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MEMOIRS
OF THE
CARNEGIE MUSEUM
VOL. XI
/
Title Page and Table of Contents Issued January, 1951
JTTSBURGH, PA.
Published by Authority of Board of Trustees of Carnegie Institute
N0VE:MBER 1927— APRIL 1937
TABLE OF CONTENTS
PAGES
I. Title pages i-ii
II. Table op Contents. ........................................ iii
III. List of Figures in Text v
IV. List of Plates. vii
V. Genera and Species described as new in this volume ix
Memoir No. L Studies on the growth of Gorilla and of other higher Primates with special reference to a Fetus of Gorilla^ preserved in the Carnegie Museum. By Dr. Adolph H. Schultz. Plates 1-8; 6 text figures; Nov. 1927. ....... 1-86
Memoir No, 2. The Brownes Park formation. By O. A. Peterson, Plates
9“15, April 1928 (Supplementary Note, p. 121) 87-130
Memoir No. 3. Osteology of Phenacoccelus typus Peterson. By 0. A.
Peterson. Plates 16-20; Dec. 1928. 131-174
Memoir No. 4. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum, By C. W, Gilmore.
Plates 21-35; Feb. 1936 175-300
Memoir No. 5. The South American Tingitid® (Hemiptera) described by Stal. By C. J. Drake and M. E. Poor. Plate 36. Jan.
1937. . 301-314
Memoir No. 6. The Spang collection of Greek and Italian vases and Etruscan Urns in the Carnegie Museum. By Henry S.
Scribner. Plates 37-43. April 1937. 315-352
Index
353-355
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LIST OF FIGURES IN TEXT
Memoir No. 1.
FIGURE PAGE
1. Table showing variations of the intermembral index 31
2. Outer ears of fetuses; human, gorilla, baboon, and howler-monkey 47
3. Deviation curves in proportions of Gorilla fetus 51
4. Deviation curves of proportions of the Ape fetus 53
5. Diagrammatic representation of the average deviation indices 57
6. Hypothetical pedigree of recent higher primates 66
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15. 20. 21. 22.
23.
24.
Memoir No. 2.
Cross-section from north to south of Brown’s Park sediments 90
Brown’s Park Formation resting on Cretaceous 91
Second view, “Post Laramie” 92
Section shown five miles north of Sunbeam, Colorado 93
Type locality of the Weller horizon 93
Bassariscops willistoni 96
“ “ 97
Ticholeptus ? 99
Cameloid allied to Stenomylus 100
Aphalops ceratorhinus Douglass 103
“ “ “ 105
“ “ “ 105
“ “ “ 106
“ “ “ 106
16, 17, 18, 19. Aphalops ceratorhinus Douglass 107-9
Chalicotheroid 110
Distal ends of radius and ulna of Serridentinus fricki Peterson 115
Serridentinus fricki Peterson 116
Parelephas washingtoni Osborn, lower left molar 119
Crown and lateral views of lower molars of P. washingtoni 119
V
MEMOmS OF THE CARNEGIE MUSEUM
vi
Memoir No. 4.
1. Geological section near Dinosaur National Monument 182
2. Geologic map of Dinosaur National Monument 183
3. Diagram of skeleton of Apatosaurus as found 186
4. Sauropod skull and lower jaws 189
5. Coalesced atlas and axis of A. louisae 192
6. Lateral view of atlas and axis 192
7. Sacral vertebrae of A. louisae 202
8. Coossified caudal vertebrae of A. louisae 208
9. Left scapula and coracoid of A. louisae 214
10. Comparative views of Apatosaurus scapulae 215
11. Left humerus of A. louisae 217
12. Left radius of A. louisae 218
13. Left ulna of A. louisae 218
14. Left scapho-lunar of A. louisae 220
15. Left fore foot of A. louisae 221
16. Left metacarpals of A. louisae 223
17. Phalanges of left fore foot of A. louisae 225
18. Ungual phalanx of digit 1 225
19. Left ilium and sacrum of A. louisae 227
20. Left coossified ischium and pubis of A. louisae 228
21. Right femur of A. louisae 230
22. Articular ends of femur 231
23. Right tibia of A. louisae 232
24. Right fibula 233
25. Articulated hind foot of A. louisae 235
26. Right astragalus of A. louisae 236
27. Left metatarsals of A. louisae. 237
28. Proximal row of phalangials of A. louisae 239
29. Phalanges of left pes 240
30. Unguals of left pes 241
31. Pelvis and sacrum of A. excelsus 250
32. Left scapula and portion of coracoid of A. excelsus 253
33. Right humerus of A. excelsus 254
34. Right radius and ulna 256
35. Right forefoot 257
36. Sacrum and pelvis 262
37. Left pubis 263
LIST OF PLATES
PLATE
I.
