassi (84) also noted these ap-

pendages, but contrary to Bütschli, who thought that they fused

to form a transitory under lip, Grassi states that they quickly

disappear without leaving a trace. Moreover, he went farther

than Bütschli in comparing them to antennze, for, as Heider has

pointed out, he says, “ Il primo paio d'arti boccali, che ha un'esis-

tenza effimera, pud forse paragonarsi ad un paio d'antenne degli

artrapode" (p. 57). Carriére ('9o) described similar rudiments

for Chalicodoma, and his statements were verified by Bürger

(97). On the whole, however, insect embryologists discard the

above evidence and agree with Heymons ('95 b) who says of the

tritocerebral segment, “Sein rudimentär Charakter gibt sich

hauptsáchlich darin zu erkennen, dass sich an ihm niemals Ex-

tremitaten entwickeln. Dies gibt wenigstens, so weit wir bisher

wissen, von den pterygoten Insecten.”

Nos. 455-456.] HEAD OF BLATTA. 789

In the face of these opinions it may seem somewhat venture-

some to assert that in Blatta there are evidences of the presence

of such appendages. And yet, I believe that the facts warrant

this conclusion. In embryos of the eleventh day, in favorable

specimens there are to be seen between the antennaland the

mandibular rudiments, in the region occupied by the tritocerebral

ganglion, small but distinct thickenings of the hypodermis (Fig.

4, 2nd ant.) . Moreover, these are to be

detected in both cross and longitudinal

sections (Fig. 5), and bear to the ganglion

of the tritocerebral segment the same rela-

tion which we should expect of appendages.

To be sure, not every hypodermal thicken-

ing is to be interpreted as the rudiment of'

an appendage. Since, however, the thick-

enings under consideration occur not only

in one but in several preparations, since

they occur in the location we should ex-

pect for such appendages and do not occur, | A a aia us

between the other pairs of appendages, embryo of eleven days. x

and since, lastly, they bear the expected zr

relation to the ganglia of the second antennal segment, I see no

other interpretation than that they represent vestiges of append-

ages such as are stil developed in some of the apterygote

insects.

The mouth-part appendages rapidly increase in size, and early

in the eleventh day the length of the rst maxillze considerably

exceeds that of the second maxilla, while a little earlier these

appendages were subequal. Measuring from the caudal angle to

the apex, we find the length of the mandibles to be about 55 p,

that of the 1st maxillae 96 p, while the 2nd maxilla are but 70 p.

There is no trace of lobulation in any of the mouth parts.

The procephalic rudiment has become prominent and, in the

latter part of the eleventh day, appears as a button-shaped swell.

ing overlying the mouth. In surface views the mouth can be

seen through the rudiment, lying slightly caudad of its center.

It is this appearance which has led to the incorrect statement

that the mouth develops zz the procephalon. In ists it of

790 THE AMERICAN NA TURALIST. [Vor. XXXVIII.

this age it will be seen that the antennz are migrating forward.

: They have increased in length so as to

almost completely overlie the mandibles.

The procephalon increases rapidly in

size and pushes caudad over the mouth

and between the antennz. In many

specimens there is to be noted a con-

striction of its caudal portion, while the

mesal notch of the caudal margin is also

distinct.

Early in the twelfth day are to be

seen signs of the lobulation of the rst

b on aet ONE c and 2nd maxille. I find no indication

days showing antennal sclerites Of the lobes of these appendages arising

ree from separate, isolated centers, as Hey-

mons (’95a) has described. E

It has been stated that the mandibles of Blatta

are always simple. I was interested to find that

ina few embryos of twelve to thirteen days they

appear to be feebly trilobed. This agrees with the

condition in Œcanthus, as described by Ayers (p.

24). Korotneff ('81) also represents the mandibles

of Gryllotalpa as feebly trilobed (see his Fig 6), m

though he says nothing about it in the text. nrbs predi

Throughout the series it is evident that there is days, showing

not a perfect fusion of the antennz with the ce- M CREE :

phalic lobes, but the point of origin of these

appendages is separated from the

lobes by a constriction which demar-

cates a more or less definite area of

attachment for the antennz. This

may be seen in Figure 6.

The lobulation of the 2nd maxillz

is well shown by Figure 7, which

shows one of these appendages from

an embryo about thirteen days old.

LAS Their mesad migration proceeds

duo Ses a ca em- rather slowly, until during the fif-

d migration, x no. — ^ teenth day they present the appear-

Fic, 7 nd ma

illa of embry

.tracheal invaginations on the plure of

Nos. 455—456.] HEAD OF BLATTA. 791

ance shown in Figure 8. They have not yet fused, but the

future labrum is clearly outlined. In the meantime they have

also migrated forward, so that the space which formerly sepa-

rated them from the rst maxillae has been considerably reduced,

and in lateral view the 2nd maxilla are more or less hidden by

the first pair.

About this time there appears a prominent invagination lying

cephalo-mesad of the base of the mandible, close to the mandib-

ular ganglion. This lies under the antenne, but can be seen

by focusing below that appendage or by removing it (Fig. 9,

a.t.) In the same embryos there are prominent ten pairs of

the meso-, and metathorax and the

first eight abdominal segments. For

the prothorax there is a pair of slight

and indistinct invaginations which have

escaped the attention of previous work-

ers on Blatta. It should be noted that

the location of the above mentioned

mandibular invaginations is in no wise

comparable to that of the trachea. In

addition to these invaginations there are am

in the mouth-part region three other fic. 9.— Embryo of about fourteen:

pairs — one pair at the cephalo-lateral shoning the ic seraa pecia

angle of the mandibles, one between the rior tentorial invagination (a. #.)

mandibles and the maxilla, close to the xt.

caudo-lateral angle of the mandibles, and one cephalo-laterad

of the base of the 2nd maxilla. The last pair of invaginations

is difficult to detect, for they are smaller, their lumen is not

prominent and is frequently hidden by the caudal margin of the

Ist maxilla. It is shown in Figure 10, 7.7. The invaginations

lying laterad of the mandibles form tendons for the mandibular

muscles. The réle of the first and the last pairs in the forma-

tion of the tentorium or endoskeleton of the head will be dis-

cussed later.

Up to this period the length of the mouth-part region has

considerably exceeded that of the combined ocular and antennal

segments. Thus, as we have seen, in an embryo of about eleven

192 THE AMERICAN NATURALIST. [Vor. XXXVIII.

days the latter region measured 275 mw in length as compared

with 350 mw for the mouth-part region. In an embryo at the

close of the twelfth day the antennae have moved forward until

the anterior section measures but 200 y. The mouth-part

region, owing to the loss of the tritocerebral segment and the

reduction also taking place in the 2nd maxillary segment, meas-

ures 280 u. A comparison early in the fourteenth day shows

that the two regions are approximately equal in length. This,

as we should suppose from the above, is due not so much to an

increase in the size of the anterior region, as to a reduction of

the mouth-part region, due to the ce-

phalad migration of the mouth parts.

The developing pronotum of the pro-

thorax is pushing rapidly forward, and

is limiting the pleural region of the

maxillary segments (Fig. 10, 7z.).

During the sixteenth day there

occurs the series of changes which

Wheeler (89) has described as the

«€ revolution of the embryo. This is

“ea en a period of very rapid growth, and at

vagination (7. 7.) and the reduction its close there has been completed

of the mouthpart region e pro-

notum (#r.) has pushed forwarduntil the circumcrescence of the yolk by

its anterior margin lies over the first

maxilla. x 53. the embryo.

At about the beginning of the six-

teenth day, just as these changes have commenced, the condi-

tions do not differ markedly from what I have just described.

The reduction of the mouth-part region is rapidly progressing,

the length of the anterior measuring 200 u as compared with

160 u for the mouth-part region. The pleural region of the

prothorax has extended forward until its most cephalic point

lies over the base of the rst maxilla, while its indistinct spirac-

ular invagination lies over the base of the 2nd maxilla. The

cephalic lobes have extended back until they lie well over the

base of the mandibles. The rst maxilla greatly exceed the 2nd,

which have all but fused to form the labium.

In ventral view of embryos about this age the appendages

completely hide the sternal region ; but on carefully dissecting

Nos. 455-456.] HEAD OF BLATTA. 793

away the antenna and the procephalon, it may be seen that the

Sternites of the mandibular and rst maxillary segments have

fused, and form a

well marked area ly-

ing between their re-

spective appendages

(Fig. 12, hyp.). We

shall refer to this

again, in the discus-

sion of the develop-

ment of the sclerites.

The procephalon is x ;

much enlarged and Fic. 12. euni Wei ef embryo

at its proximal end is sixteen days. The antenna

the AER Ts have been

Fic. 11. — Diagram showing wedged in: between moved t6 expose the hypo-

the arms to the tentorium kár X 8o.

in their relation to the the cephalic lobes in PATHS

pay 2 aad month. the manner described by ai (58).

The change which Weismann described as

a rotation of the cephalic lobes as upon an axis, occurs during

the latter part of the sixteenth and the early part of the seven-

teenth days. This change is best followed by observing the

relations of the brain lobes and of the tendon invaginations to

the rest of the cephalic region. At its close the cephalic lobes:

have pushed over the mandibles in such a way as to crowd back

the mandibular pleurites. The mouth parts assume much their

definitive position, and the head is practically formed, though it

is not until much later that the sclerites have become so chiti-

nized as to be perfectly distinct.

THE DEVELOPMENT OF THE TENTORIUM.

The tentorium or internal skeleton of the head of the adult

cockroach has been described and figured by Miall and Denny

(86) and, more fully, by Comstock and Kochi (:02). The latter

have treated of its structure in a number of forms, and have

shown that its attachments afford valuable criteria for homolo-

gizing sclerites in different insects.

In the adult Blatta the body of the tentorium consists of a

794 THE AMERICAN NATURALIST. [Vor. XXXVIII.

chitinous plate underlying the cesophagus and protecting the

sub-cesophageal ganglion. From the anterior margin of the

body of the tentorium there extends forward a pair of processes,

the anterior arms, which are attached at the cephalo-ateral

angles of the clypeus, near the bases of the mandibles. Corre-

sponding with these are the posterior arms, which are near the

margin of the occipital foramen, just above the articulation of

the maxille. In addition there passes up from the body of the

tentorium to be inserted at the margins of the antennal sclerites,

a third pair of processes, the dorsal arms. The entire structure

arises from two pairs of ectodermal invaginations. j

As early as the thirteenth day there appears cephalo-mesad of

.the base of the mandibles a prominent pair of invaginations

which give rise to the anterior arms (Fig. 9, a.t). These

invaginations lie close to the latero-anterior angles of the man-

dibular ganglion. Heymons refers to the anterior arms in For-

ficula as originating at the base of the antennze. In Blatta they

are hidden by the antennz, which must be removed in order to

see the invaginations distinctly; but they clearly belong to the

mandibular segment.

The invaginations for the posterior arms lie cephalo-laterad of

the bases of the 2nd maxillae (Fig. 10, 7.7.) They are much

less prominent than those of the anterior arms, and, more or

less hidden by the rst maxille, they are difficult to detect. They

seem to be somewhat retarded in their development, for I have

been unable to find them in embryos which clearly showed the

invaginations in the mandibular segment.

The invaginations of the anterior arms extend upwards along

the sides of the mandibular ganglion, and then, bending at an

angle, pass caudo-mesad. Ata comparatively late date the two

invaginations fuse, and thus give rise to the main body of the

tentorium. On the other hand the posterior invaginations pass

as blind sacs slightly upward and then forward, to fuse ultimately

with the anterior arms.

i While the anterior and the posterior arms thus originate 4°

Invaginations of the body-wall, the dorsal arms arise, not as in-

vaginations but as processes, from the anterior arms. These

processes gradually extend dorsad and thus come into connec-

Nos. 455-456.] HEAD OF BLATTA. - 795

tion with the body-wall in the region of the antennal sclerites.

Figure 11 shows diagrammatically the course of the tentorial

arms in an embryo of about eighteen days.

Considering the rapid growth of the embryo during the

period following their appearance, the development of these

rudiments is but slow. It is not until the twentieth day or

later that the fusion of the two pairs of rudiments has taken

place.

In view of this method of origin of the dorsal arms of the

tentorium of Blatta, it is interesting to note the condition in the

Plecoptera, a group in many respects more generalized than the

Blattidae. Of the dorsal arms in these forms Comstock and

Kochi, p. 41, say: “In the Plecoptera it appears to be merely

à chitinized tendon, the peripheral end of which is less chitinized

than the base and is only loosely attached to the skull." Believ-

ing that in other insects the anterior arms arise from a distinct

invagination, these authors continue: “It remains to be deter-

mined whether or not the dorsal arms in the Plecoptera are

homologous with the apodeme-like dorsal arms in other insects

and, if so, which type is the more generalized." As has been

seen, the evidence of embryology shows that the generalized con-

dition is that which still persists in the Plecoptera.

The most thorough study of the development of the tento-

rium was that of Heider ('89). In áddition to this investigator,

several others have devoted more or less attention to the sub-

ject. Hatschek (77) called attention to what he supposed were

tracheal invaginations in the head of lepidopterous embryos.

Tichomiroff (79) says: “There exists no head tracheze such as

Hatschek thought he discovered. The epithelial invaginations

here found I believe to be the rudiments of the inner head-skel-

eton.” Ayers ('84) noticed invaginations in the head region, oet

was unable to determine their rôle. He inclined to the belief

that they disappear altogether, “as no trace of them was to "e

found in sections of an embryo about the time of hatching.

Wheeler ('89) attributed the tentorium to five pairs of "E

tions anterior to the maxil segment. . oe

My results agree ao of Heider, who described the

è ss 2 ý . 9" y nova , in the a x

196 THE AMERICAN NATURALIST. [Vor. XXXVIII.

mandibular and one in the second maxillary segments. Heymons

(95) basing his conclusions mainly on a study of Forficula,

described essentially the same conditions, as applying to Forfi-

cula and to the Orthoptera, including the Blattidze. Carriére and

Bürger made the same observations on Chalicodoma, thus lending

strength to Heymon's surmise that such a mode of origin is

typical for the entire group of insects.

Various investigators, among whom are Palmen (77), Hatschek

(77), Wheeler (89), and especially Carriére ('90), and Bürger

(97), have regarded the tentorial invaginations as homodynamous

with the tracheal invaginations. To this Korschelt and Heider

(93) object that their rudiments do not by any means every where

agree so closely with tracheal stigmata of the following segments

as they do in Chalicodoma. This is especially true of these

rudiments in Blatta, As we have seen, the invaginations for

the anterior arms arise close to the median line, just laterad of

the nerve cord. This corresponds more nearly to the position

of the invaginations for the thoracic furca. The origin of the

posterior invaginations is more suggestive of the stigmatic

invaginations.

In considering this question it must be remembered that

Chalicodoma is a highly. specialized type with an interpolated

larval stage, while Blatta is a representative of a comparatively

generalized group of hemimetabolous insects. This being the

case, it does not seem that we should regard the location of the

tentorial rudiments in Chalicodoma as primitive, but rather I

should regard it as secondary. The fact urged by Palman, that

the chitinous lining of the tentorium, like that of the tracheze, is

shed during ecdysis, is of little weight, since the same holds

true of any hollow ectodermal invagination in insects.

However, it is not necessary to conclude that the tentorial

invaginations are metamorphosed tracheze * which have lost their

primitive function and become secondarily modified." If, with

Kennel, we derive the tracheze from dermal glands of annelidan

ancestors, there is no reason why we should not consider the

trachez and the tentorium as homologous structures. Certain of

these glands definitely localized have become modified to form

tracheze, while certain others, of the same origin though not

Nob. 455-456.] HEAD OF BLATTA. 797

homodynamous with the first, have become converted into the

tentorium. ;

THE DEVELOPMENT OF THE HEAD SCLERITES.

As before stated, it is only in the most incidental manner

that insect embryologists have touched upon the development of

the head sclerites. Before presenting my conclusions concern-

ing the development of the head capsule of Blatta, I shall briefly

review the more important of the scattered references, reserving

comment until we are ready to compare the views thus expressed

with the results of my work.

The earliest of the references was that of Zaddach (’54).

This pioneer worker has not received due credit for what he so

independently accomplished. Many general features of the

. embryological development of insects, though credited to much

_ later writers, were clearly outlined by Zaddach. Working alto-

gether with entire mounts, it is but natural that his conclusions

regarding the external changes should be the most accurate. It

was he who first pointed out the post-oral origin of the antenne,

and called attention to a special antennal segment distinct from

the cephalic lobes.: He described the procephalon (“ Vorder-

kopf”), and derived from it the labrum, the clypeus and the

front (“ hinterer und vorderer Kopfschild und Oberlippe "4 He

believed that the cephalic lobes formed the bulk of the head

capsule, while the Y-shaped suture represented the line of fusion

of the cephalic lobes and the procephalon.

Huxley (’58) gave a more accurate description of the pro-

cephalon. He also gave a fuller account of the dorsal flexure of

the cephalic region, as a consequence of which * the line of attach-

ment of the bases of the eyes and antennz is frequently alto-

gether above that of the other appendages, so that they appear

to be tergal, and not sternal, appendages.” The pfocephalic

lobes he regarded as the sternite of the first, or opthalmic, seg-

ment and though he does not make a direct statement, the

inference is that his *antennulary and antennary sterna " are

represented by the labrum and the clypeus.

Packard in his earlier work devoted some attention to the

798 THE AMERICAN NATURALIST. [Vor. XXXVIII.

study of the development of the head sclerites. In 1866 he

argued that. *since the arthropleural is the limb-bearing region

of the thorax it must follow that this region is largely developed

in the head," the sternites being reduced to a minimum and the

tergites almost completely absorbed. He believed that the bulk

of the head is formed “by the great expansions of the eye-pleu-

rites which, so to speak, are drawn back like a hood over the

basal rings."

In 1870, in the second edition of his ** Guide to the Study of

Insects (foot-note)," he reiterates these conclusions. He distin-

guishes the labrum, epipharynx, and clypeus as tergal elements

belonging to his hypothetical pre-ocellary segment. The occi-

put also is reckoned with the tergites, while the gula is sternal.

All of the other elements of the capsule are supposed to be

pleural.

Packard later completely revised his views as to the number

of primitive head segments. Instead of seven or a possible

eight, he recognized but four segments entering into the forma-

tion of this region. In 1883, in his paper on ** The Number of

Segments in the Head of Winged Insects," he thus summarizes

his views as to the development of the sclerites :

“It appears, then, that the epicranium or that piece (sclerite)

bearing the eyes, ocelli, and antennze, and in front of the clypeus

and labrum, is formed from the original procephalic lobes, and

represents the first or antennal segment ; while the remainder of

the original or primitive segments are obsolete, except in those

insects which retain traces of an occiput or fourth cephalic

tergite. All of the gular region of the head probably represents

the base of the primitive second maxillae."

Ayers (83) refers to the reduction of the maxillary and man-

dibular segments, * their dorsal portions disappearing altogether

and their ventral portions fusing with the oral region."

Viallanes (86 and '8 7b) states that the head of an insect pre-

sents three prebuccal segments and that the labrum represents:

the sternite of the third or tritocerebral segment. Beyond this

he does not enter upon a discussion of the origin of the sclerites.

Heider (’89) was the first to devote especial attention to the

development of the tentorium. His careful studies of the origin

Nos. 455-456.] HEAD OF BLATTA .199

of this structure have already been discussed. Regarding the

sclerites he states that the labrum and the clypeus arise from

the procephalon, while the remainder of the head capsule is

formed mainly by the bending over dorsally and the backward

extension of the cephalic lobes.

It is Heymons who has devoted the most attention to a study

of the embryological development of the head sclerites. Study-

ing mainly the development of various species of the Orthoptera,

he very definitely stated his conclusions in his paper on the seg-

mentation of the insect body ('95a). The labrum and the clyp-

eus he derives from the procephalon, while the front, hé con-

cludes, is developed from the fused cephalic lobes. The vertex,

occiput and gena he believes are developed from the fused

tergites of the mouth part segments, while the hypopharynx

arises from the fusion of the sternites of these same three seg-

ments.

Janet, '99, finds in the sclerites but little indication of the

primitive segments. Thus he finds that the front *appartient,

par sa partie antérieure, au somite clypéopharyngien; par sa

partie postérieure, au somite du labre; par ses parties latérales,

au somite antennaire.” Based upon years of study of the mor-

phology of certain Hymenoptera, Janet's paper presents a fund

of valuable anatomical data. From the embryological view point

it is less satisfactory. Moreover, though recognizing the con-

fusion in the use of terms to designate the head sclerites, Janet

has made no attempt to determine homologies. Until this has

been carefully done, results obtained from a study of such a

group as the Formicoidea can be of but little general application.

On the other hand, Comstock and Kochi, :02, attach very

great importance to the relation of the primary segmentation to

the sclerites of the adult insect. Their conclusions were based

` upon comparative anatomical studies of a large series of the

more generalized insects, as well as upon embryological data.

. According to their view, the typical segment is composed of two

subsegments. Ventrally the line of union of these subsegments

is determined: by the position of the furcz, while laterally it is

similarly marked by the position of the lateral apodemes. Be-

tween the segments lie the tracheal invaginations, which these

Soo THE AMERICAN NATURALIST. [Vor. XXXVIII.

authors regard as homodynamous with the invaginations of the

lateral apodemes. Thus to these invaginations of the body wall

is ascribed a prime importance in determining the homologies of

the sclerites. Another valuable criterion they found in the rela-

tions of the appendages to a typical segment.

In addition to limiting and carefully defining the sclerites

already generally recognized, Comstock and Kochi also distin-

guished four pairs of sclerites which had previously escaped

attention. These were: rst, the antennal sclerites bearing the

antennz ; 2nd, the antecoxal pieces of the mandibles, a pair of

sclerites in some species distinct from the clypeus and forming

the ventral articulations of the mandibles; 3rd and 4th, two

pairs of maxillary pleurites, narrow, chitinized bands between

which articulate the cardines of the maxilla. In addition to

these four pairs of sclerites proper, there are two pairs which

may have originally represented segments of appendages.

These are: rst, the ocular sclerites, a pair of annular sclerites.

bearing the compound eyes, and 2nd, the trochantins of the

mandibles, a pair of sclerites between the mandibles and the

gene,

Their conclusions regarding the relations of the head sclerites

to the primitive segments and to the appendages were summar-

ized by Comstock and Kochi in the following table:

SEGMENTS SCLERITES APPENDAGES

vertex and genze

- Ocular (Protocerebral) uc DAD KL Ocular sclerites.

Antennal sclerites

2. Antennal (Deutocerebral) DEED ee Antenna

Clypeus proper

2nd Antenne

3. 2nd Antennal (Tritocerebral) *

Labrum (mouth) of Campodea et al

Postgenae.

4. Mandibular - Antecoxal pieces _ Mandibles.

Pharyngral sclerites Trochantin

Nos. 455—456.] HEAD OF BLATTA. Sor

5. Superlingual Superlingual

Maxillary pleurites

6. Maxillary Maxillze

Lingua

Lateral cervical sclerites

7. Labial Dorsal cervical sclerites Labium

Vent. cerv. scl. (gula)

In each section of the middle column the dotted line indicates.

the division between the sternal and the lateral elements of the

segment.

My results have convinced me that so intimate a relation

between primary segmentation and the sclerites cannot be

shown.

I find that the front, clypeus, and the labrum are all derived

from the procephalon. This is in agreement with the conclusion

of Zaddach, '54, whose “hinterer Kopfschild" is clearly the

-clypeus posterior of Newport or the front as defined by Comstock

and Kochi.

Aside from Zaddach practically all authors agree in stating

that from the procephalon arise the clypeus and the labrum.

It has been suggested by Comstock and Kochi that the term

clypeus has thus been used in a broad sense to include both the

clypeus anterior and the clypeus posterior or the front.

That Heider used the word clypeus in a yet looser sense is.

evident. He speaks of the procephalon as giving rise to the

clypeus and the labrum. Now, in Hydrophilus, the form upon

Which he worked, as well as in any of the Hydrophili, the larva

does not possess a labrum (Schiodte, '61), or, more correctly

speaking, the clypeus and labrum have fused without. leaving any

line of demarcation between them. Thus Heider has homolo--

gized the clypeo-labral sclerite as labrum only, while the front

he has mistaken for the entire clypeus.

On the other hand, Heymons, as is shown by his figures, has

correctly distinguished the front. Yet he states that the clyp-

eus and labrum arise from the procephalon, while the front is

formed from the fused cephalic lobes. He presents no evi-

802 THE AMERICAN NATURALIST. [Vor. XXXVIII.

dence, nor have I been able to find any in support of this view.

There remains a consideration of the relation of these sclerites

to the primitive segmentation. I have called attention to the

fact that though Huxley, '58, does not make any definite state-

ment to that effect, the inference is strong that he regarded

these sclerites as representing the sternites of three preoral

segments. In fact, in referring to the Crustacea, he says that

the labrum represents the sternite of the antennary or third

somite. Viallanes, who likewise refers the labrum to the sternite

of his tritocerebral or third somite, seems to have attributed to .

these sclerites a similar definite relation to the primary segmen-

. tation.

More recently Comstock and Kochi have independently arrived

at and very definitely present the view that the front, clypeus,

and labrum represent respectively the sternites of the first three

segments. In support of this view they lay much weight on

Viallane's statement that the labrum is innervated by the trito-

cerebrum. Since the front bears one or more of the ocelli, it is

obviously derived from the first or ocular segment. The inter-

mediate sclerite, or clypeus, would then belong to the second

segment. These writers also believe that the position of the

antennal rudiments is only apparently post-oral, — that in reality

the basal pàrt of the antennal sclerites abuts against the pro-

cephalon. |

The embryological evidence, however, shows that the pro-

cephalon, and thus the sclerites derived from it, — the front, the

clypeus and the labrum, — belong to a single segment, the ocu-

lar or protocerebral.

I have discussed elsewhere the question of the primitive posi-

tion of the antennz, and have shown that in Blatta their rudi-

ments are clearly post-oral at first.

Even if we were to grant that the antennal rudiments are

only apparently post-oral, there can be no question as to the

primitive location of the second antenna and the third somite.

Yet the hypothesis derives the labrum from the sternite of this

segment.

"While I should attach great value to the innervation as a

criterion for determining the relation of sclerites to segments, I

Nos. 455—456.] HEAD OF BLATTA. 803

do not believe that it can be regarded as absolute. At any rate,

the evidence against the labrum as the sternite of the third seg-

ment cómpletely outweighs this consideration.!

The procephalon develops entirely in front óf the stomodzeal

invagination, and is already well outlined before the deutocere-

bral and tritocerebral ganglia have moved forward. Moreover,

as Prof. Comstock suggested, the labrum and the clypeus give

no satisfactory evidence of a paired origin. The absence of

median sutures in these sclerites might be readily explained by

the statement that fusion had taken place at such an early stage

as to be complete. But a more serious objection is that if they

develop post-orally and migrate to their pre-oral position, there

should be a stage in which they are paired. Such a stage does

not occur.

