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Full text of "Transactions of the Zoological Society of London"







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THIS BOOK MAY NOT BE PHOTOCOPIED . 



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, IT. „ ar — lib ^ 31— W3 iizL^). 



W 



TRANSACTIONS 



OF 



THE ZOOLOGICAL SOCIETY 



OF LONDON. 



VOLUME VI. 




LONDON: 

PRINTED FOR THE SOCIETY : 

SOLD AT THEIR HOUSE IN HANOVER- SQUAR E : 
AND BY MESSRS. LONfiMANS, GREEN, RE.ADER, AND DYER, PATERNOSTER-ROW. 



18G9. 



/■ ' 

O - Z8. 
I. 



PRINTED BY TAi'LOR AND FRANCIS, 
RED HON COURT, FLEET STREET. 



CONTENTS. 



I. On the Characters and Affinities of Potamogale, a genus of Insectivorous Mammals. 

By George J. Allman, F.B.S., Corr. Mem. Zool. Soc. Lond., Regius Professor of 
Natural History in the University of Edinburgh, and Regius Keeper of the Edin- 
burgh Museum of Natural History page 1 

II. On some Indian Cetacea collected by Walter Elliot, Es^. By Professor Owen, 

F.R.S., F.Z.S., &c 17 

III. On the Osteology of the Dodo (Didus ineptus, Linn.). By Professor Owen, E.R.S., 

F.Z.S.,&c 49 

IV. Description of the Skeleton of Inia geoffi-ensis and of the Skull of Pontoporia 

blaiiivillii, witJi Remarks on the Systematic Position of these Animals in the 
Order Cetacea. By William Henry Flower, F.R.S., F.R.C.S., F.Z.S., <&c., 
Conservator of the Museum of the Royal College of Surgeons of England . 87 

V. On a Raptorial Bird transmitted by Mr. Andeksson/?w?i Bamara Land. By J. H. 

Gurnet, i^.Z.^f H'' 

VI. On some Fossil Birds from the Zehbug Cave, Malta. By W. K. Parker, F.R.S., 

F.Z.S., &c 119 

VII. Synopsis of the species of recent Crocodilians or Emydosauriaus, chiefly founded on 

the specimens in the British Museum and the Royal College of Surgeons. By 
Dr. JonN Edward Gray, F.R.S., V.P.Z.S., F.L.S., &c 125 

VIII. Note to Memoir on the Indian Cetacea collected by Sir Walter Elliot. By Pro- 

fessor Owen, F.R.S, F.Z.S., &c 171 

IX. Contributions towards a more complete knowledge of the Skeleton of the Primates. 

By St. George Mivart, F.L.S., Lecturer on Comparative Anatomy at St. Marys 
Hospital. Part 1. The Appendicular Skeleton of Simla 175 

X. Description of the Remains of three extinct Species o/ Elephant, collected by Captain 

Spratt, C.B., R.N., in the Ossiferous Cavern of Zebbug, in the Island of Malta. 
By George Busk, F.R.S.; partly from the Notes of the late IT. Falconer, M.D., 
F.R.S 227 



iv CONTEKTS. 

XI. On a Species of Dormouse (Myoxus) occurring in the Fossil state in Malta. By 

A.'Lmivi Kvms, M.B., F.G.S page 307 

XII. On the Osteology of the Cachalot or Sperm-Whale (Physeter macrocephalus). By 

William Henry Flower, F.R.S., F.B.C.S., F.L.S., F.G.S., F.Z.S., Conservator 
of the Museum of the Boyal College of Surgeons of England 309 

XIII. On a Picture sujaposed to represent the Bidine Bird of the Island of Bourbon 
{Reunion). By Alfred Newton, M.A., F.L.S., F.Z.S., &c 373 

XIV. An account of the Fishes of the States of Central America, based on collections 

made by Capt. J. M. Dow, F. Godman, Esq., and O. Salvin, Esq. By Albert 
GtJNTiiER, M.A., M.B., Ph.D., F.E.S., F.Z.S 377 

XV. On DiNORNis (Part XI.) : containing a Description of the Integument of the Sole, 

and Tendons of a Toe, of the Foot of Dinornis robustus, Ow. By Professor 
• Owen, RR.S., F.Z.S., &c 495 

XVI. On Dinornis (Part XII.) : containing a Description of the Femur, Tibia, and Meta- 

tarsus o/ Dinornis maximus, Otven. By Professor Owen, F.B.S., F.Z.S., &c. 497 

XVII. On the Osteology of the Kagu (Rhinochetus jubatus). By W. K. Parker, 
F.R.8., F.Z.S. . '. 501 



TRANSACTIONS 



OF 



THE ZOOLOGICAL SOCIETY 



I. On tJie Characters and Affinities of Potamogale, a genus of Insectivorous Mammals. 
By George J. Allman, F.R.S., Corr. Mem. Zool. Soc. Lond., Begins Professor of 
Natural History in the University of Edinlurgh, and Begins Keeiiei' of the Edinhitrgli 
Museum of Natural History. 

Eead June 13th, 1863. 

[Plates I. & II.] 

A SMALL mammal, with which I was entirely unacquainted, was lately placed in my 
hands by Mr. Archibald Hewan, who had just returned to this country from the west 
coast of Africa, where he had been for some time residing, at Old Calabar, in the 
capacity of medical missionary. The notes which accompanied the specimen were 
scanty. It is stated to have been observed by one of the natives on the banks of a 
stream, when it was pursued, and killed, and taken to the missionary-station, where, 
after having been partly eviscerated, it was put into spirits. In this condition it was 
brought to England by Mr. Hewan, along with various other objects of uiterest fi'om 
the same quarter*. 

A little examination was sufficient to show that the Old Calabar mammal belonged 
to the order Insectivora, but that, with a well-marked insectivorous organization, it 
possessed chai'acters of a very peculiar kind, and such as separated it widely from every 
genus hitherto rei'en-ed to this order. 

While engaged in preparing a description of the new insectivore, I showed the speci- 
men to Mr. Sclater, the accomplished Secretary of the Zoological Society, who at once 
recognized it as identical with a very badly preserved skin which had been brought 
* The specimen is now preserved in the Edinburgh Museum of Natural History. 

VOL. VI. — PART I. B 



2 PEOFBSSOK ALLMAN ON THE CHARACTERS 

over by Mr. Du Chaillu from tropical Africa, and which forms at present part of the 
collection in the British Museum. This skin, however, is in a wretched condition ; the 
skull and teeth are altogether absent, and tlie specimen is otherwise mutilated ; so that 
it had been quite impossible to obtain from it characters which might enable its zoolo- 
gical affinities to be satisfactoiily determined. 

Mr. Du Chaillu, however, had already described the animal from his mutilated skin, 
aided by his recollection of it when alive ; and I am indebted to Mr. Sclater for having 
directed my attention to an article* in the ' Proceedings of the Boston Society of 
Natural History,' in which the African traveller describes, among other animals from 
equatorial Africa, that to which the skin in question belonged, referring it to the 
carnivorous genus Ci/iwgale, under the name of Cynogah veJox, Du Chaillu. Mr. Du 
Chaillu, however, is not without doubt as to the correctness of assigning his new animal 
to the genus Cynoqale ; and having in view the possibility of its being aftenvards deemed 
desirable to construct for it a separate genus, he suggests the name of Voiamocjale as a 
provisional generic appellation. 

The skin having been subsequently secured for the British Museum, it was examined 
by Dr. J. E. Gray, who disputed the justice of Du Chaillu's determination of its 
affinities, and maintained not only that it had no relation with Ci/hogale, but that it 
probably did not even belong to the order Carnivora, wliile he suspected that its real 
relations would be found with the Rodentia. Under this impression, he proposed for it 
a new generic name ; and the Ci/tiognle velux of Du Ciiaillu became the Mythomys velo.v 
of Gray f . 

Dr. Gray's characterization of his new genus is much more correct tiian that given by 
Mr. Du Chaillu ; but, as just said, not a remnant of the dentition had been left in the 
skin, which was in other respects so very imperfect that it can afford no matter of surprise 
to find so experienced and excellent a zoologist as Dr. Gray failing to discover its true 
affinities ; and it is only the chance which has thrown a comparatively well-preserved 
specimen into my hands that has enabled me to determine the real position and rela- 
tions of this remarkable mammal. 

It is not always that provisional names ought to be accepted ; they are not unfre- 
quently a mere subterfuge, in which the ignorance or incapacity of the describer of 
some new species seeks to take refuge without liis thereby abrogating his claim to be 
regarded as the original namer, though sounder views of the obvious facts may prove 
the incorrectness of his determination. But when, as in the present case, the actual 
absence of data renders it impossible to determine important characters, the describer is 
quite justified in making the best of the material at his disposal, and, by the suggestion 
of a provisional name, reserving to himself the right of giving this name to his 
discovery, if further facts rendering it expedient sliould be brought to light. 

* Du Chaillu, " Ou Animals from Equatorial Africa believed to be new," Proc. Bost. Soc. Nat. Hist., 
vol. vii. p. 353. f Proc. Zool. Soc, 1861, p. 275. 



AND AFFINITIES OF POTAMOGALE. 6 

It is exactly in this position that Mr. Du Chaillu's name of Potamogale stands : it 
has thus precedence over Gray's name of Mijthomys ; and the laws of natural-history 
nomenclature compel us to accept it. The synonomy of Mr. Du Chaillu's animal Mill 
accordingly stand as follows: — Potomogale (prov. gen.) velox, Du Chaillu,= CyHoy«/^' 
velox, Du Ch^\\\\i,-=iMytJiomys velox. Gray (gen.). 