II.
III.
IV. V.
VI.
VII.
VIII.
IX.
X.
XI.
XII.
XIII.
XIV. XV.
XVI.
XVII.
XVIII.
XIX.
XX.
XXI.
XXII.
XXIII.
XXIV.
XXV.
XXVI.
XXVII.
XXVIII.
XXIX.
Gorilla Fetus II.
Front and side views of Gorilla Fetus.
Measurements: Human Fetus, Gorilla Fetus.
Views of Gorilla and Chimpanzee anatomy.
Hands of Gorilla.
Feet of Gorilla.
Side view of heads of Anthropoids.
Front views of heads and skulls of Anthropoids.
Map showing location of Brown’s Park beds in northeastern Utah. Views of femur, scapula, and humerus of Aphalops ceratorhinus.
Limb bones of Serridentinus fricki.
Skull and jaws of Serridentinus fricki.
Cranium and jaw of Serridentinus fricki.
Facial region and teeth of Serridentinus fricki.
Restoration of Serridentinus fricki by A. Avinoff .
Vertebrae, skull, humerus, tibia, fibula, radius and ulna of Phena- coccelus typus.
Teeth, manus, pes, skull of Phenacocoelus typus and Merycoidodon culbertsoni.
Pelvis, pes, femur, skull, of P. munroensis and Cyclopidius. Articulated skeleton of Phenacocoelus typus Peterson.
Life restoration of Phenacocoelus typus by Prentice.
Views: Dinosaur National Monument Quarry.
Additional views of Quarry.
Quarry map of Dinosaur National Monument.
Cervical vertebrae of Apatosaurus louisoe.
Dorsal vertebrae of Apatosaurus louisoe.
Caudal vertebrae of A. louisoe.
Caudal vertebrae of A. louisoe.
Caudal vertebrae of Apatosaurus sp.
Dorsal ribs of right side of A. louisoe.
vii
VIU
MEMOIRS OF THE CARNEGIE MUSEUM
XXX.
XXXI.
XXXII.
XXXIII.
XXXIV.
XXXV.
XXXVI.
XXXVII.
XXXVIII.
XXXIX.
XL.
XLI.
XLII.
XLIII.
Dorsal ribs of right side of A. louisce.
Cervical vertebrae of Apatosaurus excelsus.
Dorsal vertebrae of A. excelsus.
Caudal vertebrae of A. excelsus.
Restoration of skeleton of A. louisce by S. Prentice.
Photograph of the mounted skeleton of A. louisce Figures of South American Tingitidce.
Black-figured Column Krater or Kelebe.
Reverse side of same object.
Views of Eye Kylix, Aryballos, Alabastron, Lekythos, and Kotyle. Lekythos, Bell Krater, (Enochoe, Guttus, Jug, Epichycis or Pehke, and Skyphos.
Pelike, reverse, Kylix, and Pinax.
“Canopic” Urn, and Ash Urns.
Ash urns and lids.
GENERA AND SPECIES DESCRIBED AS NEW TO SCIENCE
IN THIS VOLUME
PAGE
Order CARNIVORA
Bassariscops Peterson, gen. nov 96
Type: Phlaocyon willistoni Peterson
Order PROBOSCIDEA
Serridentinus fricki Peterson, sp. nov Ill
Order ARTIODACTYLA
Phenacocoelus munroensis Peterson, sp. nov 161
IX
Publications of the Carnegie Museum, Serial No. 134
MEMOIES
OF THE
OAENEGIE MUSEUM
VOL. XI
NO. 1
W. J. HOLLAND, Editob
STUDIES ON THE GROWTH OF GORILLA AND OF OTHER HIGHER PRIMATES WITH SPECIAL REFERENCE TO A FETUS OF - ' GORILLA, PRESERVED IN THE CARNEGIE MUSEUM
By DR. ADOLPH H. SCHULTZ
Labobatoey of Physical Anthropology, Department of Anatomy,
Johns Hopkins University
(Plates I-VIII and Six Text-figures)
PITTSBURGH
Published by the Authority of the Board of Trustees of the
CARNEGIE INSTITUTE November 14, 1927
MEMOIRS
OF THE
CARNEGIE MUSEUM
VoL. XI. No. 1.