In the Orthoptera and in the Pterygota generally, the second

antennal segment is so slightly developed and so transient as to

have been largely overlooked by investigators, yet in these forms

the labrum is usually strongly developed. On the other hand,

in the Apterygota, where the second antennal segment is

especially well developed, the labrum is comparatively insig-

nificant. :

Finally, the sutures separating the front, clypeus, and labrum

are developed at a comparatively late period in the development

of the procephalon. If these sclerites represented primary seg-

ments, we should find them developing from three distinct cen-

ters. The fact that the procephalon develops as a single piece

shows that the sclerites derived from it are not primary, but are

secondarily developed. In fact, they are not constant, but there

are many species in which one or more of the sutures is lacking,

and thus the number of sclerites reduced.

The vertex, the compound eyes, and with them the ocular

sclerites and the genze, are formed from the fused cephalic lobes

and thus, with the front, clypeus and labrum, belong to the

ocular or protocerebral segment. The median suture of the

epicranium represents the ventral line of fusion of these lobes.

I should be noted that Janet ('99) considers the labral nerves as in reality

arising from the protocerebrum. This is more in harmony with the embryological

evidence,

804 THE AMERICAN NATURALIST. [VoL. XXXVIII.

The V-shaped epicranial suture in Blatta separates the front

from the vertex. It thus represents the line of separation be-

tween the cephalic lobes and the procephalon. ‘

Heymons maintains that the front alone is derived from the

fused cephalic lobes. The relatively excessive development of

these lobes would indicate that they must enter more largely

into the structure of the head. More weighty evidence is the

fact that the front is separated from the compound eyes by the

V-shaped suture. There is, of course, no question that the

eyes belong to the first segment and are developed from the

cephalic lobes. Moreover, in following through a series of

embryos we can trace the manner in which the front develops

by the extension of the procephalon into the angle between the

fusing lobes. Thus the direction of this suture —an inverted

V with the apex continuous with the median suture of the epi-

cranium — is easily explained. Heymons emphasizes his belief

that the suture represents the line of separation between the

first and second segments. But if that were the case the

rounded caudal margins of the cephalic lobes would on fusing

present a V-shaped angle whose apex would be directed cephalad

rather than caudad, as is really the case.

On first sight it would seem obvious that the genae belonged

to the mandibular segment. Huxley (78) described the man-

dibles as articulating with this sclerite. To this, Miall and

Denny (86) take exception. Comstock and Kochi also call

attention to the fact that the chief articulation of the mandibles

is with the postgenze rather than with the genze. In the earlier

stages, after the mandibles have moved forward, it appears as

though the mandibular pleurites occupied the position of the

future gene. At the time of the rotation of the embryo, how-

ever, the cephalic lobes crowd forward over the dorsal end of

the mandibular segment, and thus the pleurites are pushed

back to occupy the position of the postgenz.

It is easy to see how, as a result of this process of displace-

ment of the mandibular segment, there should remain a portion

to serve as the ventral articulation of the mandible. In the

adult Blatta the limits of this remnant are not to be distin-

guished from the clypeus ; but, as pointed out by Comstock and

Nos. 455—456.] HEAD OF BLATTA. 805

Kochi, in Gryllus and in Corydalis there persists a definite scle-

rte. These authors have shown that this sclerite bears the

same relation to the articulation of the mandible as does the

antecoxal piece to the coxa of a thoracic leg, and they have there-

fore called it the antecoral piece of the mandible.

Along the caudal margin of the postgenz, but cephalad of the

invagination for the posterior arm of the tentorium, is a narrow

sclerite, which was first pointed out by Comstock and Kochi.

Believing that the posterior tentorial invagination belonged to

the first maxillary segment and was homodynamous with the

invaginations of the thoracic lateral apodemes, these writers

named this sclerite the anterior maxillary pleurite. A similar

narrow sclerite lying parallel with the first, but caudad of the

tentorial invagination, they called the posterior maxillary pleurite.

We have seen that the posterior arm of the tentorium belongs

to the labial segment. The position of its opening in the adult

insect — almost directly over the cardo of the first maxilla —is

due to the cephalo-mesal growth of the paired labial rudiment,

as well as to the backward pushing of the maxillae at the time

of rotation. Thus the sclerite lying caudad of this opening

belongs, not to the maxillary but to the labial segment, and

should be known as the /adial pleurite. The sclerite lying ceph-

alad of the opening may retain the name of maxillary pleurite.

However, we cannot restrict the maxillary pleurite entirely to .

this narrow sclerite. There early takes place a fusion of the

pleurites of the mouth part region. The position of the pos- .

terior tentorial invaginations serves to mark the anterior limits

of the labial pleurite, but the extent of the mandibular and the

maxillary pleurites can only be judged from the relations of their

appendages. In an embryo of about the sixteenth day, as may

be seen from Figure 10,-the maxillary pleurites exceed in size

either of the others. Though the rotation of the embryo results

in a displacement of the maxilla, I do not believe that their

pleurites become reduced to the narrow, imperfectly demarcated

sclerites which have been designated as the maxillary pleurites.

We have seen that a portion of the deutocerebral segment,

bearing the antennee, is more or less clearly marked off from the

remainder of the germ band, even in the earlier stages. I

806 THE AMERICAN NATURALIST. (VoL. XXXVIII.

believe that this persists as the antennal sclerite of Comstock

and Kochi. In embryos about ready to emerge these annular

sclerites are very clearly marked, and allow of no question as to

their equivalence to other sclerites.

There has been much discussion as to the origin and signifi-

cance of the hypopharynx, though the data have been drawn

almost solely from a study of comparative anatomy. Apparently

the majority of those who have thus studied it regard it as repre-

senting a fused pair of appendages. Vayssiere (82) who worked

upon larval Ephemeride, strongly favored this view, though he

adds that in order to settle the question studies of a number of

representatives of the various orders would be necessary.

Folsom (:00) derives the hypopharynx of Anurida from a

median unpaired evagination between the first maxillae, and two

small papilla between the mandibles. The first gives rise to

the lingua, while from the latter are developed the superlingua.

The superlinguz Folsom regards as the appendages of a dis-

tinct segment, provided with a small ganglion. He brings for-

ward much evidence from comparative anatomy to show that the

hypopharynx of Anurida is typical of the Apterygota. Further

than that, he attempts to apply the same interpretation to the

Pterygota.

The only worker who has devoted any attention to the embryo-

logical development of the hypopharynx of the Ptergota is Hey-

mons (95a). This investigator reached the surprising conclusion

that the hypopharynx represents the fused sternites of the mouth

part segments. "That this interpretation is essentially correct

my studies have convinced me. However, I would not attribute

to the labial sternite any part in this structure. I believe that

the hypopharynx of Blatta represents the fused sternites of the

mandibular and first maxillary segments (Fig. 12, 4j.) and that

of a consequence of the forward migration of the mouth parts

these fused sternites have been pushed to within the mouth

cavity.

In Blatta, then, the sternite of the labial segment is repre

sented only by the ventral cervical sclerites. It seems to me very

clear that Comstock and Kochi are right in regarding the gula

of Coleoptera and certain other insects as representing the fused .

cervical sclerites.

Nos. 455—456.] HEAD OF BLATTA. 807

My views in regard to the relations of the sclerites of the

adult Blatta to the primitive segments may be summed up in the

following table:

SEGMENT SCLERITES

Protocerebral Vertex, gene.

Front, clypeus and labrum.

Deutocerebral Antennal sclerites

Tritocerebral.

Mandibular Part of post-genz, trochantins

Part of hypopharynx

Maxillary Remainder of post-genz, maxillary pleurites

Remainder of hypopharynx

Labial : Labial pleurites; lateral cervical sclerites

Ventral cervical sclerites

From the above results it would seem that the definitive

sclerites can afford us little or no evidence as to the primary

segmentation of the insects. This is certainly true of the head

sclerites, and I see no reason why it should not apply to other

regions of the body. Sclerites originate from mechanical causes,

and do not necessarily have any relation to the primary segmen-

tation.

As an illustration might be cited the breaking up of the

annular segment into sternite, pleurites and tergite. The cir-

cumcrescence of yolk and the consequent formation of the lateral

and dorsal portions of the embryo is a process of gradual growth,

and there is no demarcation into separate areas. The distinction

into sternite, pleurites, and tergite is a secondary process.

More germane is the fact of the origin of the front, clypeus,

and labrum from the procephalon. Thus not only these three

sclerites but also the vertex and the genæ arise from the one

Segment, the protocerebral. I have shown, also, that the post-

gene are to be regarded, not as the pleurites of a single segment,

but as the fused pleurites of the mandibular and maxillary seg-

ments,

I would not depreciate the value of the sclerites as an jndex

. to relationship among insects. That they are to a marked

808 THE AMERICAN NATURALIST. (VoL. XXXVIII.

degree constant, and that they may be homologized in the dif-

ferent groups, I believe to be very true. That they are second-

ary is equally true, and thus evidence drawn from a study of the

sclerites of the adult insect is insufficient to determine the primi-

tive segmentation and the related question as to the compound -

nature of the segments.

BIBLIOGRAPHY.

BRANDT, A.

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Forsow, 5 W.

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RH s r

The Development of the Mouth- Parts of Anurida maritima. Bull.

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THE ARBOREAL ANCESTRY OF THE

MAMMALIA.

W. D. MATTHEW.

WrruriN the last few years Dollo and Bensley have adduced

strong arguments to show that the marsupials are descended

from arboreal ancestors, as indicated especially by the traces in

modern marsupials of former opposability in the first digit of the

manus and pes. The present writer has for some time been

of the opinion that this is true not only of marsupials, but of

the placentals as well. Our present knowledge of fossil Mam-

malia and of the course of evolution of the various modern

races, enables us to foreshadow with considerable detail the

characters of a common ancestral group (homogeneous in

adaptive characters, although perhaps embracing certain differ-

ences in dentition etc. of very ancient origin) from which all

known mammals, excepting the Prototheria, are descended.

That there was such a group ancestral to both metatherian

and eutherian mammals is, I believe, reasonably certain. The

evidence for it is the close uniformity of these Mammalia in

general structure in spite of their wide divergence in adaptive

specialization, and the invariable approximation towards a cen-

tral type of each race whose development is known from

palaeontology. As a preliminary to further discussion we may

point out the characters of this primitive central type.

I.— Size very small, skull of moderate length, brain case com-

pletely enclosed in bone, brain of high type compared with that of

reptilia although lower than in the modern mammals. In every

Case where we are able to trace the descent of the large mod-

ern mammals, we find their direct ancestors successively smaller

as we pass backward in time. The horse and camel have been

traced back nearly to the beginning of the Tertiary; their ear-

1 This view was expressed by Huxley in 1880.

E :

812 THE AMERICAN NATURALIST. [Vor. XXXVIII.

liest representatives are no larger than rabbits. Elephants,

rhinoceroses, tapirs, every race about whose ancestry anything is

known, exhibit a reduction in size corresponding to the distance

back through which we have been able to trace them. The

large animals of the early Tertiary are in every case early

specializations which have left no descendants.

2.— Molar teeth ** bunodont" i.e., low crowned, the crowns

composed of a few low broad rounded cones, heavily enamelcov-

ered. The molars are tubercular (crushing) teeth, the premolars

trenchant (cutting), the canines moderately large (piercing), the

incisors small spatulate (nipping) teeth. The teeth were

arranged in continuous series, except for slight gaps behind

the canines. The labors of Cope, Osborn and many other

paleontologists have amply demonstrated this as the primi-

tive type of dentition among the Mammalia. Whether we

accept the whole of the Tritubercular Theory or not, this part

of it appears to be beyond question.

3.— Neck rather short, slender and flexible, permitting quick

and easy turning of the head in all directions. Trunk slender,

flexible, ribs rather short and few in number, lumbar region long

and comparatively flexible. These features characterize all early

Tertiary mammals, without exception.

4.— Tail very long and flexible, with strong muscles towards

the base, and probably prehensible. Al primitive Tertiary mam-

mals have remarkably long and strong tails. These differ from

those of the cats and resemble those of the prehensile-tailed

monkeys in the greater breadth of the zygapophyses of the

proximal caudal vertebrae and great size and length of

the median caudals.

5.— Shoulder girdle of scapula and clavicle. No indications

of a separate coracoid have been observed in early Tertiary

mammals, but the clavicle was certainly developed in the

ancestors of several groups which do not now possess it, and

was probably generally present in the earliest types.

: 6. — Ilia narrow and rod-like, gluteal muscles long and slen-

er.

: 7 . — Upper members of limbs comparatively long and loosely

Jointed to the trunk, permitting great freedom of motion. The

Nos. 455—456.] MAMMALIA 813

great relative length of the humerus and femur is a striking

feature in the limbs of all early Tertiary mammals. In the

evolution of the limbs for running the femur and humerus have

generally been reduced in length and compacted into the flank,

while the lower legs and feet are elongated. This places the

heavy muscles high up on the limb, and by bringing its centre

of gravity near to the proximal joint, enables it to swing more

rapidly through a considerable arc. Thus the animal gains in

length of stride without losing in quickness of step and can

maintain a high speed for a long distance with less fatigue.

8.— Ulna and radius separate, equal in size, radius with

round head, permitting free supination and pronation. Tibia and

fibula separate with probably more limited motion.

9.— Wrist and ankle very flexible, all the carpals separate and

a centrale present. Astragalus with flat trochlea, distinct neck

and rounded head. This type of carpus and tarsus is found in

all Basal Eocene mammals. It is retained with but little change

in primates, insectivores, most rodents, and some other groups.

Its gradual conversion into the various types seen in other

groups is demonstrable in the Amblypoda, Condylarthra, Car-

nivora, partly so in the Proboscidea and Edentata, but not in

the Artiodactyla and Perissodactyla which have already devel-

oped their peculiar types of astragali when they first appear in

the geological record.

10. — Five digits on each foot, the joints permitting of very

Jree motion of fingers and toes, which were tipped by small

claws. The argument for the derivation of all mammals from

pentadactyl ancestors has been fully set forth by Cope and

others.. The derivation of the hoofed from clawed types is like-

wise indicated in various lines of descent as now known.

LI. — First digit more or less opposable in both manus and fes.

This is contrary to the usual assumption that the opposable

thumb found in several groups of arboreal mammals is in each

Case a new adaptation to their habits of life. But there is con-

Siderable evidence for it. In the first place, as far as we can

trace back the history of each of the arboreal groups, we find

their first ancestors with the first digit as fully opposable sm

the modern representatives (v.g. the Middle Eocene primate

814 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Notharctus). Second, in those groups which: have Bii

opposable thumb, we find as we trace back their gears wt

the trapezium, whose form and facets give the surest pre ;

on this point, approaches more and more nearly to t e ds

preserved in the Primates, etc. It becomes large, Bees

with very concave distal facet for the digit, and round-con ge

proximal faceted end abutting against the scaphoid, cre

and centrale. In the four Basal Eocene mammals isis e

Euprotogonia, Clzenodon and Dissacus) in which this part 6 t

skeleton is known, the form of the bone is surprisingly uniform,

and when the manus is put together, the first digit is pue

partly outward from the rest of the hand, and permits of ae

freer motion than the remaining digits, with a considerable

degree of opposition. With the development of the foot for

terrestrial locomotion the trapezium and first metacarpal lose

their mobility, the rounded conical proximal facet of the lees

separates into two flat facets at right angles for scaphoid 2n

trapezoid, its distal facet becomes more plane and Hs who

distal end reduced in width and closely appressed against the

reduced to a nodule and finally disappears, and the trapezium

usually follows Suit, the trapezoid either remaining —

(Perissodactyla) or being consolidated with the magnum (Artio-

dactyla). gis

The primitive Opposability of the hallux is less clearly in

cated, but the close resemblance in the form and phe

of the internal tarsals, especially the internal cuneiform, in :

intermediate stage in the evolution o

pus, but that its evolution for terrestrial locomotion began earlier

Or proceeded more rapidly

In the third place

done, that the SO-

; if we suppose, as many anatomists -—

called first metacarpal is really a proximal pha

Nos. 455-456.] MAMMALIA. 815

for the development of this anomaly, and would explain (a) the

presence of but two phalanges on digit I, three on each of the

others; (4) the epiphysis of digit I being proximal as in the pha-

langes, instead of distal as in the remaining metapodials ; (c) the

anomalous musculature of digit I on all mammals, the object of

which is clearly seen when the digit is opposable, but is quite

unexplained otherwise.

It would be beyond the purpose of this article to give at all

fully the facts which support the above characterization of the

primitive mammalia, but enough has been said to show the gen-

eral nature and force of the evidence on which it is based.

The modern orders of mammals have departed to a varying

degree from this primitive type. The arboreal primates retain

most nearly the primitive character, except that the limbs are

elongated, the face is shortened and the brain greatly increased

in size and complexity. The rodents, largely perhaps on

account of their small size, have usually retained a great

deal of the primitive skeleton character, but the dentition is

much modified —least of all in the arboreal squirrels, which

are almost unaltered in dental characters from the first known

rodentia of the early Tertiary (Paramys, pte) The Insectivora

likewise retain many primitive characters, but in most cases m

combination with one or another high specialization of an

unusual kind. The Carnivora retain the primitive character

more or less completely in the neck, trunk and legs, but show

much greater adaptive changes in the feet and especially in the

teeth. (Several of the Creodonta have a more or less opposable

pollex, but this is lost in the later carnivora.) The Ungulata

are by far the most highly altered group among the land mam-

mals, almost every part of the body having gone through great

adaptive changes, although the evolution is most marked in

teeth and feet. We are able to trace the history of these

changes more fully in this than in any other order, as its geo-

logical record is more complete. The Edentata early developed

Certain remarkable specializations which soon obscured their

primitive characters. Of the remaining orders of placental

mammals our geological record is very imperfect, and their

relationship to the central : type mainly hypothetical. The

816 THE AMERICAN NATURALIST. [Vor. XXXVIII.

marsupials, as Dollo and Bensley have shown, are probably

descended from an arboreal type which must have closely

resembled the arboreal ancestors of the placentals, but with

distinctions in the number and succession of the teeth, the

origin of which is not yet clear. In these, as in the placental

mammals, the modern arboreal forms are the most primitive.

But it should be remembered that we know comparatively little

about the paleontology of the marsupials earlier than the Qua-

ternary, while we are able in most of the families of Ungulata,

Carnivora, Rodentia and some other orders of placentals, to

trace back their ancestry into the Middle or Lower Tertiary,

and find them in every case converging toward the type char-

acterized above, and quite closely converging as we come into

the Basal Tertiary. We are thus enabled to place the earliest

divergence from this central type in the Middle or Upper Cre-

taceous.

It may almost be taken for granted that if the characters of

this central type were in all respects as stated above, it must

have been of arboreal habitat. All its adaptations would be

suitable for such a mode of life, and some would be more or less

unsuitable for any other. The various modern groups (monkeys,

squirrels, arboreal insectivores, opossums) which have retained

this habitat are the least altered in structure, while the amount

of structural change in other groups, as shown by their known

paleontology, is proportioned to the change in their mode of

life, the Ungulata exhibiting the greatest changes.

The hypothesis may be stated as follows :

The Cretaceous ancestors of the Tertiary mammals were

small arboreal animals of very uniform skeletal characters, but

probably somewhat differentiated in dentition according as

fruit, seeds and nuts, or insects, formed the staple of their :

diet. At the beginning of the Mesozoic the available modes of

life for land vertebrates were chiefly the amphibious-aquatic,

1A further develo

distinguished the marsupials, i

is this feature that is regarde

habits.

Nos. 455-450-] MAMMALIA. 817

the arboreal and the aerial, the terrestrial habitat being sub-

ordinate because the upland Flora was largely undeveloped or

inedible as compared with its present condition. The three

available provinces were occupied by reptiles, mammals and

birds respectively. In the later Cretaceous the spread of a

great and varied upland flora vastly extended the terrestrial

province, and opened a new and constantly widening field for

the expansion of the mammalia. These then commenced a

great evolution, the new terrestrial groups expanding con-

tinually and becoming adapted to various modes of life. The

arboreal types maintained or increased their lead in intelligence,

but changed comparatively little in other respects. The terres-

trial types became far more numerous and dominant, adapting

their primitively arboreal organization to their various modes

of life, yet retaining, in spite of extensive changes, a certain

fixity of type which had been impressed upon them by their

long arboreal residence.

The great extension of the terrestrial province in the Ter-

tiary may be supposed to have opened a correspondingly large

field for the expansion of the birds, but these, retaining in the

main their aerial life, suffered but little change in organization,

and the vast majority are today as homogeneous in skeleton

structure as the mammals were at the beginning of the Ter-

tiary. Coincident with the expansion of the terrestrial prov-

ince was a great contraction of the amphibious-aquatic prov-

ince. The extensive swamps and deltas and great inland seas

of the Cretaceous, shrank at its close to small proportions,

and the Reptilia underwent a corresponding diminution, some

groups completely disappearing, others surviving through the

Tertiary in continually decreasing numbers, the land reptiles

. (lizards and snakes) alone prospering. Both birds. and rep-

tiles, so far as they have retained their typical habitats, have

changed but little structurally since the Mesozoic; only the

few terrestrial reptiles and terrestrial or aquatic birds cel an

amount of change comparable with that in the terrestrial mam-

malia. It would seem therefore that all the facts accord with

the explanation of the evolution of mammals during the Ceno-

zoic as caused by their invasion of a new province and change -

of habitat from arboreal to dominantly terrestrial.

818 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The little that is known of the Mesozoic Mammalia fits in

with our hypothesis of their arboreal habitat but adds little

to the evidence in its favor. Practically nothing is known of

their skeletal structure; they are all of small or minute size,

with teeth of insectivorous or granivorous type. They have

been referred to monotremes, marsupials and insectivores, in

each case on very insufficient evidence, but their ordinal

relationships have little to do with the question of their habi-

tat and need not be considered here. Their minute size, and

association, in strata of fresh or brackish water origin, with

large amphibious and aquatic reptiles and with abundance of

fossil wood, suggest that the deposits in which they occur

were laid down in extensive forest-clad river deltas and coastal

swamps, and that the minute mammalia represent the arboreal

fauna of these forests.

I am not well acquainted with the embryologic evidence

which might bear upon the hypothesis advocated in this paper.

Most investigations into mammalian ontogeny have had other

problems in view. The deviation of placentals from mar-

supials or vice versa, and the ultimate origin of the mammalia

from amphibian or reptilian ancestors, do not directly affect the

question of the habitat of the Mesozoic ancestors of the Ter-

tiary mammals. But undoubtedly important evidence on this

point could be obtained from the ontogeny of their modern

descendants, although not, perhaps, of the same force as the

more direct evidence from palzeontology.

AMERICAN MUSEUM OF NATURAL HISTORY

June 15th, 1904.

LOCALIZED STAGES IN COMMON ROADSIDE

PLANTS.

JOSEPH A. CUSHMAN.

THE plants considered in the present study are of common

occurrence along country roadsides. With, perhaps, the excep-

tion of some of the early stages of development, the leaf forms

noted may be found on country roads in the spring or summer.

Although among our commonest plants, some actually classed

as weeds, the stages shown in their development are as striking

as could be found in any plants. The stages in development of

other common plants have been described (Amer. Nat., Vol. 36,

. No. 361), as well as some exotic plants of the Botanic Gardens

of Harvard University (demi, Vol. 37, No. 436). My thanks

are due to Dr. R. T. Jackson for corrections and helpful sugges-

tions. Each species is treated separately and its stages described

in detail. ,

Thalictrum polygamum Muhl. Figures 1, 2.

(Tall Meadow Rue.)

Although this plant does not become conspicuous by its flow-

ering until midsummer, its early growth from the rootstock and

seedlings must be looked for in moist ground very early in the

Spring.

In the seedling (Fig. 1) the first nepionic leaf is simple, with

three lobes. Such a first leaf is frequently found, but fully as

often the first leaf is like the second leaf in the figure. The

second leaf is trifoliolate, each leaflet being like the whole of

the first nepionic leaf as figured. From this point the leaves of

the young plant, as added, increase in complexity, the plant

finally producing the 2—3 ternately compound leaves characteris-

tic of the adult.

819

820 THE AMERICAN NATURALIST. [Vor. XXXVIII.

As the plant is a perennial its early spring growth may be

easily studied.

When first noted it usually has complicated leaves, but if the

very earliest growth is looked for, leaves will

be found exactly comparable to those of the

seedling. The trifoliolate form is common-

est, but in some cases, especially where the

plants grow in sandy soil, the first leaf will

be simple, as in the first leaf of the seedling.

The steps in the further complication of the

compound leaf follow the same stages as in

the seedling, but usually more rapidly, arriv-

ing at the typical leaf of the species by fewer

stages than in the seedling. This is a good

example of acceleration of development seen

in localized stages.

The plant produces great masses of the

Fi. 1—Seedling of 74a- small flowers, and the strength thus used up -

showing cotyledons, e, 18 deducted from the amount which goes to

“ and first and second leaf development, with the result that below

first with 3,the second the flower there is always shown localized

mith s lolita Senescence, exhibiting regressive develop-

ment. Figure 2 shows leaves in the series below the flower

panicles. There is a gradual reduc-

tion of the proximal portion as shown

in leaves 1 and 2, until a leaf is pro-

duced (Leaf 3 of Fig. 2), in which the

leaf is trifoliolate, as in leaf 2 of

Figure 1, but having no lobes on the

lateral leaflets. Next, the lobes of

the terminal leaflet are reduced and

the outline of all the individual leaf-

lets thus becomes entire. Still later

the two lateral leaflets disappear and

a very small, simple, entire léaf is Fic. 2.—Flowering stalk of a

found (Leaves 5 and 6), simpler than i ie en

anything seen in progressive develop-

ment, either in the straight develop-

ment of the seedling or the localized development in early spring

flower. Leaves 1-6 enlarged to

natural size.

Nos. 455-456] COMMON ROADSIDE PLANTS. 821

growth. The stages below the flower are very noticeable and

are easily made out.

Daucus carota L. Figures 3-13.

(Wild Carrot.)

In this common weed very marked localized stages are found.

The direct development of the seedling shows

much acceleration as the first leaf of the ordi-

nary seedling is considerably complex. The

first nepionic leaf of the seedling of this spe-

cies is shown in Figure 3. It is a decidedly

compound leaf. The second leaf is still more

.. complex.

The spring growth is to some degree diffi-

cult of investigation, as the plant starts its

growth very early. The first leaves seen

under usual conditions of growth are very

complex. It was therefore a matter of some

trouble to obtain specimens showing rever- mc e jen

sionary characters. Finally plants growing $% s coni amc

under unfavorable conditions were sought.

There the results were most gratifying. The, plants shown in

Figures 4-6 were found growing in sand along a railroad

embankment where the conditions were decidedly adverse.