External Characters and Teeth. 
Potamogale velox (Plate I.) is somewhat larger than a stoat ; it has very much the 
aspect of a small otter, but is rendered especially striking by its broad, almost spatuli- 
form muzzle and its very large laterally compressed tail. Both fore and hind limbs are 
short and nearly equal to one another in length. The body is clothed with somewhat 
coarse but soft hair, which projects from a shorter dense coat of very fine silky hairs ; 
and the same kind of clothing covers the base of the tail as far as an oblique line which 
terminates below at about an inch behind the vent, and above at about an inch still 
further back; the whole of the rest of the tail is covered with short, coarse, closely 
appressed hairs. The sides of the upper lip give origin to stiff bristle-like whiskers, 
which commence at the point of the nose, and continue to be borne as far back as a 
point nearly vertically over each angle of the mouth, increasing in length and thickness 
from before backwards ; the most anterior are short and incline forwards, and they then 
acquire more and more of a backward direction until we find the most posterior 

Fig. 1. 




Hair from the body of Potamogale velox.— A, one of the longer hairs, magnified about ten diameters ; B, a por- 
tion of one of the shorter and ner hairs, magnified about 40 diameters to show its structure ; a\ a piece 
from near the middle of the narrow basal portion of A ; a% from the middle of the broad terminal lamina : 
and a', the terminal portion of the lamina : the last three magnified about 40 diameters. 

attaining a length of nearly two inches, and inserted so obliquely that their tips are 
nearly an inch behind the ears; a few stiff bristles also arise from the cheeks, a little 
below and in front of the ears. The underside of the muzzle is clothed towards its 



PROFES.SOR ALLMAN ON THE CHARACTERS 



extremity with very short hairs, which gradually increase in length as they approach the 
angles of the mouth. The upper side of the head, with the back and the entire tail, 
and the outer side of the fore and hind limbs are dark brown. The whole of the 
underside of the body, from the extremity of the nose to the vent, is brownish yellow. 

The fine hairs which constitute the shorter and denser coat are seen under the 
microscope to be of uniform thickness, -with the cortical substance presenting an im- 
bricated structure, and the cells composing the medullary substance so disposed as to 
give a septate appearance to the interior of the hair (fig. 1 b). The long hairs, A, which 
project from this coat have a remarkable form : commencing very thin at the bulb, 
they gradually increase in thickness for about a third of the entire length of the hair, 
then suddenly contract, and immediately after expand into a broad lanceolate lamina, 
which terminates in a fine point. The basal portion of these hairs has a thin imbri- 
cated cortical investment and medullary contents, which consist of an aggregation of 
small spherical cells («'). In the broad lamina, the cortical portion has acquired greater 
thickness, has lost its imbricated character, and is seen to be composed of minute longi- 
tudinally arranged fusiform cells ; the medullary substance is here composed of aggre- 
gated spherical cells like those of the basal portion of the hair, and dies out before it 
reaches the point (or, a'). The remarkable difference thus observed between the two 
kinds of hair presents us with a condition not unusual among the Insectivora, and one 
which finds its maximum in the aculeate genera of this order. 

Fig. 2. Fij;. 3. 




Head viewed from above. Head viewed from below. 

The muzzle is long and broad, and so much appressed as to acquire a somewhat spatu- 
late form (figs. 2, 3). It projects in front for half an inch beyond the extremity of the 
lower jaw, and for more than a quarter of an inch beyond the jaw at its sides ; the angles 
of the mouth are situated at about a quarter of an inch in front of a vertical line from 
the eyes. Each nostril opens beneath the external edge of a cartilaginous valve, which 



AND AFFINITIES OF P0TA:\I0GALE. 




Right ear, enlarged. 



extends over it from the septum, and by which it may be completely closed ; the two 
valves form together a heart-shaped, naked shield by which the muzzle is terminated. 

The ears (fig. 4) are inserted about half an inch behind the eyes, and project for about 
the same distance from the head. They are rounded, Fig. 4. 

the breadth being about two-thirds of the height ; 
and at about one-fourth from the summit they have 
a deep notch on their posterior edge : the upper 
fourth is quite naked, but the rest is clothed with 
silky hairs. The helix is distinct anteriorly and 
posteriorly, but is obsolete towards the tip ; the 
anthelix is represented by a short, nearly trans- 
verse ridge ; tlie tragus is indistinct, but the anti- 
tragus is well developed. 

The eyes are very small. The opening of the 
eyelids, when fully expanded, is one-tenth of an 
inch in its antero-posterior, and a little less in its 
vertical diameter; it leads into an oval palpebral 
chamber, which extends for some distance beneath the anterior and posterior margins, 
somewhat further posteriorly than anteriorly. In the specimen, the globe of the eye was 
retracted into this cavity, and thus rendered difficult to detect ; it is about one-twelfth of 
an inch in diameter, and, so far as could be determined from the state of the specimen, 
is completely developed, and receives an optic nerve fully proportioned to its size. 

The fore limbs, as far as the wrist, are clothed with moderately long hair, which on the 
back of the metacarpal bones becomes very short and appressed, and is thus continued 
over the back of the fingers as far as the claws ; the whole of the palm, with the under- 
side of the fingers, is naked (fig. 5 a). The fingers are five in number, and are connected 
at their bases by a ^'ery narrow extension of the skin, but nothing like a distinct web is 
developed ; they gradually increase in length from the outer finger to the middle, which is 
the longest of all ; the index is a very little shorter than the 
annularis ; and the pollex, which is inserted a little further 
back than the index, is the shortest. The claws are of mo- 
derate size, nearly equal on all the fingers, compressed, curved, 
and with a funow on the underside. 

The hind limbs are clothed, as far as the tarsus, with 
moderately long hair, which becomes short and appressed 
upon the entire back of the foot as far as the claws ; the entire 
sole of the foot is naked (fig. 5 b). The toes arc five in 
number ; the outer and inner toes are the shortest, the 
inner being a little shorter than the outer; the second, 
third, and fourth are nearly equal to one another in length ; the second and third are 





Feet, jjlantar surface. — 

A. Kight anterior. 

B. Eight posterior. 



6 PEOFESSOE ALLMAN ON THE CHAEACTEES 

united by their opposed surfaces for the entire length of the first phalanx, a very narrow 
extension of the skin existing at the base of the other digital intervals ; the claws are 
of the same form as those of the fore limbs, but are a little longer. The outer edge of 
the sole projects as a narrow membranous border along the whole of the metatarsal 
region. 

The length of the tail, measured from the posterior margin of the vent, is equal to 
the distance from the same point to the middle of the throat. It is so thick at its 
base that the trunk seems uninterruptedly continued into it ; but it soon becomes 
laterally compressed, and then grows gradually thinner and narrower towards the tip ; 
immediately in front of the vent it is nearly cylindrical, with a diameter of about l^^ 
inch ; from this point it gradually thins away, and, at an inch beyond it, its vertical 
height is 1^^ inch, and its breadth ^q- inch; while at three inches from the same 
point its height is 1 inch, and its thickness -^ inch ; its lower edge is rounded, and 
its upper is continued into a membranous crest of about |^ inch in height, and clothed 
with the same short, stiff, appressed hairs with which the distal part of the tail is 
covered. 

Teeth.- — The determination of the dental formula is not without difficulty. . The incisor 
teeth of the upper jaw (PI. II. & fig. 7, p. 10) can be easily ascertained by the limits of the 
premaxillary, whose suture with the maxillary continues very distinct. They will be found 
to be three on each side, though one of them closely resembles a large projecting canine ; 
so also to the first three teeth on each side in the lower jaw the same significance must 
be assigned. A difficulty, however, lies in the tooth which in each jaw succeeds to the 
incisors. In position it is a canine, but in form it is a premolar. It follows close upon the 
third incisor, without the intervention of any distinct diastema ; but in the upper jaw it is 
two-fanged, and in other respects is entirely similar to the premolar which follows it ; 
in the lower jaw, however, it is implanted by a single fang, and does not so entirely 
resemble the succeeding premolar as in the upper jaw ; this lower tooth passes imme- 
diately in front of the upper one when the jaw is closed, and must certamly be regarded 
as its equivalent. Considering, therefore, that in the upper jaw the tooth in question 
is absolutely similar both in its root and in its crown to an indubitable premolar, 
I believe we may safely regard it and its corresponding tooth in the lower jaw as 
premolars rather than canines; and the dentition of Potamogale will then present 
a series in which the canine teeth are suppressed, and which may be formulated as 
follows : — 

. 3_3 0-0 3-3 3-3 5^ 
'■3Z3'^-030'^-3=3''"-3=3'=^'^- 

In the upper jaw the first incisor resembles a canine ; it projects more than any other 
■tootli in the jaw ; it is conical and pointed, converging above from its base toward its 
fellow, and then diverging below so as to form a curve whose concavity looks outwards ; 
it is curved also in another direction, having the concavity looking backwards. The 



AND AFFINITIES OF POTAMOGALE. 7 

second incisor is separated from the first by a narrow space which receives the second 
incisor of the lower jaw when the mouth is closed ; it is triangular, compressed, with a 
sharp anterior and a sharp posterior edge — the anterior edge being convex, and the 
posterior slightly concave. The third incisor is of the same form as the second, but a 
little smaller. The incisors are each implanted by a single fang. 

