a

JOURNAL

OF

MORPHOLOGY:

EDITED BY

C. O. WHITMAN,

With the Cooperation of

EDWARD PHELPS ALLIS, Jr,

MILWAUKEE.

Vien ae

BOS TOW: GINN & COMPANY. 1889.

II.

|i

IV.

II.

CONTENTSHOP VOLT,

—————— ->-- ——_——— No. 1.— July, 1888.

FREDERICK TUCKERMAN, M.D., Amherst, Mass. Observations on the Structure of the Gus-

tatory Organs of the Bat (Vespertilio subulatus) 5

E. D. Cope. On the Tritubercular Molar in Human Dentition

C. O. WuiTMaAn, Director of the Lake Labora- tory, Milwaukee, Wis. The Seat of Formative and Regenerative Energy .

Pror. HENRY FAIRFIELD Osporv, Princeton Col- lege. A Contribution to the Internal Structure of the Amphibian Brain

Dr. WILLIAM PaATTEN, Assistant in the Lake Laboratory, Milwaukee, Wis. Studies on the Eyes of Arthropods

No. 2. November, 1888.

Henry V. Witsovn, Fellow of the Johns Hopkins University. On the Development of Manicina Areolata,

Joun M. CLARKE. The Structure and Development of the Vis- ual Area in the Trilobite, Phacops Rana,

PAGES

I-6

7-26

27-49

51-96

97-190

IQI-252

Green . : 5 : : , » 253-270

{Whi

IV.

fT,

ITy,

CONTENTS.

R. W. SHuFeEtpt, M.D., C.M.Z.S. Further Studies on Grammicolepis Bra- chiusculus, Poey .

Eo COPE: On the Relation of the Hyotd and Otic Elements of the Skeleton tn the Batra-

chia

R. W. SHuFELDT, M.D., C.M.Z.S. On the Affinities of Aphriza Virgata

No. 3.— April, 1889.

CHARLES-SEDGWICK Minot, Harvard Medical School, Boston, Mass. Uterus and Embryo: 1. Rabbit; I. Man,

EDWARD PHELPS ALLIS, JR. The Anatomy and Development of the Lateral Line System in Amita Calva.

Pror. A. E. DotBeEar, College Hill, Mass. On the Organization of Atoms and Mole- cules

C. O. WHITMAN. Some New Facts about the Hirudinea

Dr. WILLIAM PATTEN. Segmental Sense-Organs of Arthropods .

PAGES

271-296

297-310

311-340

341-462

463-568

569-585

586-599

600-602

Volume II. Fuly, r88s. Number 1.

JOURNAL

OF

MOK PHOROGY.

OBSERVATIONS ON THE STRUCTURE OF WHE GUSTATORY ORGANS OF THE BAT (Vesper-

tilio subulatus).

FREDERICK TUCKERMAN, M.D.,

' Amuerst, Mass,

THE present paper contains a description of the anatomy of the taste organs of a single species of Chiroptera. It is highly probable that further study of these organs in other species of this interesting group of animals will reveal important varia- tions, respecting both position and structural characters, from the results embraced in this short memoir.

It will be of interest first to notice briefly the form and gen- eral appearance of the tongue of this mammal.

GENERAL DESCRIPTION OF THE TONGUE.

The organ measures 13.5 mm. in length, its greatest transverse diameter is 5 mm., and at its thickest part it measures 4 mm. Anteriorly, it is free from the floor of the mouth for 6 mm., or nearly half its length. The upper posterior surface is slightly convex, and has a nearly uniform breadth. In the anterior half of the organ the lateral margins gradually converge, blending at the tip in a slightly rounded or pointed extremity. The upper surface of this portion of the tongue is marked by several sub- parallel, transverse rugze or folds, with corresponding depressions between them. These folds decrease in size as they approach the anterior extremity of the organ, and cease altogether at 1.5 mm. from its apex. The dorsal surface is unmarked by any

2 TUCKERMAN. [Vot. II.

median groove or raphé, except at the extreme posterior region. Here there is a wide and rather deep mesial groove, 2 mm. in length, beginning in front of the epiglottis and terminating midway between the two circumvallate papillee.

In one of my specimens, near the line of union of the poste- rior and middle third of the tongue, is a rounded eminence, showing a tendency to a raised posterior part, as seen in the tongue of the Rodentia. This feature, however, is wanting in other tongues of Vesfertzlio which I examined.

The upper surface, including the lateral margins, is covered with closely-set tactile and mechanical papilla, the points of which are directed backwards and inwards.

The fungiform papillz are only fairly numerous, and are dis- tributed with some degree of uniformity over the dorsal surface and upon the sides of the tongue. Posteriorly they terminate in front of the gustatory area, and they cease anteriorly a short distance from the tip. Those scattered over the anterior third of the dorsum are usually larger than those occurring elsewhere.

On each lateral half of the tongue, 2 mm. from the base, is situated a circumvallate papilla. The two papille are placed quite near the median line, the distance between them being only 0.6mm. They are oval in form, of nearly equal size, and are placed obliquely to the long axis of the tongue, their anterior extremity being directed outwards.