They are of course weak

specimens, although the

size of the stem shows

them to be adult plants

of at least one year’s

growth.

ics. 1-6. — spring growths of Dancer ys, $8 [n 90. In Figure 4, the first

weak growths; 4-6 (1-4), more vigorous growths. leaf of spring growth is

simple and entire, the second leaf is similar but with a lobe on

one side, the third leaf is trilobate and the fourth also. The

fifth-and sixth leaves have five lobes. In Figure 5, the stages

except in the second leaf are exactly similar. From the plant

822 THE AMERICAN NATURALIST. [Vor. XXXVIII.

shown in Figure 6 two buds started. The series from the lat-

eral weak bud, 1a—4a, are in the main like those of Figure 5, but

the series from the stronger bud, 1—4, show a greater accelera-

tion of development. The first leaf, No. 1, is comparable to

the third leaf of Figures 4 and 5 and the third leaf of weaker

growth on this same plant. The second leaf, No. 2, is compara-

ble to the fifth leaf of Figure 4, and the third leaf is more com-

plex than anything shown in the other cases of spring growth

figured, but is closely comparable to the first nepionic leaf shown

in the seedling (Fig. 3). In the localized development shown in

early spring growth of weak plants, therefore, stages are found

which are simpler than anything found in the direct development

of the seedling. In vigorous plants spring growth is so much

accelerated that these simpler stages are not found.

In Figures 7—13 are shown the leaves beneath the flower. Fig-

| 9 12

IO H

, Fics. 7-13. — Series of reduced leaves (“involucral bracts ") found beneath the flower of

mucus creía, 7-8, upper leaves on the stalk ;- 10-13, clustered bracts beneath the flower,

13 being the upper one of the series.

ure 7 shows the typical adult leaf of the stem, Figure 8 the upper

leaf on the stem in the specimen examined. By comparing Figure

8 with the second nepionic leaf of the seedling (Fig. 3), it will

be seen that the two are very similar except that the leaf tips are

more acute, almost spiniform. Below the flower cluster there

are what are usually termed “cleft involucral bracts.’ These

Nos. 455-456.) COMMON ROADSIDE PLANTS. 823

are really reduced leaves as they are the morphological equiva-

lent of such. Figure 10 is, in the number of its main divi-

sion, comparable to the first nepionic leaf of the seedling (Fig.

3), and to leaf 3 of Figure 6 in the early spring growth. It dif-

fers however in not showing secondary lobes as seen in the other

cases. Leaves of the next succeeding whorl have five main

lobes, Figure 11, comparable to leaf 2 of Figure 6, or to leaf 5

of Figure 4. The one shown in Figure 12 is comparable to leaf

3 of Figure 4, or to the second stage in early spring growth.

Figure 13 shows the ultimate reduction and a simple leaf com-

parable to the simplest first stage of spring growth. his series

below the flower repeats in the reverse order, the steps shown

in the progressive localized development of early spring growth

and in the direct development of the seedling. In other words,

it is regressive development repeating progressive development,

but in the reverse order.

Baptisia tinctoria R. Br. Figures 14-16.

(Wild Indigo.)

This plant shows localized stages ina simple manner, yet very

completely. In the direct development fre-

quently starts off with the first nepionic leaf

simple, broad and rotund, distally emarginate,

the stipules not large but nevertheless notice-

able (Fig. 14). The first leaf is

broader and more rounded than

the type leaf of the species (Leaf

I of Fig. 16) in which the tip B

dn m ri. slightly mucronate instead of

Minia tinct lightly emarginate. The sec

ledons, c, c, the ond nepionic leaf is trifoliolate,

nepionicleaves,the the leaflets rounded, wedge- .

angi obovate. The succeeding leaves sto ~

are all trifoliolate, the stipules comparable to di-

becoming smaller and less persistent.

Comparing these stages in direct development — . :

with localized stages indevelopment seen in early sprog growth,

Fic. 15.-Early spring

growth of Baptisia

824 THE AMERICAN NATURALIST. (VoL XXXV IHI.:

the similarity is at once apparent. In some plants, probably but

a year old, there were found many cases where the first leaf of

spring growth was simple and emarginate, but like that of the

first nepionic, much broader than the later

leaves (Fig. 15). The stipules were also

larger than in succeeding stages and in this

again comparable to the first nepionic leaf of

the seedling, showing a more accelerated con-

dition. In all cases the second leaf of spring

growth was trifoliolate. Those which were

found with the spring growth starting off

with a simple leaf were obtained in a situa-

tion unfavorable to the plant and thus favor-

ing such retarded conditions of growth. The

overlying loam had been removed from a

glacial sand plain, leaving the sand exposed

Vu rM a south slope. This gave a poor supply of

of Baptisia tinctoria, mOisture and exposed the plants to the full

sowing the reduction of force of the sun. As a result, the plants

2 Pug: belowthe showed a retarded development.

In the adult plants, when in flower, cases

of localized senescence are not infrequent. Below the flowers,

the last leaf is often simple, comparable to the first nepionic

leaf of the seedling (Fig. 14), and to the first leaf of spring

growth in feeble or retarded plants (Fig. 15). This reversion-

ary condition below the flower is not usually shown, however.

Rumex acetosella L. Figures 17-109.

(Sheep Sorrel.) |

The peculiar auricled form of the leaves of this plant fur-

nishes a means for study of stages, although the leaf is not

compound as in the preceding cases. The seedling (Fig. 17)

has the first and second nepionic leaves ovate, tapering some-

what at the base and with acute apices. In the third leaf the

base is broader and the tip more blunt. The fourth leaf has a

much broader base and more blunt apex so that it assumes à

Nos.455-456] COMMON ROADSIDE PLANTS. 825

roughly oblong shape. The fifth leaf has the auricled base

Characteristic of the species. Later leaves hold to this char-

acter. The early leaves of the seedling are red while later ones

are green.

Turning to the early spring growth, very similar stages are :

noted (Fig. 18). In this f Z

weak plant the stages are

exactly the same, leaf for

leaf, as in the seedling after T

Z 2

the cotyledons. In Figure

19, the early growth of a

somewhat stronger plant, the Fics. 17-19. — Seedling and early spring growths of

first two leaves are similar to pecs tp Mieres pide ji

the first two of the seedling. early spring growths showing stages by whi

à auricled form of base is reached.

The auricled form appears

here in a vigorous plant in the fourth leaf, one stage earlier than

in the seedling. The leaves in the early spring growth also

resemble the seedling in their reddish color, but this is a super-

ficial resemblance and not one of true repetition. It is depend-

€nt upon other causes.

On the flowering stalk there is a reduction of the base in the

same manner, but in the reverse order, until below the flower

there are linear leaves with narrow apices and bases. Thus the

first leaves of the direct development in the seedling, the first

leaves in the localized development in early spring growth and

the last leaves in the reversionary localized. senescence below

the flower aye without the auricled bases. In all other parts of

the plant the leaves have the auricled base typical of the

Species.

Potentilla canadensis L. Figures 20-23.

(Common Cinquefoil.)

In this plant the seedling usually has the first nepionic leaf

Simple, the second ternate (Fig. 20). Succeeding denves are

ternate but apparently five palmate, due to the splitting of the

lateral leaflets into two parts as shown in leaf 4 of Figure 21, a

Spring growth.

826

THE AMERICAN NATURALIST. [Vor. XXXVIII.

The early growth of this plant from adult roots is usually

strong, but some plants have simpler leaves than characterize the

adult. Over a hundred young plants were examined, all in an

FiG. 20.- Seed-

c,and first and

second nepi

onic leaves,

the first si

. of the early spring growth.

early condition of spring growth. Of these, the

great majority had already the dissected ternate leaf,

typical of the plant. Between fifteen and twenty of

the smaller plants had the early leaves ternate, with-

out dissection of the lateral leaflets (Fig. 22, leaf 2).

- In this specimen the first two of the early leaves had

d this character. In only one case in the hundred

or more examined was a simple leaf found as the first

In this specimen (Fig.

21) the first leaf was simple, the second ternate, and

m- E *

- the succeeding ones of the typical dissected ternate

orm. This shows the rarity of this stage both in

seedlings and spring growth, it being usually skipped

by acceleration. ;

In the flowering there is usually little reduction, for the flower

is comparatively very small and not I

confined to the end of the shoot. j

The leaf from the axil of which the — 7

flower comes, may be reduced to »

simple ternate one, but in the speci- 3

mens examined no further reduction

2374

Fics. 21-22. — Early spring growths

of Potentilla canaden 21, show-

was found. In the late :

ing exceptional case where first leaf

season’s growth, how-

ever, very definite sen- ;

escence is seen (Fig. 23). Plants were obtained in

FtG. 23. — Final

rare cas in

which the last

leaf, no. €, is

simple, due to

lack of growth

force at the end

of the growing

season. *

late November, when the leaves were nearly or

quite dead, where the last leaves were reduced very

frequently, to a simple ternate form, as shown in

leaf 5, Figure 23. In one case out of those of this

late growth which were examined, there was a sim-

ple leaf as the last leaf of the season's growth (No.

6, Fig. 23). This leaf, the last of the senescent

ones of the season's growth, is comparable to the

simple leaf seen in the first stage in the direct

development of the seedling, and also to the first leaf of the

Nos.455-456] COMMON ROADSIDE PLANTS. 827

localized development, seen in the early spring growth. This is

due to the lack of strength in the latest part of the season's

growth and a consequent reduction follows. In this senescence

then, the same stages are found, but they are developed in the

reverse order.

Sambucus canadensis L. Figures 24-25.

(Common Elder.)

This seedling of this plant was not obtained, but one of

another species, S. zégra, is figured by Lubbock (Seedlings, vol.

II, p. 51, Fig. 420). In this species the leaves are in pairs.

The two first pairs of nepionic leaves are simple, toothed, and

the third pair is trifoliolate.

In the early spring growth of our species there are first, leaf-

like bracts shown in Fig-

ures 24 and 25. These

are notched or toothed

at the distal end. In

most cases there are

three of these teeth or in

some cases five. These

teeth seem to represent

the reduced leaflets of a

compound leaf — in the Frcs. 24-25. Ears spring growths of Sambucus c

commonest case a trifo- 24, showing the five toothed bract followed by , m

liolate one. In such a wih a

case the first true leaf is trifoliolate or it may due five leaflets.

In some cases, after the three-toothed bract there is a five-toothed

. One, usually more expanded and leaflike. Such a form is not

followed by a trifoliolate leaf, but, in all the cases noted, by a

leaf with five leaflets. These teeth then appear to represent the

reduced leaflets, for if they did not there would be no reason

against producing a trifoliolate leaf directly after the five-

toothed bract. These bracts might be thought of as the stage

representing a simple toothed leaf, but this definite suctession

of a leaf with five leaflets after a bract with five teeth seems to

indicate them as reduced leaflets and not true teeth. :

528 THE AMERICAN NATURALIST. (Vor. XXXVIII.

Beneath the flower there may be a very slight reduction in

the number of leaflets, but as a rule it is not carried to any

noticeable extent.

Chenopodium album L. Figures 26—36.

(Lamb's-Quarters. Pigweed.)

This very common weed shows excellent localized stages. It

' is an annual and therefore shows it simply in

the localized senescence below the flower.

The seedling (Fig. 26), shows the direct

development. The first two nepionic leaves

are narrowly oval, tapering at the base and

apex, with the basal portion not broader

than the similar apical portion. The third

leaf has a broader base and in the fourth it

has become decidedly angled. In succeeding

leaves the base becomes angulate-toothed (leaf

Fs wena d 9). This leads quickly to the adult character

Chenopodium album, (Fig. 27), and is continued until flowering.

e ma oyid Then comes the reduction shown in the series

c, and several of the

nepionic leaves. (Figs. 27—36). This series represents leaves

in the axils of which branches or flowers are produced. The

27 ^ 28 29 oO " «| n uU

FiGs. 27-36. — Series of leaves on the stalk of Chenopodium album,

showing the st ges in th duction from 27, the typical species leaf,

to 36 the linear, reduced leaf below the flower.

wine

5 y»

typical many toothed condition shown in Figure 27, 1s, in

succeeding leaves, reduced until finally the simple leaves shown

m Figures 35 and 36 are produced below the flower. The

Nos.455-456] COMMON ROADSIDE PLANTS. 829

stages passed through are very similar to those in the direct

development of the seedling, but are in the reversed order of

sequence.

Achillea millefolium L. Figures 37—42.

(Common Yarrow.)

The seedlings of this plant are very similar to those of

Tanacetum vulgare L., as may be seen e a cesat of the

figures of the seedlings of that :

plant (Amer. Nat., vol. XXXVI,

p. 881, Nov. 1902) with those

of Achillea given here (Figs.

37-40). Both plants belong to "rer de ta mir

the same tribe of the Composite. poai individuals. 37 and 4o showing

There is much variation in the rine eee oe

seedlings as was also noted in

Tanacetum (oc. cit. p. 871). The first pair of nepionic leaves

are usually three toothed ; the second pair five toothed (Fig. 40).

As shown by the figures this may be decidedly varied. Figure

37 shows the typical first pair of leaves; Figure 40 in addition the

typical second pair. Figure 38 shows a retarded individual where

the character of a three-toothed leaf, usually taken on in the

» 9 first pair of nepionic leaves has not -

2 appeared, except as indicated by a

single tooth of one leaf. Figure

39 shows an accelerated individual

where the character of a five-

toothed leaf, characteristic of the

second pair of leaves, is shoved

back and appears in one leaf of the

Fics. 41 ma ede mea DUM pair. |

de mii ate In the early spring growth better

ment of two individuals, dai temas results were obtained than in the

more retarded than those usually found. case of Tanacetum. Specimens

of Achillea were found growing in the sandy railroad embank-

ment which has been previously referred to. As in Tanacetum,

the simplest leaves as a rule seem to be five toothed as in Figure

Al 42

830 THE AMERICAN NATURALIST. [Vor. XXXVIII.

42. In rare cases, however, plants were found which had the

first leaf with three teeth. In the specimen (Fig. 41), the first

leaf was simple like one of the first nepionic leaves shown in

Figure 38; the second was three toothed like the first nepionic

leaves of the typical seedling (Figs. 37 and 40), and the third

leaf has five teeth as was so commonly found as the first stage

from rootstocks in the early spring growth. This then is a very

retarded individual and repeats in its localized development, all

the main stages seen in the whole series of seedlings representing

the direct development.

Beneath the flower reduced leaves are also found. At the

junctions of the stems of the compound corymb, leaves are found

which are like those of the early spring growth and seedling in

that they are simple and but few toothed. At the upper portion

near the ultimate branchings are found leaves with five and

three teeth and in some cases, simple entire leaves. These

leaves are seen to repeat the same stages as in the localized

development shown in spring growth and in the direct develop-

ment of the seedling but in the reverse order. It is still another

case of the localized reversionary senescence already referred to.

Aster acuminatus Michx. Figures 43-48.

(White Wood-aster.)

The seedling of this species was not obtained but a compari-

son of the early spring growth of an

adult plant with the portion below

the flower heads shows striking simi-

larities. The stages are very simple,

varying mainly in the number of the

lateral teeth.

In the early spring growth of a

a d 5 plant a year or more old, the radical

Fiaa. aat Losies of ANE ai leaves are at first entire and some-

showing three of the stages what spathulate (Fig. 43). The next

typical species leaf, 45. leaves have two lateral teeth and

succeeding leaves have two lateral

pairs (Fig. 44). In further development three lateral pairs are

Nos.455-456] COMMON ROADSIDE PLANTS. 831

produced and in the mature condition the leaves have four pairs,

as in Fig. 45, which represents the typical leaf of the species.

Leaves like that shown in Fig. 45 are produced until the time

of flowering. Then a reduction in the num-

ber of teeth takes place. Leaves with four,

then three, pairs of lateral teeth are produced

and nearer the flower heads, leaves with two

pairs are shown in Fig. 46, later with the

single pair (Fig. 47), and finally below the

flower, the simple leaf shown in Figure 48. "eS, e lu cree

The reduction here is carried on in a definite ^ ized senescence below the

manner and retraces exactly, in reversed mys showing ian ^u

order, the same stages that are passed ‘sein thereduction.

through in the early spring growth, so that, using numbers for

the lateral teeth, we may represent the stages graphically, call-

ing the entire leaf number 1, as follows:

Early spring growth to Flowering

a ee ee ee > d y ai CD Jr a

Eupatorium perfoliatum L.

(Thoroughwort. Boneset.)

This well known plant shows in its leaf characters certain

things which may be noted here as in the same line with the

preceding cases. The main leaf character of this plant is the

connate-perfoliate form of the leaf. This, however, by an exam-

ination of a flowering specimen, will be seen to occur on only a

part of the stem. In the early spring growth of the plant, the

bases of the opposite leaves are free and narrow. Later the

coalescent form is taken on and kept until well toward the flow-

ering. Then the early form is again taken on, the base of the

leaves being narrowed and failing to meet.

A variety of this species, var. cuneatum Engel., is apparently

a more primitive form. It keeps the early character throughout

its life history, the leaves failing to coalesce at their bases.

This variety also has fewer flowered heads than the typical form

of the species,

832 THE AMERICAN NATURALIST. [Vor. XXXVIII.

The commonest plants illustrate well the principle of localized

stages in development seen in the direct progressive develop-

ment of the seedling, in localized stages of spring growth and in

the localized senescence at flowering. It may be shown in a

variety of ways, as noted in the present paper. In Thalictrum,

Baptisia, etc., it was shown especially by change in the number

of leaflets; in Rumex, by the changes in the auricled base; in

Daucus, etc., by the change in the number of the teeth, and

finally in Eupatorium, by the presence or absence of the connate

form of the leaf:

Different individuals show variations due to differences in

acceleration of development. There is a differential acceleration

in individual seedlings, shown where the first nepionic leaf in

various individuals is more progressive or more retarded than in

the typical cases. In spring growth it is also shown in much

the same way, and with a still wider range of variation. These

differences in acceleration may be caused by poor conditions,

such as poor soil, lack of sufficient moisture, cold, etc., and by

internal causes not accounted for in the above.

Regressive development, seen in the localized senescence

below the flower, is shown in many cases to be more reversion-

ary than stages in the usual seedling. When a large number of

seedlings are examined, however, such extreme cases of varia-

tion may be seen which would otherwise be overlooked.

Boston SOCIETY OF NATURAL History,

May, 1904.

AN ARRANGEMENT OF THE FAMILIES AND THE

HIGHER GROUPS OF BIRDS.

R. W. SHUFELDT.

Birps constitute a class of the phylum Vertebrata, and

belong to the branch Craniata. Above the latter subphylum

they are associated with the Reptilia in the tribe Sauropsida,

"which last in time has divided into two now distinct classes,

namely, Reptilia and Aves (see A Classification of Birds, postea).

Of all the existing vertebrate groups, birds are, morphologically

speaking, the most homogeneous; probably none of the phyla

recognized by zoólogists are more so. There is a very con-

siderable gap between a thrush and an ostrich, but it in no way

compares, in the matter of profundity, with the gap that stands

between man and the duckbill, or between an elephant tortoise

anda garter snake. This morphological homogeneity in birds by

no means renders their taxonomy any the less difficult for us ;

indeed, for very obvious reasons it greatly tends to enhance the

intricacies of the problem. This fact is now so generally appre-

ciated by avian classifiers that it is quite needless to discuss it

in the present connection.

To classify birds correctly and to point out the natural rela-

tionships and interrelationships of all the species and subspecies

now in existence we resort to various lines of research and

employ data of widely different nature. In the field of palzon-

tology we meet with a mass of material, the comparative study

of which has led to the conviction that Aves and Reptilia have

arisen from a common stock. The indications of this have by

no means died out in certain existing representatives of these

two classes of the Sauropsida. For example, it is quite appar-

ent when we come to trace the ancestry of the existing ostrich

and its surviving allies in various quarters of the globe. The

trend backward in time is distinctly reptile-wards and eventually |

brings one to the consideration of a long-extinct assemblage of.

833

834 THE AMERICAN NATURALIST. (VoL. XXXVIII.

forms from which not only have the ostriches arisen, but both

modern birds and reptiles have been derived.

In the classification of existing birds, again, we have recourse

to their anatomical structure or morphology, and incidentally their

physiology to assist us, the first of these being a very powerful

aid. By morphology is meant the science of the outer form and

internal structure, and to be of practical value in taxonomy it

must be made thoroughly comparative. As far as possible we

also study the morphogenesis of birds or the genesis of form in

their case through evolution. Of great assistance is the knowl-

edge we derive from our researches into the comparative ptilosis

and pterylography of the group and all that pertains to these

related sciences. In our efforts, too, to seek out the true rela-

tionships of birds in order to arrive at a natural classification we

must take into consideration, in the broadest possible sense,

their embryology; their oólogy and nidiology ; their habits and

comparative longevity, and even their various notes and songs

must be given due weight. Their geographical distribution, a

very important factor to be studied in their taxonomy, is to be

considered not only from the viewpoint of the present distribu-

tion of the species of the class over the earth's surface, but like-

wise a comprehension, as far as possible, of the question as to

how that distribution came about. The data for the latter, in

the case of birds, are extremely meagre, as it takes into consid-

eration the migration and original habitats of various forms of

the class during geologic times, and as but very few fossil

remains of birds have been discovered in any part of the world,

there has been in consequence but very little light thrown upon

this latter side of the question. So faras it goes, however, it

has its value, as has also, to some extent, a consideration of the

migrations, er se, of existing species and subspecies.

In some of these sciences, as in the case of ptilosis and ptery-

lography for example, not only must both sexes be studied and

compared, but the young at all stages of their existence. Both

ptilosis and pterylography furnish very useful data to assist us

in the classification of Aves, and as exponents of a bird's topo-

graphical anatomy, for this purpose they are quite co-equal in the

matter of importance with the osteological system of the internal

FAMILIES AND HIGHER GROUPS.

structure ; indeed, ptilosis, pterylography, and osteology stand

among the most important factors at our command, in the

Kio. 1. — keleton of the Great Bustard. O£is tarda Linn.

Y The enlargement on the upper

SEQUI E ,

mandible is abnormal. Coll. Natl. Mus. No. 12,315.

economy of this group of vertebrates, that become available in

their classification. Right here it will be as well to state, how-

836 THE AMERICAN NATURALIST. [Vor. XXXVII.

ever, that we will never arrive at the true and natural classifica-

tion of birds, through the employment for that purpose of any

single set of morphological characters.

A study of the skeleton in birds, for example, carries us a long

way toward an understanding of their various alliances and rela-

tionships ; this is particularly the case on account of what is to

be derived from avian paleontology, — the bones being, with

few exceptions, the parts preserved. It is extremely unsafe and

dangerous to the science of this subject, however, to thus employ

the osseous system alone. The taxonomical scheme based upon

such knowledge should be modified and corrected by the employ-

ment of every other fact, every particle of information that has

been made known in the matter. To this end various char-

acters that have been found to exist in the muscular, the arterial,

the digestive, the nervous and other systems have proved to be

of considerable use.

A comparative study of the beaks and feet ; the various char-

acters presented on the part of the plumage, especially as to

color, structure, and style, often constitute admirable checks on

a classificatory scheme of the class based on osteological data.

Food and other habits when properly studied, and the results

applied, are also efficient aids to establishing many of the prim-

ary divisions in our scheme ; and a consideration of such data

without reference to facts of any other kind, is sufficient to

enable us to refer a duck, a hawk, or a thrush to their proper

places in a taxonomical arrangement. In other particulars nidi-

ological and oólogical studies are of value taken in connection

with the anatomical and other ones enumerated. Some birds

build no nest at all, others construct them of all manner of shapes

and sizes and out of all manner of materials. Some birds lay

but a single egg, while others may lay a dozen or more; some

eggs are plain white and unspotted, others are .of other colors

and unspotted, still others are variously marked or possess other

peculiar characters, — all such facts, however, when properly

comprehended and assorted, may be employed in classification

with distinct advantage. For example, certain birds possessing

an association of osteological characters in their skeletons lay

but two white eggs in an elaborately constructed arboreal nest,

Nos. 455-456.] FAMILIES AND HIGHER GROUPS. 837

while no other existing birds of the class Aves have a similar

association of characters and habits. Here then nidiological and

odlogical facts support and emphasize osteological ones in estab-

—

lishing affinities. Again, were we told that we had before us the

Coll. Natl. Mus. No. 11,416.

Fic. 2, — Skeleton of the Kiwi, A fteryx australis Shaw.

2m 4 - ^ * . C DEC S >» SQ > qt j-

Skeletons of two small birds zz each ot which the osseous mandi

bles were slender and short ;

pterygoid processes absent ; while in the sternum the keel was

the nasals holorhinal ; the basi-

838 THE AMERICAN NATURALIST. [Vor. XXXVIII.

well developed, and the posterior margin of the body of the bone

presented zo notches whatever, we would be quite at a loss to say

what the two species were, but the moment that we added to

these osteological characters the fact that it was also known that

in the case of one bird it laid its s¢zg7e white egg in a burrow in

the ground, while the other laid several white eggs in a little

basket-like nest built by itself and found within the cavity of

some great hollow tree, we would not hesitate to say but what

the skeleton of the first belonged to some one of the smaller

petrels, and that of the second to a swift, and very possibly a

Cheetura. Your opinion is considerably strengthened when you

are told that the bird laying the single white egg had webbed

feet, and was strictly a marine species, while the other possessed

no such character of the feet, and was a typically insectivorus

aérial land bird. When still other characters from other ana-

tomical systems and parts are added, the true relations of the

two species can be fixed with absolute certainty. In the pres-

ent instance they chance to be very remote, although this by

no means seemed to be the case when only the few osteological

characters were mentioned. Such researches place forms in

their proper groups, but to decide upon, or to discover the true

relationships of, the families and main groups to each other is an

entirely different matter and infinitely more difficult.

It is very important indeed that we should thoroughly com-

prehend the origin and evolution of such an assemblage of

vertebrate forms as birds, and it is highly important, too, that

we classify existing birds in such a manner that our classification

conveys to the mind not only an orderly arrangement of our

knowledge upon this subject, but a scheme representing aS

near às possible the actual and natural relationship of the major

and minor groups of birds as they now exist. To convey such à

Scheme to the mind various plans and methods have been pro-

posed and adopted by a number of ornithological taxonomers at

wa of the history of the science. It is not neces-

all of these be described here,— a few of the more

ensis examples will answer our purpose. In his memoir

ada i Osteology of Gallinaceous Birds and Tinamous,

Ctore the Zodlogical Society of London on the 25th of

Nos. 455-456.] FAMILIES AND HIGHER GROUPS.

639

" ` I . Mus. No.

Fic. 5. — Skeleton of the spotted Tinamou, Nothura maculosa Temn. Coll. Natl. Mus

17,949.

840 THE AMERICAN NATURALIST. [Vor. XXXVIII.

November, 1862, Professor Wm. Kitchen Parker gives us one

of the earliest methods of setting forth in print the relation-

ships of a number of birds treated of in the work named.