Since in the view here adopted the canine is supposed to be absent, the first pre- 
molar follows immediately on the third incisor, with an interval so slight as to have no 
claim to be regarded as a distinct diastema ; it is inserted by two fangs ; its crown is 
slio-htly larger than that of the third incisor, but otherwise it resembles it. The second 
premolar is also implanted by two fangs, and is otlierwise similar to the first. The third 
has the form of a triangular pyramid, with a small cusp developed from the posterior 
internal basal angle, and another from the posterior external ; it is implanted by three 

fangs. 

The first, second, and third true molars are similar to one another : they are trian- 
o-ular in horizontal section, with the apex of the triangle situated internally; the 
greatest antero-posterior diameter of the crown is to its transverse diameter as 2 : 3 ; 
the internal- angle of the crown presents a single cusp ; the centre, two ; and the 
external side projects downwards as a tuberculate ridge ; they are each implanted by 
three fangs. 

In the lower jaw (PI. II. & fig. 8, p. 11) the incisors present, as in the upper, the 
usual single-fanged insertion. The first is very small, chisel-shaped, and with its crown 
converging to that of its fellow. The second incisor is high, conical, curved, with the 
concavity of the curve looking backwards, and presenting from its base to its apex two 
surfaces separated by a sharp ridge ; it is sharp-pointed, and resembles a canine ; it is the 
most projecting tooth in the jaw. The third incisor is small, irregularly conical, convex 
anteriorly, concave posteriorly ; it is the smallest of the teeth, except the first incisor. 

The first premolar is triangular, compressed, with a sharp convex anterior edge and a 
sharp concave posterior edge ; it has but a single fang. The second premolar is trian- 
gular, compressed, with sharp anterior and posterior edges ; it is implanted by two 
fangs ; its crown is a little lower than that of the first, but it otherwise resembles it. 
The third premolar is a little larger than the first, triangular, compressed, with sharp 
anterior and posterior edges ; the anterior and posterior basal angles have each a small 
tubercle ; it is implanted by two fangs. 

The first, second, and third true molars are prismatic, and equal in height to the 
second premolar ; the crown is furnished with three cusps in fi-ont and a single one on 
a lower level behind. They are each implanted by two fangs. 

A very striking aspect is given to the dental series by the form of the crowns of the 
second, third, fourth, and fifth teeth of the upper jaw, which are all triangular, much 
compressed, with sharp anterior and posterior edges, thus vi\'idly reminding us of the 
teeth of certain sharks. In this respect they resemble the premolar teeth of the viver- 



8 PEOFESSOK ALLMAN ON THE CHARACTERS 

ridan genus CynofiaJe — a resemblance which did not escape Mr. Du Chaillu, and which, 
doubtless, decided him in referring his animal to that genus. 

The characters thus presented by Potamogale velox would justify a belief -in the 
aquatic habits of the animal. Indeed it is scarcely possible to connect with any other 
mode of life the valvular nostrils, and broad, strong, vertically flattened tail, with (as 
will be presently seen) its greatly developed hsemal arches. The trenchant incisors and 
premolars, so like the teeth of a shark, also point to the same conclusion, and indicate 
a diet exceptional among the Insectivora*. 

Skeleton. (Plate II.) 

Cervical Beyion. — The transverse process of the atlas is broad and flattened horizon- 
tally. The neural spine is reduced to a mere tubercle. The body of the axis is carinated 
below ; its transverse processes are short, narrow, and directed backwards, while the 
neural spine forms a large, vertical, laterally compressed, sharp-edged, and hatchet- 
shaped plate. In the third cervical, the transverse process is longer and thicker. In 
the fourth and fifth, the pleurapophysis forms a flat process coalescent with the diapo- 
physis, and extends forwards with a sharp angle, so as to slightly underlap the vertebra 
in front ; while in the sixth it becomes much larger and hatched-shaped, and extends 
backwards so as to underlap the seventh. The transverse process of the seventh has 
no canal for the vertebral artery, and consists of a simple stiliform diapophysis. The 
neural spines of the third, fourth, and fifth cervical are very short, those of the sixth 
and seventh longer. From the inferior surface of the body of the third, fourth, and 
fifth cervical a prominent hypapophysis is developed, which becomes smaller in the sixth 
and seventh. 

Dorsal and lumbar Rerjion.—ThQve are sixteen, dorsal and five lumbar ^•erteb£e. The 
commencement of a metapophysis shows itself in the second dorsal, acquires greater 
length in the third, still greater in the fourth, and then continues of equal length as a 
long blunt process on every vertebra as far as the twelfth ; on the thirteenth dorsal it 
becomes shorter, and is here associated with a short anapophysis, and then continues 
of the same length, but broader, on the fourteenth, fifteenth, and sixteenth dorsal, and 
on the whole of the lumbar. On the last three dorsal vertebra; the anapophysis 

* Tlie account wliicli Mr. Du Chaillu ijue. cit.) has given us of the habits of his Potamognle relax is entirely 
in accordance with what the structure of the auimal would suggest. " This extraordinary animal," he says, 
"is found in the mountains of the interior, or in the hilly country explored by me north and south of the 
equator. It is found along the water-courses of limpid and clear streams, where fish are abundant ; it hides 
under rocks along these streams, lying in wait for fish. It swims through the water with a rapidity which 
astonished me ; before the fish has time to move, it is caught. On account of the rapidity of its movements, 
I have given it the specific name of velox. The animal returns to land with its prey almost as rapidly as it 
started from its place of concealment. The great motive power of the animal in the water seems to be in 
its tail." 



AND AFFINITIES OF POTAMOGALE. 9 

becomes separated by a wide intei-val from the metapophysis, and then disappears on 
the first lumbar. No diapophyses are developed on the fifteenth and sixteenth dorsal ; 
but they reappear on the first lumbar, and constitute broad, flat processes, directed 
downward and outward and a little forward, on all the lumbar. The diapophysis of 
the first lumbar is terminated by a flat, nearly square pleurapophysis, which appears 
as a simple continuation of the diapophysis, but whose line of junction Avith it still 
remains distinct. 

The neui-al spine of the first dorsal is nearly vertical, long, and slightly compressed ; 
those of the second, third, and fourth are equal to it in length, and of a similar form, 
but incline more backwards, the inclination gradually increasing to the fourth ; from 
the fifth to the tenth, the neural spines are stiliform, gradually decreasing in length, 
and incline so much backwards that the anterior rests upon the posterior; and the 
vertebrae here present a remarkably imbricated appearance ; from the eleventh to the 
thirteenth, the neural spines are shorter, and incline less backwards; they then assume 
the form of laterally compressed vertical plates, gradually increasing in size to the fii-st 
lumbar, whence they continue of nearly the same form and size to the fifth lumbar. 
From the body of the first dorsal a small hypapophysis is developed ; it becomes some- 
what larger on the second, third, and fourth, is reduced to a nearly obsolete keel on 
the fifth, and then entirely disappears. 

The ribs are sixteen in number, of which the first nine articulate with the sternum ; 
the remainder are free. The first is the shortest and stoutest ; its cartilage is broad 
and flat, and articulates with the manubrium ; of the remaining ribs, the last two arti- 
culate with the bodies only of their respective vertebrae, while the others articulate 
also with the transverse processes. The sternum is composed of eight pieces : the 
manubrium is spade-shaped ; from the second to the sixth, they form quadilateral prisms, 
gradually decreasing in length and increasing in breadth ; the seventh is nearly cubical, 
with two surfaces posteriorly for the articulation of the cartilage of the eighth pair of 
ribs ; the eighth piece is long and appressed, and carries the small xiphoid cartilage on 
its extremity. 

Sacral and caudal Regions. — There are three sacral and thirty-three caudal vertebrse, 
the ossa innominata being united to the first and second sacral. The neural spines, 
metapophyses, and diapophyses continue to be well developed on the sacral and for 
some distance on the caudal vertebrae; they then gradually diminish on the caudal 
vertebrte with the diminishing size of these, until towards the end of the tail they dis- 
appear, and the vertebrae become reduced to minute centra. All the caudal vertebrae, 
from the second to about the twenty-thu-d, are provided with chevron bones : towards 
the proximal end of the tail these bones are remarkably large; they then gradually 
diminish in size, and become mere rumdients before their final disappearance. They 
are each articulated in an intervertebral space ; most of them develope a short haemal 
spine, and send off at each side from their lower surface a broad horizontal plate. 

VOL. VI. PART I. c 



10 



PROFESSOE ALLMAX ON THE CHARACTERS 



FiK. 6. 



The Skull— Yie^yeA in its vertical aspect, the skull presents a piriform shape between 
the occiput and a line immediately behind the orbits, and then, becoming suddenly con- 
tracted, it is bounded by parallel sides as far as the end of the muzzle, interrupted, 
however, by the projection of the posterior part of the alveolar 
border of the maxillaries. The profile contour of the skull, from the 
lambdoidal crest to the nostril, is nearly a straight line. 

The basioccipital is thin and flat, broader than long, and extends 
forwards as far as the junction of the posterior and middle thirds of 
the tympanic bulla3. The occipital condyles are large, about a line 
distant from one another below ; and thence extending upwards and 
outwards, they reach a point a little above the level of the superior 
margin of the foramen magnum. The foramen magnum is trans- 
versely oval ; its plane extends upwards and backwards at an angle of 
about 100° with the base of the skull. The supraoccipital extends 
upwards and forwards, and forms by its upper and outer edge a well- 
marked, sharp, lambdoidal ridge. The paramastoids constitute two 
small but well-marked processes, which extend horizontally backwards. 
The anterior condyloid foramina are very large. 