STUDIES ON THE GROWTH OF GORILLA AND OF OTHER HIGHER PRIMATES WITH SPECIAL REFERENCE TO A FETUS OF GORILLA, PRESERVED IN THE CARNEGIE MUSEUM
By Dr. Adolph H. Schultz
Laboratory of Physical Anthropology, Department of Anatomy,
Johns Hopkins University.
(Eight plates and six text-figures.)
CONTENTS
Part 1: Introductory Chapters
Chapter 1. Introduction: Scarcity of fetal ape material 3-5
Chapter 2. Material: Gorillas described in literature and those exam- ined by author, grouped according to relative age. Specimens of other higher primates used for comparison 5-10
Chapter 3. Technique: Description of measurements taken 10-13
Chapter 4. Gorilla fetus H: Notes on fetus of gorilla described for first time; comparison of its measurements with those of human fetuses of corresponding development 13-15
Part H: Growth of Gorilla
Chapter 5. Rate of Growth: General rate of growth before and
after birth in gorilla, chimpanzee, and man 15-18
Chapter 6. Pigmentation: First appearance of pigment of skin in higher primates. Notes on changes due to age in color of skin, eye, and hair 18-20
1
2
Chapter 7.
Chapter 8.
Chapter 9.
Chapter 10.
Chapter 11.
MEMOIRS OF THE CARNEGIE MUSEUM.
Hair; Earl}^ development of hair in gorilla. Hair of head in older fetal, new-born, and infant gorillas and chimpanzees 20-22
Trunk: Changes in proportions of trunk of gorilla due to growth. Coccygeal tubercle in higher primates. Ex- ternal female genitalia in fetuses and adults of apes.
Ischial callosities in anthropoids and gibbons 22-30
Limbs: Growth changes in the proportions of the limbs of the gorilla. Difference between Mountain Gorilla and Lowland Gorilla in regard to inter-membral index.
Relative length of forearm in gorilla and man. Vari- ability of proportions of limbs in the adult gorilla.
Growth changes in hand and thumb of the gorilla. Callosities on middle segments of fingers of the gorilla and the chimpanzee. Growth changes in foot of the gor- illa; relative length of toes; the sole and its flexure-folds; the heel. Differences in formation of foot between Mountain Gorilla and Lowland Gorilla. Loss of arboreal characters in the foot of the adult gorilla .... 30-44
Head: Growth changes in the proportions of the head of the gorilla. The nose of the gorilla compared with the nose of other higher primates. The lips and the ear of fetal gorillas 44-48
Part III: Comparative Studies.
Body proportions of higher primates in fetal and in ADULT life: Explanation and discussion of deviation indices and curves. Relative position of human pro- portions in the range of proportions of higher primates. Differences between primates in limb proportions con- trasted with differences in trunk proportions. Degrees of similarity of higher primates in fetal and in adult life. Relative degrees of difference among higher primates. Relative amounts of ontogenetic change in proportions of apes and man 48-61
SCHULTZ: STUDIES ON THE GROWTH OF GORILLA.
3
Part IV: Chief Results and Their Interpretation.
Chapter 12. Summary and conclusions: Enumeration of findings ap- pertaining to general conditions of growth; body surface; proportions of body; comparisons based upon deviation indices. Phylogenetic conclusions. Pedigree of higher primates 61-67
PART I.
Chapter 1. Introduction.
The inseparable problems of human evolution and of man’s exact place among the primates can be expected to be solved only by a consideration of the findings from all the many different fields of science which have evidence to offer for the reconstruction of the pedigree of the primates. The contributions from the various sciences to our knowledge of the higher primates still differ markedly in amount and significance. The prenatal development of the apes ranks probably first among the phases of which least is known to-day. Indeed, much less informa- tion is available as to embryos and fetuses of the anthropoids than in regard to fossil apes. Few fetuses of apes have been collected and some of these have either never been described at all, or are merely mentioned in a few words. ^ Still other fetuses have been erroneously identified. Breschet (1845)* was the first author to deal with what he supposed to be the fetus of a higher ape; he labelled this specimen “ Semnopithecus Hilobates. Gibbon?”. The writer is firmly convinced that this fetus is not that of a gibbon, but one of some short-tailed macaque.^ On the other hand, the fetus pictured by Darwin (1874) is a gibbon fetus and not that of an orang, as Darwin states, nor that of a chimpanzee, as Schmidt (1892)
* The dates in brackets refer to the Bibliography, which is appended.