These had to do with genera and not with families, and he

conveyed his ideas on their relationships in two ways. One

of these consisted in a method of what might be termed

grouping, and the other the columnar method, or as it is some-

times called, the “linear,” for the fact that the genera, or fami-

lies, or the higher groups are printed in a linear sequence fol-

lowing in an order determined by what is supposed to be their

relationships. Thus Parker said in the above quoted memoir on

page 235, *I will first show, in two parallel columns, how both

the Fowls and the Rails run insensibly through certain leading

genera into the lowest (reptilian) types of diving-birds." |

Notornis Gallus

Brachypteryx Crax

Ocydromus Talegalla

Tribonyx Palemedea

Crex Anseranas

Rallus ; Plectropterus

Gallinula Anser

Porphyris Anas

Fulica Fuligula

Podilymbus Harelda

Podiceps Biziura

Podica Merganser

Aptenodytes Phalacrocorax Colymbus — Alca

This method has its advantages, also its many disadvantages,

and Parker felt the weig

ht of some of these when he placed at

the foot of the first column Aptenodytes aside from but near to

Phalacrocorax, and in the second column Colymbus aside from

Alca. Soon the very next page or two (236, 237) he resorts

to the grouping method and uses it in the case of Pluvialis,

Talegalla, Hemipodius, Syrrhaptes, and Tinamus. This plan is

well shown in the case of the last named genus,— thus :

ors ius f ie

455-156] FAMILIES AND HIGHER GROUPS. 841

mel. Coll. Natl

Pic. 4. — Skeleton o :

keleton of the Hoatzin, Opzsthocomus cristatus G

842 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Gallus

Charadrius Dendrortyx Ocydromus

Syrrhaptes Hemipodius

TINAMUS

Apteryx Rhea

Casuarius

MAMMALIA REPTILIA

It is very evident that such a method of grouping could never

be adopted, for to treat in general a// the genera in any such a

manner would certainly fill the subject with confusion, and the

same would apply to the families being dealt with and printed

according to any such an arrangement. It answers fairly well

in the case of exhibiting the relationships of a single genus or

other group, and for this purpose it has been adopted by some.

A modification of this scheme consists in joining the names by

straight lines in order to render the kinships more evident. This.

doubtless led to the use of the phylogenetic tree, a plan which

will be more fully described farther on. In the matter of the

linear method, with various modifications it has been employed

by not a few avian taxonomers. Sharpe arranged Huxley's

(1867) classification of Birds in this way in his now famous

paper entitled, “ A Review of Recent Attempts to Classify

Birds" (1891, p. 4); Garrod resorted to it, and so did Sclater.

Alfred Newton did to a limited extent, while Reichenow,

Stejneger, and Fürbringer have presented complete schemes of

avian classification in this manner, — Reichenow and Fürbringer

employing the phylogenetic tree in connection with it. To some

extent Seebohm also used the linear method, and Sharpe illus-

trating the latter employed a “diagrammatic map” giving the

Seebohmian “Orders” in small circles, these circles bearing cer-

tain relations to each other, which relations were supposed to

represent in a way, and convey to the mind, the relations the

groups of birds themselves bore to each other. Finally these

orders were surrounded by broken lines in such a manner as to

show the kinships and connections of the subclasses found in

Seebohm's scheme (oc. cit., PP. 44-48). Sharpe also in his

“Review” gives us an admirable example of the linear scheme

^ E * p - Deo 843

AMILIES AND HIGHER GROUE

c—456.] FA; HES i

Nos. 455-456.]

Coll. Natl.

Fic

g 2 pides emn.

I l Buceros rhinocero

—S h n eros Horn i l,

: : oc

. Skeleton of the Rhi

Mus. No. 18 755.

844 THE AMERICAN NATURALIST. [Vor. XXXVIII.

of classification, and this is rendered vastly more useful from the

fact that many of the characters of the suborders are con-

veniently footnoted in the same arrangement. He also em-

ployed the ‘circle system’ in several plates in his memoir, as

well as a modification of the ‘radiating line system,’ and both

with more or less effect (Plates IX-XII). This plan of pre-

senting group characters is excellent in many particulars, and in

connection with a modified linear scheme it has been employed

by Cope in his Classification of Aves (Amer. Nat., Vol. XXIII,

No. 274, Oct., 1889, p. 869), and also by Hans Gadow (P x,

1889). Still earlier than these it was used by Garrod (P. Z. 5.

1874). i

Turning to the plan of the ‘Phylogenetic Tree, it has,

among other authors, notably been employed by Reichenow

(Vögel. der Zoologichen Gärten, 1882) and Max Fiirbringer

(“ Untersuchungen zur Morphologie und Systematik der Vogel” :

‘ Bijdragen tot de Dierkunde, Amsterdam, 1888. Tafln XXVI,

XXVII) Reichenow’s * Tree " is a very crude representation,

and needs no special description in this place. It has, however,

an historical interest for us, and a good copy of it may be found

in Dr. Sharpe's ‘Review’ on page 23. A far more elaborate

and widely known representation is Fürbringer's ‘ Phylogenetic

Tree of Birds.’ This brings up the main stem from the rep-

tilian root-stock, from which, as it ascends, are thrown off the

now dead limbs of a number of more or less known or even

hypothetical groups of avireptilian or reptilioavian forms. Far-

ther up, more extensive branches carry out the evolution of the

main ostrich types, and soon above these the generous and com-

plete development of the tree,— its many more or less closely

dividing and branching limbs, boughs and stems, showing all the

main modern or existing groups as they are supposed to have

been given off from each other, or else to have arisen from the

mam trunk. This tree is horizontally divided by imaginary

planes at certain points, dividing it into upper, lower and middle

sections or horizons. Viewing the projections of these ver-

tically we have the circular sections of the various branchings

ribet, bs us and these will again give the relations to

€ various groups. Three such projections are

Nos. 455-456.] FAMILIES AND HIGHER GROUPS. 845

obtained, which Fiirbringer has represented for us upon three

plates. The horizontally divided stems produced by these cut-

ting planes are also grouped by a system of three kinds of

6. — Skeleton of the Red-billed Toucan, Rhamphastus carinatis

Mus. No. 17,315

FE ta t eost `

surrounding lines which associate, in the opinion of this —

—

aut] i . : >j :ucl Ir as to

authority, the minor and major groups in such a Manner à

846 THE AMERICAN NATURALIST. [Vor. XXXVIII.

have them furnish the requisite data upon which the linear

scheme of this classification has been based. Without any man-

ner of doubt this is the most elaborate provisional scheme of

the classification of Birds extant, and beyond all question, in

most particulars, the relationships of the groups as therein rep-

resented are correct, and, as far as possible, convey to our minds

what has taken place in their evolution.

In setting forth a scheme of classification, the principal objec-

tion to the linear method is that it does not properly represent

the branching or the derivation of new groups in time as the

taxonomer has them in his mind. It brings some groups too

close together, throws others too far apart, and in some instances

in the lineal descent interpolates groups among others in such a

way as to either do violence or misrepresent the true affinities

and relationships of the forms as they exist or have existed in

nature. By the employment of the phylogenetic tree we obviate

a great many of these objectionable features so dangerous to

the credit of a natural classification. Still notwithstanding all

this, the “near method has been adopted in the present memoir,

and for the reason that it is believed that the time is not yet at

hand again, when the making of another elaborate phylogenetic

tree for birds would be considered justifiable as we are still in

the provisional stage. Therefore it has been adopted here.

Further, it has been thought better to weave into this scheme

such fossils of extinct types and species of birds as have come to

hand, and this has likewise been done. For the rest, I have

been influenced in the classification here set forth by my having

read the literature of ornithology since boyhood; by my having

studied everything that pertains to birds for a period extending

over thirty years. This study has been constant, and has been

undertaken in the field, in the closet and in the dissecting room.

It has been devoted to every department of ornithology and of

ornithotomy and morphology.

These studies have been supplemented by similar studies in

the structure and habits of many mammals, reptiles, batrachians,

fishes and other groups. These latter researches have convinced

me that to render the matter of classification more homogeneous

with respect to the vertebrate in general, or in fact to express

Nos. 455-456] FAMILIES AND HIGHER GROUPS. 847

the conditions as they really occur in nature, the Class Aves is

susceptible of being divided into but zwo orders, — the first or

Order I, including the SauruR#, and the second, or Order II,

including all other bird-forms, whether extinct or existing, that

do not belong to Order I. This Order II is known as the

ORNITHUR&. There can be no possible doubt but what in the

early history of birds these two orders arose from the same gen-

eral ancestral stock, but during the geologic ages that have fol-

lowed since, myriads of species have become extinct and the

remains of this vast host have never, save in a few isolated

instances, ever been found or seen byus. Hence the profundity

of the gap now existing between the Saururz and the Ornithurze.

As wide as this gap is, however, the discovery of a very few of

the extinct and intermediate types would tend to greatly lessen

its width. Nevertheless, we must classify the forms as we have

them, and as we find them, and to do this consistently we must

recognize the two orders aforesaid.

Fürbringer in his classification considers the Saururz and the

Ornithurze each to represent a subclass. In this I cannot agree

With him. Birds offer no such division, and are, as a matter of

fact, too homogeneous in their structure to admit of it.

The Saururze, as is well known, are at present represented by

that unique, if it be unique, form Archzeopteryx of the family

Archzopterygide. There is every reason to believe that there

once existed higher divisional groups of this family, therefore

the order Saurure is here subdivided into the supersuborder

Archornithiformes and the suborder Archornithes (see “A

Classification of Birds," postea). As thus created this order is

capable of admitting into it any other fossil genus or genera of

birds allied to Archaeopteryx, whether they come from the

Jurassic age of the Mesozoic epoch of Bavaria or from any other

geologic horizon in any other part of the world. So far as 1s at

present known they represent the oldest avian types in the hands

of science, | i

The classification of birds set forth in the present memoir

carries the arrangement down to include the families only, while

for the higher groups, intermediate between the order and the

family, I employ the superfamily, the suborder and the super-

548 THE AMERICAN NATURALIST. [Vor. XXXVIII.

suborder. To my mind such divisions can be very profitably

employed to express the normal relationships of birds as they

exist, and the relative nearness to each other of the various

groups as compared with other natural assemblages of the

Vertebrata.

Passing now to the consideration of the second order of birds

(Order II, Ornithuree) we meet at first with the array of the

so-called ostrich forms it contains, or supersuborder II. These

represent some of the lowest types of existing bird forms, and a

number of extinct species allied to them have been discovered.

It will not be necessary to discuss the systematic position of the

Dromzognathze (supersuborder II) for the reason that I have

already recently done so in a paper published in 77e American

Naturalist (Vol. XXXVII, No. 433, January, 1903, pp. 33-64:

2 figures), and this likewise applies to the supersuborder III, the

Odontoholcee. The Odontoholcz are placed next in order after

the ostrich-forms not for the reason that they possessed anything in

their osseous systems that in any way connected them with the

Dromzognathz, but because they represent the archaic ances-

tral stock from which has been derived the existing Supersub-

order Colymbo-Podicipitiformes (IV) containing the loons, divers,

and grebes, and these last in the matter of organization stand

among the lowest of known types of modern birds.

This group has been thoroughly treated in my published

papers. A few of the more important groups, however, are still

Ih Press, as for example the “Osteology of the Accipitres "

(Carnegie Museum) ; « Osteology of the Anseres” (Carnegie

Museum) ; and the “Osteology of the Lariformes ” (complete

In manuscript). I now offer my scheme for A Classification of

Birds. This classification requires in some of the groups illus-

trations in the way of bird skeletons that I have heretofore been

unable to publish, and which will throw additional light upon

the subject. These have been kindly photographed for me at

the U. S. National Museum at Washington, and reproductions

of those photographs are herewith presented, with the necessary

descriptions,

Nos. 455-456.] FAMILIES AND HIGHER GROUPS.

A CLASSIFICATION OF BIRDS.

PHYLUM : —

Vertebrata.

BRANCH : —

Craniata.

TRIBE : —

Sauropsida.

CLASSES : —

Reptilia.

Aves.

Criass AVES.

Order I. SAURUR.

Supersuborder i. ARCHORNITHIFORMES.

‘Suborder I. ARCHORNITHES.

Family E Archeopterygide.

Order II. ORNITHUR.

Supersuborder Ik DROMOGNATHE.

Suborder H. STRUTHIONITHES.

. Family I. Struthionidæ.

Suborder III. | RHEORNITHES.

Family E Rheidz.

Suborder IV. CASUARIORNITHES.

Family I Dromaiide.

Hu Casuariidæ.

III. Dromornithide.

Suborder We DINORNITHES.

Family I Dinornithidæ.

Suborder VL ÆPYORNITHES.

Family T Æpyornithidæ.

Supersuborder III. ODONTOHOLC#.

Suborder VII. PyGOPOFORMES.

Superfamily L Hesperornithoidea.

Family I. Enaliornithide.

T. Hesperornithide.

849

850 THE AMERICAN NATURALIST, (Vor. XXXVII].

Supersuborder

Suborder

Superfamily

Family

Superfamily

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Superfamily

Family

Supersuborder

Suborder

F amily

IV.

VIII.

II.

Stercorariidae.-

COLYMBO-PODICIPITIFORMES.

PYGOPODES.

Podicipoidea.

Podicipidz.

Urinatoroidea.

Urinatoride.

APTENODYTIFORMES.

IMPENNES.

Spheniseidze.

Cladornithidze.

PROCELI ORMES.

TUBINARES.

Procellariidæ.

Puffinidæ.

Pelecanoididæ.

Diomedeidæ.

PELECANIFORMES.

STEGANOPODES.

Pelecanoidea.

Pelecanidze.

Pelagornithide.

Phalacrocoracidze.

Odontopterygide.

Anhingidze.

Sulidze.

Phaéthonitoidea.

Phaéthontidae.

Fregatoidea.

Fregatidze.

IcHTHYORNITHIFORMES.

ICHTHYORNITHES.

Ichthyornithidze.

A patornithidze.

LARIFORMES.

LONGIPENNES.

Rhynchopide.

Laridz.

Nos. 455-456.] FAMILIES AND HIGHER GROUPS.

Suborder

Family

Suborder

Family

Supersuborder

Suborder

Family

Superfamily

Family

Suborder

Family

Superfamily

Family

Supersuborder

Suborder

Family

_ Supersuborder

Suborder

Superfamily

Family

Superfamil y

; Family

e Superfamily

| Family

XIV.

ALCA.

Alcide.

CHIONIDES.

Chionidide.

CHARADRIIFORMES.

LIMICOLÆ.

Charadriidæ.

Arenariidæ.

Hæmatopodidæ.

Aphrizidæ.

Scolopacidæ.

Phalaropodidæ.

Recurvirostridæ.

Jacanoidea.

Jacanidæ.

CURSORÆ.

Thinocoridæ.

Dromadidæ.

Glareolidæ.

Cursoriidæ.

Otidoidea.

(Edicnemide.

Otididz.

STEREORNITHIFORMES.

STEREORNITHES.

Phororhacidz.

GRUIFORMES.

GRUES.

Gruioidea.

Gruidea.

Psophiide.

Cariamoidea.

Rhineko

Mesitidæ. mM

Aptornithide. p. : : » MA is RU.

851

852 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Supersuborder

Suborder -

Superfamily

Family

Superfamily

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Suborder

Family

Suborder

Family

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Suborder

Family

Supersuborder

XIII.

XX.

RALLIFORMES.

FULICARLE.

Heliornithoidea.

Heliornithidz.

Ralloidea.

Rallida.

Aramidae.

APTERYGIFORMES.

APTERYGES.

Apterygidae.

GALLIFORMES.

HeEmIPopI.

Hemipodide.

CRYPTURI.

Crypturidz.

GALLINA.

Megapodide..

Cracide.

Phasianidz.

Tetraonidz.

Odontophoridze.

Numididz.

Meleagridz.

OPISTHOCOMI.

Opisthocomide.

PTEROCLIDIFORMES.

PTEROCLETES,

Pteroclididze.

COLUMBIFORMES.

COLUMBÆ.

Treronidæ.

Columbidæ.

Peristeridæ.

Gouridæ.

Didunculidæ.

Dipr.

Dididz.

PALAMEDEIFORMES.

Nos. 455-456.] FAMILIES AND HIGHER GROUPS.

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Superfamily

Family

Superfamily

Family

Supersuborder

Suborder

Superfamily

Family

Superfamily

Family

XXIX.

PALAMEDEJE.

Palamedeide.

ANSERIFORMES.

ANSERES.

Gastornithidze.

Anatide.

PHOENICOPTERIFORMES.

PHCENICOPTERI.

Palzeolodidze.

Phoenicopteridae.

PELARGIFORMES.

HERODIONES.

Ibididz.

Plataleidz.

Ciconiidz.

Scopide.

Ardeidz.

Balzenicipitide.

ACCIPITRIFORMES.

ACCIPITRES.

Falconoidea.

Serpentariidz.

Falconidez.

Milvidz.

Pandionidz.

Vulturide.

Cathartoidea.

Cathartidae.

PsITTACIFORMES.

PsITTACI.

Psittacoidea.

Nestoridz.

Loriidz.

Cyclopsittacidz.

Cacatuidz. :

Psittacidae.

Stringopoidea.

Stringopidee.

854

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

Suborder

Family J.

Suborder XXXVIII.

Family

Supersuborder

Suborder

Family

"Suborder

Family

Supersuborder

Suborder

Family

Suborder

Family

Suborder

Family

Suborder

Family

Suborder

Family

Suborder

Family

Supersuborder

Suborder

Family

Supersuborder

Suborder

Family

XXIV.

XXXV.

II.

XXV.

XXXVI.

XXXVII.

THE AMERICAN NATURALIST. [Vor. XXXVIII.

STRIGIFORMES.

STRIGES.

Bubonide.

Strigidz.

CAPRIMULGIFORMES.

STEATORNITHES.

Steatornithide.

PODARGI.

Podargide.

CAPRIMULGI.

Caprimulgide.

CORACIIFORMES.

LEPTOSOMATI.

Leptosomatidae.

CORACLE.

Coraciidae.

HALCYONIFORMES.

HALCYONES.

Dacelonidz.

Alcedinidz.

BUCEROTES.

Bucerotidz.

Upup2.

Upupide.

Irrsoride.

MEROPES.

Meropide.

Moworir.

Momotidze.

Topi.

Todidæ.

TROCHILIFORMES.

TROCHILI.

Trochilidez.

J ACAMARIFORMES.

GALBULJE.

Bucconide.

Galbulidz.

Nos. 455-456] FAMILES AND HIGHER GROUPS.

Supersuborder XXX.

Suborder XLIX.

Family I.

Supersuborder. XXXI.

Suborder oF

Family I.

Suborder LL

Family L

Supersuborder XXXII

Suborder Lil:

Family Li

Supersuborder XXXIII.

Suborder LII,

Family le

Suborder LIV.:

Family li

Suborder LV.

Family I.

Supersuborder XXXIV.

Suborder LVI.

Family I.

Supersuborder XXXV.

Suborder LYH.

Family L

Supersuborder XXXVI.

Suborder LVIII.

Family Ll.

Supersuborder XXXVII.

Suborder LIX.

Family m

Supersuborder XXXVIII.

Suborder LX.

Family L

Supersuborder XXXIX.

Suborder LXI

Family L

II.

. TROGONIFORMES.

TROGONES.

Trogonidze.

COCCYGIFORMES.

MUSOPHAGI.

Musophagidæ.

COCCYGES.

Cuculidæ.

COLIFORMES.

PAMPRODACTYLÆ.

Coliidae.

PICARIFORMES.

CAPITONES.

Capitonidæ.

RHAMPHASTIDES.

Rhamphastidæ.

INDICATORES.

Indicatoridz.

PICIFORMES.

Pict.

Picidz.

CYPSELIFORMES.

CYPSELI.

Macropterygidae.

Cypselidz.

EURYL/EMIFORMES.

EURYLÆMI.

Eurylæmidæ.

MENURIFORMES.

MENURI.

Menuridæ.

ATRICHONITHIFORMES.

ATRICHONITHES.

Atrichonithidz.

PASSERIFORMES.

MESOMYODI.

Pteroptochidz.

Conopophagide.

855

856

THE AMERICAN NATURALIST. [Vor. XXXVIII.

Suborder

Family

XV.

XVI.

XVII.

XVIII.

XIX.

XX.

XXI.

XXII.

XXIII.

XXIV.

XXV.

XXVI.

XXVII.

XXVIII.

Formicariidae.

Dendrocolapide.

Tyrannide.

Pipridz.

Cotingide.

Phytotomide.

Pittidae.

Philepittidae.

Xenicide.

ACROMYODI.

Hirundinide.

Muscicapidee.

Campophagidae.

Pycnonotidze.

Timelidz.

Mimide.

Troglodytide.

Cinclide.

Turdidz.

Sylviide.

Vireonide.

Ampelidz.

Prionopide.

Lanide.

Sittidae.

Paride.

Zosteropidz.

Dicæidæ.

Nectariniidæ.

Meliphagidæ.

Certhiidæ.

Mniotiltidæ.

Motacillidæ.

Alaudidæ.

Fringillidæ.

Drepanidæ.

Cærebidæ.

Tanagridæ.

os. 455-436] FAMILES AND HIGHER GROUPS. 857

Family XXIX. Ploceidae.

XX. Icteridze

XXXI. Oriolidz.

XXXI. . Dicruride, Er |

"m MANE Eulabetidae | Do f

XXXIV. Sturnide. ee

XXXV. Paradise.

XXXVI.. Comi

OBSERVATIONS ON HEARING AND SMELL IN

SPIDERS.

ANNIE H. PRITCHETT.

OnLy a small amount of previous work has been done upon

the senses of hearing and smell in spiders. :

According to Campbell ('80) spiders are well provided with

the means of feeling the slightest movements of their webs or

other near objects. On their legs are long, slender, silken hairs

which differ from other hairs in that they are attached to a disc

on the integument.

Dahl (83) found these hairs to vibrate to the tones of a violin

and designated them as auditory hairs. Later (84) he distin-

guished certain of these as organs of smell and attempted a

Classification of spiders according to the distribution of the

various hairs.

Bertkau (85) compared the so-called taste organs on the ends

of the palps with the flask-shaped bodies on the antenna of ants

and spoke of them as organs of smell.

J. W. and E. S. Peckham (87) found that with three excep-

tions all the spiders experimented upon by them gave responses

to strong smelling substances, but the Epeiridee alone seemed

capable of hearing the vibrations of a tuning fork. Orb-making

Spiders were the most sensitive to these vibrations, while those

that make no web gave not the slightest heed to the sounds,

and in the former the sense seemed to be distributed generally

over the whole epidermis.

Wagner (88) insisted that the so-called auditory hairs are

only capable of perceiving tactile sensations.

Gaubert ('9o) considered the lyriform organs as organs of

hearing.

exas, No.

! Contributions from the Zodlogical Laboratory of the University of T ,

59. A thesis submitted to the Faculty of the Department of Literature, Science

and Arts of the University of Texas for the degree of Master of Science.

859

860 THE AMERICAN NATURALIST. [Vor. XXXVIII.

McCook ('90) concludes that the senses of smell and hearing

are very rudimentary and are distributed over the entire body.

They are located in the delicate hairs which constitute the cov-

ering and armature, so that the nervous system receives through

these organs or appendages impressions that may be considered

analogous to hearing and smell in the higher animals. Further,

the sense of hearing can scarcely be distinguished from that of

touch as it is known to us. He thinks that even in the orb-

weavers there is no true sense of hearing, but that the web fila-

ments transmit the vibrations and the sensation is tactile rather

than auditory.

Pocock ('93) found spiders sensitive to heat, provided with

acute sense of taste, with defective sight and no hearing what-

ever.

Dahl (:04) designates the auditory hairs as trichobothria and

makes an elaborate classification of suborders according to their

distribution.

The two species used by me for experiment are Geolycosa

texana Montg. and Pardosa mercurialis Montg. and are found

in great abundance in Austin, Texas, and its vicinity. The

large Geolycosa terana lives in underground holes of 4 to 1}

inches in diameter and digs its home on creek banks or in the

fields wherever the ground is comparatively clear except for a

close sod of native grass. Pardosa mercurialis has been found

mainly under small surface stones along the banks of streams

or in the dry beds during droughts, but sometimes occurs in

great abundance far from water.

The specimens used for the observations were kept in cages

made by fastening glass plates together at their edges with

gummed linen, as described by Montgomery (: 03).

I. EXPERIMENTS ON HEARING.

The apparatus used for these experiments were tuning forks

of 128, 256, 320, 384 and 512 vibrations and a specially pre

pared, isolated cage. The cage was made of three glass plates

fastened with summed linen and covered with black paper

Nos. 455-456] HEARING AND SMELL IN SPIDERS. 861

except on the bottom. A small hole was left in the paper of

one side to admit light and one in the top for observations.

The bottom was of mosquito netting which served to admit

the sound waves from below. The cage, supported at the three

corners by wooden posts 10 cm. high, rested on a wooden float,

the float in a stone jar filled with water and attached to it by

rubber bands. A layer of sawdust three inches thick isolated

the jar and stool upon which it stood from vibrations from

without. The apparatus, though placed in a quiet, darkened

corner of the laboratory, was used only when there were as

few outside disturbances as possible.

a. Geolycosa texana Montg.

? No. r3 was placed in the hearing cage and left unmolested

one hour. At the end of that time successive trials were made

With tuning forks of 128, 256, 320 and 512 vibrations, but no

response whatever was obtained. At two later dates the

experiments were repeated but with no more definite results.

The same results were obtained with three other females, in

each case using all the tuning forks. Also a male, after

remaining quiet one hour in the hearing cage was tested with

all the forks but gave no response.

b. Pardosa mercurialis Montg.

? No. 5 was placed in the hearing cage and after an hour's

intermission was tested. with the forks but remained perfectly

quiet. She was left in the cage over night and tested next

morning but gave no response.

? No. 6 was tested by three separate series of experiments,

the first after two intervals of an hour and the third after a half

hour had elapsed, but she did not respond to.any of the sounds.

During the first series she cleaned her palps and legs in the

normal way as if nothing unusual were happening. No

responses were obtained on similar experiments with two

other females and a male.

At another time a steel bar which gave a pitch of approxi-

862 THE AMERICAN NATURALIST. [Vou. XXXVIII.

mately 2300 vibrations was suspended above the hearing cage

and struck repeatedly with a metal hammer. Eight fresh speci-

mens were placed successively in the cage but no spider made

any motion that could be interpreted as a response to the

sound.

Again, a mason’s trowel was held near the cage and struck

several times with a metal bar producing a loud, crashing noise,

but the spiders made no motion whatever.

2. EXPERIMENTS ON SMELL.

These experiments were undertaken to ascertain first whether

the spiders possessed the sense of smell and second to localize

the same, if possible, in case it was found to exist. The experi-

ments were made mostly upon Pardosa mercurialis.

Two classes of the essential oils were used as tests. Those

of the first class were the non-irritants, lavender, cedar, winter-

green, almond, juniper-berries, bergamot, cassia and cloves; of

the second class, those irritants that will cause sneezing in man

were oils of mustard and black pepper. The results from the

use of the two irritating oils were essentially the same as those

for the stronger oils of the non-irritating class, and the reac-

tions were the same in every case only that they were more

intensified with the oils of stronger odors.

The hearing cage was used for the smell experiments.

Small glass rods were dipped into the oil and the drop that

adhered was held immediately below the cage. In each case

the spider seemed repelled to a greater or less degree corre-

sponding to the strength of the oil, and in no instance was it

attracted toward the stimulus. The males and females gave

exactly the same response, thus showing that neither sex has a

superior sense of smell.

The responses were (1) turning away and moving as far as

possible from the odor, (2) vibratory movements of the palps

and mandibles, (3) raising each leg as the oil is applied beneath

i D some cases all the responses were given to a single

stimulus, in others only one, or a combination of either two.

Experiments were made first on unmutilated specimens to

Nos. 455-456] HEARING AND SMELL IN SPIDERS. 863

ascertain the normal reaction to odors, then upon individuals

variously mutilated in order to localize the olfactory sense.

a. Observations on Unmutilated Individuals.