A papilla of similar type to those just mentioned, but less developed and apparently in a transitional stage, is present at the posterior limit of each lateral border. Further investigation will be necessary to determine whether these papille are con- stant or not. In the specimens which I have examined I have always found them, although exhibiting striking variations in form, general appearance, and structure, from normal circum- vallate papillae. No papilla foliata was found.

The under surface of the tongue is perfectly smooth. Ante- riorly there is a median ridge, with sloping sides, extending from the fraenum to the tip.

GUSTATORY STRUCTURES.

The Circumvallate Papille. These papilla show no indica- tions of lobation. Their upper surfaces are rounded, and they measure 0.30 mm. in their transverse diameter, and are 0.22 mm.

No. 1.] GUSTATORY ORGANS OF THE BAT. 3

in height. Where they join the tongue, the transverse diame- ter is only 0.12 mm. Each papilla is encircled by a rather shal- low and very wide trench. In some sections this extreme width of the trench (as shown in Fig. 1) is confined to its upper part, the lower portion curving beneath the papilla and becoming quite narrow. The ridge surrounding the trench, and forming its outer wall, has elongated tactile papillae projecting from its surface (Fig. 1). The general surface adjoining this gustatory area is covered with large and small papillz, quite symmetrical in arrangement, but presenting a great variety of forms. Serous glands are fairly numerous in the gustatory area, but none were found within the papillary body itself. The ducts of the serous glands open into the trench at its base and sides. The papilla at its upper part bears many secondary papilla, the depressions between which are filled by the epithelium. The nerves are chiefly non-medullated and ramify throughout the papilla, but I was unable to trace their terminal branches with any distinct- ness. The large ganglion described by Poulton! in the circum- vallate papilla of Pevameles, and observed by me? in the cir- cumvallate papilla of F7der, I failed to detect any indications of here. -

The two lateral circumvallate papillz are asymmetrical, the right one being much less developed than the left. The latter, as seen in vertical section, is elliptical in shape, and joins the tongue by a narrow pedicel. The trench which surrounds this papilla is very wide at its upper part, and narrow and of uni- form breadth at its lower. Serous glands are sparingly scattered through this region, and are entirely wanting within the papil- lary body.

The taste-bulbs are not very numerous in the circumvallate papilla of Vespfertzlio. They are disposed at the sides in a girdle of seven or eight tiers, the uppermost tier being nearly on a level with the top of the trench. From horizontal sections, made at different levels, I estimated the average number of bulbs in a tier at fifty. If we allow for eight tiers, we shall have four hundred bulbs for each papilla. I did not succeed in finding bulbs in the epithelium investing the upper surface of the papilla, nor was I able to detect them in the outer wall of

1 Quart. Journ. Micr. Sci., Vol. XXIII., 1883, p. 73. 2 Journ. of Anat. and Physiol., Vol. XXII., 1888, p. 136.

4 TUCKERMAN. [Vot. II.

the trench. The bulbs vary somewhat in shape, and in point of size they are the smallest I have yet observed, the nearest approach to them in this respect being those of the circumval- late papilla of the mouse. They vary in length from 0.025 to 0.030 mm., and their breadth is about 0.015 mm. Usually in a bulb of this area, the diameter of the peripheral end of the bulb exceeds that of the central. The latter is also curved slightly downwards. This form of bulb I have not observed before, although it is seen in those figured by Loven in the circumval- late papilla of the calf. I did not succeed in finding a bulb with the peripheral processes of its gustatory cells projecting beyond the pore. One bulb (in vertical section) shows a fissure, 0.015 mm. long and 0.0015 mm. wide, caused by a separation of the edges of two adjoining peripheral cells.

I was unable to isolate the central or gustatory cells of the bulbs sufficiently well for study, but the peripheral cells do not differ materially from those already described in the taste organs of other mammals. They are elongated, slightly flattened, nucle- ated cells, with their two extremities tapering gradually to a point.

The number of bulbs in the left lateral circumvallate papilla could not be estimated from my sections with any degree of accuracy. The most noteworthy thing about them in this region is their very unusual arrangement (Fig. 3). They occur only on one side of the papilla; but here they form a continu- ous chain, seventeen tiers deep, extending from the base of the papilla nearly to its summit. Bulbs are likewise present in the lower half of the outer wall of the trench. Here I counted eight tiers. The bulbs of this region measure 0.024 mm. in length and 0.015 mm. in breadth, being thus a little smaller than those of the normal circumvallate papilla.

The Fungiform Papille. These papille are distributed quite regularly over the dorsum and sides of the tongue, from the gustatory area nearly to the tip. Interspersed among those of the posterior part of the dorsum are a few which appear to be undergoing transition to the circumvallate type of papilla (Fig. 6).

In several instances taste-bulbs were present in the epithe- lium at the upper part of these papillae. They are usually placed vertically, directly in the long axis of the papilla. By

Non: ] GUSTATORY ORGANS OF THE BAT. 5

far the most interesting specimens (which are shown in Fig. 6) were found in a papilla from the posterior region of the tongue. In this papilla there are two well-formed bulbs, and placed between them is a third, which is either of a low order or unde- veloped. The largest bulb of the three measures 0.036 mm. in length and 0.016 mm. in breadth, and its apex appears to reach the free surface of the epithelium, its base penetrating the mucosa. Some of the isolated bulbs met with elsewhere in these papillae, particularly those of the anterior dorsal surface, are even larger than those shown in Fig. 6. Neither serous nor - mucous glands were observed near the fungiform papillz.