* For instance, Lonnberg (1917) lists a chimpanzee fetus in a report on mammals collected in Central Africa; and Prince William of Sweden (1923) mentions that a gorilla fetus was obtained on his expedition, but, as far as the writer is aware, no description of these valuable specimens has appeared as yet. Huxley (1864) devoted only a few words to a chimpanzee fetus; and Anthony (1918), in his detailed description of the brain of a gorilla fetus and of a chimpanzee fetus, gives merely a photograph and a few measurements of the entire specimens.
^ The upper extremity of this specimen is much too short for a gibbon fetus and the tail would no longer be visible in a gibbon at this stage of development (See Chapter 8, coccygeal tubercle). Several other points support the author’s diagnosis; such as the formation of the forehead, of the outer ear, of the foot, and of the placenta.
4
MEMOIRS OF THE CARNEGIE MUSEUM.
tried to prove later on.^ The first thorough description of an anthropoid fetus (orang-utan) is given in the excellent paper by Trinchese (1870). This was followed by Deniker’s (1885) classic monograph on a gorilla fetus. To these have since been added a few more or less extensive reports on other ape fetuses; most note- worthy among which are the papers by Schv'-albe (1911) and by Bolk (1926 b). The rapid and fundamental changes occurring during intra-uterine life necessitate the study of a great many specimens at different stages of development in order to gain even the barest outlines of prenatal growth. Information on the embryonic period in apes is practically restricted to the one gibbon and one orang embryo described by Keibel (1906). In writing a review of the fetal growth of primates the author (1926 b) collected the few scattered data from the literature and com- bined these Avith his own observations in order to give at least a preliminary sketch of some of the fetal conditions in apes.
There can be no doubt that investigations on the ontogeny of the higher primates will be of most essential help in the solution of the problems of primate phylogeny in general and of human evolution in particular, but many more reports on additional material are needed.
In the present paper it is intended first of all to contribute some observations on a new fetus of one of the apes. The author is indebted for this opportunity to Dr. W. J. Holland, who has generously entrusted him with the description of the fetus of a gorilla (Plate I). Since there exist publications on several other gorilla fetuses it seems desirable to discuss them all together and to add to this study of fetal life such data on postnatal growth in the gorilla as are available in the literature and are made possible through an examination of material in various collections. In this attempt to establish the more important changes, which take place during growth in one of the anthropoid apes, it became necessary for a correct interpreta- tion of the findings to compare the conditions in gorilla with those in other repre- sentatives of the higher primates. However, these comparisons could not be made as extensive, as might seem desii-able, on account of tlie scarcity of corresponding data on other apes, which v-ill first have to be augmented and subjected to separate study.
^ Deniker had suspected that this specimen might be a gibbon fetus and not an orang-utan fetus, but Schmidt objected to the “gibbon” determination chiefly on the ground that the ischial callosities were missing. The writer has studied a considerable number of gibbon fetuses and is thus in a position to state that these callosities appear very late in the gibbon (See Chapter 8). The illustration and measure- ments by Schmidt of the specimen in question agree so closely with the author’s data on gibbon fetuses that there can be no doubt of its being a gibbon. Schwalbe was evidently misled by the form of the ear, (which, as Schmidt showed, is due to artificial deformation) in stating that the gibbon fetus, pictured by Darwin, was in his opinion the fetus of a macaque (G. Schwalbe in “Evolution in Modern Thought,” Modern Library, New York, p. 119).
SCHULTZ: STUDIES ON THE GROWTH OF GORILLA.
The entire investigation is restricted to the characters observable on the outer body and will deal particularly with the body proportions, which are of such outstanding interest in studies of growth. Most of the literature appertaining to growth in gorilla deals with the outer body and its proportions. Of the fetuses of gorilla, for instance, only the specimen of Deniker was actually dissected. The fetus, described for the first time in this paper, belonging to the Carnegie Museum has not been dissected. It was hoped at first that the skeletal system of this specimen could be studied by means of X-ray photographs, but since it had been preserved in formalin for the past sixteen years no satisfactory results could be obtained in that way.^
Chapter 2. Material.