The basispenoid is broad behind at its junction with the basioccipital, and then 

rapidly contracts as it passes forwards, forming on the cerebral aspect a narrow vertical 

crest between the internal openings of the foramina lacera antcriora : there are no 

clinoid processes. 

Fis. 7. 




Skiill, vertical as- 
pect: nat. size. 




Skull, basal aspect : twice the nat. size. 

The tympanic and petrosal bones unite to form tymj)ano-petrosal bullae of moderate 
size. 

The sagittal suture is obliterated, its place being taken by a nearly obsolete sagittal 
crest. 

The coronal suture is very faintly indicated by a line which forms an arch, very con- 
cave in front, where it embraces the posterior margin of the frontal bones. These are 



AXD AFFINITIES OF POTAMOGALE. 



11 



nairoM', forming by their union a very convex margin posteriorly, which is received 
between the parietals and a deep notch anteriorly, which receives the nasal bones. 
The frontals are entirely excluded from the orbits by the anterior extension of the 
parietals, which, passing between them and the lachrymals, are separated from the 
maxillaries by a very narrow extension of the lachrymals, which ascends to unite with 
the frontal. The frontal suture is obliterated posteriorly, but anteriorly it continues as 
an harmonia. The nasal bones are long and flat, forming a very convex edge posteriorly, 
where they are received between the frontals, while theii- anterior free edge presents a 
wide semicircular notch. The nasal suture, except for a short distance posteriorly, is 
entirely obliterated ; the external edges of the single bone, thus formed, are nearly 
straight and parallel. 

The zygomatic process of the squamosal forms a small, horizontal, triangular plate, 
whose lower side affords a surface for the glenoid cavity. This cavity is bounded behind 
by a broad vertical process, which checks the retraction of the mandible ; the axis of 
the cavity is inwards and slightly forwards. 

The facial plate of the maxillary is united internally with the premaxillary, tlie 
nasal, and the fi-ontal, anteriorly with the premaxillary, and j,. 

posteriorly with the lachrymal. The alveolar margin for the 
hindmost four teeth projects outwards and backwards, form- 
ing) by its coalescence with the rudimental malar, a com- 
pressed, sharp-edged process. There is no zygomatic arcli. 
The antorbital foramen is very large. The orbits are very 
badly defined ; they are marked by no postorbital process, 
and are continued without interruption into the wide tem- 
poral fossa. 

The palatine plates of the maxillary form the greater por- 
tion of the palate ; the palatines form the posterior third, and 
the premaxillaries about a sixth. Two large incisive notches 
exist in the premaxillary, and are completed into foramina 
by the anterior edge of the palatine plate of the maxillary. 

The pteiygoid ridges converge from before backwards, 
and enclose a deep, narrow interpterygoid fossa, whose roof 
is continued without interruption into the inferior surface 
of the basisphenoid and basiocciptal. 

The horizontal ramus of the mandible is straight, with its upper and lower edge 
parallel ; it forms with its fellow an acute angle, with a rather long and very oblique 
symphysis. The condyle is borne on a distinct neck ; its axis is directed inwards and 
slightly downwards and forwards. The posterior margin of the ascending ramus is thin, 
and runs from the neck of the condyle upwards and slightly forwards to the coronoid 
process, and downwards and backwards to the prominent hook-like angle. The anterior 

c2 




Lower jaw, twice the nat. size. 



12 PROFESSOK ALLMAX ON THE CHARACTERS 

edo-e of the coronoid process runs downwards and slightly forwards, with a convex curve ; 
it meets the horizontal ramus at about a line behind the posterior molar. 

Anterior Extremities — The scapula measures one inch in length, and is half an inch 
broad at its base, which forms a uniform convex curve. From the angles of the base 
the superior and inferior costse converge towards the anterior end of the spine, where 
the scapula becomes contracted into a neck, whose superior margin is continued into a 
slightly prominent coracoid. The supraspinal fossa is posteriorly about twice as broad 
as the infraspinal fossa ; but it rapidly narrows towards the neck of the scapula, and then 
disappears, while the infraspinal fossa continues still distinct. The long free edge of 
the spine is continued forwards as a very slender acromion. The glenoid cavity is ovo- 
triangular, with its apex directed downwards. The subscapular surface is smooth and 
slightly concave. 

The clavicles are entirely absent. 

The humerus, measured from the upper surface of its head to the lower end of the 
bone, is 1^ inch in length. The head is nearly hemispherical ; the lesser tuberosity 
forms a slightly elevated prominence ; while the greater tuberosity forms a strong 
pyramidal projection, by which the axis of the shaft is continued for about t'o^'is of an 
inch beyond the head. The shaft of the humerus presents a sharp edge in front, and 
is smooth and rounded behind. The anconeal fossa is imperforate, and there is no 
foramen above the internal condyle. Almost the whole of the front of the elbow-joint 
is formed by the surface for the radius. 

The ulna, measured from the superior margin of the great sigmoid cavity to the lower 
end of the bone, is 1 inch in length ; the olecranon process is -njths of an inch. The 
radius and ulna are quite distinct ; but the radius cannot be rotated on the ulna so as 
to effect supination. 

There are eight bones in the carpus, arranged in the usual proximal and distal series, 
with four bones in each series. The pisiform bone is large and subcylindrical ; it 
projects backwards from the outer side of the wrist, so as to form a sort of carpal heel. 
The metacarpal bone of the poUex is the shortest ; that of the minimus comes next to 
it in length ; those of the index and annularis come next, and are equal to one another, 
while that of the medius is the longest. 

Posterior Extremities. — The pelvis is narrow. The ossa innominata articulate with 
the first and second sacral vertebrse. The ilium is a narrow bone, nearly semicylindrical 
in shape, convex on its outer surface, and with its superior or anterior end slightly 
everted. The ischium nearly continues the axis of the ilium as far as the thin tuber- 
osity, and then turns vertically downwards to form the posterior boundary of the oval 
obturator foramen. The pubic bones form an angle of about 188° with the iliac, being 
thus almost on a line with them. The two pubic bones converge towards one another, at 
an angle of 40° ; but they form no true symphysis, being separated from each other at 
their posterior and inferior angle by a space of about -roth of an inch wide, wliich is 



AND AFFINITIES OF POTAMOGALE. 13 

occupied by a ligament admitting of considerable motion between the two bones at 
this spot. 

The femur is of the same length as the humerus, measured in each case from the 
upper surface of the head to the distal extremity of the bone ; it has a prominent 
tubercle, with a rough surface, upon the middle of the outer side of the shaft. 

The tibia is 1^ inch in length, measured from its upper to its lower articular 
surface. The tibia and fibula are confluent with one another for the lower third of 
their length. The tibia is curved, so as to present in its upper two-thirds an arch, 
convex forwards. The fibula is a slender bone, formmg the cord of the arch produced 
by the curvature of the proximal two-thirds of the tibia. 

The tarsus is composed of seven bones. The calcaneum is large, and projects for 
about one half its length behind the tibia. 

The metatarsal bone of the liallux is the shortest ; that of the outer toe is next in 
length; and the metatarsal bones of the three middle toes are the longest, and are 
nearly equal to one another. 

Anatomy of the Soft Parts. 

The imperfect state of preservation of the viscera, combined with the small amount 
of time which it was possible for me to spare from other avocations, has not allowed of 
more than a fragmentary description of the anatomy of the soft parts of the animal. 

The stomach and the whole of the organs of digestion between this and the vent, 
with the exception of about an inch of the terminal portion of the rectum, had been 
removed before the specimen was placed in my hands; so that certain important 
characters, such as that derived from the presence or absence of a caecum, could not 
be ascertained. The terminal portion of the canal, however, which escaped (fig. 9) 
presents several points of interest. The rectum, instead of opening directly on the 
surface of the body, opens into a sort of cloacal or postanal chamber, which also 
receives the orifices of the vagina and urethra, and those of the ducts of a pair of 
large anal glands. 

These glands are oval, about half an inch in their longer diameter. They are 
situated immediately beneath the skin, one on each side of the postanal chamber, 
into which each discharges its secretion by a single orifice. The excretory orifice of 
each gland opens into the bottom of a little pouch formed by a fold of the lining 
membrane of the postanal chamber at each side immediately within its margin. 
Just behind the line where the cavity of the rectum becomes continuous with the post- 
anal chamber, may be seen several very oblique pores in the mucous membrane of the 
chamber — apparently the outlets of small submucous glands. 

The uterus and its appendages and the urinary bladder were also left behind in the 
specimen ; but the kidneys had been cut away with the other viscera. The fundus of 



14 



PROFESSOR ALLMAJf ON THE CHARACTERS 



the Uterus is continued at each side into a long, curved, cylindrical cornu, which gives 
off the oviduct from its distal extremity. The ovaries are situated at a short distance 

Fig. 9. 