The entire upper surface of the tongue is covered with papillz of mechanical and tactile (?) function. They are quite closely set, except at the basal region, are largest at the posterior part of the dorsum, and gradually decrease in size as they approach the anterior extremity. These papilla, when near the tip, en- large slightly again. One from the posterior third of the tongue measured 0.11 mm. in height and 0.04 mm. in breadth. Behind the circumvallate papilla, and also about the tip, are numerous rather coarse, retroverted, conical papillae. Each papilla is seated upon a single papillary upgrowth of the mucous mem- brane, and is invested by epithelium of a uniform thickness. The outer layers of epithelium covering the upper surface and sides are usually partly, and occasionally wholly, cornified. These papillae vary much in shape and general appearance. Many of them are cone-shaped, while others resemble, in ex- ternal structure, minute fungiform papillae. The upper surface is now and then flat or slightly convex, but usually the papilla terminates in a retroverted, horny spinule.

1 Figure 7 represents a vertical section through a fungiform papilla, from the ante- rior dorsal surface of the tongue of a pig, containing eight taste-bulbs.

6 TUCKERMAN. [VoL. Il.

EXPLANATION OF PLATE I. List OF REFERENCE LETTERS.

c.e. Columnar epithelium. g.. Gustatory pore. g/. ad. Duct of serous gland. m.m. Mucous membrane. Z. Elongated papilla. f. e. Pavement epithelium. 2. £. Papillary processes of mucous membrane. ». Ridge. s. e. Stratified epithelium. s. p. Secondary papillz. ¢. Trench. 7. 4. Taste-bulb. 7. 6’. Taste-bulb of outer wall of trench. 7¢. 4.’ Taste-bulb undeveloped or of low type.

Fic. 1.— Vertical section through one of the circumvallate papillz. (x 100 diam.)

Fic. 2. Vertical section through the base of the same papilla, showing over- hanging side with four lowermost tiers of taste-bulbs. (Xx 480 diam.)

Fic. 3. Vertical section through the left lateral circumvallate papilla. (xX 125 diam.)

Fic. 4.— Horizontal section through the lower part of one of the circumvallate papillz, representing the taste-bulbs arranged ina zone. /. d. Ducts of the serous glands which open into the trench at this level. (xX 160 diam.)

Fic. 5.— Vertical section through a fungiform papilla of the mid-dorsal surface of the tongue, showing a single taste-bulb at its upper part. (XX 400 diam.)

Fic. 6.— Vertical section through a fungiform papilla of the posterior dorsal sur- face of the tongue, which is probably undergoing transition to the circumvallate type. (X 480 diam.)

Fic. 7.— Vertical section through a fungiform papilla of the anterior lateral dor- sal surface of the tongue of a pig. (X 200 diam.)

Journ. Morph. Vol.It.

Cf Hall, del

sahil

Pik

Pe Nee PAL an . ft

bil he

ON THE TRITUBERCULAR MOLAR. IN| HUMAN DENTITION.

E. D: COPE:

Descriptions of the molar teeth of man, given by anato- mists, differ in important respects. Thus, F. Cuvier (‘ Dents des Mammifers’’) states that, while the crown of the first superior true molar consists of four tubercles, those of the second and third superior true molars consist of but three tubercles. In the American edition of ‘“Sharpey and Quain’s Anatomy” it is stated that the crowns of the superior true molars of man consist of four tubercles; and the same state- ment is made in Allen’s late work on human anatomy.

My observations having shown me that both of these descrip- tions apply correctly to certain types of dentition, I determined to examine for myself, to ascertain, if possible, the extent and value of the variations thus indicated. My interest in the sub- ject had been especially stimulated by the researches among the extinct mammalia, and the results which I had derived from them. These are, in brief, as follows: first, the quad- ritubercular type of molar crown, illustrated by the first supe- rior true molar of man, belongs to the primitive form from which all the crest-crowned (lophodont) molars of the hoofed placental mammals have been derived; and second, this quad- ritubercular type of molar has itself been derived from’a still earlier, tritubercular crown, by the addition of a cusp at the posterior internal part of it. This tritubercular molar in the upper series has given origin directly to the superior secto- rial teeth of the creodonta and carnivora. In the inferior series, I have shown that in known placental mammalia at least, the primitive molar crown is quinquetubercular, or tritu- bercular with a posterior heel; that this form gave origin to the inferior sectorial tooth of carnivora by modification, and to the quadritubercular type corresponding to the superior quad- ritubercular crown —by a loss of the anterior inner cusp and

8 COPE. [Vo. II.

connecting crest. And from the quinque- and quadrituber- cular types of molar crown, the various specialized types of the ungulates have been derived.

Considerable significance, therefore, attaches to the question as to whether the superior true molars of Homo sapiens are quadritubercular or tritubercular. The inferior molars are also either quadritubercular or quinquetubercular; but less significance attaches to this modification than to that of the superior true molars. This is owing to two facts; viz., the fifth tubercle is not the anterior inner which completes the anterior triangle of the primitive inferior molar, but is a median posterior, such as is not uncommon in mammalia of Puerco and Eocene age; and second, because this tuber- cle is of quite small size, and is therefore more liable to variation from insignificant causes.