As mentioned above, a special effort was made in this study to collect from the widely scattered literature those data which have a bearing upon the problem of growth in gorilla. It will be helpful to other students to enumerate here these sources of information and to establish also the age sequence of all the available material. This sequence is chiefh'' based upon the size of the specimens. Since frequently no measurement of total size has been recorded, the length of the limbs has to serve in such instances as the only possible criterion for the relative age of the specimen.
I. ^ The youngest gorilla fetus on record is the specimen described by Duck- worth (1904 a). This fetus is male, and belongs to one of the subspecies of Gorilla gorilla. The distance from vertex to coccyx (straight) amounts to 71 mm., the length of the upper arm is 21 mm., and the combined length of thigh and leg equals 30 mm. In its state of development this specimen corresponds most closely to human fetuses of the twelfth or thirteenth week (menstrual age).
II. The gorilla fetus described for the first time in this paper is considerably older than Duckworth’s specimen. This fetus is female and corresponds in develop- ment to human fetuses of the beginning of the fifteenth week. The distance from vertex to coccyx measures 85 mm., the sitting height is 88.5 mm., the length of the upper arm is 25 mm., and the total length of thigh and leg amounts to 40 mm.
^ The writer wishes to express his sincere thanks to Dr. E. C. Hill for his persistent and most skilful efforts to obtain good X-ray photographs of this fetus.
The placenta, uterus, and ovaries, which Dr. Holland kindly sent with the gorilla fetus, have been studied by the author’s colleagues. Dr. G. B. Wislocki and Dr. C. Hartman, who will report else- where on this rare material. (A discussion of this gorilla placenta will appear in a paper by Wislocki, 1927, which is now in press).
^ Wherever a particular specimen is referred to hereafter these numbers of age sequence will be given. All the specific names in this paper are according to Elliot (1913).
6
MEMOIRS OF THE CARNEGIE MUSEUM.
All these surpass the corresponding dimensions of fetus I. This fetus belongs to the Carnegie Museum (Acc. No. 7698) and was collected by the medical missionary, Dr. Hymen L. Weber, in the vicinity of Efulen, Cameroon, in 1910. Dr. Holland has kindly informed the author that this specimen belongs either to Gorilla gorilla diehli or to Gorilla gorilla matschiei, but that it is assigned with greater probability to the former subspecies.
HI. According to its size the gorilla fetus of Deniker (1885) ranks third, corresponding in development to human fetuses of the eighteenth or nineteenth week. This fetus is female, was obtained in West Africa, and hence belongs in all probability to the species Gorilla gorilla. The distance from vertex to coccyx measures 135 mm., the upper arm has a length of 53 mm., and the total length of thigh and leg amounts to 82 mm.
IV. The fourth gorilla fetus is that described recently by Bolk (1926 b). Since this author does not state in which part of Africa his specimen was obtained but mentions that it was brought to Amsterdam a long time ago, it can be assumed that it belongs most likely to the Gorilla gorilla group and not to the much rarer and more recently discovered East African Gorilla beringei. This fetus is male and corresponds in size to human fetuses of the beginning of the sixth month, but in regard to certain other characters of development it seems better comparable to human fetuses of the middle, if not end, of the sixth month. The straight distance from the vertex to the anus amounts in this gorilla fetus to 170 mm., the upper arm is 54 mm. long, and femur and tibia combined have a length of 92 mm.
V. The fifth gorilla fetus is in all probability of the same stage of develop- ment as Bolk’s specimen; it may be even slightly younger. Unfortunately, Anthony (1918), who described the brain of this fetus, gives only a photograph of the entire specimen (Photograph reproduced on PL II, fig. 1) and one of a plaster cast thereof. This fetus is male, came from the French Congo, and belongs, therefore, to one of the subspecies of Gorilla gorilla. The distance from the vertex to the coccyx (taken with a tape) is given as approximately 190 mm.; it is to be regretted that no other measurements have been published.
No information of any sort is available in regard to the last part of fetal growth in gorilla.
VI. The next stage in the development of gorilla is represented by the inter- esting photographs of a gorilla baby published by Reichenow (1921). These pictures were taken at intervals from the age of one month to that of seven months. Reichenow’s paper contains no measurements. This specimen was obtained in Cam- eroon and belongs to the species Gorilla gorilla.
SCHULTZ: STUDIES ON THE GROWTH OF GORILLA.