Terminal portion of intestine with the adjacent structures, slightly enlarged : a, rectum ; h, margin of anus ; 
c, postanal chamber laid open from bcliind ; d, vulva ; e, orifice of uretha ; /, anal glands ; ff, pouches into 
which their ducts open ; h, mucous pores ; i, uterus ; 7.-, cornu of uterus ; I, ovary ; m, oviduct ; n, vagina ; 
0, urinary bladder ; j), ureters. 

from this extremity, to which they are attached by a narrow cord-like ligament, which 
accompanies the o^dduct ; they are surrounded by a hood-like covering of peritoneum. 
From tlie uterus a wide, straight vagina passes backwards to open into tlie vulva, which 
also receives the orifice of the urethra, and is situated on the walls of the postanal 
chamber. 

The position of the mammse was unfortunately neglected to be ascertained before the 
specimen had been skinned, and it is now impossible to find any indication of them in 
the dried skin. They are probably uropygial as in Solenodon. 

The brain was in a very bad state of preservation: the cerebellum and medulla 
oblongata were entirely broken down ; but the cerebral hemispheres were sufficiently 
well preserved to show that they are destitute of distinct convolutions. The coi'pora 
quadrigemina were also preserved ; they are large, and are exposed behind the posterior 
margin of the hemispheres ; the posterior pair are larger than the anterior. The olfac- 
toiy lobes are rather large, and project in front of the cerebral hemispheres. 



From the details given above, certain characters, as perhaps eminently distinctive, 
may be selected and embraced under the following diagnosis : — 



AND AFFIMTIES OF POTAMOGALE. 15 

PoTAMOGALE, Du ChaiUu. 

„ ^, . 3-3 0-0 3-3 3-3 
^'''^'' '■ 3=3' '■ 0-0' P- 3=3' '"^ 3-=3 = 3«- 

Superior — first incisors laniariform ; second and third incisors and first and second premolars 
triangular, compressed, with sharp anterior and posterior edges ; third premolar pyramidal ■ true 
molars prismatic : inferior — first incisor very small, chisel-shaped ; second large and laniariform ■ 
third small, conical ; first, second, and third premolars triangular, compressed, sharp-edged • true 
molars prismatic. Muzzle broad, appressed. External ears well developed. Eyes very small. 
Nostrils valvular. Limbs of moderate length, plantigrade, pentadactyle. Second and third toes of 
hind feet syndactyle for the length of the first phalanx. Tail large, compressed ; its distal portion 
covered with short, stiff, closely appressed hairs, while the haii- covering the proximal portion re- 
sembles that upon the body. Body clothed with soft, rather coarse, hair, which projects from a 
dense covering of very fine, short, silky hairs. Anal glands two. Anus, vidva, urethra, and duets of 
anal glands opening into a postanal chamber. Zygomatic arches absent. Clavicles absent. Radius 
and ulna separate. Tibia and fibula adnate. 

From the description now given, it will probably be conceded that Potamogale is 
more nearly allied to Solenodon than to any other known genus of Insectivora. The 
absence of zygomatic arches, small eyes, well-developed ears, and large tail are all so 
many points of direct affinity. On the other hand, the remarkably compressed, 
triangular teeth, the compressed form of the tail, the broad appressed muzzle, the 
presence of anal glands, the coalescence of tibia and fibula, and, above all, the absence 
of clavicles are points of marked divergence from the West-Indian genus. 

On the whole I am of opinion that the genus Potamogale ought to be assumed 
as the type of a distinct family of Insectivora, to which the name of Potamogalidce may 
be given. 



The above paper had been already printed when I became acquainted with a descrip- 
tion of Potamogale velox, contained in a communication presented to the Zooloo-ical 
Society on the 25th of April, 18G5, by Professor J. V. Barboza du Socage, "On certain 
rare and little-known Mammifers from Western Africa, preserved in the Lisbon 
Museum," * as well as with another and more extended memoir, on the same animal, 
read by Professor Barboza du Bocage at a meeting of the Lisbon Academy, on the 27th 
of April, 1865. 

The specimen from which the Lisbon Professor's description had been drawn up was 
sufficiently well preserved to enable him to recognize the true insectivorous relations of 
the animal, and to give a detailed account of its external characters and osteology. He 
will not, liowever, accept the generic name of either Du Chaillu or Gray, but constructs 

* See Proo. Zool. Soc, 1865, p. 401. 



16 POFESSOR ALLMAN ON THE CHABACTEBS AND AFFINITIES OF POTAMOGALE. 

a new one of his own, and proposes to call the West African insectivore by the name 
of Bayonia veJox. For the reasons, however, already stated, I must still adhere to the 
claims of '' Potamofjale" over all other synonyms. 



EXPLANATION OF PLATES I. & II. 

Plate I. PotamogaJe velox, size of life. 

Plate II. Skeleton of Potamogale vdox, of the natural size. 






j'^ 

J 




a 
> 

< 

O 

< 
Eh 
O 




g 

^ 



\ 






o 



[ 17 ] 



II. On some Indian Cetacea collected by Walter Elliot, Esq. 
By Professor Owen, F.R.S., F.Z.S., &c. 

Bead June 26th, 1865. 

[Plates III— XIV.] 

Contributions to our knowledge of the singular and interesting order of 
Cetacean mammals {Cetacea vera, Cuv.) are so desirable, and acquisitions of evidences 
of exotic kinds are so few and far between, that I am induced to think the following 
may be deemed acceptable and worthy of publication by the Zoological Society. 

The materials chiefly consist of coloured drawings and skulls of species captured or 
cast ashore on the east coast of the Indian peninsula, in the vicinity of the harbour of 
Vizagapatam, in the northern circars of the Madras Presidency. 

Special care was taken by Walter Elliot, Esq., of Wolfelee^ when resident at that loca- 
lity, to have all such " stray waifs" from the whale-family brought directly to his cogni- 
zance ; and he availed himself of the skill of a native artist, for whose accuracy he vouches, 
to make drawings of the specimens while recent ; and these, for the most part, were 
executed under Mr. Elliot's own eyes. A selection from the drawings and some skulls 
of the Vizagapatam Cetacea have been confided to me by my friend for comparison and 
description ; and the results of this labour, as respects what seemed " new to science," 
I have now the pleasure to communicate. 

Family DELPHINID^. 

Genus Delphinus, Ciivier. 
Delphinus (subgenus Steno, Gray) gadamu, Owen. 
The " Gadamu" Dolphin. (PI. III. figs. 1 & 2.) 

This species is known to the Vizagapatam fishermen by the name of " Gadamu." It 
averages about 7 feet in length. The specimen figured is a female of 6 feet 10 inches 
in length. 

The body is fusiform, gaining its greatest diameter at the fore part of the dorsal fin, 
where the girth is .3 feet 9 inches. From this point the body decreases forward to the head, 
by straight converging lines laterally (fig. 2), and with a gentle convex curve superiorly 
(fig. 3), to the eyes and blow-hole; thence the sides of the head converge more acutely to the 

' Now Sir "Walter Elliot, K.C.S.I. 

VOL. VI. —PART I. D 



18 PROFESSOE OWEX OX INDIAX CETACEA. 

base of the snout, while the forehead descends with a bold convex curve to the same 
part. The snout, which is divided from the forehead by a transverse groove, extending 
almost horizontally nearly to the angles of the mouth, equals in length the distance 
from its base to the eyes, which is five inches and a half. Its vertical diameter at the 
base rather exceeds the transverse diameter : it gradually decreases to an obtuse apex. 
The lower jaw projects a little beyond the upper: the "rictus oris" extends backward 
to very near the eye. This opens at the junction of the lower with the middle third of the 
vertical diameter of that part of the head. The " blow-hole" is on the same transverse 
line with the eyes, symmetrically situated on the middle of the vertex, of a crescentic 
form, with the cresses bent forward (iig. 2, b). The pectoral and dorsal fins are fal- 
cate, of nearly similar size. The pectorals commence at the beginning of the second 
fourth part of the entire body : the extent of their base (i. e. from the attached fore 
part to the angle at which the concave hind border begins) is about 9 inches ; their 
length, following the anterior marginal curve, is 1 foot 6 inches : they are attached 
low down. 

The dorsal fin commences 3 feet from the end of the snout (in a straight line) : the 
extent of the attached base is 13 inches; that of the convex anterior border, following 
the curve, is 1 foot 4 inches. 

From the dorsal fin the trunk diminishes in size to tlie root of the tail-fin, more 
rapidly laterally than vertically; from the dorsal to the end of the caudal measures 
24 inches. The antero-posterior extent of the middle of the tail-fin is 7 inches ; the 
extreme breadth of the fin is 1 foot 10 inches; the circumference of the base or pedicle 
of the tail-fin is 10 inches. The vent is situated on the mid line below, in the interval 
between the vertical parallels of the dorsal and caudal fins, and nearer the dorsal, being 
2 feet 6 inches from the hind border of the caudal fin : about 2 inches in advance of the 
vent is the vulva. 

The colour of the body is a dark plumbeous grey, almost black upon the fins, 
especially at their fore part, becoming vei-y gradually lighter to the longitudinal 
parallel of the attachment of the pectorals, below which the body, from beneath the 
base of the snout and eye to below the base of the tail, is of a pinkish ashy-grey tint, 
with a few small irregular blotches of light plumbeous grey. The length of the snout, 
from the frontal groove, is 5 inches 6 lines ; that of the " rictus oris," in a straight line 
lengthwise, is 11 lines; the eye is about equidistant from the end of the snout and 
the beginning of the pectoral fin. The greatest vertical diameter of the body is 1 foot 
5 inches ; the greatest transverse diameter is the same ; the greatest girth is 3 feet 10 
inches; the vertical diameter of the base of the snout is 3 inches, the transverse 
diameter 2 inches G lines. The number of teeth, as noted by Mr. Elliot in one specimen, 
'■'^'^^ 17^ = 108; in a secondspecimen,||^=96; in the skull transmitted, |fEi= 101. 