In the nearest allies of man, the anthropoid apes, the supe- rior true molars are quadritubercular, the posterior internal tubercle of the last or third molar being usually smaller than in the other molars in the chimpanzee. The inferior molars are quinquetubercular, in the human sense, the gorilla not infrequently adding a sixth lobe on the external posterior margin of the crown. The molars of both series are quad- ritubercular, with an occasional posterior fifth in the inferior molars in the Cercopithecidz and Cebidz, excepting the genus Pithecia of the latter, where the superior molars are tritubercular. The superior molars of the Hapalidz are tri- tubercular. In the Lemuridz the second and third, and fre- quently the first, superior true molars are tritubercular. In the Tarsiidze the superior true molars are tritubercular throughout. The superior molars of the extinct lemuroids differ, like those of the recent forms. Thus, in Adapis and its allies they are quadritubercular, but in Necrolemur they are tritubercular. In Chriacus (whose reference to the Lemuroidea is uncertain) they are tritubercular, as is the case, also, with Indrodon. In Anap, tomophus they are of the true tritubercular type. This is the genus of Lemuroidea, which in its dental character most nearly approaches the anthropoid apes and man. I have elsewhere? pointed out that the formula is I. - Cc. ; Pm. =; M. ; The canines are small, and there is no diastema in either jaw.

1 Report U. S. Geol. Survey, Terrs., F. V. Hayden, Vol. II1., 1885, p. 245, Pl. xxv. fig. 10, and Pl. xxive. fig. 1.

No. 1.] TRITUBERCULAR MOLAR. 9

It may be readily seen, in consideration of these facts, that the appearance of tritubercular superior molars in the genus Homo constitutes a reversion to the lemurs, and not to the anthropoid apes or to the monkeys proper. And among lemurs the reversion is most probably to that type which presents the closest resemblance to Homo in other parts of the dentition. The genus which answers most nearly to this requirement among those at present known, is Anaptomorphus.

Figure 1.—Two species of Anaptomorphus. Fig. a, A. evulus, lower jaw from above X2. Figs. 4, c, d, A. homunculus; 6, c, natural size; d, 3 natural size. Both Po

4

from Eocenes of the Rocky Mountains.

In studying the dentition of man, I have examined the crania contained in the following six collections: those of the Academy of Natural Sciences of Philadelphia; of the Army Medical Mu- seum of Washington; of the College of Physicians of Philadel- phia; of the University of Pennsylvania; of the Boston Society of Natural History; and of my own museum. The first of these is especially valuable on account of the negro, Egyptian, and Hindu crania it contains. My acknowledgments are due to the Board of Curators, of which Professor Leidy is chairman, for the opportunity of studying it. I am also indebted to the Boston Society of Natural History, and its learned curator, Professor

10 CORE. PVOL. Hie

Hyatt, for the opportunity of examining Hindu and Chinese crania in their museum. The collection of the Army Medical Museum at Washington is especially important on account of the Kanakas, Esquimaux, Peruvians, and North American In- dians which it possesses. I am under great obligations to its distinguished director, Dr. J. S. Billings, for the facilities which he placed at my disposal. The museum of the Philadelphia College of Physicians contains the collection made by the late Professor Hyrtl of Vienna, of crania of Eastern and Mediterra- nean Europeans. In this department it is unrivalled, and I am greatly indebted to the council of the college, and its curator, Dr. Guy Hinsdale, for the opportunity of examining it. Some French skulls in the University of Pennsylvania were of value in the investigation. My own collection, though small, contains a number of Maoris, Australians, Tahitians,! and North Ameri- can Indians, which have proved to be of importance. Of Eng- lish and Europeo-American crania, I have been able to examine but few of what might be termed the thoroughly amalgamated race. Of the latter there are probably many crania in the war collection of the Army Medical Museum, but how free the race of each may be from foreign intermixture, of course it is impos- sible to know. In selecting such as are supposed to be “stock Americans,” those of persons with English names have been preferred, although many now true Americans are of German ancestry. In order to increase the list of this class of examina- tions, I have imposed on the forbearance of my friends by fre- quent inspections of their dentitions in ove aperto.

I suspect that the characters thus obtained will prove of im- portance in a zodlogical and ethnological sense. They have been already found to be of great fixity, and hence significance, in the lower mammalia. The only reason why they should be less so in man is, that the modification in reverting to the tritubercular molar is a process of degeneracy, and may be hence supposed to be less regular in its action than was the opposite process of building up, or addition of the posterior internal cusp. Some justification for a light estimate of its value may

1 For the Maori and Australian skulls, I am indebted to Mr. Speechey Gotch of Melbourne; and for the Tahitians to Dr. Chassaniol, Chef de la departement de Santé de Taiti, of Paris. My best acknowledgments to these gentlemen are hereby ex- pressed.