7
VII. The youngest gorilla infant, of which there exist measurements, is the specimen described by Ehlers (1881). It is a male, undoubtedly of the Gorilla gorilla group, according to Ehlers less than one year of age, and measures from vertex to coccyx 308 mm. The upper arm length is 114 mm. and the length of thigh and leg amounts to 172 mm.
VIII. Next in age is the female infant (G. gorilla) measured by Mollison (1911). From the height of the trunk (208 mm.) and the various indices given by Mollison the upper arm length of this specimen is calculated as 146 mm. and the thigh-plus-leg length as 231 mm. These values are very considerably above the corresponding measurements of Ehlers’ infant, thus indicating a marked difference in age between specimens VII and VIII.
IX. The infant described by Deniker (1885) is but little older, i.e., larger, than the infant just mentioned. This specimen is a female with a distance between vertex and coccyx of 405 mm., an upper arm length of 165 mm., and a thigh-plus- leg length of 259 mm. That it must be of the species Gorilla gorilla is evident from the fact that the Mountain Gorilla was still unknown at the time of Deni- ker’s publication.
X. Famelart (1883) has published some few measurements on a male gorilla infant, which he kept alive for a few months. According to this author it was captured when about seven months old and had then a sitting height of 320 mm. which is about the same as in infant VII with its distance from vertex to coccyx (somewhat less than sitting height!) of 308 mm. The head measurements were taken by Famelart about three months later, when the animal had attained a size slightly above that of infant IX, judging by its sitting height which had increased to 410 mm. or more (?). This specimen is assigned to the species Gorilla mayema Alix and Bouvier, or, as called by Elliot (1913), Pseudogorilla mayema.
XI. Information on this stage of growth is available in regard to the pro- portions of the limbs. These proportions were obtained from measurements on three infantile skeletons of Gorilla gorilla. Data for one of these were published by Deniker (1885) and for another by Bolk (1926 b). A third specimen was kindly measured for the author by Dr. W. L. Straus, Jr. in the collection of the Department of Anatomy of Western Reserve University. The lengths of the humeri of these infants average 168 mm. A detailed enumeration of the limb proportions of all the gorilla skeletons, used for this paper, will be found in Table 5. In the tables on growth only the average proportions of these skeletons will be quoted.
XII. The last specimen which could properly be called an infant is the second gorilla {G. gorilla) measured by Mollison (1911). From the trunk height
8
MEMOIRS OF THE CARNEGIE MUSEUM.
and the proportions of this young female the length of the upper arm is calculated as 212 mm. and the thigh-plus-leg length as 322 mm.
XIII. The juvenile male Gorilla gorilla, measured by Bolau (1876), must be considerably older than gorilla XII, since its thigh-plus-leg length amounts to 385 mm. The only other measurement, given by Bolau, which may be used for a determination of relative age, is the combined length of upper arm and forearm amounting to 430 mm. This dimension is calculated as 386 mm. for gorilla XII and amounts to 445 mm. in the next specimen.
XIV. The second juvenile is a male Gorilla heringei which the author has studied at the American Museum of Natural History. This specimen was unfor- tunately skinned and eviscerated so that some measurements could not be taken at all and for the others certain corrections had to be made to allow for the thickness of the skin. These corrections were based upon comparisons between the author’s measurements on a chimpanzee of similar age before and after it was skinned. The sitting height of this gorilla amounts to 619 mm., the distance from vertex to coccyx to 587 mm., the upper arm length equals 254 mm., and the thigh-plus-leg length 387 mm.
XV. The average limb proportions of three juvenile skeletons of Gorilla gorilla represent the next stage. Data on one of these skeletons were given by Alollison (1911) and on another by Bolk (1926 b). The third skeleton belongs to the collection of the Laboratory of Physical Anthropology, Johns Hopkins University. The average length of the humerus of this series, which is 261 mm., stands between the length of the upper arm of gorilla XIV (254 mm.) and that of gorilla XVI (275 mm.).
XVI. Sommer (1907) has published some measurements on an adult female gorilla {“Gorilla castaneiceps Slack” = G. gorilla castaneiceps) . However, this specimen, which had lived in captivity for seven years, is probably not of normal size for its age, since its sitting height, for instance, measures only 610 mm. The upper arm length of 275 mm. is very little above the humerus length of the juvenile skeletons. The thigh-plus-leg length equals 475 mm. The last two dimensions fall below the range of variation of these measurements in other adult female gorillas (XVII and part of series XVIII). Further remarks on the size of this specimen will be found in Chapter 5. Good photographs of this animal, when alive, have been published by Grabowsky (1906).