This Dolphin would probably belong to that section which Dr. Gray has cha- 
racterized, under the name of Stem, as having the symphysis of the lower jaw 



iiK 






PEOFESSOE OWEIV OX INDIAN CETACEA. 19 

/ 
" elongate, about j the length"'^; but the definition of the term of comparison being 

omitted, whether it may be " length of the dental series," " of the mandibular ramus," 

or " of the entire skull," detracts from my means of testing this osteological character, 

whatever may be its value in regard to the variation in length of the " symphysis man- 

dibulae" of the restricted Delphini of Cuvier's system. 

In the skull, no. 423, of the " Gadamu" (PI. IV.), the symphysis mandibulse (figs. 
3& 4,5,5) is more than jth the length of the entire dental series, and about -g^th the 
length of the entire ramus. 

Assuming, however, the section or subgenus of the present Dolphin to be Steno, it 
then belongs to that subsection which is characterized as having the " ** Beak sepa- 
rated from the forehead hy a cross groove"". 

In this section the present species diifers from the Delphinus {Steno) malayanus in 
colour, iu number of teeth, and perhaps also in size. The D. malayanus is " greyish 
above and below;" the dental formula 3g3^ = 144. From the Delphinus (Steno) 

frontatus of the Indian Ocean, ^vith teeth 1^^=86 or ^j^~^j^ =:84, I), gadamu differs 
in the greater number of teeth. From Belpihinus {Steno) compressus the present species 
differs in the minor compression of the head, the shorter and less attenuated snout. 
The D. {Steno) attenuatus. Gray, departs still further from I), gadamu in the length 
and slenderness of the snout and the more numerous teeth, the formula being jqh^, 
= 160. 

In the skull of I). {Steno) gadamu (PI. IV.) the maxillo-premaxillary part of the rostrum 
is broader and lower than in D. {Steno) frontatus, the premaxillaries rise above the max- 
illaries, at the middle of the rostrum, with a more abrupt transverse convexity, and the 
maxillaries slope therefrom outward and less steeply downward to the alveolar border. 
Behind the dental series the bony palate, there formed by the back part of the maxil- 
laries, by the palatines, and pter3goids, forms a longitudinal bar convex across and 
increasing in depth as it recedes ; the sides of the bar are continued into channels of 
the same length, concave transversely, and impressing the sides of the posterior palatal 
surface of the maxillaries. This undulating disposition of the bony palate subsides 
opposite the penultimate or antepenultimate teeth, in advance of which the bony palate 
is nearly flat, with a strip, 2 inches long, of the vomer at the mid line, and iu advance 
of this is slightly hollow transversely, or canaliculate. 

The sockets of the teeth are in contact, about 4 lines in diameter. In the skull 
transmitted, and here noticed and figured (PI. IV.), I count 23 — 23 in the upper 
jaw, and 27 — 28 in the lower jaw. The teeth have a long and large rounded base and 
a short enamelled crown, slightly incurved, not very sharply pointed ; about ten anterior 
alveoli are coextensive with the symphysis. 

' Zoology of the Voyage of H.M.S. Erebus and Terror: " Cetacea." 4to. 18-14, p. 43. Not any of the 
figures of the skulls of Steno, Gray, illustrate the symphysial character in question. In a specimen of Sienn 
frontatus in the British Museum the mandibular symphysis is about one-fourth of the entire length of the 
skull. ' Ibid. 

D 2 



20 PROFESSOR OWEX ON INDIAN CETACEA. 

The specimen of the Gadamu Dolphin here figured was taken on the 20th March, 
1853, at Waltair, the civil station at Vizagapatam ; the posterior margin of the dorsal 
fin had been accidentally slit. 

Delphinus (StbnoI) lentiginosus, Owen. 
Freckled Dolphin. (PI. V. figs. 2 & 3.) 

By the same general fusiform character of the body, diminishing to the ends from 
the greatest girth at the fore part of the dorsal fin, and by the small size of this fin 
and especially of the pectorals, I am induced to place this Dolphin in the same 
section with the preceding. From the Gadamu it differs, not only in colour, but in 
the size of the fins, the pectorals and dorsals being relatively smaller, the caudal fin 
larger. The body is narrower, being subcompressed ; the vertical diameter at the 
deepest part (fig. 2) exceeds the transverse (fig. 3). The back is rounded in front of 
the dorsal fin, but is sharp, or keeled, behind it for about half the distance to the 
caudal, where it again becomes convex until near the root of the tail-fin, which is 
compressed and sharp above. The forehead is higher and more convex than in D. 
fusiformis (PI. V. fig. 1), but is continued by an alteration of curve more du'ectly into the 
rostrum than it is in D. gadamu (PI. III. fig. 1). The transverse groove, as indicated in 
the drawing (PI. V. fig. 6, c), is defined at the sides of the base of the beak, but above 
it is less deep or definite than in the two above-named species. The contour-line from 
the dorsal fin to the foreliead is nearly straight, very slightly undulated, not convexly 
curved as in B. gadamu. 

The specimen figured (PI. V. figs. 2, 3) was a female, captured at Waltair, Sep- 
tember 18, 1854. She measured 7 feet 10 inches in length, and 4 feet in greatest 
circumference, being probably pregnant. The colour is pretty uniformly bluish 
cinereous, or slaty, freckled with irregular small spots or streaks of brown or plumbeous 
pigment, the streaks longitudinal and flecked with white ; the under surface is a shade 
lighter than the rest of the body. The snout is 6 inches in length, 3| inches in 
depth at the base, and 3 inches there across; the skull shows better the pre- 
dominance of tlie vertical over the transverse diameter of the rostral production of 
the jaws. The " rictus oris," 1 foot in length, bends gently upward from the base of 
the snout to \vithin 2 inches of the eye. This is situated just above the middle of the 
vertical line crossing that part of the head. From the end of the snout to the eye is 
142- inches. The blow-hole, median in position and shaped as in the foregoing species, 
is a little in advance of the vertical parallel of the eyes ; in the male specimen it was 
on the same parallel. From the end of the snout to the pectoral fin is 2 feet ; the 
attachment of this fin is subpedunculate, the antero-posterior extent of the peduncle 
being only 3 inches, while the breadth of the fin, at the posterior basal angle, is 5 
inches ; the length of the anterior margin, following its very slight convex cui've, is 
12 inches. The dorsal fin is relatively lower than in I), fusiformis, much more so than 



PKOFESSOR OWEN ON INDIAN CETACEA. 21 

in B. gadamu ; the hind border slopes away gradually to an extensive base of attach- 
ment, which is continued as a ridge halfway between the dorsal and caudal fins : the 
leno-th of the dorsal at its front margin is 1 foot 1 inch ; from the end of the snout to 
the dorsal fin is 3 feet 4 inches ; from the front border of the fin's base to the mid fissure 
of the tail-fin is 4 feet 2 inches ; the fin is rather more posterior in position than in 
D. fusifonnis, and is more obtusely terminated than in that species or in B. (jadamu. 
From the hind border of the caudal fin to the vent is 2 feet 5 inches : the vulva is 
2^ inches in advance of the vent. The upper part of the pedicle of the caudal fin is 
obtusely ridged ; the middle of the posterior margin of the fin is notched, as in the two 
foreo-oino- species ; the antero-posterior breadth of the fin, near the notch, is 7 inches 
6 lines ; the transverse breadth of the entire fin is 1 foot 9 lines. 

A profile-view of the head and pectoral fin of a male I), lentiginosus, taken also at 
Waltair, which was of a rather darker bluish slate-colour than the female, shows 
the feeble indication of the fronto-rostral groove beyond the lateral indentations ; the 
interruption of the convex curve of the forehead, before reaching the snout, is rather 
more marked. The mouth is represented a little open, indicating the relative size of 
the teeth so exposed; they were ||=|=129. As in the female specimen, the pectoral 
fin is not falciform, but has rather the shape of a scalene triangle, the two shorter sides 
straight. 

The skull of Delphinus (Steno) lentiginosus is rather narrower in proportion to its 
length than in B. gadamu ; the occipital condyles are larger, the superoccipital surface 
is narrower, the temporal fossae more squared above ; the premaxillaries do not rise to 
form a distmct convexity at the upper part of the rostrum, as in B. gadamu, but con- 
tinue upwards the roof-like slope, begun by the maxillaries, which gives a triangular 
transverse section to the middle and fore part of the rostrum. The breadth of the 
rostrum at the antorbital notches is the same in both species, viz. 4 inches ; the length 
of the rostrum, from the notches, is IQi mches in B. gadamu, 11 inches in B. lentigi- 
nosus. But the chief distinction is in the number of the teeth : in the skull here 
noticed and figured there are, in the upper jaw, 33—33, in the lower jaw, 32—32 
= 130, and the teeth are smaller. The extent of the dental series of the upper jaw in 
B. lentiginosus is 9 inches 9 lines, but is not more than 8 inches 6 lines in B. gadamu. 

The B. lentiginosus is known to the Waltair and Vizagapatam fishermen by the 
Telugu name of " BoUa Gadimi." 

Delphinus (Steno?) macuuventeb. 
Spot-bellied Dolphin. (PI. VI. figs. 1 & 2.) 
In the degree of convexity of the forehead the present species resembles the B. 
fusiformis (PI. V. fig. 1); but the head is relatively larger, and the body is deeper in 
proportion to its length, than in either B. fusiformis or B. gadamu. 