Lavender.— Tests were made with eight females and seven

males and each responded by running from the stimulus. In

four cases this was accompanied by motions of the palps and

mandibles.

Wintergreen. — Seven females and four males. In five cases

the palps gave definite vibratory responses and in the others the

spiders turned away.

Bergamot. — Six males and six females. Each time the

spiders turned away, and in seven instances responded with

motions of the palps.

Cassia. Six males and six females. All but one turned

quickly away, and eight gave vibratory motions of the palps.

Clove. — Six females and five males. One female gave no

response, six gave movements of palps and mandibles, while ten

responded also by turning away.

Cedar.— Eight females and six males. The responses were

quite indefinite, palpal vibration in two cases, and in four there

was no motion whatever.

Almond.— Seven females. No palpal movements were given

and the spiders turned away indifferently.

Juniper-berries. — Eight females and six males. The re-

sponses were quite slow and consisted generally in turning

away. The palps were moved slightly but not in a vibratory

manner.

Mustard.— Six males and eight females. The responses

were no more decided than those given to strong oils of the

non-irritating class. In only one instance were there definite

palpal movements, and the spiders usually turned quickly away

at first, then later more slowly, as if overcome by the odor.

Black pepper. — Six males and six females. The responses

were quite slow and given after prolonged stimulation. Vibra-

tory motions of the palps were entirely lacking.

864 THE AMERICAN NATURALIST. [Vor. XXXVIII.

b. Observations on Individuals with Palps extracted.

It next seemed desirable to determine the location of the

olfactory sense, and for this purpose the oils that produce defi-

nite results with the unmutilated individuals were used exclu-

sively.

Both palps were extracted with forceps at the femoro-tro-

chanteral joint from twelve females and six males, and after a

week had elapsed experiments were made with three nonrri-

tating oils, lavender, wintergreen and bergamot.

Lavender.— In every case the spider gave definite responses.

to the stimulus. When the drop of oil was held directly beneath

each leg successively, each responded by a slight, quick, upward

jerk. Almost without exception the first pair were raised

highest and held poised in the air several moments or until the

other legs were stimulated. The second pair were not raised so

high as the first, nor the third pair so high as the second, and

rarely ever both at the same time. The fourth pair were usu-

ally raised only a little distance and then drawn up closer to the

body while the spider moved away.

Wintergreen. — The same specimens, twelve females and six

males, were used and practically the same results were obtained.

Bergamot.— A simtilar series of experiments was performed

on the twelve females and responses corresponding to the

above were given, though bergamot appears to offer a weaker

stimulus than the other two oils used.

Cassia.— The palps were extracted from twelve other males.

and six other females and experiments were performed after

twenty-four hours. In only one instance did the spider fail to

respond, all others showing the usual reactions — the first pair

of legs responding most definitely and the others less so. The

spiders usually turned away after a few applications of the

stimulus. :

c. Observations with the first Pair of Legs Removed.

In five females and six males the palps were left intact but

the first pair of legs were similarly removed with forceps. The

Nos. 455-456.] HEARING AND SMELL IN SPIDERS. 865

responses were exactly the same as in the foregoing series

Thus it was shown that the sense of smell is not localized in

the palps nor in any one pair of legs, but that each is capable of

perceiving the stimulus to a certain degree and of giving a defi-

nite response to it.

d. Observations on Individuals with the Sense Hairs removed.

The theory has several times been advanced that the long,

spiny hairs on the legs of these spiders are organs of special

senses, accordingly all these hairs were cut off the tarsi and

tibia of male No. 94 and after twenty-one hours it was tested

with lavender oil. Each leg responded as do those from

which no spines were removed.

Female No. 95 had the hairs cut from the entire leg and each

leg sand-papered, still it gave the normal response to bergamot

twenty-one hours after the operation.

Female No. 96 had all the legs sandpapered, yet after only

an hour’s time it responded normally to bergamot, raising each

leg in succession.

Male No. 97 had the tarsus of each leg removed and the

spider seemed to suffer more than when an entire leg was cut

off. However after twenty-two hours it responded normally to

bergamot, raising each leg as it was stimulated. It was not

considered safe to cut the legs-at any higher joint, but the

experiments seem to indicate that the entire leg possesses the

ability to perceive sensations of odors. '

3. CONCLUSIONS.

1. Neither Geolycosa nor Pardosa respond to tuning forks

Of 128, 256, 320, 384 and 512 vibrations. No responses.

were given to a metal bar of approximately 2300 vibrations

nor to the crashing sound of a metal plate when struck with a

bar. Therefore it is quite probable that these spiders do not

hear at all.

2. It might be supposed that possibly this extreme quiescence

was itself an indication that the spiders perceived the tones and

responded in this way; however the fact that these spiders.

. 566 THE AMERICAN NATURALIST. [Vor. XXXVIII.

remain motionless in their cages for hours together seems to

prove that the sound is not perceived in any measure, or at

least that it produces no response. All parts of the body are

extremely sensitive to touch and the spider responds immedi-

ately if it or the cage comes in contact with the vibrating

tuning fork.

3. Males and females respond exactly alike to stimuli of

smell and hearing.

4. Pardosa responds normally to the essential oils by turning

away, making vibratory movements with palpa and mandibles,

and raising each leg as it is stimulated.

5. Individuals from which the palps have been removed

respond to odors normally with each leg.

6. Individuals with the first pair of legs removed respond

normally with the palps and the remaining three pairs of legs.

7. Spiders that have had the sense hairs cut off or sand-

papered respond as do normal specimens.

8. Specimens with the tarsal joint cut from each leg respond

as usual by raising each leg when stimulated.

9. From the foregoing results it seems quite evident that

the ability to perceive odors is distributed over the whole

integument of the spider and there is no definitely localized

olfactory organ.

The work for this paper has been done entirely under the

direction of Dr. T. H. Montgomery, Jr., and the writer is

indebted to him for many valuable suggestions as well as

sympathetic interest and encouragement.

BIBLIOGRAPHY.

BERTKAU, P.

'85. Ueber die Augen und ein als Gehórorgan gedeutetes Organ der

Spinnen. Situngsb. Niederrhein. Ges. 1885.

CAMPBELL, F. M.

'80. Observations on Spiders. Hertfordshire Nat. Hist. Soc. I.

DAHL, F.

':83. Ueber die Hórhaare bei den Arachniden. Zool. Anz. VI.

Nos. 455-456.] HEARING AND SMELL IN SPIDERS. 867

Danr, F.

':8&. Das Gehór-und Geruchsorgan der Spinnen. Arch. mikr. Anat.

XXIV

DAHL, F.

:03. Ueber das System der Spinnen. Sitzungs-Berichte der Gesell-

schaft naturforschender Freunde. Jahrg. 1904, Ar. 5.

GAUBERT, P.

'90. Note sur les Organs Lyriformes des Arachnides. Bull Soc.

Philom. (8) 1I.

GAUBERT, P.

'92. Recherches sur les organs des sens et sur les systémes, integu-

mentaires, glandulaires et musculaires des appendices des

Arachnides. Ann. Sci. Nat.

McCook, H. C.

'90. American Spiders and their Spinning Work. Vol. II. ' Philadelphia.

MENGE, A.

'43. Ueber die Lebensweise der Arachniden. Neueste Schr. Naturf.

Gesell. 2: 4.

MONTGOMERY, T.

Studies on the idem of Spiders, particularly those of the Mating

Period. Proc. Acad. Nat. Sc. Phila.

PECKHAM, G. W. AND E. G.

'87. Some Observations on the Mental Powers of Spiders. Journ.

Morph. I

Pocock, R.

'93. Further Notes and Observations upon the Instincts of some com-

mon English Spiders. Nature XLIX.

AMITOSIS IN THE EMBRYO OF FASCIOLARIA.

HENRY LESLIE OSBORN. ,

A NUMBER of years ago, while studying the embryology of

Fasciolaria, I noticed evidences of amitosis in three different

locations. As biological opinion has not reached a state of rest

in regard to amitosis and its relation to mitosis, I have felt that

a study of the facts in this case would perhaps prove of interest.

A few brief references to the literature of the subject will show

the unsettled condition of public opinion in this matter. Wal-

deyer ('90) said that “ nuclear division is a single process with

Remak's simple amitotic division as the fundamental form."

Johnson ('92), on the contrary, said that he was “ convinced that

it (amitosis) is not derived from mitosis, and on the other hand

is not a forerunner of the more complicated process.” Johnson

considers it a different type which along with karyokinesis has

been transmitted from the simplest forms of life to the most

highly organized. Nor is there unanimous consent as to the

significance of amitosis. The bulk of observation is to the

effect that mitosis is found in growing (7. €. regenerative) tissue

and amitosis is found in senescent, metabolic or pathologic tissue.

In 1891 Ziegler contended that amitosis is due to senescence or

highly active secretory action in cells, and vom. Rath (94)

espoused this view and defended it strongly. According to

these writers amitosis is never regenerative (i. 4, found in devel-

oping tissue), but seals the fate of a nucleus which can never

subsequently divide mitotically. E. B. Wilson in his work, 77e

Cell, appears to be less favorably disposed to accept this view in

the 1900 edition of his work than in the edition of 1896; in the

1900 edition he says that the view is “too extreme," though

“there can be no doubt but that Flemming's hypothesis in a

general way represents the truth, and that in the vast majority

of cases amitosis is a secondary process which does not fail in

the generative series of cell-divisions."

869

870 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Some observations exceptional to this law may be briefly cited.

Frenzel (85) found in the intestine of certain decapods that the

epithelium is regenerated from basal cells which show all stages

of amitosis, while in other closely related genera the growth is

from mitotic nuclei. Wheeler (89) found that in the early blas-

toderm of Blatta all the nuclei divide amitotically at a certain

stage, and that this is followed later by mitosis. Goppert (91)

found amitosis in the leucocytes of the salamander's liver which

he regards as regenerative. Verson (91) found that in the

testis of the silkworm single large nuclei divide amitotically

while their daughter cells give rise to spermatocytes mitotically.

Meves ('91) found that in the salamander certain of the cells of

the testis divided amitotically in winter and mitotically in sum-

mer. Preusse (95) found mitosis in the youngest egg compart-

ments of the ovaries of hemiptera, which consist of only a few

cells and where active cell multiplication must be taking place.

This writer cites observations of Carnoy and Claus in support of

his claim that amitosis is regenerative in some cases. Pfeffer

(99, Wilson, : 00) found that if *spirogyra be placed in water

containing 0.5—1.00 56 of ether active growth and division con-

tinue, but only by amitosis. If, however, the same individuals

. be replaced in water mitotic division is resumed and an entirely

normal growth continues."

' These various observations upon amitosis all indicate a close

relation between amitosis and mitosis. The fact that nuclei

divide in one way in winter and in the other in summer, that in

one genus one method prevails and in another related genus the

other, and that mitosis can be replaced in the same cell by ami-

tosis under the infiuence of ether, all of them suggest a process

at bottom the same but operating differently under the influence

of different circumstances. If so, we should regard amitosis as

the more primitive mode and mitosis as a more highly special-

ized mode derived from it. And we need not then be surprised

if the more primitive mode should occasionally occur in a place

where on general principles we should look for the more highly

specialized one. Though we generally find cell multiplication to

be associated with mitosis and special metabolic activity to be

associated with amitosis, we might in special cases find that cells

Nos. 455-456.] FASCIOLARIA. | 871

were multiplying by amitosis, the simpler mode having been

adopted for local reasons.

The material on which this paper is based was collected at

Beaufort, North Carolina (Osborn, '85), and preserved in either

cold aqueous corrosive sublimate, picronitric or chromic acid

solution. Sections were cut serially and stained on the slide in

borax-carmine, in the days before iron-haematoxylin had been

Fic. 1, — Section of embryo in plane of the throat, showing the very thin ectoderm, partial en-

doderm and food-ova. Scale o.1 mm., cam. luc. X 60.

discovered. I should have been glad to check these results by

more recent methods had access to fresh material been possible ;

however, the technique is fairly adequate for my purpose, as

shown by the condition of the mitotic nuclei present and many

other delicate cytological details. A brief account of the struc-

ture of the embryo will facilitate orientation, especially as this

form is considerably aberrant. My studies at Beaufort were

broken off before I had gotten the earliest stages. Sections of

my earliest embryo are shown in Figs. ! and 2. It is already

past the gastrula stage, and consists of an extremely attenuated

ectoderm enclosing a great number of small spherules each of

approximately 0.15 mm. in diameter. There is a throat and a

small amount of ectoderm, not nearly enough to enclose the

spherules. The spherules are made up of a small mass of cyto-

872 THE AMERICAN NATURALIST. (Vor. XXXVIII.

plasm and a great many yolk granules enclosed by a sharp cell

wall. In the centre of the cytoplasm there is a nucleus or more

than one. These structures are not the ordinary yolk cells

resulting from segmentation of the endodermal part of the origi-

nal ovum, but according to MacMurrich (87) they are unferti-

lized ova which were discharged into the egg capsule by the

mother at the time of its formation together with the true eggs.

He says *each egg capsule contains a large number, perhaps

two hundred, of ova measuring about 0.25 mm. across ; of these

Fic. 2. — More highly magnified view of part of figure r, scale o.1 mm., cam. luc. X .30.

only four or six ever develop, the rest being swallowed by the

developing embryos and used as food. The non-developing ova

do not ever segment nor push out polar globules, nor do they

break down or disintegrate, but are ingested by the embryos so

that at an early Stage one of these appears to consist of a large

number of ova bound together into a ball" The embryo of

Fasciolaria is thus very unlike that of related prosobranchs such

as Nassa, in which the yolk is a part of the original structure of

the egg cell, and hence part of the endoderm of the embryo,

while in Fasciolaria the egg segments and reaches the gastrula

Stage and then swallows the other eggs itself having no supply

of food. The ova thus swallowed I shall call food ova.

The larval kidneys are conspicuous organs in many proso-

Nos. 455-456.] FASCIOLARIA. 873

branch embryos, but they do not reach such large dimensions or

become such conspicuous objects in any other described forms

as in Fasciolaria. They are called *sub-velar masses" in my

paper of '85, and are shown in Figs. 1, 2, 3, 4, and 5. They

are very early of development and are already conspicuous

organs when the shell gland first shows. The velum subse-

quently arises and spreads out over them. In the earliest

embryos the ectoderm cells of the area on each side of the

mouth undergoes a peculiar change (Fig. 3), the boundaries of

the cells cannot be seen, vacuoles

of various sizes appear in the cyto-

plasm, and some of the larger vacu-

oles push the nuclei aside compress-

ing the nucleus, much as in the fat

cells of vertebrates. This is the

first step in the development of the

larval kidney. The nuclei at this

time still show mitosis, two of the

cells of the figure are in stages of

active karyokinesis, there is no evi-

dence as yet of amitosis. As the

accumulation of material within the

cell goes on, the outer end is pushed

up above the general level of the

ectoderm and becomes the swollen

organ we see. The cells become Fic. 3.— Surface view (tangential section)

of ectoderm at one side of the mouth,

wedge-shaped, each retains its place showing first stages in the formation of

on the ectoderm by its base, the meni Dono

tytoplasm is pushed out into the outer end of the cell, which

becomes rounded there as if through the influence of pressure

from within. The content of the cell is chiefly a material which

in life is fluid and highly refractive, but itis coagulated by pre-

servative reagents; it is faintly stainable and not dissolved by

absolute alcohol or oil. The cytoplasm at the outer end of the

cell (Fig. 4) contains one or more nuclei. The relation of the

cells to the general ectoderm is shown in Fig. 4; the organ is

considerably constricted at its base.

There are no evidences of cell division among the older cells ;

874 THE AMERICAN NATURALIST. [VOL XXXVIII.

additions are evidently made at the periphery of the organ at the

expense of the ordinary ectoderm cells. But though the older

cells are not dividing, there are many of them in which the cyto-

plasm at the outer end contains two nuclei, so that we infer that

nuclear division has been going on. No mitotic figures are to

be found in these cells but there are evidences of amitosis. Fig.

Fic. 4.— Section of the larval kidney ata laterstage than figure 3, showing the ectodermal

cells greatly enlarged by the secreted material, and the cytoplasm occupying the outer end

of the cell, cam. luc. x 211. Scale — `

5 shows three stages of this: in the lower figure the initial con-

striction, in the middle one the completion of the division, and in

the upper one the two nuclei have moved apart.

The physiological significance of this organ is indicated by its-

name. As usual with prosobranchs, the embryo develops inside

an impervious capsule ; consequently it is of great importance

that the wastes from its actively developing tissues be kept out

of contact with them. This is done by intracellular storage.

The barrier of living cytoplasm at the outer end of the cell

keeps the cell the better from disintegrating there and the con-

tained material from escaping. This device reminds one of the

lepidoptera and their scales as reservoirs for waste nitrogen dur-

ing metamorphosis.

Amitosis here is clearly in accordance with the view that

senescence and amitosis are closely related, for these cells are

Nos. 455-456.] "ASCIOLARIA. 875.

plainly reaching the end of their career. Though they are for

the time a part of a very young

embryo, the organ is no more an

integral part of the embryo than

are the embryonic membranes of

arachnids and insects, in which

amitosis is well known to be asso-

ciated with senescence. It is really

merely an embryonic structure.

It is not, however, clear that the .

active secretory function of these M6 5-— Outer ends of three oe

` : same situation as figure 4, showing nu-

cells is a cause of the amitosis, If cleiin different stages of division, cam.

it were we should expect that in T |

cells where it was taking place as actively as in the one in Fig.

3at the bottom of the drawing should show amitosis, whereas

it and the one over it are dividing amitotically.

The food-ova present nuclear appearances of sufficient interest

to deserve mention. In the earliest embryos (Figs. 1, 2) they

are intact and still manifest cell activities of their own notwith-

standing their having been

swallowed as food. Each has

a definite egg-membrane en-

closing a mass of cytoplasm

often located near the surface

of the cell, its branches reach-

ing out among the very nu-

merous grains of yolk. Many

of these ova exhibit peculiar

nuclear activities. In some

one finds a single nucleus

mear the centre of the cyto-

IG. 6.— The nucleus i immediately adjacent lasm of large dem and con-

Fx a me lated con- taining within its membrane

supe Uu. not the more customary chro-

matine network but instead a mass of bubbly material (fig. 6) as

if the chromatine had been converted into a froth. Other cases

are met in which there are numerous small nuclei, as in Fig. 8.

876 THE AMERICAN NATURALIST. (Vor. XXXVIII.

In such cases the centre of the protoplasmic area often contains

a somewhat more deeply stained material not limited by any dis-

tinct boundary from the general cytoplasm, but occupying the

position of the single nucleus of other ova and having the same

size, and the cytoplasm often radiates from this as a centre of

influence. The lesser nuclei do not usually show active division,

but are in the resting stage. They show evidences of disorgani-

zation in various respects. Some are distinctly vacuolated as

the parent nucleus was ; others contain bits of chromatine of no

regular shape or position, resembling very closely the * spore-like

bodies" found by Herrick ('92) in the “degenerating nuclei "

of the yolk cells of the egg nauplius of Alpheus. In a few

instances two of these smaller nuclei are still connected by a

thread of stainable material as in Fig. 7, which I interpret as

the last stage in amitotic division, and an indication of the proc-

Fic. 7.— Nuclei and surrounding cytoplasm, showing the last stage in a nuclear division, cam.

X 600.

ess by which the multiple nuclei of the food-ova have been

formed. The cells in which the nuclei have undergone these

changes are on the road to complete breakdown and these

changes are the last events in their lives. The process is a

Nos. 455-456.] FASCIOLA RIA. 877

futile attempt at segmentation where normally we should find

mitosis, but in this case the cell having the impulse to divide

but being powerless to do so by mitosis falls back on the easier

mode and does so by amitosis. We may call it cell senescence.

The gastrulas before they have swallowed the food-ova are

(according to information and drawings access to which I owe to

the kindness of Prof. MacMurrich) very queer looking objects

on account of the very ample folds of the ectoderm to allow for

the distension which is to follow. One would expect that the

endoderm would be equally so, in order to receive the ova into

an endoderm-lined cavity, but after much study of this point I

am convinced that unusual as it is there is not enough endoderm ~

to enclose these ova, but only a very small amount reaching out

a short distance from the throat in all directions, as in Fig. 2.

Fic. 8.— Multi-nucleate condition of older food-ovum, the nuclei showing vacuolation, cam.

luc. X 600

A study of various series agrees in showing only one very thin

layer the ectoderm in contact with the ova except near the

Stomodzeum. As development progresses the endoderm pushes

Out around the food-ova, at first with very thin flat cells, which

later still become cubical and finally differentiate into the large

and vacuolated definitive cells of the liver, or the columnar cells

878 THE AMERICAN NATURALIST. (Vou. XXXVIII.

of the intestine. During the early part of this process, while

the endoderm cells are still flat or cubical, and before their final

differentiation has taken place, their nuclei exhibit unmistakable

evidences of amitosis. Fig. 9 is taken from such an embryo.

The food ovum, part of which is shown in the figure, is still

intact; it has not yet undergone the breakdown of structure

which is soon to take place. The endoderm is composed of

Fig. 9. — Section vertical to the ecto-

derm and underlying endoderm in Fic. 10. — More highly magnified

very young embryo, scale —.or mm., view of part of fig. 9, X 600.

cam. luc. X 211.

cubical cells in which dividing nuclei are to be seen. The endo-

derm cells do not show any walls at this time between the cells

but has the appearance in this view and in surface views of a

Syncytium. Walls appear at a later date and distinct cells are

clearly seen in older embryos. A more highly magnified view

of the cells of Fig. 9 (Fig. 10) shows nuclei at different stages

of amitotic division. These views are very characteristic and

found in many different places in several series of embryos of

this age. They are further corroborated by flat views from

places where sections are cut tangentially through the endoderm

(Fig. 11), in Which the syncytial character of the layer is

Nos. 455-456. FA.SCIOLARIA. 879

shown. Large nuclei of irregular shape (Fig. 12) are also found

in the endoderm of this stage; they are apparently also forms

of amitotic division. Their exact relation to the more typical

cases could not be determined

from the material which I had.

They are quite commonly to be

found in embryos of a little

older stage but before the endo-

derm has assumed its definitive

form. Nuclei of both these forms

predominate decidedly during the

development of the embryo, but

in addition to them there are

occasional nuclei showing un-

mistakable karyokinetic figures.

The latter are too infrequent to

be regarded as the nuclei from

which the rapidly growing en-

doderm is being chiefly regener-

ated, but there is nothing toshow + Mee exit 3

that they do not give rise to en- ric. i. — Surface view (tangential section) of

doderm cells as well as the ami. Wi, eedem. shoning mle, i dt

totic nuclei. The presence of

the mitotic figure is a guarantee of the adequacy of the tech-

nique, whatever we may think of the amitosis in these cases, at

least it is not possible to dispose of them as artifacts. We

recall in this connection the cases of amitosis in the intestinal

cells of isopods observed by Ryder and Miss Pennington, as

referred to by Conklin ('97). These were shown by him to be

artifacts due to pressure in manipulation and resultant distor-

tion. In that case the amitosis was in adult cells and the pres-

sure was brought about by instruments and not in embryonic

tissue and in a course of nature. The food-ova do not compress

the endoderm, for at the time they are swallowed the layer has

not as yet been formed ; they grow out into the cavity after the

ova have been swallowed. After a study of the available mate-

rial I have not been able to resist the conviction that in Fascio-

. laria we find a true case of embryonic cells multiplying, contrary

880 THE AMERICAN NATURALIST. [Vor. XXXVIII.

to the very general rule, by amitosis. A renewed study of this

and kindred forms with reference to this point is much to be

desired.

These cases of amitosis are sufficiently out of the common

order to require a further study in the light of the general prin-

ciples relating to the

amitosis question. That

of the larval kidney could

perhaps be attributed to

senescence but hardly to

metabolism, since cells

are found dividing by

mitosis after secretory

activity has been going

on to such an extent

that the nucleus is dis-

torted and crowded into

one side of the cell.

The cells after they

have reached a stage in

which amitosis is found

have apparently reached

the end of their career

and do not multiply

Fic. 12. — Nuclei in the TUE MR plasmodial ids cam peines i pe = ue

luc. X 600. "TT are senescent:

It is highly remark-

able that the food-ova nuclei should not divide mitotically, for

the food-ova are to be interpreted as unsegmented eggs. The

divisions of the nucleus in ordinary eggs both in maturation and

segmentation are mitotic, and we are justified in expecting

karyokinesis of the food-ova nuclei. But instead their nuclei

are queer from the first and seem to divide amitotically. This

amitosis cannot be assigned to either senescence or metabolism,

the cells show no signs of secretory activity, and are not senes-

cent. The case of Spirogyra, according to Pfeffer, and of the

salamander, according to Meves, are somewhat analogous in that

Nos. 455-456.] FASCIOLA RIA. 881

in both of them mitosis has been replaced by amitosis through

the influence of an extraordinary circumstance, cold weather or

ether. We know of no parallel to.the case of the food-ova

among the gasteropods. In the trematodes the yolk cells of

the egg are perhaps somewhat similar; their nuclei do not divide

at all.

The nuclei of the young endoderm are also out of the ordinary

course. We should expect them to divide by mitosis. While

in a small percentage of cases they do divide in that manner,

in most cases the indications are strongly in favor of amitosis.

Here neither senescence nor metabolism can be the cause of the

amitosis. The cells have not yet arrived at their definitive

form, much less become old, and secretion has not yet com-

menced ; but, on the contrary, the cells are actively multiplying,

so that they quite contradict the theory proposed by Ziegler and

vom Rath.

We see, then, that while the amitosis of Fasciolaria can be

attributed to senescence in one case, it cannot be attributed to

either senescence or metabolism in the other two cases; and in

the endoderm the amitosis is even regenerative. We have seen

that senescence and metabolism cannot be regarded as causes

in certain other instances of amitosis, such as those reported by

Fresnel, Meves and Pfeffer. Upon the hypothesis that mitosis

and amitosis are independent and non-related processes, cases

of this sort are wholly unintelligible, and consequently indicate

a defect in the hypothesis, if not that it is erroneous. It is

true that we do not find intermediate modes of nuclear division

between mitosis and amitosis, and yet @ priori we should sup-

pose them to be related and mitosis to have been derived from

amitosis, The exceptions that are being found are all of them

cases in which mitosis would be looked for and amitosis is

found. The frequency of mitosis in embryonic tissue does not

prove that embryonic tissue cannot multiply by amitosis ; it

only shows that there is some reason why in most cases it does

not. Since mitosis is such a delicately adjusted and compli-

cated process, may we not suppose that a nucleus to be capable

of it should be in the best possible condition, —that is, young

and vigorous, — and to that end that the surrounding conditions

882 THE AMERICAN NATURALIST. [Vor. XXXVIII.

should be as favorable as possible? And may we not further

suppose that in the presence of conditions lowering the vigor of

a nucleus, even if it were.in a case where we should ordinarily

expect mitosis, it would then revert to the more primitive mode

of amitosis? We are not unfamiliar with the principle of physi-

ological reversion in organs and tissues. Can we extend the

principle to nuclei? If so, all the cases where amitosis occurs

in a place where mitosis is expected to occur would perhaps be

capable of being brought within its scope. Possibly both senes-

cence and secretion as causes of amitosis could be interpreted

in the same way. Thus senescence as a cause of amitosis, as it

undoubtedly is, would be in reality due to the lowered vitality

of the cell consequent on age. Metabolism perhaps could be

supposed to cause it, where there is not at the same time, as in

fat cells, milk cells and others, senescence, through the presence

of the secretion which may exercise an unfavorable influence on |

the nucleus chemically. Pathologic tissues would also find an

easy explanation on this basis to account for the common occur-

rence of amitosis in them. Summer by inducing optimal well-

being of the cell would put it in condition for mitosis ; winter

by lowering its vitality would render it unable to divide by

mitosis but not by amitosis. Pfeffer's observation on the effect

of ether would also fall in line, the cell tending to divide by

mitosis but being prevented by the interference with its powers

consequent on the influence of the ether. In Fasciola the

larval kidney cells while young still divide by mitosis and are

consequently vigorous, notwithstanding the presence of consid-

erable metabolic material, but as they become old they lose this

power and divide by amitosis. The food-ova with their ten-

dency to mitosis are placed in conditions so unfavorable to

their well-being that cell division does not take place at all and

nuclear division is only by amitosis. The deeply staining gran-

ular particles in some of them may be imperfectly formed chro-

mosomes, as if mitosis had been attempted but without success.