XVII. The female Gorilla gorilla, described by Ehlers (1881), is the only adult gorilla of typical size in the flesh of which detailed and reliable measure- ments have been published as far as the author could ascertain. This specimen
SCHULTZ: STUDIES ON THE GROWTH OF GORILLA.
9
had a complete second dentition and all the epiphyseal lines of the long bones were obliterated. The distance between vertex and coccyx is given as 700 mm., the upper arm length as 339 mm., and the thigh-plus-leg length as 535 mm.
XVIII. In order to place at least some of the proportions in adult gorilla on a more representative basis than is afforded by the above specimens alone, the average limb proportions of 38 adult skeletons of West African gorillas {G. gorilla and its various subspecies?) have been figured from data in the literature and from measurements of specimens in various collections. Publications of the following authors were consulted in assembling this series: Du Chaillu (1861) one skeleton; Deniker (1885) one skeleton; Mollison (1911) sixteen skeletons; Lorenz v. Liburnau (1917) three skeletons; and Bolk (1926 b) two skeletons. In addition measurements by Dr. W. L. Straus, Jr. on five skeletons in the collec- tion of the Western Reserve University were incorporated and also the results of the author’s examination of nine skeletons in the collections of the U. S. National Museum, the American Museum of Natural History, the Department of Zoology of Columbia University, and the Anthropological Laboratory of Johns Hopkins University, as well as of one skeleton from West Africa, measured some time ago in a dealer’s store in Berlin.
In order to test the question whether there are any specific differences between the proportions of Gorilla gorilla and Gorilla beringei, five adult skeletons of the latter type were measured by the author and their limb proportions compared with those of the above series XVIII. Four of these rare skeletons of the Mountain Gorilla are from the American Museum of Natural History and one from the National Museum. For the purpose of certain further comparisons use was made of measurements by Lorenz v. Liburnau (1917) on the limb bones of five adult Mountain Gorillas of the species Gorilla graueri Matschie.
Two stages of the growth of gorilla were selected for detailed comparison with representatives of the other higher primates at corresponding ages. The first stage is given by the fetus (H) of the Carnegie Museum, the only specimen of prenatal growth of which a complete set of measurements is available. The second stage was chosen to represent the completion of growth, i.e., adult life. The special technique employed for these comparisons required observations on a considerable number of individuals of at least one of the higher primates in order to establish some measure for range of variation. A larger series of cases of the same age is available for man only. The following material, or papers describing certain material, was assembled for these comparisons:
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MEMOIRS OF THE CARNEGIE MUSEUM.
For the stage of fetal growth :
I . Twenty human fetuses (white) from the end of the fourteenth and the beginning of the fifteenth week, ranging in sitting height from 85 to 92 mm., with an average sitting height of 88.5 mm., i.e., exactly the same as in gorilla fetus II.
2. The youngest chimpanzee fetus on record is that described by Friedenthal (1914). This specimen is somewhat larger than gorilla fetus II, but is the only one which can serve for comparison. The distance from the vertex to the symphysis pubis is obtained as 99 mm. by adding certain measurements given by Friedenthal. This dimension corresponds to a sitting height of approximately no mm. Judging by the photograph of this specimen, its stage of development can not differ very markedly from that of gorilla fetus II.
3. Of the four orang-utan fetuses, of which measurements have been published by Schwalbe (1911), specimen No. 3 agrees in its size most closely with gorilla II. The vertex-coccyx length, taken with a tape, amounts in both to 115 mm., the greatest head length is in the orang 35 mm. and in the gorilla 34.1 mm. It can be assumed that the sitting height (straight measurement) of this orang fetus must be also very similar to that of the gorilla fetus, i.e., approximately 88.5 mm.
4. A gibbon fetus {Hylobates concolor) of a stage of development corresponding to that of gorilla II was measured by the author. This specimen belongs to the collection of the late Prof. E. Selenka (No. 24 a). It has a sitting height of 56 mm.
For the stage of adult growth:
5. Twenty-four adult white men from eighteen to twenty-five years of age. These adults, as well as the above mentioned human fetuses (Series i), had been measured by the author in connection with his extensive investigations on human growth. Full details on these series will be published in a later paper.
6. Measurements on a fully adult chimpanzee have been published by Friedenthal (1914). This specimen is a female, weighing 55 kgm.