22 PROFESSOE OWEN ON INDIAN CETACEA. 

In colour it presents a well-marked distinctive character from all the Vizagapatam 
species; it is of a deep, shining, plumbeous black on the upper part, becoming 
paler near the belly, which, from the under part of the jaw to the perineum, is 
ashy grey, with irregular spots or blotches, whence the specific name maculivenfer. 
The specimen from which figs. 1 and 2 were taken was a female, 6 feet 11 inches in 
length, found at Waltair, 2Gth April, 1854. It is called by the fishermen " Suwa." 

The fronto-rostral groove is well marked, but short; the " rictus oris" slightly rises as it 
extends back, to about 3 inches below the fore part of the eye ; the under jaw extends be- 
yond the upper, and chiefly forms the obtuse end of the rostrum ; this is 5 inches in length, 
and higher at its base than it is broad. The blow-hole resembles in position and shape 
that of the previously described species. Both pectoral and dorsal are falcate, but small ; 
the length of the front border of the pectoral, following the curve, is 1 foot 3 inches ; 
from the end of the snout to the setting-on of this fin measures 1 foot 9 inches. The 
greatest circumference of the body is just in advance of the dorsal fin ; the height of 
this fin is 8 inches, the extent of its basal attachment 18 inches ; to the fore part of the 
dorsal from the end of the snout, in a straight line, is 3 feet 4 indues ; from the back 
part of the dorsal to the hind border of the base of the caudal fin is 3 feet. The body 
is more compressed than in J), lentiginosus (PI. V. fig. 3). The girth of the pedicle of 
the caudal fin is 1 foot 2 inches ; the fore-and-aft diameter of the fin is 7 inches, the 
extreme breadth is 1 foot 8 iuclies ; from the median notch of the caudal to the vent is 
2 feet 3 inches ; extent of perineum (or between the vent and vulva) 3 inches. 

The dentition of this species is 3^^q=114. It appears not to be rare. Specimens 
were taken in March 1853 and April 1854, all showing the character of colour given 
in the female figured in PI. VI. figs. I & 2. 

Delphinus (Lagenorhynchus) fusiformis, Owen. 

Spindle-shaped Dolphin. (Plate V. fig. 1.) 

The present species is more slender in proportion to its length, has a less elevated and less 
convex forehead, a proportionally thicker, broader, and more obtusely terminated snout, 
a deeper mandible or under jaw, especially posteriorly, and smaller dorsal and pectoral fins, 
especially the latter, than in the foregoing species of Delphinus. It appears, likewise, to 
be a smaller species. The specimen figured, which was the largest taken (at Waltair, on 

the 23rd August, 1853), was a female, 6 feet in length: the dentition 2iEfi^86. The 
greatest girth of the body is at the fore part of the dorsal fin ; from this the body 
tapers to both ends, and, through the lower forehead and thicker snout, more regularly 
than in I), rjadamu, and presenting a truer spindle-shape of the whole animal, whence the 
specific name. The " rictus oris" bends upward as it recedes, and does not approach so 
near the eye as in 2>. gadrniut. Both the angle of the mouth and the eye are more 
elevated in position ; the blow-hole is medial, symmetrical, on the same vertical parallel 



PKOFBSSOR OWEN ON INDIAN CETACEA. 23 

with the eyes ; crescentic, with the angles bent forward. The length of the snout is 
6 inches, of the " rictus oris" 10 inches; from the end of the snout to the eye 1 foot; 
from tlie same to the setting-on of the pectoral fin 1 foot 7^ inches ; from the same to 
the setting-on of the dorsal fin 2 feet 7 inches ; from the hind part of the base of the 
dorsal fin to the hind border of the caudal fin 2 feet 8 inches. The pectoral fin measures 
5 inches across the broadest part of its base, and is 1 foot in length, following the curve 
of the front border, which curve is much less than in the Gadamu. The dorsal fin is 
lower in proportion to the length of its base ; its anterior border also shows a minor 
degree of convexity ; the extent, following the curve, is 10 inches ; the line of attach- 
ment measures 11 inches. The fore-and-aft extent of the mid part of the caudal fin is 
5 inches; the extreme breadth of the fin is 1 foot 4 inches. The vent is 1 foot 9 inches 
m advance of the mid notch of the caudal fin ; the Milva is 5 inches in advance of the 
vent, the interspace being relatively greater than in the Gadamu. 

The colour of the "Spindle-shaped Dolphin" is less darkly plumbeous than in the 
Gadamu, and becomes more gradually lighter towards the belly; the dorsal fin, the 
fore part of the pectoral and caudal fins, and the snout have the darkest pigment ; the 
light ashy-grey belly shows no spots. 

The difference from any of the three preceding species is still more marked in the skull 
(PI. VII.), which presents the general characters of that section of Delphinidce to which the 
term ^^ Lagenorhynchus " has been attached. It resembles in size and general characters the 
skull of Lagenorhynchus elecfra. Gray ; but the occipital condyles are more approximate 
below the foramen magnum, the presphenoid is narrower, the longitudinal channel 
foi-med by it and the pterygoid is deeper and narrower : the rostrum is of equal length in 
the two species, viz. 9 inches 8 lines from the antorbital notches {k) ; but the breadth 
there is 5| inches in Lagenorhynchus electra and 5 inches in Lagenorhynchus fusifonnis. 
In this species a narrow slip of the vomer (fig. 4, is), about an inch in length, appears 
on the bony palate, 3 inches from the anterior end. 

In Lagenorhynchus (PI. VII.) the skull is broader in proportion to its length, and the 
mandibular symphysis shorter, than in Steno (PI. IV.) ; the transverse undulation of the 
hind part of the palate is less marked, the middle convex tract being broader and lower, 
and the lateral channels wider and shallower. 

Delphinus pomeegea, Owen. 

The Pomeegra Dolphin. (PI. VI. fig. 3.) 
This species belongs to the same section of Belphinus as the Black Dolphin of the Cape 
and Ceylon (Delphinus longirostris, Gray^) and the Belphinus forsterP of the Pacific. 

' Schlegel. Mr. Blyth has inserted a note on this species in the ' Journal of the Asiatic Society of Bengal.' 
1848, pp. 249, 250. 

2 Forster, " Descriptio Animalium," drawing no. 24 (copied by Dr. Gray, in the ' Zoology of the Erebus and 
Terror,' " Cetacea," 4to. 1845, plate 24). 



24 PROFESSOR OWEN ON INDIAN CETACEA. 

It was taken off the coast of Madras, and is known to the fishermen there as the " Po- 
meegra." It is of a very deep plumbeous shining coloiu-, almost black, with a rather 
lighter shade at the under part of the belly. Mr. ElHot, who was indebted to Mr. Blyth 
for the specimen, notes it as " a small Cetaceous species;" but the length is not given. 
The proportions of the snout, of the rictus oris, of the fins, and the form of the forehead 
(which rises from the base of the snout with a low convexity) are characters in which the 
D. pomeegra resembles the D. longirostris. Gray. It chiefly differs in the larger propor- 
tional size and smaller number of the teeth, viz. ^i^= 173. The blow-hole is crescentic, 
and on the same vertical parallel as the eye. The body enlarges more gradually to the 
origin of the dorsal fin than in D.forsteri, the greatest circumference being at the fore part 
of that fin. It is more slender iu proportion to its length than any of the above-described 
fusiform Dolphins belonging to the subsection Steno, Gray. The symphysis mandibulae 
(PL VIII. fig. 4) is less than |^th the entire length of the ramus. The hinder half of the 
palate (ib fig. 2), is widely and deeply channelled on each side. This is, however, but an 
extension of the modification already pointed out in the hind part of the palate of D. ya- 
damu (PI. IV.), and it is subject to varieties in species which, from the brevity of the 
mandibular symphysis, the great number and small size of the teeth, and the transversely 
convex rising of the premaxillaries along a considerable part of the rostrum, would be 
retained among the iJelphini as restricted by Dr. Gray. In Delphinus eiqjhrosyne, e. g. 
(PL VIII. fig. 5 : no. 15, p. 251, ' Catalogue of Cetacea in the Br. Mus.'), the hinder middle 
tract of the bony palate is not longer, deeper, nor more convex transversely than in Steno 
and Lagenorhynchus, and the lateral channels show the same proportions as in the latter 
subgenus. The prominent mid tract of the palate is too broad and obtusely convex to 
be regarded as a " ridge," in any species of Delphinus proper that has come under my 
observation. 

Sp. dub. Delphinapterus molagan, Owen. 

Mr. Elliot wi'ites, " I have (or rather ' had,' for I cannot find it) a drawing of a small 
Cetacean, copied from one made in the Chief Engineer's Office at Madras for Col. 
Monteith, which was taken from an individual, 32 inches long, of a uniform black 
colour, with a rounded obtuse head, small mouth, and no dorsal. The Tamil fishermen 
called it ' Molagan.'" 

Genus Phoc^na, Cuvier. 
Phoc.<ena (Orca, Gray, ReinJiardt) brevirostris, Owen. 

Short-snouted Porpoise (skull). (PL IX. figs. 1, 2, 3.) 