A careful cytological study of these food-ova would be very

likely to repay study, as it might throw light on the relation

between the two modes of nuclear division. The endoderm

cells are developing in the amitotic manner and this is the chief

Nos. 455-456.] FASCIOLARIA. 883

mode by which the embryonic gut enclosure is taking place, if

not the only mode. In their case neither senescence nor metab-

olism can be urged as the cause of the amitosis. Under ordi-

nary circumstances we should find the endoderm increasing by

mitosis, but here the mode of growth is very extraordinary, and

the case only finds explanation on the principle I have béen

indicating. I do not know just what particular cause should be

assigned in the case of the endoderm, which is to be considered

as interfering with the mitosis ; perhaps it is the pressure condi-

tions existing in the enteron owing to the presence of such a

large amount of inert and foreign matter; perhaps it is the

absence of the most favorable food for the endoderm, since the

food-ova have not yet begun to be consumed.

Note.— Since revising the last proof of this article, the writer

has read with interest a passage in Dr. Davis' article (Am. Nat.

38, p. 434) containing much the same idea vid.— * It is possible

that direct division in the higher plants is in a sense a reversion

to early ancestral conditions, a reversion ‘hat comes on only when

Jor some reason the normal activities of the germ id are in abey-

ance or have ceased."

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Series.

ZIEGLER, H. E.

'91. Die biologische Bedeutung der amitotische dica cer, im Tier-

reich. Biol. Cent. vol. 11.

BIOLOGICAL LABORATORY OF HAMLINE UNIVERSITY,

SAINT PAUL, MINN.

April 25, 1904.

THE TRANSPLANTING OF TROUT IN IHE

STREAMS OF THE SIERRA NEVADA.

DAVID STARR JORDAN.

THE rivers of California contain four well marked species of

trout: (1) The rainbow trout, Salmo zrideus, in the streams of

the Sierra foothills, and in all the streams of the coast range ;

(2) the steel-head trout, for which the oldest and therefore cor-

rect name seems to be Samo rivularis, rather than Salmo gaird-

neri. This is found in the coastwise streams only, whence it

runs regularly to the sea, the rainbow trout running only when

the sea is conveniently near; (3) the cut-throat trout, Salmo

clarki, found only in the streams of the northern and northwest-

ern counties; and (4) the Tahoe trout, Salmo henshawi, a deriva-

tive of the cut-throat found on the east flank of the Sierras, in

the tributaries of the extinct Lake Lahontan.

Subspecies or variants have developed from these, especially

from Salmo irideus. The typical rainbow trout is found in the

streams about San Francisco Bay. A slight variant called var.

shasta is found in the upper Sacramento and in the northern

Sierras. With it, but rare, is var. s/ozez, the Nissui trout. In

the Kings and Kern rivers is the smaller scaled variety, gilberti,

and from gilberti is descended the dwarf golden trout, var. agua-

bonita, found only in Volcano (Whitney) Creek on the banks of

Mount Whitney, above the high waterfall sometimes called

Agua Bonita. The typical Tahoe trout, Sa/mo henshawi,

spawns in the streams, but in the depths of the lake is a well

marked subspecies óf large size spawning in the lake, called var.

tahoénsis.

The distribution of these species is well made out, but certain

anomalies were noticed, especially in the distribution of the Tahoe

trout. Allthese anomalies have been found to be due to trans-

planting by the hand of man. For the benefit of future stu-

dents of trout I place these facts on record.

885

886 THE AMERICAN NATURALIST. (Vou. XXXVIII.

The Stanislaus, Tuolumne and Mokelumne Rivers flowing

westward from the Sierra Nevada had in the early days no trout

above the falls in their cafions. To these streams the trout

were carried over the Divide, and the Tahoe trout is now abun-

dant in the upper waters of these rivers.

For the details of the transplanting of these trout I am in-

debted to the kindness of Mr. H. S. Blood of Angels, the owner

of Bear Valley, at the head of the north fork of the Stanislaus,

on the “Toll Road” from the Calaveras Big Trees to Carson

Valley.

The earliest plant was made in 18 59, the Tahoe trout having

been taken from Hope Valley, on the east slope of the Sierras,

by the Union Water Company to stock their reservoir at the

head of the north fork of the Stanislaus.

In 1861, Abram Ritchie of Big Trees and John Christy of

Jenny Lind brought Tahoe trout from Hope Valley and planted

them in the head waters of Mokelumne River.

In 1863, H. S. Blood of Angel's and J. C. Curtis of Murphy's

brought Tahoe trout from Wolf Creek, Carson Valley, and

stocked Highland Lakes, at the head of the Stanislaus.

In 1864, O. S. Boardman of Galt took Tahoe trout fróm Hope

Valley, planting them in the Blue Lakes, near the head of the

Mokelumne. At about the same time H. S. Blood, Mark

M'Cormick and Herman Tyrie stocked Highland Creek with

Tahoe trout from Hope Valley.

The middle fork of the Stanislaus was stocked with Tahoe

trout from Walker River, Nevada.

In 1873, Joseph Emery of San Francisco, brought trout from

the north fork of the Mokelumne, stocking the Blue Lakes of

Amador with the Tahoe trout.

The Tahoe trout was planted in Feather River by Mr. Pratt,

founder of the village of Prattsville in Plumas County, the fish

being taken from Truckee River.

The golden trout, agua-bonita, has been taken from Volcano

Creek and successfully planted in Lone Pine Creek, on the east

side of the Sierra Nevada.

The rainbow trout, var. shasta, has been planted by the State

Fish Commission in the Truckee River, where it is now abun-

Nos.455-456] TRANSPLANTING OF TROUT. 887

dant. With it was taken the Sacramento perch, Archoplites

interruptus. The present writer lately obtained a small speci-

men of this species from an Indian ditch at Wadsworth, Nevada.

The eastern brook trout, Salvelinus fontinalis, has been

planted in many California streams, which it seems to find

congenial.

A NEW SPECIES OF DIAPTOMUS FROM MEXICO.

A. 8. PEARSE.

Diaptomus lehmeri, n. sp.

Cephalothorax, broadest at the middle; with four distinct

ally and armed with two

prominent spines. The

domen, armed on each

side with a strong sharp

spine. The first seg-

Pic. -: Juge ai domen is tapering and

meri n. sp. Abdomen about as long as the re-

v sanie mainder of the abdomen.

The second segment is short, being only

one-fourth the length of the third. The

furcal rami are almost twice as long as

broad, and hairy on both margins in the

female (Fig. 1), but on the inner margin

only in the male.

Antenne twenty-five segmented aid

reaching beyond the tips of the furcal

rami. The right male antenna (Fig. 4)

is moderately swollen and its antepenulti-

mate segment bears a pointed curved hook,

which is not as long as the ultimate seg-

ment.

The male fifth feet (Fig. 2) are rather

robust. The first basal segment of the

first segment of the ab-

ment of the female ab- :

sutures. The last segment, produced later-

Fic. 2.— Diaptomus lehmeri

n.sp. Fifth feet of male

X 205.

right foot has a tubercle near its outer margin which bears a

long sharp spine. The second basal segment is almost as

889

890 THE AMERICAN NATURALIST. [Vor. XXXVIII.

broad as long and besides the usual hair on the outer margin

outer distal

angle. The

, second seg-

Fic. 3. — Diaptomus lehmeri ment has a

Ls Fifth foot of female large tubercle

on its poste-

rior surface. The marginal spine

is strong, but slightly curved, longer

than the segment, two-thirds the

length of the terminal hook, and is

roughened on its inner margin.

The terminal hook is strong, arcu-

ate and minutely denticulate at the

middle of its inner margin. The

inner ramus is: one segmented, |

short, (about as long as the first

Segment of the outer ramus) and

hairy at the tip.

In the left male fifth foot (Fig.

2) the first basal segment is armed

with a tubercle and spine like that

of the right. Second basal seg-

ment is three-fourths as broad as

long and bears the usual hair on

its outer margin. First segment

of the outer ramus curved on outer

margin, the inner margin notched

itis provided with a characteristic process

on its posterior surface. There is a small

hyaline projection on the distal margin of

this segment and also on that of the first

segment of the outer ramus. The first

segment of the outer ramus is very short,

being one-third as long as the second seg-

ment and is produced into a tubercle at its

Fic. 4.— Diaptomus lehmeri n. sp. Tip

of right antenna of male X 128.

and ciliated. The terminal segment is triangular and ends in a

blunt process and proximad to this is a slender hair; the

Nos.455-456] WEW SPECIES OF DIAPTOMUS. 891

inner surface is ciliated. The inner ramus is short and hairy

at the tip.

The female fifth foot (Fig. 3) bears a large tapering spine at

the outer distal angle of the first basal segment. The second

basal segment is triangular and provided with the usual lateral

hair. The outer ramus is three segmented. The first and

second segments are about equal in length. The second

segment is tapering and along the center of its inner side is

armed with strong teeth and is usually with two or three small

teeth on the outside. The second segment has a skort spine

above the third segment. The third segment is small and is

armed with a long and a short spine. The inner ramus is not

half as long as the first segment of the outer ramus and is two

segmented. The second segment is armed with two terminal

spines with delicate hairs between them and a lateral spine on

the inner side.

Length: Female 1.69-1.83 mm. Male 1.61—1.67 mm.

The specimens from which this species is described were col-

lected by Mrs. Frank Lehmer, July 22 to August 7, 1904, in

Mexico City, Mexico. It was apparently abundant at that time,

as it occurs in most of the dredgings made.

This species was associated with Cyclops viridis insectus

Forbes, Cyclops albidus Jurine, Canthocamptus sp., and an

ostracod. ;

HYLA ANDERSONII AND RANA VIRGATIPES

AT LAKEHURST, NEW JERSEY.

WILLIAM T. DAVIS.

On the 5th of last September while looking for insects near

a swamp at Lakehurst in the pine barrens of New Jersey, the

writer was fortunate in finding a specimen of the rare tree frog

Hyla andersonii Baird. The frog was in a small oak tree

standing but a few feet from the swamp. At the time Cope's -

work on “ The Batrachia of North America" was published in

1889, but three specimens had been recorded, namely the type

from Anderson, South Carolina; the one collected at Jackson,

Camden Co., New Jersey, in 1863 by Leidy, and the third exam-

ple found by Dr. John E. Peters at May's Landing, Atlantic Co.,

New Jersey on June first, 1888. A record of this last is to be

found in the American Naturalist for January, 1889. In the

American Naturalist for December, 1894, J. P. Moore gives an

account of two of these tree frogs collected in June, 1889, at

Pleasant Mills, New Jersey, and of the many others heard at

the time. The frogs, however, disappeared shortly and no

others could be found on subsequent visits to the locality.

Lakehurst is considerably farther North than the three New

Jersey stations mentioned above. It may be well to mention

at this time that Rana virgatipes Cope, is also to be found at

Lakehurst. The species was originally described from Cape

May County, New Jersey.

According to Cope, in Myla andersonii, * The green of the

back and extremities is everywhere margined with pure white,

except posteriorly on the femur and tibia, and anteriorly on the

former where saffron takes its place.’ My living specimen,

however, differs from this description and the colored figure, by

having the band of pure white extend along the end of the

body and a short distance on the posterior margin of each

femur.

New BRIGHTON, STATEN ISLAND, N. Y.

. 893

NOTES AND LITERATURE.

PSYCHOLOGY.

Natural Science and Psychology. — For the convenience of

those who do not have access to his * Grundzüge der physiologischen

Psychologie" Prof. Wundt has reprinted under the above title the last

two chapters of the third volume of his great work. In the space of

a hundred pages there is given a clear statement of what the author

believes concerning : (1) The logical foundations of natural science,

(2) Mechanics and energetics, (3) Mechanism and vitalism, (4)

Causality and teleology of the psychophysical life-processes, (5) The

concept of mind, (6) The principles of psychical causality.

Of special interest to biologists are the discussions of mechanics and

energetics, and of the mechanistic and the vitalistic theories. Such

a book is valuable to most natural sciences in that it serves to call

attention to the too much neglected presuppositions on which special-

istic work in the natural sciences rests, as well as to the principles of

the sciences. Wundt writes in a patient, plodding, persistent man-

ner; his sanity is all the more evident because of the lack of any

brilliancy of treatment.

In view of the appearance of this reprint it is worth while to notice

the interest, especially among the German biologists, in the psychic

«as an elemental factor in nature,” as Driesch in a recent book has

stated it. Whether one agrees with Wundt's conception of the rela-

tions of the physical and the psychical, with Driesch's, with Ost-

wald's, with Schneider's may not seem of much consequence to

most students of natural science; nevertheless, there can be no

doubt that only those who hold theories and see or seek principles

can do more than assemble facts whose meaning is clear only in the

light of the generalizations which are called laws. No one can study

Wundt's book without gaining an interest in the various forms of

organic processes that the mere search for facts would not give.

To state the content of the book would defeat my purpose to arouse

an interest which may lead many to the work itself.

ROBERT M. YERKES.

1 Wundt, Wilhelm. Maturwissenchaft und Psychologie. Leipzig, 1903,

8vo, 126 pp.

895

896 THE AMERICAN NATURALIST. [Vou. XXXVIII.

* The Criminal Classes.” ! — Rev. D. R. Miller, D.D., has been

chaplain of the Ohio Penitentiary and superintendent of the Girls'

Industrial Home of Ohio. In * The Criminal Classes. Causes and

Cures " he has collected material of various sorts connected with his.

service in these positions. -

At the beginning; near the middle, and near the end of the volume,

are chapters briefly treating in a general way the extent of crime, its

cost to the community, causes of crime, and methods for the preven-

tion of crime. "These discussions are somewhat vague and common-

place and are not fully adjusted to the results of recent scientific

study. "This sentence from the chapter on “ Innate and Psychologi-

cal Causes" may serve as an illustration of the nonagreement with

conceptions of present day physiological psychology: * Regarding

moral conduct, in this class there seem to be indications of a broken

or missing link between the brain tissues and the moral monitor."

There are evidences also of lack of familiarity on the part of the

author with recent literature in penology. For instance, he writes,

" As agencies in the prevention of crime and the reformation of young

criminals, too much importance cannot well be attached to the pri-

vate and public institutions denominated ‘Child-saving institutions,”

‘Schools for the little ones,’ and ‘ Homes for children.” He then

fails to mention the movement now in force in many parts of this

country, which recognizes that care for children in institutions has

not generally prepared them well for life outside of institutions, and

which substitutes for the institutional method the methods of pro-

bation - and placing out, that is, the supervision of the rearing of

delinquent and dependent children, while they live as members of

their own or other families. Many of the figures on which arguments.

are based are given without dates but as taken from books and

reports now many years old. On the whole these general chapters.

cannot be said to give an adequate summary of facts known and

theories held to-day on the topics which they treat.

A few chapters offer meagre outlines of the careers of noted crimi-

nals. Others give brief accounts of the lives and crimes of Ohio

convicts. “In the introduction, however,” as Dr. Miller says in the

preface, * of these narratives, sketches, and illustrations, special care

has been exercised to avoid sensational details of the crimes alleged

because it is the opinion of the author that all such details serve as

! Miller, D. R. The Criminal Classes. Causes and Cures. Dayton, United

Brethren Publishing House, 1903. 8vo, ix + 227 pp., portraits. -

Nos. 455-456.] | WOTES AND LITERATURE. 897

factors to turn into like crime those who may read them." Some

of these chapters close with reflections, which, it must be said are in

part obvious, in part without special applicability to the cases to

which they are attached.

About half the chapters of the book are devoted to incidents that

occurred in the author's experience as chaplain, and to writings by

prisoners, in prose and poetry, on religious and other topics, some

apparently sincere, others with a suggestion of cant. In religious

work for the prisoner results can doubtless be gotten easiest by

appeals which recall religious instruction received during childhood.

At the same time nothing in a man's work as chaplain forces him to

adapt himself to changes in religious thought in the world at large.

It is natural, therefore, that the weight of the religious interest of the

author should be, as it is, such as belongs with religious beliefs of a

conservative type. ;

There is an introduction by Ex-Governor Foster of Ohio. A chap-

ter by Professor Krohn of the University of Illinois compares crimi-

nals in whom strange combinations of traits are found with cases of

double personality, but ten to call attention to some important dif-

ferences.

Apparently Dr. Miller is a man who has done earnest and helpful

personal work with the criminals placed in his care, but who in his

devotion to his own special task has failed to gain the broad acquaint-

ance with current thought in criminology and related subjects which

would be needed for writing a very useful book on the causes and

cures of crime.

ZOOLOGY.

Fixation of the Eggs of the Crab. — Dr. H. Charles Williamson,

in the 22d Annual Report of the Fishery Board of Scotland, adds con-

siderably to our knowledge of the life history of the shore crab of

Europe, Cancer pagurus. Possibly the most interesting is his discov-

ery of the way in which the eggs are fastened to the swimmerets.

These, like other eggs, have double envelopes, the outer chorion and

the delicate inner vitelline membrane, and between these a small

perivitelline space. When the eggs are extruded from the oviducts

they pass into the brood chamber between the abdomen and thorax

898 THE AMERICAN NATURALIST. [Vor. XXXVIII.

and here are held.in position by the exopods of the swimmerets

while the endopods, covered by slender filiform hairs, extend into the

mass of eggs. The pleopoda have powers of motion in two planes,

one fore and aft, the other oblique. By these motions the ends of

the hairs are brought in contact with the eggs which are thus pene-

trated by them, the hair passing in and out again. The egg itself

rotates and escapes injury. In this way numbers of eggs are impaled

or skewered on each hair. As development proceeds the chorion

stretches and gives rise to a stalk, which increases in length with

time. This process is also aided by the perivitelline fluid, which is

adhesive and sticks the chorion to the hair. After the egg is hatched

the egg shells and their stalks are cast off by molting the integument,

the outer layer of the hairs, together with the adhering shells being

cast with the rest of the ‘skin.’ The eggs are fastened only to the

hairs of the endopods, as these alone are smooth and filiform. The

hairs of the exopods are plumose or serrate, a condition which pre-

vents their perforating the eggs.

Gardiner's Maldives.!— The third part of the second volume of

Gardiner’s Report maintains the unusually high quality of its prede-

cessors. It contains six papers, of which three are written by mem-

bers of the expedition and the three others by well known specialists.

The Isopoda are treated by Stebbing in 23 pages. Thirteen

species are described, representing eight families. While some are

from the sea at moderate depths, two (a Corallana and a Limnoria)

are found in rotten wood in the lagoon, one (Cirolana) lives in tenta-

cles of a large tubicolous polychaete. A Cymothoa lives on the gills

of a parrot-fish. Tylokepon, n.g., is a bopyrid living on the gills of.

crustacea. Finally, a sphaeromid (Exosphzroma) and a Ligia are

representatives of two semi-terrestrial groups. :

The Hydromeduse are treated by Browne. The discussion of

the Anthomedusz leads to a revision of the family Williade with

two genera, Willia and Proboscidactyla, of which the latter only is

in the collection from the Maldives. It occurs also on the east

coast of North America and has several species notable for their

production of medusa buds. Among the Leptomedusz is a five-

rayed representative of Mayer's genus Pseudoclytia which Browne,

following Mayer, believes to have arisen as a sport. The Tracho-

medusz are treated in synoptic fashion. A new species and a new

1 Gardiner, G.S. Th Fauna and Geography of the Maldive and Laccadive

Archipelagoes, etc. Vol. II, part III, pp. 589-698, pls. 35-48. :

Nos. 455-456.] NOTES AND LITERATURE. 899

genus are created. Hermaphroditism is described in one case. One

Narcomedusa, a long known species of the Indian Ocean, is described.

Five species of Siphonophores were collected. The author states that

he could find no copy of the Mark Anniversary Volume in London.

Let him apply to Henry Holt & Co., New York.

The reptant decapods of the suborders Hippidea, Thalassindea,

and Scyllaridea, are treated by Borradaile. The first two groups are

taken between tide marks, the latter in the reefs or on sandy bottom.

There are enumerated two albuneids and three hippids (Remipes),

one axiid, a gebiid, four 'callianassas, two palinurids and one

Scyllarus.

The Madreporaria are treated by Gardiner, who lays especial stress

on their variation, distinguishing vegetative, continuous, and discon-

tinuous or specific. This extensive paper is accompanied by well

executed half-tone plates from photographs of the dry corals.

The Antipatharia are described in a few pages by Forster Cooper.

They are chiefly Indian Ocean species.

The Arachnida of the archipelagoes, treated by Pocock, raise the

question of their origin. “Some of the species, such as Isometrus

europzeus, Heteropoda regia and Uloborus geniculatus, which fre-

quent human dwellings, have doubtless been introduced by human

agency; but it is probable that the ancestors of the majority of the

Spiders reached these islands on floating gossamer threads." “The

presence of only one species of Scorpion [Isometrus europzus], and

that a form notoriously liable to dispersal by man's instrumentality,

suggests that, unless in very remote times, there has been no connec-

tion between the archipelagoes and the mainland of India . . .. and

this is further borne out by the almost complete specific identity

between the two faunas." f

CRD

CLIMATOLOGY.

Ward's Translation of Hann's Climatology.!— Dr. Hann, pro-

fessor of cosmical physics at the University of Vienna and formerly

director of the Austrian Meteorological Bureau, published in 1883

n, Dr. Justius. — Handbook of Climatology. Part I, General Climatology.

aoe s Robert DeCourcy Ward. The Macmillan Company, 1903. 8vo,

437 pages

goo THE AMERICAN NATURALIST. [Vor. XXXVIII.

his Handbuch der Klimatologie which he reissued in 1897 in three ^

volumes. This is the standard work on the subject of climate and

no book in the English language is so authoritative or covers so wide

a field. Consequently, Professor Ward's translation of the volume

relating to General Climatology, which was made primarily for the

use of his classes at Harvard University, confers a great boom on all

American and English students of meteorology and physical geog-

raphy. :

In his title, Professor Ward has been too modest, for his book is

itself revised and enlarged, considerable new matter relating to

America having been substituted for certain topics which pertained

more particularly to Europe. While it might be wished that this new

material were distinguished from the author's German text, it is

stated that all the additions have been approved by Dr. Hann, so

that the translation is virtually a third edition of this part of the origi-

nal work. Numerous references to recent periodical and other

literature, in various languages, have been added and all the old

references have been verified, so far as possible, by the translator.

An added convenience is the summary of the contents of each chap-

ter, while a copious index of subjects and authors facilitates looking

up either. The metric system and the Centigrade scale of tempera-

ture are used throughout and should aid in familiarizing our students

with these units, which are destined to supplant the English system

in scientific writings. At present, however, the conversion tables in

the Appendix will, no doubt, be found necessary by many readers-

The German text has been accurately rendered into elegant English,

and almost no typographical or other errors have been noted. As was

said, only the first volume of Dr. Hann's work has been translated

and this evidently possesses the most general interest, since the

climatology of special regions, discussed by Dr. Hann in his other

two volumes and consisting largely of statistics, can be more easilv

consulted by persons not familiar with the German language.

In conclusion, the reviewer expresses the hope that this admirable

and disinterested work may have a wide circulation in our high

schools and colleges and indeed among all English speaking persons-

who desire to understand the climatic conditions of the globe and

their cause.

A. LAWRENCE ROTCH-

Nos.455-456] | NOTES AND LITERATURE. gol

BOTANY.

Bog-Trotting for Orchids! is the title of a popular book dealing `

with the swamp flora of the Hoosac Valley. It gives a pleasant,

diary-like account of the author’s rambles for three seasons, in the

course of which are given descriptions and other information regard-

ing her various finds. An appendix of 44 pages presents a system-

atic account of the genera and species of New England Orchidacez,

and there is an admirably full index. The illustrations are photo-

graphic, many of them excellent; some are colored, not always to

advantage, and a considerable proportion are of landscape views or

objects other than orchids. As one reads the rambling narrative

one cannot but sympathize with the writer’s enthusiasm, and is sure

to get refreshing whiffs of out-door air. Despite occasional inac-

curacies, showing the lack of expert revision, and the inclusion of

considerable material of rather more personal than general interest,

the book has the charm and value which belong to a faithful record

of first-hand impressions, and which will doubtless render it an

inspiring help to many amateurs.

P.L.

Agriculture for Schools.— Two interesting attempts to meet the

demand for instruction in matters of special importance to farm-

ing communities, are found in a handbook of exercises edited by

Prof. Hays? and an elementary text-book by Profs. Burkett, Stevens

and Hill.3

The hand-book consists of 237 exercises of an eminently practical

nature as may be judged from the following sample titles taken at

tandom: Food stored in seeds; Making maps of public roads ;

Transpiration of water by plants; Teeth of domestic animals; Bow-

line knot; Sharpening pocket knives; Classes of chickens ; Classi-.

! Niles, Grace oe —Bog-Trotting for Orchids. New York, C. P. Put-

nam's Sons, 1904. v + 310 pp., 72 pls. dk

* Hays, Willet M. xe School Agriculture. Bulletin No. 1. Exercises in

Agriculture and Housekeeping for Rural Schools.. St. Anthony Park, Minn.,

Department of Agriculture, University of Minnesota, 1905. — 12mo, viii + 196

79 gs.

? Burkett,.C. W. --Stepdan. F. L., and Hill, D. H.—Agriculture for Beginners.

Boston, Ginn & Congmdj, 1903. 12mo, xii + 267, 215 figs.

902 THE AMERICAN NATURALIST. [Vor. XXXVIII.

fication of soils; Crossing corn; Emergencies; Removing ink from

white goods; To make liquid yeast; Chicory in coffee; Cash ac-

count; Tent caterpillars ; Planting trees; Farm gate lesson; Sewed

seam; Overhand patch: Three-cornered darn; Cleft grafting.