No. 1.] TRITUBERCULAR MOLAR. II

be found in the following tables. But it must be remembered that it is not always possible to determine exactly the race of the person represented by a skull, even when care in its identi- fication has been exercised. Emigration and war have con- stantly rendered races impure, and transplantation on a large scale has in some parts of the earth produced hybrid races. The results of a study of human crania are sure to be more or less vitiated by these circumstances. We obtain averages rather than exact definitions. Nevertheless, the extremes of the series of variations are likely to be found to be characteristic of estab- lished forms of man, and will thus justify my belief in the value of the characters presented. To ascertain the relation of these variations to the races is the object of the present inquiry.

The cause of the tritubercular reversion belongs to the class of agencies active in evolution of organic types, of whose real nature we know little. It cannot be said to be due to a con- traction of the maxillary arcade, for the Esquimaux and some other peoples which display the tritubercular dentition are not deficient in this respect. Nor do tritubercular molars require less space than the quadritubercular, for the external width of the crown is the same in both cases. They generally require less material however than a quadritubercular crown, since a triangle is smaller than a square drawn on the same base line; however, in some men of the lower races who present the tritu- bercular molars, their outline is nearly square. The hypothesis advanced to account for the reduction of the number and quality of human teeth observed in the higher races, as well as for the replacement of the prognathous jaw by the orthognathous, is that such changes are due to a transference of material and of growth energy from these parts to the superior part of the skull and its contents. The relative superiority of the dimensions of these parts in the higher races is thus accounted for.

In the following tables the tubercular formule are represented by numbers. Only the last three, or the true molars, in each jaw are considered. Tubercles of reduced size are represented

by fractions. Thus eae indicates that each superior molar is quadritubercular, and each inferior molar quinquetubercular. This represents the extreme of the series represented by the 4a 3 A 44

lowest races. The formula

indicates that the true molars

12 COPE. [VoL. lI.

have four, three, and three, tubercles respectively, and that the inferior true molars have four each. This represents the ex- treme common among the higher races. In the table which follows, the numbers attached to crania in the respective collec- tions are appended, and initials indicating the collection follow. Thus A. M. M. refers to the Army Medical Museum, Washing- ton; C. P., the College of Physicians of Philadelphia; U. P., the University of Pennsylvania; B. S., the Boston Society of Natural History; E. D. C., my private collection; no initials follow the numbers of the Academy of Natural Sciences of Philadelphia.

4-4-4

Meg re

Malay! of Madura, *1339; Malay, 425 ; Negroes, 63 and 258, C. P. ;

S. Sea Islander, 86, A. M. M.

4-4-4 5 a ara Malays, 47 and 433 (latter of Sumbawa) ; Negroid Egyptians, 43, 798, 852*, 869.

4-4-4 4-4-4 Malay, 1338 (of Amboyna). 4-4-4 is

Tahitian, E. D. C.; Kanakas, 143 and 1308, C. P.; Malay, 425 (Java) ; Negro, 963; Hindu, 1070, B.S.

dices paeie roy) Marquesas Ids., 1531, C. P.; Greek of the Morea, 55, C. P. I ory Sa Si) Si our ee Peruvian, 932 (Arica). A Aa ee ama an

Malay, 430 (Amboyna) ; Italian, 114, C. P. (Elba) ; Peruvian, 2300, A. M. M.

1 Just what is meant by the Malays” of the Philadelphia Academy collection I do not know.

No. I.] TRITUBERCULAR MOLAR. 13

Yeni ea 3 Broan © Negro, 4122, C. P.; Kaffir, 13585 Pessahs, 1095-*7; Gipsy, 31, C. P.; Montenegrin, 29, C. P.; Tablunka, 106, C. P. ; Germans, 1063-4 (Tiibingen) ; Feejees, 292-3, A. M. M.

4—4— 3% ?

Australian, E. D. C.; Tahitian, E. D. C.; Negroes, 901, 903, 914, 920, 921, 927, 964; American Negro, 980, A. M. M.; Chatham Ako 1557, A. M. M.; Peruvians, 68, 452; 2 N. Amer. Indians, E. D. Cxs Ponka India, 487, A. M. M.; Comanche, 6563, A. M. M. ; Sicilian, 110, C. P.; Esquimaux, 1859, A. M. M.; French, 1579, A. M. M.; Anglo-Americans (Wilderness Battle-Field, Va.), 6305-6-7, A. M. M.; Kanakas, 286, 434, and 584, A. M. M.; Hottentot, Be Ss

A Aer Die core Madagascar, two (Hovas) ; Samoan Id.; N. Amer. Indians, 1345 (Lipan), and Hudson’s Bay Ind.; Mexican, 714 (Ancient) ; Peruvians, 72, 1373, 1465 ; Italian, 118, C. P.; Lombard, 117, C. P.; Czech, 131, C. P.; German, 239, C. P.

roa Seg) ew arghe NG N. Amer. Indians, 530 and 1804 (Pawnees), A. M. M. A483 aaa. O

Italians, 2 (Giurgevo and Piedmont), C. P. ; Gipsy, 18, C. P.; Ural, 33, C. P.; Bulgar, 94, C. P.; Slav, 84, CyB:

4 Sie 3 5 ena N. Amer. Indian (Spokane), E. D. C.; Italians, 46 and 47 (Terra- cina and Ragusa), C. P.; Albanian, 123, C. P.; Viennese, 9 59, Cre:

Aa Aes 4—4—4 Negroid Egyptian, 885 ; Swede, 1549. A= AS 8 ?