7. An adult orang-utan in a perfect state of preservation was measured by the author at the American Museum of Natural History (Department of Comparative Anatomy, No. 563). This ape is a female and has a sitting height of 717 mm.
8. The adult gibbon used in this paper was also measured by the author. This specimen is of the same species {Hylobates concolor) as fetus 4 and belongs also to the Selenka collection (No. 16 a). It has a sitting height of 237 mm.
The author wishes to take this opportunity to express his sincere thanks to Mrs. E. Selenka, Prof. W. K. Gregory, Prof. J. H. McGregor, and Mr. G. S. Miller, Jr., for their ready and generous permission to study the valuable material mentioned above. The writer is furthermore indebted to his former student. Dr. W. L. Straus, Jr., who kindly measured for him the gorilla skeletons in the collection of Western Reserve University, and to Prof. T. W. Todd, who very obligingly permitted this.
Chapter 3. Technique.
The measurements taken on this primate material correspond, wherever possible, to those generally used in anthropometry (See Martin, 1914). Many measurements have to be taken in order to express accurately the dimensions
SCHULTZ: STUDIES ON THE GROWTH OF GORILLA.
11
and proportions of every part of the body. It is very essential that such measure- ments are defined precisely, so that the recorded data are not open to misinter- pretation. Unfortunately it is often very difficult, and sometimes impossible, to make use of some measurements given in the literature, simply because authors have omitted to state their methods of measuring.
The following list of measurements explains the technique adopted by the author. The diagram on PL III, fig. 1 obviates a detailed description of many dimensions. The specimen, no matter of what age, is first placed flat on its back with its spine practically straight and its head posed so that an imaginary plane, determined by the ear openings and the lowest point of one orbit (ear-eye horizon), stands perpendicular to the body axis. All the “height” measurements on the trunk and the head are taken strictly parallel to the body axis, as represented by the table on which the specimen rests.
General measurements:^
1. Sitting height: from vertex to buttocks over ischial tuberosities (See PI. Ill, fig. i).
2. Stature: can be taken directly only in man after birth. In all primate fetuses and in all apes the knees are flexed so that the approximate stature is best obtained by the addition of measurements 3, 1 6, and 17.
3. Cephalo-thoraco-abdominal height: vertex to upper edge of symphysis pubis (symphysion).
Trunk measurements:
4. Anterior trunk height: suprasternal notch to symphysion.
5. Acromion height: middle of line connecting the lateral points on acromial processes of shoulder blades (acromion) to symphysion.
6. Shoulder height: projective height of acromion over suprasternal notch (obtained by subtraction of m. 4 from m. 5).
7. Nipple height: middle of line connecting centers of nipples to symphysion.
8. Umbilicus height: center of attachment of umbilical cord or center of umbilical scar to symphysion. This measurement can not be taken in many adult apes, since they frequently have no trace of an umbilical scar left.
9. Shoulder breadth: distance between right and left acromion.
10. Hip breadth: distance between most lateral points over the great trochanters.
11. Nipple breadth: distance between centers of nipples.
12. Transverse chest diameter: at level of sternal attachment of fourth pair of ribs.
13. Sagittal chest diameter: at same level as m. 12.
14. Chest circumference: at same level as m. 12 and m. 13.
Limb measurements:
15. Total lower limb length: obtained by addition of m. 16 and m. 17.
16. Thigh length: length of femur taken laterally from top of great trochanter.
17. Knee — sole length: length of tibia plus foot height, taken medially.
18. Leg length: length of tibia to tip of internal malleolus.
® Wherever the absolute measurements are referred to hereafter their serial number will be given together with the letter m to distinguish them from the relative measurements, or indices, the numbers of which will be accompanied by an i.
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MEMOIRS OF THE CARNEGIE MUSEUM.
19. Foot length: from heel to tip of longest toe.
20. Great toe length: from heel to tip of hallux (hallux adduced).
21. Foot breadth: greatest width between metatarso-phalangeal joints of toes II to V.
22. Total upper limb length: obtained by addition of m. 23, m. 24, and m. 25.
23. Upper arm length: length of humerus from acromion to humero-radial joint.
24. Forearm length: length of radius to tip of styloid process.
25. Hand length: from middle of line connecting tips of ulnar and radial styloid processes to tip of middle finger.
26. Thumb length: from styloid process of radius to tip of thumb (thumb adduced).
27. Hand breadth: greatest width between metacarpo-phalangeal joints of fingers II to V.
Head