These exercises are arranged somewhat in the order of difficulty and

in related groups, but the extent to which they shall elucidate princi-

ples or develop general ideas must very largely depend upon the

teacher. If skilfully incorporated with other work and their bear-

ings broadly discussed many of these exercises would have real

educational value in fitting pupils to meet intelligently the actual

problems of rural life.

The text-book proceeds in a more orderly way, establishing gen-

eral principles and showing their application to problems of soil,

culture, protection, selection, stock-raising and dairying. Sugges-

tions for profitable observations or experiments are freely introduced.

The illustrations are of unusual excellence, and the style clear and

direct. Such a book cannot fail to advance the best interests of

practical education in much needed directions. Both books might

well be used together. |

FREDERICK LEROY SARGENT.

Notes.— André Michaux's “Travels into Kentucky," F. 4.

Michaux's “Travels west of Alleghany mountains," and Harris’s

“Journal of a tour northwest of Alleghany mountains," are repub-

lished, with annotations, by Thwaites as vol. 3 of his series of Zar/y

Western Travels, issued by the Arthur H. Clark Company, of Cleve-

land. 1t is regrettable that the volume is not provided with an index

making its wealth of detailed information available.

A small list of arctic. plants is given in Schrader's * Reconnais-

sance in Northern Alaska," published as Professional Paper No. 20

of the U. S. Geological Survey.

A polyglot suggestion of amendments to the Paris code of botani-

cal nomenclature, addressed to the Vienna Congress of 1905, has

been published by the botanists of the Gray Herbarium, the Crypto-

gamic Herbarium and the Botanical Museum of Harvard University,

under date of June 9.

The annual species commonly referred to Polypteris have been

segregated under Rafinesque's generic name Othake, by Bush, in vol.

14, no. 6, of the Transactions of the Academ yy of Science of St. Louis.

Nos. 455-456] NOTES AND LITERATURE. 903

Three new exotic orchids ascribed to Rolfe are described by Ames

in Proceedings of the Biological Society of Washington, of June 9.

Two new Jamaican species of Polypodium are described by Maxon

in no. 13740f Proceedings of the U. S. National Museum.

Blakeslee has an interesting note on the sexuality of Rhizopus in

Science of June 3.

A paper on vitality and germination of seeds, by Duvel, forms

Bulletin no. 58 of the Bureau of Plant Industry of the U. S. Depart-

ment of Agriculture.

A résumé of ecological work in 1903 is given by Cowles in Science

of June roth.

A healthy sane tone pervades an article by Holt on the value of

field and herbarium work in high school botany, published in Schoo/

Science for June.

Ecological experiments on Rumex Acetosella are described by

Transeau in Science of June 3.

A critical study of Lepidium is being published by Schinz in the

Bulletin de P Herbier Boissier.

A popular account of the native Papaw is given by Viola McCohn

in Country Life im America for July.

A further list of Eastern species of Cratzgus is published by

Ashe, under date of June 15, in vol. 20, no. 2, of the Journal of the

Elisha Mitchell Scientific Society.

Professor Bessey calls attention, in .Sezence of June 24, to the early

falling of the aberrant lowermost leaves of Acer Wegundo each year.

The July number of Arboriculture, printed on paper made from :

the wood of Catalpa, contains data as to the availability of this wood

for paper purposes.

A review of Californian Polemoniacez, by Jessie Milliken, forms

vol. 2, no. 1, of the University of California Publications, Botany,

issued May ro.

Heft 19 of Engler’s “ Pflanzenreich,” is devoted to Betulacez, by

Winkler.

An account of Persea gratissima as cultivated in Florida is pub-

lished by Rolfs as Bulletin no. ór of the Bureau of Plant Industry of

the U. S. Department of Agriculture.

904 THE AMERICAN NATURALIST. [Vor. XXXVIII.

A developmental account of Juncacez is contributed by Laurent

to no. 2—3 of the current volume of Annales des Sciences Naturelles,

Botanique.

Anatomico-ecological studies of Tillandsia are published by Mez

in Vol. 4o, Heft 2, of Jahrbücher Jür wissenschaftliche Botanik.

The Germination of Amaryllidacez is considered by Worsley in

the Journal of the Royal Horticultural Society of May.

A reprint of Bradbury’s “Travels in the Interior of America,”

second edition, London, 1819, which forms vol. 5 of Thwaites’

“Early Western Travels,” contains a list of plants collected about

St. Louis and on the Missouri River.

Under the title “ Contributions to our knowledge of the flora of the

Rocky Mountains,” Aven Nelson has collected as a thesis for the

Doctor’s degree his more important papers published in various jour-

nals between 1898 and 1904.

The opening fascicle of vol. 5 of Urban's “Symbolæ Antillanz"

contains revisions of Smilax (by Schulz), Celastracez (by Urban),

and Sapotacez, in part (by Pierre and Urban).

Under the heading “T rabajos de la Oficina de Historia Natural,"

the Colombian government is publishing a series of papers on the

botany of that country, by Cortes.

The first fascicle of vol. 2 of Arechavaleta’s “ Flora Uraguayana,”

has been issued by the Museo Nacional de Montevideo, as a part of

its Anales.

Volume 2 of Wood's * Natal Plants ” is completed by the issuance

of part 4, dealing with grasses.

No. 28 of the new series of “ Contributions from the Gray Her-

barium of Harvard University,” forming vol. 4o, no. 1, of the Zro-

ceedings of the American Academy of Arts and Sciences, and issued on

July 18th, contains the following papers: Robinson and Greenman,

Revision of the Genus Sabazia, Revision of the Mexican and Cen-

tral American Species of Trixis, and Revision of the Mexican and

Central American Species of Hieracium ; Fernald, Synopsis of the

Mexican and Central American Species of Alnus, and Some New

Species of Mexican and Nicaraguan Dicotyledons ; and Greenman,

Diagnoses and Synonymy of Mexican and Central American Sperma-

tophytes. ;

Nos. 455-456.] NOTES AND LITERATURE. 905

A companion to M. van den Bossche's “Icones Selectae Horti

Thenensis” has been started, also with the botanical collaboration

of de Wildeman and the delineating skill of d'Apreval, under the

title * Plantae Nove vel Minus Cognite ex Herbario Horti Thenen-

sis." The first part bears date March, r904.

An account of the Harvard Botanical Station in Cuba, with ex-

tracts from Superintendent Grey's report, is given by Dr. Goodale in

The American Journal of Science for July.

A quarto treatise on the Phytoplankton of the Atlantic and its

tributaries, by P. T. Cleve, is published by the author at Upsala.

A large number of economic topics are well treated by Halsted in

his recently issued Report of the Botanical Department of the New

Jersey Agricultural Experiment Station, for 1903.

An account of the principal commercial plant fibers, by Dewey, is

separately printed from the Yearbook of the U. S. Department of

Agriculture, for 1903.

An illustrated account of some old trees, by Pollmer, is contained

in Die Gartenwelt of June 25.

Professor Peck’s Report of the State Botanist for 1903, published

as Bulletin 75 of the New York State Museum, contains among other

things a paper on edible fungi and one on the species of Crategus

found in the State.

Experiments in the heredity of peas are detailed by Hurst in the

Journal of the Royal Horticultural Society of May.

A second edition of E. G. Paris’ “ Index Bryologicus," reaching to

the end of 1900, is being issued in fascicles from the Hermann Press,

of Paris.

A further addition to the nomenclature literature to be laid before

the Vienna Congress of 1905, by Harms, forms appendix 13 to the

current WVotizblatt des K. botanischen Gartens und Museums zu Berlin,

and is dated June 20, 1904.

A bacterial rot of the Calla is described by Townsend in Bulletin

No. 60 of the Bureau of Plant Industry of the U. S. Department of

Agriculture.

An account of “fungoid pests of the garden,” by Cooke, is con-

tained in current numbers of the Journal of the Royal Horticultural

Society.

906 THE AMERICAN NATURALIST. [Vor. XXXVII].

Cultural experiments with biologic forms of the Erysiphacez are

detailed by Salmon in a paper separately printed from the Philosoph-

«al Transactions of the Royal Society of London, Series B, vol. 197.

“Investigations of Rusts” is the title of Bulletin No. 63 of the

Bureau of Plant Industry of the U. S. Department of Agriculture,

by Carleton.

A paper by Cooke on edible fungi is contained in the Journal of the

Royal Horticultural Society of May.

The botanical articles of greatest interest in the Year-book of the

United States Department of Agriculture for 1903 are: — Kebler,

“The Adulteration of Drugs”; Taylor, “Promising New Fruits”;

Shepard, “ Macaroni Wheat”; True, “Cultivation of Drug Plants in

the United States”; Dewey. “ Principal Commercial Fibers” ; and

the reports on plant diseases forming part of the appendix.

Vol. 11 of the Proceedings of the Jowa Academy of Sciences contains

the following articles of botanical interest : — Buchanan, * A Contri-

bution to our Knowledge of the Development of Prunus Americana”

Miller, “The Lichen Flora of the ‘Ledges,’ Boone County, Iowa”

Martin, “A Chemical Study of Rhus glabra”; Cratty, “The Flora of

Emmet County”; and Mueller, “A Preliminary List of the Flowering

Plants of Madison County.”

A number of interesting botanical papers are contained in the

recently issued Vol. 36 of the Zransactions and Proceedings of the

New Zealand Institute.

Vol. 1 of Rendle’s “Classification of Flowering Plants” (Cam-

bridge, 1904) deals with Gymnosperms and Monocotyledons.

An account of Pzonia, as now popularly cultivated, is contributed

by Miller to Country Life in America for September.

The first part of a discussion of the variability of Eucalyptus

under cultivation is published by Maiden in No. 112 of the Zro-

ceedings of the Linnean Society of New South Wales.

An illustrated account of the cultivation of Acacia mollissima, by

Fairchild, forms Buletin no. 5r, part 4 of the Bureau of Plant Indus-

try of the U. S. Department of Agriculture.

Vol. 10, part 1, of the Annals of the Royal Botanic Garden, Calcutta,

consists of a revision of the species of Dalbergia of S. E. Asia, by

rain.

Nos. 455—456.] NOTES AND LITERATURE. 907

Opuntia rutila is figured in the Monatsschrift für Kakteenkunde of

July 1o. |

Ailantus Vilmoriniana, a Chinese species, recently introduced into

France, is described and figured by Dode in the Revue Horticole of

Sept. 16.

Two segregates of Rubus argutus Randii are described by Blanch-

ard in ZZe American Botanist of July. :

A very large tree of Quercus alba is described and figured by

Rothrock in Forest Leaves for August.

The fruit of Melocanna, a viviparous grass without endosperm, is

described by Stapf in Vol. 6, part 9, of the current botanical series of

Transactions of the Linnean Society of London.

An important paper on moneecious and dicecious grasses, with

illustrations, is published by Pilger in ere s Botanische Jahrbiicher

of August 16.

The aerating nodal roots of Haahi are discussed by Monte

martini, in Vol. 3, fascicle 2, of the Contribusioni alla Biologia Vege

tale of the Palermo botanical institute.

. The root structure of North American terrestrial orchids is con-

sidered by Holm in Zhe American Journal of Science for September.

A study of the Corbularia group of Narcissus is published by

Bureau in No. 1—2 of the current volume of the Bulletin de ča Société

des Sciences Naturelles de P Quest de la France.

A synopsis of the varieties of pineapple cultivated in Florida is

given by Hume and Miller in Buletin 70 of the Experiment Station

of that State.

No. 23 of Holm's *Studies in the em published in Zhe

American Fournal of Science for October, deals with morphological

aspects of the inflorescence.

No. 2 of Plante Nove vel Minus Coginte ex Herbario Horti

Thenensis, dated in June last, contains a considerable number of

Cyperacez by Clarke.

An exhaustive paper entitled * Contributions to the Knowledge of

the Life History of Pinus with Special Reference to Sporogenesis,

the Development of the Gametophytes and Fertilization," by Miss

Ferguson, forms a thick brochure of Vol. 6 of the Proceedings of

the Washington Academy of Sciences, issued on October 4th.

908 THE AMERICAN NATURALIST. [Vor. XXXVIII.

In the Gardeners’ Chronicle of August 6, Mr. Shaw restores two

lost names of Morelet (1855) for later names in Pinus by Grisebac

and himself.

A general account of the vegetation of Missouri, by Duggar, is con-

tained in Williams’ “The State of Missouri,” printed at Columbia,

Mo.

A physiographic and ecological study of the Lake Eagle region of

Indiana, by Mills, is contained in the 28th Annual Report of the

Department of Geology and Natural Resources of that State.

Separates of Dr. Kennedy’s “ Flora of Willoughby, Vt.,” have been

issued, in attractive binding, by the author, from Rhodora.

An account of the flora of the Peace River region of Canada, is

given by Macoun in Zhe Ottawa Naturalist of September.

Botanical items are included in Notes on the Falkland Islands, by

Vallentin, in Vol. 48, part 3, of the Memoirs and Proceedings of the

Manchester Literary and Philosophical Society, issued in July last.

A “Novus Conspectus Flore Europe,’ by Gandoger, is in course

of publication in the Buletin de ’ Académie Internationale de Géo-

graphie Botanique.

Ascherson and Grabner's “ Synopsis der Mitteleuropaischen Flora,”

in Lieferung 31-2 concludes the Spathiflorz.

Coste's “ Flore descriptive et illustrée de la France, etc.,” in Vol.

3, fascicle 2 reaches into Polygonacez.

The concluding third volume of Halácsy's *Conspectus Flore

Grace,” has recently been issued from the Engelmann press of

Leipzig.

Vol. 4, section 2, part 2, of the * Flora Capensis,” under the editor-

ship of Sir William T. Thiselton-Dyer, is occupied with Scrophu-

lariacez.

Cooke’s “ Flora of the Presidency of Bombay,” in Vol. 2, part 1,

reaches well into Boraginaceæ,— on the Bentham and Hooker

sequence of families.

An account of the vegetation of the district of Minbu, in Upper

_ Burma, by Gage, forms Vol. 3, part 1, of the Records of the Botanical

Survey of India,

Nos. 455—-456.] NOTES AND LITERATURE. 909

A comprehensive biological account of the island of La Mocha, by

Reiche and others, forms No. 16 of the Anales del Museo Nacional de

Chile.

Notes by Bessey in Science of July 22 show that each plumed akene

of Taraxacum weighs on an average .00044 gm., and that a medium

sized tree of Populus deltoides bears not far from 28,000,000 seeds,

of an average weight of .ooo65 gm. each.

An interesting paper on the adventitious epiphytic occurrence of

terrestrial plants in Norway has been distributed by Holmboe from

No. 6 of the current volume of the Christiania Videnskabs-Selskabs

Lorhandlingar.

The thermotropism of Rhododendron is described and illustrated

by Johnson in Country Life in America for November.

A preliminary paper on electrotropism of roots, by Plowman, is

published in Zhe American Journal of Science for August and Sep-

tember.

A popular account of carnivorous plants, by Shreve, is contained

in Zhe Popular Science Monthly for September.

A new and enlarged edition of Pammel’s “ Flower Ecology” has

been published from the J. B. Hungerford Press of Carroll, Iowa,

under the title “Ecology.” Seventeen chapters deal with the chief

topics.

Professor Kraemer has issued separates of a paper on the origin

and nature of color in plants from Vol. 43 of the Proceedings of the

American Philosophical Society.

A note on the pollination of Calepogon pulchellus is published by

Klugh in Zhe Ottawa Naturalist of August.

A syllabus of the economic plants of the temperate zones repre-

sented in the Dahlem garden is given by Engler in Appendix 14 of

the Wotizblatt des K. botanischen Gartens und Museums zu Berlin.

A key to the genera of the forest trees of Indiana, based chiefly

upon leaf characters, by Coulter and Dorner, has been issued by the

authors from Lafayette, Ind. i

Under the title “ Getting Acquainted with the Trees,” a tasty and

beautifully illustrated little book by McFarland has recently been

issued by The Outlook Company, of New York.

9IO THE AMERICAN NATURALIST. (VoL. XXXVIII.

An excellent little illustrated pocket guide to British trees is Step’s

“Wayside and Woodland Trees” (Warne, London and New York,

1904).

Stone's “ Timbers of Commerce and their Identification ” (Rider,

London, 1904) is a descriptive catalogue — rather than handbook —

of 247 species, with phototype illustrations of the wood sections.

The forest resources of Texas are discussed by Bray in Bulletin 47

of the Bureau of Forestry, U. S. Department of Agriculture; and

the forests of the Hawaiian Islands are discussed by Hall in Bulletin

48 of the same Bureau.

“The Forest Manual,” containing the Forest Act (No. 1148),

extracts from other laws of the Philippine Commission relating to

the forest service, and the forest regulations prepared in accordance

with the provisions of the Forest Act, is a little pamphlet recently

issued by the insular Bureau of Forestry.

An account of basket willows and their insects is given by

Hubbard and Chittenden in Buletin No. 46 of the Bureau of For-

estry, U. S. Department of Agriculture.

The chestnut in southern Maryland is discussed by Zon in Buletin

No.53 of the Bureau of Forestry of the U. S. Department of Agri-

culture.

Hooper reports on Indian gums yielded by species of Acacia in

The Indian Forester of September.

A considerable account of Gutta Percha and Rubber is contained,

with illustrations, in the recently issued Report of the Superintendent

of Government Laboratories in the Philippine Islands, for the year

ending Sept. 1, 1903.

Economic articles on the Sisal plants and the West Indian

anthracnose of cotton [caused by Colletotrichum Gossypii Barba-

dense] are contained in Vol. 5, no. 2, of the West Indian Bulletin.

An account of the propagation and marketing of Oranges in Porto

Rico is given by Hendricksen in Buletin JV». 4 of the Porto Rico

Agricultural Experiment Station.

An account of the anatomy of edible berries, by Winton, is pub-

lished in the American Journal of Pharmacy for September.

Native ornamental plauts of New Mexico are considered by

Nos. 455—-456.] NOTES AND LITERATURE. QII

Wooton in Bulletin No. 5r of the New Mexico Agricultural Experi-

ment Station.

Nestler, in a pamphlet on “ Hautreizende Primeln ” (Berlin, Born-

traeger, 1904), shows that Primula Sinensis, P. Sieboldii and P. Cor-

tusoides have the poisonous properties now well known as possessed

by 2. obconica, which he finds not shared by P. officinalis, P. mega-

seafolia, P. floribunda, P. Auricula, P. capitata, P. farinosa, P. Ja-

ponica, P. hirsuta, P. Clusiana, P. minima or P. rosea.

The cultivation of mushrooms, including the “tissue-culture ” |

growth of spawn, is considered by Duggar in Farmers’ Bulletin No.

204 of the U. S. Department of Agriculture.

An important paper on sexual reproduction in the Mucorineæ, by

Blakeslee, forming No. 58 of the “Contributions from the Crypto-

gamic Laboratory of Harvard University,” is published as Vol. 40,

no. 4, of the Proceedings of the American Academy of Arts and

Sciences.

' Nos. 15 to 18 of Lloyd’s Mycological Notes deal mainly with puff-

balls, especially the typical material of European and North Ameri-

can herbaria.

An article on Uredinez, with far-reaching conclusions on sexuality,

by Blackman, is published in the Annals of Botany for. July.

An account of Bacillus violaceus Manile, a pathogenic species, is

given by Woolley in Publication 75 of the Bureau of Government

Laboratories of Manila.

An account of Coniothyrium Diplodiella is given by Montemartini

in Z’ falia Agricola of September 30.

Conidial fruit of Morchella, similar to if not identical with Costan-

tinella cristata, is described by Molliard in the Revue générale de

Botanique of June 15.

Three fascicles of Lindau’s account of Hyphomycetes (“ Raben-

horst’s Kryptogamen-Flora von Deutschland etc., Vol. 1, Abtheilung

8”) have recently appeared.

A continuation of Rehm's “ Ascomycetes America Borealis” is

contained i in the July number of Annales Mycologict.

Diseases of Ginseng are discussed by Van Hook in Bulletin 219 of

the Cornell University Agricultural Experiment Station, issued in

June.

912 THE AMERICAN NATURALIST. [Vor. XXXVIII.

An excellent series of folio views in the Botanic Garden at Brus-

sels has recently been issued by the Ministére de l'Agriculture of

Belgium.

A portrait of Sir Joseph Hooker is published in Vol. 6, part 5, of

the Zransactions of the Natural History Society of Glasgow.

A medallion portrait of A. P. de Candolle forms the frontispiece of

Vol. 4, no. 8, of the Bulletin de Herbier Boissier.

The Journals.— ourna! of Mycology, May : — Morgan, “ Zubercu-

laria fasciculata”; Smith, “A New Egg-Plant Fungus [Ascochyta

Lycopersici|”; Durand, “Three New Species of Discomycetes” ;

Christman, “ Variability in our Common Species of Dictyophora” ;

Cockerell,“ A New Hypholoma” ; Clements, * Saccardo, De Diagnos-

tica et Nomenclatura Mycologica, Admonita Quadam"; Keller-

man, “A New Species of N amosphoera [ JV. lactucicola|” ; Kellerman,

“Minor Mycological Notes—IV”; Kellerman, “Index to North

American Mycology ” [continued]; Kellerman, “ Elementary Mycol-

ogy” [continued]; Kellerman and Ricker, “ New Genera and Species

of Fungi Published since the year 1900”; and Kellerman, “ Notes

from Mycological Literature — X."

Botanical Gazette, June: — Thaxter, “ Notes on the Myxobacteria-

cee”; Smith, *Undescribed Plants from Guatemala and other Cen-

tral American Republics -— XXVI ”; Kearney, “Are Plants of Sea

Beaches and Dunes trie Halophytes ? ”; Eastwood, * Some New Spe-

cies of Western Polemoniacez ”; Peirce, “Notes on the Monterey

Pine”; Chrysler, “ Anatomical Notes on Certain Strand Plants" ;

and Allen, * Chromosome Reduction in Lilium Canadense.”

Botanical Gazette, July : — Land, “ Spermatogenesis and Oogenesis

in Ephedra trifurca” ; Smith, “ Water-Relation of Puccinia Asparagi” ;

MacDougal, “Delta and Desert Vegetation ”; Arthur, * The ZEcid-

ium of Maize Rust”; Livingston and Jensen, *An Experiment on

the Relation of Soil Physics to Plant Growth "; and Eastwood, “A

New Gilia.”

The Bryologist, July: — Fink, “Further Notes on Cladonias —

IIL”; Britton, E. G., * Further Notes on Sematophyllum " ; Gilbert,

“Mounting. Mosses " : and Grout, “ Zortula pagorum in Georgia."

Bulletin of the Torrey Botanical Club, June : — Salmon, “ A Revi-

sion of some Species of Ectropothecium ”; Murrill, “The Polypora-

Nos.455-456] NOTES AND LITERATURE. 913

cee of North America — VII ”; Cook, “The Nomenclature of the

Royal Palms”; and Osterhout, “ Notes on Colorado Plants."

Journal of Mycology, July: — Morgan, * New Species of Pyreno-

mycetes ”; Holway, “ Notes on Uredinez — II”; Ricker, “ Notes

on Fungi — I, New or Interesting American Uredineæ ” ; Ellis and

Everhart, “ New Species of Fungi from Various Localities” ; Keller-

man, “A New Species of Peronospora”; Kellerman, “Cultures of

Puccinia Thompsonii” ; Kellerman, “ Elementary Mycology ” (contin-

ued) ; Kellerman, “Index to North American Mycology ” (contin-

ued); Kellerman, “Notes from Mycological Literature — XI s

and Kellerman and Ricker, “ New Genera of Fungi Published since

the Year 1900 ” (continued).

Journal of the New York Botanical Garden, July : — Britton,

* Explorations in Florida and the Bahamas”; Nash, “ Effects of the

Past Winter on Shrubs.”

Ohio Naturalist, June: — York, “The Embryo-Sac and Embryo

of Nelumbo; Kellerman and Jennings, “Flora of Cedar Pome”:

Kellerman, “ Flora of Hen and Chicken Islands, 1903”; and Schaff-

ner, “The Jacket Layer in Sassafras.”

The Plant World, June : —Nehrling, “The Beginning of Spring in

Florida — III”; Safford, “ Extracts from the Note-Book of a Natur-

alist on the Island of Guam — XIX "; Schofield, “The Glumes of

. * Beardless' Barley "; and Barrett, “ The Lleren, a Rare Root Crop."

The Plant World, July : — Safford, “ Extracts from the Note-Book

of a Naturalist on the Island of Guam— XX”; Goetting, “On

Lonely Rocks and Sand-edged Bluffs”; Shear, “The Black Fungi”;

and Bailey, “Some Unusual Woody Plants.”

Rhodora, June: — Kennedy, “Flora of Willoughby, Vermont” ;

Fernald, “Identity of Michaux’s Lycopus uniflorus” ; Eggleston,

“ Addenda to the Flora of Vermont”; Hervey, “ Plants New to the

Flora of New Bedford "; Collins, * Some Maine Mosses ?: Sheldon,

“Some Introduced Weeds of Connecticut”; , Robinson, * James

Lawrence Bennett”; Knight, ^A New Sunflower for Maine "s and

Kennedy, * Additional Notes from Willoughby."

Rhodora, July: — Collins, “Some Interesting Rhode Island

Bogs”; Deane, “Preliminary Lists of New England Plants — XVII

[Polemoniacez — Acanthacez] ”. Fernald, “The Green Alders of

New England”; Clark, “ An Interesting Specimen of Arisema tri-

914 THE AMERICAN NATURALIST. (Vor. XXXVIII.

phyllum”; and Parlin, “A New Station for Myssa sylvatica in

Maine.”

Torreya, June: — Campbell, “Resistance of Drought by Liver-

worts”; Lloyd, “The Pollen Tube in the Cucurbitacez and Rubi-

aceæ” ; MacDougal, “Evening Primroses"; Berry, “Teratology of

Seedling Bean”; and Britton, “ Scirpus Coloradensis sp. nov.”

Torreya, July: — Jelliffe, * Additions to ‘The Flora of Long

Island’”; Nash, “A Collecting Trip to Haiti”; Berry, “ Two-

bracted Dogwood”; Britton, “ Savia Bahamensis, n. sp.; and

Shafer, “Notes on Cuban Plants.”

Botanical Gazette, August : — Davis, *Oogenesis in Vaucheria qe

Billings, “A Study of ZZ/andsia usneoides” ; Spalding, “ Biological

Relations of certain Desert Shrubs. r, The Creosote Bush ( Covz//ea

tridentata) in its Relation to Water Supply”; Hitchcock, '*Notes

on North American Grasses — III"; Perkins, “Carl Schumann a

(with portrait) ; Chamberlain, * A Correction."

The Botanical Gazette, for September: — Chrysler, * Development

of the Central Cylinder of Aracez and Liliaceze"; Johnson, “ Devel-

opment and Relationship of Monoclea”; Coker, “On the Spores of `

Certain Conifere”; Peirce, “ Artificial Parasitism” ; and Herre,

“Growth of Ramalina reticulata.”

The Botanical Gazette, October: — Davis, “The Relationships of

Sexual Organs in Plants”; Fink, “A Lichen Society of a Sandstone

Riprap"; Bergen, “Transpiration of Sun Leaves and Shade Leaves

of Olea Europea and other broad leaved Evergreens "; Hitchcock,

“Notes on North American Grasses — IV ” ; Stevens, “ Oogenesis

and Fertilization in A/bugo Lpomee-Pandurate.”