2 Tahitians, E. D. C.; Kanakas, 291, 451, 455, 570, 588, 844, 1057, A. M. M.; Chatham Id., 5721, A. M.M.; Esquimaux, 1232, A.M. M.; N. Amer. Indian, 380 (Chippewa), A. M.M.; ? Anglo-Americans, 5721,

14 COPE. [VonvIL

32654, 6785, A. M. M.; Jew, 3, C: P.; Rouman, 96, C.PoeiChinese, Sagi. Os) (EO, Te 7e BS. oe maa f

Kanakas, 79, 280, 283, 421, 423, 427, 431, 457, 463, 599, 709, 862, A. M. M.; Chinese, 958, 5130, 5139, A. M. M.; Peruvian, 1765 (An- con), A. M. M.; Peruvian, 2302, A. M. M.; S. Sea Islander, 344, A. M. M.; Indian of Yucatan, 629, A. M. M.; Mojave, 209, A. M. M.; Hindu, 123, B. S.

A Ota A Aur Circassian, 762. 4 33 33 STE Gate Negro, 1343 (Archipelago) ; American Negro; S. German, 1289. Mama Bagh Dh 4 Fic

- N. Amer. Indian, 1794 (Ute), A. M. M.; Hindu, *1344, Bengal ; Gipsy, 98, C. P.; S. German, 1158, Italians, 2, 8, and 109 (Tessino and Roveredo), C. P.; Greek, 53 (Cephalonia), C. P.; Bosniak, 100, CoP:; Cossackienh Don; cor, ©.50.

A 3h = 35 OSS od Patagonian, 1232 (last infer. mol. 44) ; Negro lunatic, 55. Weephis Aue =)

African Negroes, 580, 685, 902, 917; Kanaka, 861, A.M. M.; Ameri- can Negro, 411, A. M. M.; Peruvian, 1326; 2 N. Amer. Indians, E. D. C.; Mound Builders, 1049-1123, A. M. M.; Piegan, 6486, A. M.M.; Cheyenne, 5560, A. M.M.; ? Anglo-Americans, 2, A. M. M. ; Esquimaux, 1182, 1194,1250; Hollander, 9 10,C. P.; Hindu, 124, B.S.

Mises ee DONS Malay, 1340 (Macassar), 1341 (Java) ; Finn, 1539 ; Czech, 79, C. P. ; Wirennese,/61,)\C.P. 3 French. Ws Ps

/ pemesr aa inh Algerine, 3900, C. P.; Uskoke of Banjaluka, 90, C. P.; Styrian, 85, C.'P. ;’ Hindu, 432; Latin, 115, 'C. P.; Albanian, \o7,"C. Rae nGermean of Siebenbiirgen, 76, C. P.

NO.;1-] TRITUBERCULAR MOLAR. 15

Ai Brr3 bo 4a A Hungarian, 14,C. P.; Russian, 35,C. P.; Australian, (1327) ; Negroes (Benguela), 421, (Mozambique), 423; Hottentot, 1351; Aztec; Cir- cassian ; Moravia, 9 135, C. P.; Croat, § 134, C. P.; Galician, 139, COPS; Serb;.96, iC. P..s * Pizgau? Go C.P. > Saltzbure, 65, C..P.

4 aais 3

ee ae Swedes, 1487-50.

43h = 3

? ? ?

Negro (Dey), 1100; Peruvian; Kanaka, 576, A. M.M.; Dutch, 434; Malay, 39; Bengal, 20, A. N.S.; Hindu, 1330; Fellah, 999; Kabar- dine Caucasus, 38, C. P.; Ital. Piedmont, 45, C. P.; N. Amer. Indians (Flathead), E.D.C., (Cheyenne), 6525, A.M. M., (Ponka), 486, A.M.M., (Sioux), 9 2049, A. M. M.; Mound Builders, 169, A. M. M.; Esqui- maux, 1183, 1189, 1245, A. M. M.; ? Anglo-Americans, 6847, 5922, A. M. M.; German, 1066; Chinese, 879, B. S.

de Se ? Kanakas, 564, 569, 571, 591, A. M. M.; Peruvian, 2308, A. M. M.; Arucanian, 970, A. M. M.; Chuktchi, 277, A.M. M.; Esquimaux, 1779, A. M. M.

AS 3h ? Chukchi, 263, A. M. M. Ae Se 3a ? ? German-American, 6445, A. M. M. ATONE SO. Bee ame

Dalmatian, 132, C. P.: Greek (Trieste), 56, C. P.; Indians (Santa Barbara, Cal.), 3.

‘ire OS Sea ae Gipsy, 130, C. P.; Trieste; 120,,C..P.; Apache, 1168, A. M. M- Aly) onan 5 4p 4s

Modoc, E. D. C.

16 COPE. [VoL. II.

1 Rep & RAS) Sra eater Druse, 122, °C. P.; Idria, 89, C. P.; ‘Carniola, $7,,1C. Pees. siyrol 121, C. P.; Paris, 3916, U. P.; 4 Anglo-Americans, E. D. C.; French, 867 (Paris), U. P., 1620, A.M.M. Kurd, 28, C. P.; Thug Hindu, 128 (A. N. S.); \ Esquimaux, 676 (trace of 5 on 2 and 3) ; Italian (Calabria), 48, C. P.; Greeks (Candia), 54 and 51, C. P.; Slovak, v2. C. P.; Czech, 4,:C. P.; ‘Lithuania,/9 25, Cooke bokovina. 90 ae, CG. Ps Tyrol,"9'67, CC. P. 5. Upp: Austria, 625. B) ivorayia, Goi e Hollander, 9, C. P.

4 Saree Yay aaa) = “Krakuse” ((Czech or Pole, large), $2,0C. P. 3) * Kumanie!” i(riiaun- gary), 71, C. P.; Pole, 24, C. P. 5. Volhyman p26, C2. Me odolanve C. P.; Hindu (Bengal), 1344; Tatra (large), 83, C. P.; Wallach, 95, Ge Ps. French; 105 55U.P-

A ai nea 4-4-4

Tchuktchi; Zagrebin, 126, C. P:; Croat, 77, C. P.; Mixed German and Croat, 17, C. P.; French, 1082; do., 1801, A. M. M.; Anglo- American, 1108.

Coe Ieee) ?

Peruvian, 1478; do., 2299, /A: ‘M. M:; Turkish ‘grave; a2%) 1G. Pos Crimean, 34, C: P.; \Cossack, 132.4) Ruthenian, 103, /C. P.; Armeman 99, C. P.; Esquimaux, 675, 679, .A..N. S.; Esquimaux, 12r1,\ 1258, L219, 1206, 1221, 1225, 1226, 1230, T2321, 124%, 1244, F250, L278, E7or, 1782, 2113, A.M. M.; Chuktchis, 256-7, A. M. M.; N. Amer. Indians, 5550 (Pawnee), 6561, 6565 (Comanches), 7023 (Cheyenne), 170, (Mound Builder), A. M. M.; Kanakas, 580, 841, A. M. M.; Indians of Yucatan, 626-628, A. M. M.; American Negroes, 981-983 (Rich- mond, Va.), 6615 °(S. Carolina; m. 2 on one side is 33); 6 Anglo- Americans, E. D. C.; ? Anglo-Americans, 95, 6305—6—7-8, 6640, 739, 2699, 1768, 5116, 5145, 5708 9, 5940, 6550, A. M. M.; Hindu, 425, leASe

hase Span

44 a Hungarian, 15, C. P.

No. I.] TRITUBERCULAR MOLAR. 17

Neolithic Man.

Through the kindness of Mr. Thomas Wilson, honorary Curator of the Department of Prehistoric Anthropology in the United States National Museum, I have had the opportunity of examining a number of dentitions of men of the Neolithic period of France. The best preserved of these includes both series in place in both jaws, the last two inferior molars of one side and the last inferior of one side being absent. This man was disinterred from a cemetery on the island of Thinic, on the coast of Brittany. Teeth of a considerable number of men, in a separate condition, were obtained by Mr. Wilson from the dolmen of Poulzongue, near the town of Gramat, in the depart- ment of Lot, Central France, and have also been examined by me.

Ai) 3, 445 The second superior molar is a perfect quadritubercular, while

the third is an equally perfect tritubercular, of somewhat reduced size. This dentition is, then, exactly intermediate between extremes. It is further advanced than that of the lowest existing races, but has not reached the final modification of the most specialized. The dental characters of the men of Poulzongue vary between the type of the man of Thinic and 43 3, 444 molar plainly tritubercular, and perhaps as many have the

fourth tubercle imperfectly developed. But no indication of a tritubercular first superior molar is observed, such as occurs in a few Esquimaux. The resemblance of perhaps half of the superior molars of Poulzongue to those of Esquimaux and the more specialized Indo-Europeans is distinct. Pointing in the direction of the latter, are the small size of the teeth and the frequent occurrence of caries.

The dental tubercular formula of the man of Thinic is

the formula Several of them have the second superior

The Accessory Anterior Internal Tubercle.

This tubercle is characteristic of the genus Lemur (Fig. 11) and some of its extinct allies. Such are the Chriacus pelvidens and C. truncatus Cope, the former figured in Tertiary Vertebrata (Report U. S.-Geol. Survey Terrs. III), Vol. XXIII, d, Fig. 7.

13 COPE. [Vot. Il.

I have found it well developed in the Malay (Fig. 2, aaz), No. 1339. It is present in three Tahitian crania in my collection, and I have observed it in another Micronesian cranium, and in some others. This character is decidedly lemurine, but whether to be properly termed a reversion or not may be ques- tioned in view of the fact that it always occurs, so far as yet known, in dentitions most remote from that type in other respects. It might be rather regarded as a survival or as a character which has persisted from the “‘ protanthropos”’ which was itself immediately derived from lemurine ancestors.

Anomalous dentitions.

I have observed the following examples of aberrant molars. In an Esquimaux (A. M. M.) the formula of the superior molars of one size is 3—4—0; in another (B. S.) it is 3—-3—4. Ina French skull (A. M. M.) we observe on one side 4—4—3; on the other side, 33—3—33. The posterior superior molar is, by reason of its late and retarded development, the most liable to exhibit abnormalities. Ina Tahitian skull in my collection, it is tritubercular on one side and obtusely haplodont (conic) on the other.