The Bryologist, September: — Britton, * Hyophila, a New Genus

to the United States ”; Harris, “ Lichens — Stereocaulon, Pilopho-

rus and Thamnolia”; Clarke, * Curbstone Mosses.”

Bulletin of the Torrey Botanical Club, July : — Gruenberg and Gies,

* Chemical Notes on Bastard Logwood " ; Bicknell, *Studies in Sisy-

rinchium — X, The Species of California”; Cushman, * Notes on

Micrasterias from Southeastern Mass." ; Rydberg, *Studies on the

Rocky Mountain Flora — XI."

Bulletin of the Torrey Botanical Club, August: — Murrill, “ Poly-

poracez of North America”; Shaw, “ Note on the Sexual Generation

and the Development of the Seed-Coats in Certain of the Papavera-

Nos. 455-456.) NOTES AND LITERATURE. 915

cee”; Cannon, “Observations on the Germination of Phoradendron

villosum and P. Californicum”; Griggs, “Two New Species of

American Wild Bananas, with a Revision of the Generic Name.”

Bulletin of the Torrey Botanical Club, September : —Vail, “ Studies

in the Asclepiadacez — VIII; Cockerell, * North American Species

of Hymenoxys."

The Fern Bulletin, July : — Price, “ Contribution toward the Fern

Flora of Kentucky”; Clute, “The Star Fern”; Prince, “Some

Ferns of the Cave Region of Stone County, Mo. ” ; Eaton, * The Cali-

fornia Gold Fern”; Clute, *A New Form of the Christmas Fern”

Eaton, “Is Asplenium lanceum American?”; House, “Some Rare

Ferns of Central New Jersey”; Parish, “ Additions to the California

Fern Flora”; Clute, “ Raising Prothallia of Botrychium and Lyco-

podium ” + Chats, “ Concerning Forms and Hybrids"; Eaton, “ Zsoetes

Amesti, a Cona ; Osmun, “ Eguisetum R in Connec-

ticut.”

Journal of Mycology, September: — * Benjamin Matlack Everhart,

Obituary ” [with portrait]; Morgan, ^ Pyrenomycetes Scarcely

Known in North America "; Holway, * Notes on Uredineæ — III” ;

Fairman, “Some New Fungi from Western New York”; Ellis and

Kellerman, *A New Phyllachora from Mexico"; Kellerman and

Ricker, *New Genera of Fungi published since the year 1900”

(continued) ; Kellerman, *Index to North American Mycology "

(continued); Kellerman, * Notes from Mycological Literature— XII."

Journal of the New York Botanical Garden, August : EU

* Report upon Further Exploration of Southern Florida"; Howe,

“Collections of Marine Algz from Florida and the Bahamas”

Journal of the New York Botanical Garden, September : — Lloyd,

“A Visit to the Desert Botanical Laboratory”; Nash, “An Agave

[A. Palmeri] in Flower." :

Journal of the New York Botanical Garden, October : — Robinson,

“A Visit to the Botanical Laboratory at Cinchona, Jamaica” ;

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— STONE, H. The Timbers of Commerce and their Identification. London,

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Chiroptera, Insectivora, Carnivora, Pinnipedia, yo e Sas Friedlænder,

917

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1904. 8vo,iv + 546 pp. 24 marks.— West, G. S. A Treatise on the British

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AMEs, O. Three New Orchid Species. ge - Biol. Soc. Wash. Vol. xvii, pp.

119-120.— AMES, O. Additions to the Orchid Flora of Florida. Proc. Biol. Soc.

Wash. Vol. xvii, pp. 115-118.— BAKER, F.C. The Molluscan Fauna of the Dells

of Wisconsin. Trans. Acad. Set. St. Louis. Vol. xiv, No. 2, pp. 99-105.--

BAKER, F. C. Notes on Planorbis truncatus Miles. 7Zrans. Acad. Sci. St. Louis.

Vol. xiv, No. 3, pp. 107-110.— BANCROFT, F. W. Estivation of Botrylloides

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Two New Subspecies of Tropical American T yrant Birds. Proc. Biol. Soc.

Wash. Vol xvii, pp. 113-114.— BA&ks, N. ^A Treatise on the Acarina or

Mites. Proc. U. S. Natl. Mus. Vol. xxviii, pp. 1-114, 20t figs.-— BENNETT,

ARY E. Are Roots Aerotropic? Bot. Gas. Vol. xxxvii, pp. 241-259, 5 figs.—

BOWNOCKER, J. A. The Occurrence and Exploitation of Petrolium and Natural

Gas in Ohio. Bull. Geol. Surv. Ohio. Ser. iv, No. 1, 325 pp., maps.—- BRAY,

W. L. Forest Resources of Texas. U. S. Dept. Agr, Bull. Bur. Forestry.

No. 47,70 pp., 8 St Bray, W. L. The Timber of the Edwards Plateau of

Texas, etc. U.S. Dept. Agr., Bull. Bur. forestry, No. 49. 30 pp., 5 pls.—

BROLMANN, H. W. Chilopodes monégasques. Aull. Mus Oceanogr. Monaco.

No. 15. pp. rs.— Burrirt, C. H. The Coal Measures of the Philippines, etc.

Rept. to U. n Military Governor in the Philippines. 269 pp.. maps, etc.— BUSCK, A.

Tineid Moths from British Columbia with Descriptions of New Species. roc.

U. S. Natl. Mus. Vol. xxvii, pp. 745-778.— CAUDELL, A. N. Two Orthoptera .

hitherto unrecorded from the United States. Proc. U. S. Natl. Mus. Vol. xxvii,

9-952.— CHITTENDEN, F. H. Insects Injurious to Basket Willow. U.

S. ha Agr., Bull. Bur. Forestry, No. 46. pp. 63-801, figs. 11-27.— CLA

L. The Echinoderms of the Wood’s Hole Region. Buil. U. S. Fish Pn.

for 1902. pp. 545-576, pls. 1- 14.— COCKERELL, T. D. A. Notes on Tetraneuris

linearifola. Proc. Biol. Soc. Wash. Vol. xvii, pp 1r11-112.— Dow inc, D. B.

eport on an Exploration of Ekwan River, Sutton Mill Lakes and Part of the

West Coast of James Bay. Ann. Rept. Geol. Surv. Canada. Vol. xiv, 60 pp.,

2 pls, 7 figs—Dvyar, H. G. The Lepidoptera of the Kootenai District of

British Columbia. Proc. U. S. Natl. Mus. Vol. xxvii i, pp. 779-938.— ELLIOT,

D. € Land and Sea Mammals of Middle America and the West Indies.

Field Columb. Mus. Publ., Zool. Ser. Vol. iv, Pts. i-ii, xx + xlix + 850 pp., 68

pls. & 329 text figs. — Evans, H. M. A New Cestraciont Spine from the Lower

Triassic of Idaho. Univ. Cal. Publ., Bull. Dept. Geol. Vol. iii, No. 18, pp. 397-

402, pl. 47.— FARRINGTON, O. C. Observations on the Geology and "ipie

of Western Mexico including an Account of Cerro Mountain. Publ.

Columb. Mus., Geol. Vol. ii, pp. 197—228, pls. 55-70, map and 4 figs— F eges

E. P. and JRvKkEL, L. H. Monograph of the Genus Saperda. Bull. N. Y. State

Mus.: No. 74. 86 pp., 14 pls.— FERGUSON, MARGARET. resets to the

Knowledge of the Life History of Pinus with Special Reference to Sporogenesis-

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pods of the Waters in the Vicinity of the Gulf Biologic Station Louisiana.

Nos. 455—456.] PUBLICATIONS RECEIVED. 919

Second Rept. Gulf Biol. Sta. pp. 67-79.— GiRTY, G. H. New Molluscan Genera

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45-47.— Harriss, W. P. and C. W. Lichens and Mosses of Montana. Bull.

Univ. Mont. No. 19, pp. 313-330, pls. 58-64.— HERRERA, A. L. La Citogéne-

sis Experimental y ea Oftamologia. Ann. de Oftamologia. Tom. 4, pp. 425-429,

2 figs.— HOLDEN, P. G. Selecting and Preparing Seed Corn. Bull. lowa. Agr.

Exp. Sta., No. 77. pp. 169-334, 47 figs.— HousER, G. L. The Animal Cell in

the Light of Recent Work. Proc. Jowa. Acad. Sci. Vol. xi, pp. 39-53.—

HUBBARD, W. F. The Basket Willow. U.S. Dept. Agr., Bull. Bur. Forestry,

o. 46. 61 pp., 7 pls.— IJ1tMA, I. Studies on the Hexactinellida. Contribution

IV. (Rosselide). Jour. Coll. Sci. Imp. Univ. Tokyo. Vol. xviii, 307 pp., 23 pls.

— ILLINOIS STATE LABORATORY OF NATURAL History. Studies of the Food

of Birds, Insects, and Fishes. Second Edition. Bull. I. State Lab. Nat. Hist.

Voli, No. 3,176 pp. JENNINGS, H. S. Asymmetry in Certain Lower Organisms

and its Biological Significance. Mark Anniversary Vol. pp. 315-337.—

JorpDAN, D. S. and Snyper, J. O. Notes on Collections of Fishes from Oahu

Island and Laysan Island, Hawaii with Descriptions of Four New Species. Prec.

U. S. Natl. Mus. Vol. xxvii, pp. 939-948. Jorpan, D. S. and SNYDER, J. O.

Description of a New Species of Fish (Apogon evermanni) from the reete

Islands, with Notes on other Species. Proc. U. S. Natl. Mus. Vol. xxviii, pp.

123-126.— Jupay, C. The Diurnal Movement of Plankton Crustacea. Zyans.

Wis. Acad. Sci. Vol. 14, n 534-568. — KELLO R. S. Forest Planting in

Western Kansas. U.S. Dept. Agr., Bull Bur. wegen No. 52. 51 pp., 7 pls.—

Kempton, H. B. The e of White Pine in New England. U. S. Dept

Agr., Bull. Dept. Forestry, No. 45. 40 pp., 15 pls., 2 figs.— LAMB, L. M. On

Dryptosaurus incrassatus (Cope), from the Edmonton Series of the North West

Territory. eol. Surv. Canada, Kubiprs to Canadian Paleontology. Vol.

iii, 27 pp., 8 pls. LAMSON-SCRIBNER, F. List of Philippin che Prod-

ucts and Fiber Plants. Bull. Bur. Agr. pie fds. No.5, 47 pp. -- MARsH, C

D. The Plankton of Lake Winnebago and Green Lake. Bull. Wis. Geol. &

Nat. Hist. Surv. No. 12, vi + 94 pp., 22 pls. — Maxon, W.R. Two New Ferns .

of the Genus Polypodium, gis Jamaica. Proc. U, S: Natl. Mus. Vol. xxvii,

pP. 741-744.— Maxon, W. R. A New Fete, Goniophebium etat from

Mexico. Proc. U. S. Natl. Mus. Vol. xxvii, pp. 953-954, pl. 48.—

G. Medusz of the Bahamas. Mem. Brooklyn Inst. Arts. & Sci. eat i, "d h

33 pp. 7 pls.— McCaskey, H. D. Report on a Geological Reconnoissance of

the Iron Region of Angat Bulucan. Bull. Manila Mining Pur. O05; 67

pp., 56 pls., maps.— McGnEcon, R. C. Notes on Hawaiian Replies from the

Island of Maui. Proc. U. S. Natl. Mus. Vol. xviii. pp. 115-118.— MoENKHAUS,

W. J. The Development of the Hybrids between Fundulus heteroclitus and

Menidia notata with especial Reference to the Behavior of the Maternal and

Paternal Chromatin. Amer. Jour. Anat. Vol. iii, " 29-65, 4 pls.— Monaco,

PRINCE ALBERT DE. Progres de la Biologie Marin Butl. Mus. Océanogr-

Monaco. No. 14, pp. 7.— MonGaN, W. C. and T RN. M. C. A Fossil Egg

from Arizona. Univ. Cal. Publ., Bull. Dept. Geol. Vol. iii, No. 19, pp. 403-410,

pls. 48-49.— Morse, M. Batrachians and Reptiles of Ohio. Bull. Ohio State

Univ. Ser. 8, No. 18, pp. 96-144, 1 pl, 3 figs.— MERRIAM, C. H. Four New

Grasshopper Mice, Genus Onychomus. Proc. Biol. Soc. Wash. Vol. xvii, pp.

123-126.— MERRIAM, C. H. New and Little Known Kangaroo Rats of the

920 THE AMERICAN NATURALIST. [Vor. XXXVIII.

Genus d Proc. Biol. Soc. Wash. Vol. xvii, pp. 139-146.— MERRIAM

CH. o New Squirrels of the Aberti Group. Proc. Biol. Soc. Wash. Vol.

xvii, pp. 129-1 eps eq: C. H. Unrecognized fae Rabbits of the Lepus

texianus Group - Biol. Soc. Wash. Vol. xvii, pp. 135-138.— MERRIAM,

C. H

Vol. xvii, pp. 131-134.— NEEDHAM, J. G. New Dragon- Fly Nymphs in the

United States National Museum. Proc. U. S. Natl. Mus. Vol. xvii, pp. 685-720,

pls. 38-44.— NELson, E. W. Descriptions of Seven New Rabbits from Mexico.

Proc. Biol. Soc. Wash. Vol. xvii, pp. 103-110.— NuTTING, C. C. American

Hydroids. Part II. The Sertularide. Special Bull. Smith. Inst. I51 pp., 41

pls.— OBERHOLSER, H. C. Description of Two New Birds from Somali Land.

Proc. U. S. Natl. Mus. Vol. xxvii, pp. 737—739. — Oscoop, W. H. Two New

Pocket Mice of the Genus Perognathus. Proc. Biol. Soc. Wash. Vol. xvii, pp.

127-128. — PEABODY, C. and MOORHEAD, W. K. The Exploration of Jacobs

Cavern, McDonald County, je Bull. Dept. Archeol. Phillips Acad. No.

I, 29 pp, map & 11 pls.— Rice, W. N. The Proper Scope of Geological Teach-

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853-856.— Rick, W. N. The Physical Geography and Geology of Connecticut.

Rept. Com. Board Agr. for 1903. pp. 94-112, map.— RICHARD, J. Campagne

scientifique du yacht “ Princesse-Alice” en 1903. Observations sur la Sardine,

sur le Hunc sur les Cétacés, sur des filets nouveaux, etc. Aull. Mus.

Oceanogr. Monaco, No. 11, pp. 29. Route, L. La Place des Antipathaires

dans la Saas et la classification des anthozoaires. Bull. Mus. Oceanogr.

Monaco, No. 16, pp. 3.— SAITO, K, gss japonica sp. nov. Jour. Coll.

Sci. Imp. Univ. Tokyo. Vol. xix, 8 pp., — SAITO, K. Uebr das Vork-

ommen von Saccharomyces BONS te E Jour. Coll. Sci. Imp.

Univ. Tokyo. Vol. xix, 14 pp., 4 figs.— SANDERs, J. G. Coccida of Ohio, I

Contrib. Dept. Zool. & Ent. Mit State Univ. No. 18, pp. 27-80, 9 p

SCHRENK, H. von. Cross-Tie Forms and Rail Fastenings, with Spécial Refer-

ence to Treated Timbers. U. S. Dept. Agr, Bull. Bur. Forestry. No. 50, 70

Pp. 71 figs. — SCHRENK, H. von. Recent Progress in Timber Preservation.

Yearbook U. S. Dept. Agr. for 1903, pp. 427-440, pls. 53- 55.— SINCLAIR, W. J.

Euceratherium, a New Ungulate from the Quaternary Caves of California.

Univ. Cal. Publ. Bull. Dept. Geol. Vol. ili, pp. 411—418, pls. "r on E.

ys and McCALLEY. Index to the Mineral Resources of Alabama. Geol.

79 pp., map and figs.—SMrrH, H. M. Description of a New sem ia

ino Eel of the Genus Anguilla. Proc. Biol. Soc. Wash. Vol. xvii, pp. 121-

122, I fig. STEVENSON, W. H. and ieri J. "d Draining Conditions in

Iowa. Bull. lowa Agr. Exp. Sta. No. 78, 239-263, -— STEWART, J. H.

and HITE, B. H. Commercial Faan p Va. Agr. Exp. Sta. No. 92,

59 pp.— TassiN, W. The Persimmon Creek Metearite. Proc. U. S. Natl. Mus

Vol. xxvii, pp. 955-959, pls. 49-50.— THOULET, J. Mesures des conu

marines au moyen de l’ analyse physique et Sp Wisin d'eaux

récoltés en séries. Bull. Mus. Oceanogr. Monaco. No. pp. 8.— TOPSENT,

- Sarostegia oculata. Hexactinellides bodies pw i di cap-vert. Budi.

Mus. Oceanogr. Monaco. No. 10, pp. 8, 3 figs.— Toumry, J. W. The Relations

of Forests to Stream Flow. Yearbook U. S. Dept. Agr. for 1903. pp. 279-288.

— WARD, H. A. Willamette Meteorite. Proc. Rochester Acad. Sci. Vol. x.

PP- 137-148, pls. 13-18.— WEBSTER, E. M. Studies of the Habits and Devel-

Nos. 455-456.] PUBLICATIONS RECEIVED. 921

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vii, pp. 15-25, pl. 3.— WEBSTER, F. M. Studies of the Life History, Habits,

and Taxonomic Relations of a New ees of Oberea (Oberea ulmicola Chit-

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MAN, S. The Barboo Iron-Bearing ois of Wisconsin. Bull. Wis. Geol. and

Nat. Hist. Surv. No. 13, x + 190 pp., 23 pls. WEIDMAN, S. Preliminary

Report on the Soils and Agricultural Conditions of Central Wisconsin. Bul.

Wis. Geol. & Nat. Hist. Surv. No. 11, 68 pp., 10 aS & map.-- Wooton, E. O.

Native nacio Plants of New Mexia: Bull. N. Mex. Agr. Exp. Sta. No.

51, 40 pp., 12 pls.— YENDO, K. -A Study of the Genicula of Corallinz. Jour.

Coll. Sct. Imp. Univ. Tokyo. Vol. xix, 44 pp., 1 pl.— Zon, R. Chestnut in South-

ern Maryland. U.S. Dept. Agr. Bull. Bur. Forestry. No. 53, 31 pp., 5 pls.

Actes de la Societé Scientifique de Chili. Tom. xii, Liv. 4-5; Tom. xiii, Liv.

1-3.— Anales del Museo Nacional de Montevideo. Serie ii, Pt. r.-— Botanical

Gazette, Tis Vol. 37, Nos. 4-6, Vol. 38, Nos. 1-3.— Bulletin of the Johns

Hopkins Hospital. Vol. 15, Nos. 157-163.— Bulletin du Musée Océanographique

de Monaco. Nos. 13 and 17.— Condor, The. Vol. 6, Nos. 3-4.— Higher Science,

May.— Journal Cincinnati Society of Natural Histor Vol. xx, No. 4.— Journal

of Geology, The. Vol. 12, No. 4.— National Chai Magazine, The. ol.

xv, No. 7.— JVotarisía, La Nuova. Ser. 15, Apr. and July.— Periodische Blatter

the Iowa Academy of Sciences for 1903. Vol. xi, 282 pp., 15 pls. Proceedings of

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ings of the Rhodesia Scientific Association. Vols. i-iii, 1899-1902. Proceedings

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I chart.— Retort of the U. S. Fish Commission for the year ending June 30, 1903.

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Sedit Tom. v, No. 36.— Verhandlung s deutschen SHIT. Vereins

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INDEX.

ALLEN, G. M. Rorquals of di.

ern Susféobdian d ‘ . 613

Amblystoma, Meali of 88

American Society of Zodlogists,

Proceedings of . . . . . . AGS

Amitosis in embryo of Fasciolaria 869

Ameeba, Physical imitations of

Ancestry of Mammalia . . 811

Breeding habits

Andrews, E. A.

of crayfish «a MONS

Apparatus for Manus inida € 4 ORS

Arboreal ancestry of Mammalia . 811

BEECHER, C. E. . 407

Bigelow, H. B. idi se f hearing

in goldfi -27

Birds, Bermudian EA

—— Classification of 311, 833

Blatta, Deveiopment of head of . 777

itc Bermudian birds . 555

Breathing valves of teleosts 405

Breeding habits of crayfish . 165

CAMPBELL, D. H. Affinities of

hioglossacez and Marsiliaceze 761

Cicindila, Distribution of

Classification of birds

Cone Flower, Variation in

Coniferales, Anatomy of

331, 243, 523, 671

Correspondence . 88, 241, 405, 689

cent uae. habits of . . 16

Cushman, J. A. Localized stages

~= m e x

Cursorial ipia 1 in habits i

mammals RU.

E rO

311, 833

"WI

Davis, B. M. . Studies of the plan

cell - 367, 431, eni 725

LI *

iii

Davis, W. T.

"ur Rana virgatipes at Lake-

hurst, N. J.

Desmognathus fines, Early jue

opment

Mesas P Desmognathus

fusca .

Diaptomus, Now species =

EASTMAN, C. Dentition of

Rhynchodus and other fossil

fishes i. ^

Fossil hee ;

Vernacular names oi ub

Undetermined fac-

Environment,

tors in

FASCIOLARIA, Amitosis in embryo

OR uo i T

Fossil fishes, Dentition of

Fossil plumage

Gi 4 pues Vernacular names of

dE eg et bearing | in

-HAMINEA, Natural history of .

Hatcher, J. B. Princeton Pata-

gonia expedition . à

Head, Development of i in Blatta

Hearing, Sense of in goldfish .

in spiders

Heredity, Undetermined factors in

Herrera, Correspondence

icka, A. Malar division À

Parietal division

Hyla andersonii at Laan N. x

JacksoN, R. T. Charles Emerson

Beecher. > kee

PAG

Hyla andersonii

iv INDEX.

PAGE

Jaggar, T. A., Jr. The visant

of Mount Pelée i in 1851

Jennings, H. S. TS imita-

tions of Amoeba

Jordan D. S. nies on recent fish

literature . - 471, 599

fanspiakting ou trout . . 885

Ligtins, Apparatus for aérating

Localized stages in plants :

ucas, F. C. Variation in ray

flowers of Rudbeckia . . . . 427

Lull, R. S. a Sory

in mamma

. 655

. 819

MALAR division.. . . 36

Mammalia, Arboreal akoy uL. 811

Mammals, Cursorial SRS In.

habits o

Mammals and reptiles in Taie

gonia, Distribution ER

Man, Influence of on aie

tion of € and mammals in

Patagon E cux

Monaco: Affinities PI : -761

Mast, S. O. Apparatus for aérat- -

ing liquids ..

Matthew, W. D. MESTRE

try of the Mammalia Bri

Metamorphosis of PEE A | oB

Metcalf, M. M. Neritina . - 565

Mitchell, Evelyn G. Oral bea:

ing valves of teleosts . 153. 405

Mount Pelée, The eruption’ of in

1851 ee has

NERITINA virginica .

Notes and Literature :

‘Biology » 321, 397

Botany, E EI Mine veg 474, 608,

679 9o

Clar ee uo NA

Palzontology 605, 677

Psychology . < - 895

75» 324, 399, 471, 595, 673, 897.

OPHIOGLOSSACEÆ, Affinities of . 761

Osborn, mitosis in em-

bryo of Fasciolaria . . 869

Osborn, is F. Reclassification a

the : Wi I ear Oe

PNRA p Psyopodes Pre eras i

PARIETAL division .

Pearse, A. S. New Net of

Diaptomus V-r« er da a NE

Peirce, J. J. Uadron iiis

in environment and heredity .

Penhallow, D. P. Anatomy of the

orth American Coniferales

243. 331, 523, 691

Photinus marg nales, Light €

Plant Ceil, Studies of P 431, re

Plants, Localized stages in .

lumage, Fossil . pee

Pritchett, Annie H. Heart aud

smell in spiders .

Proceedings inb Society at

Zoülogists - 495

Publications ved E RS 917

Pygopodes, oe and ie

atic position of . .

RANA virgatipes at Lakehurst,

NL... 05. 0

Reclassification of the Reptilia. 93

Reptilia, Reclassification of . 93

Reptiles and mammals in Pata-

gonia, bei ams. of. + + 524

Review

P and Robinson's

Fasciculi Malaye mses... . 399

Bateson and Saunder's Report

on Roy. Soc.. Evolution

Committee . . . . "M

Boehm's Sino-Australian Con-

tinent . .6

Boulenger on n Ciani ation of

Fishes

Burkitt, Savers "d Hill's

Agriculture for Beginners . gol

Campbell’s Fer . 60

Coues’s Key to 'N. A. Birds .

Coulter and Chamberlain’s

sri Ai Angiosperms 76

Covi and MacDougal’s

ed Laboratory... . «31

INDEX. V

i PAGE

Davison’s Anatomy of the Cat 4co

Dodge's General Zoólo

Eastman's Translation of Zittel 605

Gardiner's Maldives and Lacca-

dives . 326, 898

Grant's Origin of Mision of

CANTO UA i 677

Guyer’s Spermatogenesis of

Pigeon

Hays’ Kasela] in Kaa gol

Jenning’s Monograph of the

Rattulidze ant

Miller’s Criminal Cima 896

Montgomery’s Gordiacea . . 326

Niles Bog Trotting for Or-

i OI

CHIES — = 4 ee Ps

Natural Selection. ; 3PT

Pompeckji’s Life of Zittel . ; HE

' Porter's Flora of Pennsylvania 608

Redfield's Control of Heredity 228

Reese's Vertebrate Embry-

ology . -07

Walther’s Solpheten oe O77

ita. Wil

Walton’s

Friend 675

Ward’s "Tenasiibn af Mann. 899

Weed and Dearborn's Birds

in their Relation to Man 75

Whiting's aks in New

Engiand incre o. 321

Williamson's Fixation of

Crab’ 7

s). d d es

Wundt's Natural Science and

Psychology <<. . « 5 s 998

RHYNCHODUS, Dentition of. . . 295

Riley, W. A. Embryological e

velopment of the skeleton of the

head of Blatta IPS

Rorquals of VH CUR “Newfound.

nd ee

an ;

Rudbeckia, Vanation 1 Hn A 07

SHUFELDT, R. W. Classification

of birds 311, 833

Families aa kaha groups of

birds . 833

Dstology. ini tenia ti: pint

tion of Pygopodes . . . . .

Smallwood, W. M. Natural his-

tory of Haminea solitaria Say . 207

Smell, Sense of in spiders . . . 859

Spiders, Hearing and smell in.

TELEOSTS, Oral breathing valves

ye Mu LT MI UI

Townsend, Anne B. The Histol-

ogy of the light organs of Pho-

tinus marginalis . : uc cura

Transplanting of trout

Trout, Transplanting of

VARIATION in Neritina :

Vernacular names of T 241, 689

WıckHaM, H. F. Influence of

sten of Pleistocene Lakes .

on distribution of Cicindila . . 643

Wilder, H. H. he early devel-

opment of Desmognathus fusca 117

ZOÓLoGISTS, American

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Proceedings o ous l

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