General Conclusions.

For clearer understanding of these characters, they are arranged in the form of a table. Only the principal races are represented, and hybrids, when determinable, are omitted. The characters of the superior molar teeth only are referred to. These are classified under four heads, viz.: first, tubercles 4—4—4); second, tubercles 4—4—34 or 4—34—34; third, tubercles 4—34—3; fourth, tubercles 4—3—3. As already remarked, the extreme types of the series give the most precise ‘indications of race, while the intermediate conditions have a various range.

In the first table the most obvious results are, that only the three lowest races present four tubercles on all the superior molars, and that of those with tritubercular second and third molars, Europeans and their American descendants greatly predominate. Also that of uncivilized races, the Malays and

1 Mulattoes, Mestizoes, Half-breed Indians, Gipsies, etc., are omitted.

NOW i] TRITUBERCULAR MOLAR. 19

Negroes never, and Micronesians very rarely, present this type of dentition, while in the Esquimaux it considerably predomi- nates, examples of tritubercular first molars even occurring.

& P a A |e uo) ros Pe | telaee 3 I ae a. ers [il aS) bl 45) cS S38 | STS AS A a iI * un .g = Br | a fs Sy Bea | ok a) = tata estas a |S] o 2 ut leas anced: Ih me le 6 los |) oe ae1g2| Ss / 82 aa| a A = a tare (aa cei Wee Sa ya laoreet She UP ices Ee Batis pega tinge (lee etal ees lanier (pea) ees alte (20) ae |4e/OS! B eo le aie Doi ae A Oh aaa I 18 (4-4 —?) 13 2 Eig I 20 1 4—4 32) Mesias Stic Te 28. \0 274). .Oy 2a Ee le Sui inOr (eae pees 4-4-3 J 1 5 Soe ames. 3 = 5 r ced labios z 3 | 24 | 45 Las eNom 2 2 3 | 19 | 56 | 90 Motaligaih: EON A429 MIA eat | RS rx) Zou EnGi son

I now give a table of the characters of the superior molars in the Europeans and Europeo-Americans examined. The number is not sufficient for final conclusions; nevertheless there are some indications of value. Some of the one hundred and nineteen dentitions examined are referred with doubt to their respective races. Thus the Europeo-Americans may have been in many instances immigrants, as many such left their bones on the battle-fields of the American civil war, where many of the crania were picked up. The supposed Germans are largely Austrians, so that some of them may be more or less Slavic or Magyar. Allowance for these would reduce the number of tritubercular molars.

1Ten crania in the Museum of the Cincinnati Soc. Nat. History, mostly Mound- Builders, have the formula 4—4— 3.

20 COPE, [ VoL: If

Ee

Z S .

ao] S oS 7)

a i Gi | ea Bae $5 D yi ae O24 | ME | SB as, ass a ag BB g jas | eS | Es 5 Sa Nes S.S Ss is} aes ole 3 8 a) ete | SO | Ba eee ea eames Be, 2 3 2 I 3 II A 35 3% Z 2 3 3 2 ro Neon eee 5 Pa es 35) me 4—3+—-3 I I 6 2 3 8 I 2 24 4—3 —3 2 ged 4 7 6 6 20 56 otal ior I 3 25 7 23 22 8/320 119

As results we have the following: the tritubercular dentition appears in II out of 25 Slavs; in 7 out of 23 Greeks and Ital- ians ; in 6 out of 22 Germans and Scandinavians; in 6 out of 8 French; and in 20 out of 30 Europeo-Americans. The only great race which presents a similar high percentage of trituber- cular molars is the Esquimaux, where they occur in 21 out of 30 dentitions. The tendency is most marked in Slavs, French, and Europeo-Americans, and is least marked in Greeks and Italians and in Germans. The former subrace stands in the series between the intermediate type of the North American Indians and the other Europeans. In the Germans the number with tubercles 4—34—3 is large. If these be added to the number with 4 3 3, we have 16 out of 22.

It is important to remember in this connection that the dis- tinguished ethnologist and archzologist, W. Boyd Dawkins, affirms that the earliest inhabitants of Britain and some other parts of Europe were Esquimaux. He refers especially to the men of the caves, whose implements and arts he declares to be identical with those used by the Esquimaux of the present day.? As it is evident that the lemurine or tritubercular reversion commenced with the Esquimaux, it may be that in some in- stances at least its appearance in men of Anglo-Saxon and other European races is due to inheritance alone. But it is also rea- sonable to suppose that in this case as in other evolutions, the

1 Perhaps improperly included in this table. 2 Early Man in Britain, 1880, p. 233.

No. 1.] TRITUBERCULAR MOLAR. 21

cause which produced this modification of the Esquimaux den- tition is still active, and its recent appearance in the most civil- ized races must be due to this unknown cause. The progressive character of the French dentition in this respect is in broad contrast with the primitive character of that of Italians and Greeks. The characters seen in the latter go far towards sus- taining Professor Huxley’s hypothesis, that the dark Mediterra- nean sub-races consist of a mixture of Egyptian