Of this Cetacean I possess only the cranium ; but, as it presents the characters of 
maturity, it is too small for the species represented by the drawings already described, 



PKOFESSOK OWEN ON INDIAN CETACEA. 25 

if even the proportions of the rostral part of the skull (PI. IX. fig. 1, 21' 22) did not show 
that it belongs to a different section of Belfhinidce^ . The present part of a Cetacean 
skeleton, as the skulls of those species, figured in Pis. IV. VII. VIII. demonstrate, affords 
better grounds for comparison and specific determination than do coloured drawings of 
the entire animal, however accurate, — the number of skulls of ascertained species in 
home-museums, or otherwise accessible, being much greater than entire and stuffed spe- 
cimens of the Cetacea, which rarely give the natural contour of head or body. 

The animal from which this skull was taken was thrown ashore in the harbour of 
Vizagapatam in too decayed a state to be figured, and was noted as a " small kind of 
Porpoise " by Mr. Elliot, who fortunately secured the present evidence of the species, 
which is now preserved in the British Museum. 

The following are the dimensions of the skull : — 

inches, lines. 

Length H 

Breadth, greatest, across zygomata 7 4 

From the back of occipital condyle to antorbital process of malar bone 7 6 

From the antorbital process of malar to anterior end of premaxillary 4 8 

From the back part of nostrils to do. do. 7 6 

These dimensions show that in the shortness of the " facial " as compared with the 

" cranial " part of the skull the species agrees with the section of Del])hinid(e, including the 

Grampuses and Porpoises, for which Cuvier proposed the subgeneric name PJiocwna-, 

and which, in his ' Ossemens Fossiles,' tome v. part i. (1823), he distinguished as 

" § 2. Les Dauphins a tete obtuse" (p. 280), from " § 1. Les Dauphins a bee" (p. 275) 

{Belpliinns, proper)^ 

The number of Belphinidce with obtuse heads or short jaws, which have since been 
observed, have manifested so many minor modifications in the relative size, shape, and 
number of the teeth, in the relative size and length of the jaws, in the formation of the 
bony palate, in the extent of anchylosis, and the forms of processes, &c., of the cervical ver- 
tebrtE, that numerous subgenera have been founded on these characters. Nevertheless, 
as each additional kind of blunt-headed Dolphin tends to exemplify the gradational 
tendency of these modifications, the benefit to zoology of the additional g'Wfm-generic 
names is doubtful ; and I shall refer the present skull, which appears to me to belong 
to an undescribed species, to the PJioccena brevirostris, as a member of the section of 
Cuvier's PhocmncB, charactei-ized by conical teeth, in which its nearest alliance appears 
to be with the Phoccena gloMceps, Cuv.* 

' The following is Mr. Elliot's note respecting this specimen ; — " August 1852. Got the skull of a poi-poise 
which one of the fishermen found dead at the mouth of the Vizagapatam river. He caUed it ' Ganumu,' and 
described it as having a rounded head, without beak, colour black or dark above, white below ; perhaps a 
Phoccena or Glohicephalus." 

' Eegne Anim. tome i. p. 290 (1829). ^ Ibid. p. 287. 

' Ibid. p. 290 ; Annales du Museum, tome xix. ; Oesem. Foss. tome v. part i. p. 290, tab. 21. pis. 1, 2, 3, figs- 

11, 12,13. 

VOL. VI. PABT I. ^ 



26 PROFESSOR OWEN ON INDIAN CETACEA. 

The elements of tlie occipital have coalesced. The basioccipital (PI. IX. fig. 3, i ) 
forms the lower fifth of tlie foramen magnum, intervening for an extent, measured in a 
straight line, of 10^ lines between the lower ends of the occipital condyles (ib. 2') : it is 
liere thick and concave transversely : it becomes thinner vertically and expanded later- 
ally as it advances to join the basisphenoid (ib. s), with which it has coalesced: a slight 
median longitudinal obtuse rising divides two large shallow concavities, from the sides 
of which the aliform expansions of the basisphenoid extend, which bend slightly down- 
ward to form the lower and inner or mesial wall of the otocrane (ib. or). The occipital 
condyles (figs. 1 & .3, 2', 2') are narrow, vertically elongate, oval convexities, wider at 
their lower half, with the mesial margin gently convex, the lateral or outer margin 
sinuous, through a slight concavity marking oif the upper third of the condyle : the 
length of the condyle in a straight line is 2" 1'", the greatest breadth 1" 11'" : the 
upper ends of the condyles are 1" 3'" apart ; they are low and sessile. The foramen 
magnum is vertically oval, widest above, and notched at the middle of the upper 
border; its length, to the end of the last notch, is 2", its breadth 1" 3'"; the breadth 
across the broadest parts of both condyles is 2" 9'". The paroccipital (figs. 1 & 3, 4 ) 
an exogenous growth of the exoccipital, forms the back part of the otocrane, towards 
which it is sinuous or slightly concave, and terminates below in a thick rough border, 
4'" across the thickest part (figs. 3, i'" ) ; this border is divided by a notch from the 
otocranial plate (&') of the basisphenoid, and just within the bottom of that notch' 
opens the canal for the nervus vagus. The supcroccipital (figs. 1,2, 3) rises and ex- 
pands, as in other Delphinidce, into a broad and lofty convex plate reaching the vertex, 
and there articulating with the parietals ( 7 ) and interparietal ( /* ) ; a low median 
ridge (fig. 2, 3' ) divides vertically the upper half of the supcroccipital. On the inner 
surface, 1" 6'" above the foramen magnum, a vertical triangular plate of bone descends 
into the falx ; it is thickest behind, where its base is groo\'ed transversely by the lateral 
sinus. 

The alisphenoids (PI. IX. figs. 1 , 3, 6 ) coalesce with the fore part of the lateral borders 
of the basisphenoid, in advance of the otocrane (fig. 3, or), of which it forms the anterior 
wall or boundary : the base of the alisphenoid is notched posteriorly for the third, and 
anteriorly for the second, division of the trigeminal ; it expands as it passes outward, 
slightly rising (fig. 1, 6) to join the parietal (?), and frontal (n), and to overlap the 
process of the squamosal (fig. 3, ir ), continued, mesiad, from the glenoid cavity {g). The 
suture between the interparietal' (fig. 2, ^* ) and supcroccipital ( 3) is obliterated, and that 
with the parietals is partially so. The suture between the parietal and superoccipital 
remains at its lower half (fig. 1, 7 ), showing that a narrow strip of the parietal appears 
on the external surface of the cranium, extending backward, between the squamosal 
( 27 ) and superoccipital (3 ) to the exoccipital (2 ), and slightly expanding at its junction 
therewith. 

The presphenoid (ib. fig. 3,9) is distinct from the basisphenoid (s), and extends in 
the form of a compressed rostrum forward, contracting, to be enclosed by the pos- 
terior sheath-shaped part of tlie vomer (13). The orbitosphenoids (ibid. 10) extend 



PROFESSOR OAVBX OX IXDIAN CETACEA. 27 

outward, overlapping the pterygoids ( 24 ), contract where they form the fore part of the 
foramen lacerum anterius and tlie optic foramina, beyond which they expand to 
support the orbital plate (fig. 3, n' ) of the frontal.' 

The frontals (PI. IX. figs. 1 & 2, 11, ir ), in great part overlapped, as in other Cetacea, by 
the maxillaries ( 21 ), sliow at their narrow exposed strip, extending transversely across the 
summit of the cranium, the persistant frontal suture, half an inch in length ; from this 
suture the strip curves outward and backward, expanding beyond the interparietal ( /• ]. 
and then downward and forward, contracting and again expanding, to form the post- 
orbital process (figs. 1, 2, 12): this is triangular and three-sided, one facet being a 
continuation of the exposed strip, a second contributing to the temporal fossa, and a 
third to the orbit (or). In the temporal fossa, the frontal (fig. 1, 11 ) articulates with 
the parietal (7) and alisphenoid (g); in the orbit (ib. or), with the orbito-sphenoid 
(fig. 3, 10 ) and malar ( 26' ) ; then, arching forward from the postorbital process, the 
frontal forms the superorbital ridge (fig. 1, 11), and articulates anteriorly by a kind of 
gomphosis with the malar ( 20' ) ; it is overlapped here, as on the cranium, by the max- 
illary (21"). The medial parts of the frontals (fig. 2, 11) are united posteriorly with 
the interpai'ietal ( ;» ), anteriorly with the nasals ( is ). 

The vomer (ib. fig. 3, 13 ) extends forward to within an inch and a half of the end of 
the premaxillaries, and behind these it intervenes upon the bony palate between the 
maxillaries, along a strip of two inches extent and three lines across the broadest part. 
This palatal part of the vomer (13) is the lower convexity of the canal formed by the 
spout-shaped bone ; the hollow of the canal is exposed at the upper interspace of the 
premaxillaries. Here, also, is seen, tw^o inches behind the fore end of the vomer, the 
rough thick anterior border of the coalesced prefrontals (fig. 2, u ), which contracts as 
it passes into their upper border, forming the septum of the nostrils, expanding below 
and behind to form the back wall of the nasal passages ( u' ). At this part a trace of 
the suture between these foremost neurapophyses of the skull remains. Their bifid spine 
— the small transversely extended subquadrate nasals ( la ) — intervenes between the. 
frontals (11) and prefrontals ( 14' ). The palatine bones appear on the palate as narrow 
strips (fig. 3, 20 ) wedged between the maxillaries, (21) and pterygoids (24), and united 
together beneath the vomer by a longitudinal suture of 3'" extent : then, passing out- 
ward and forward, after a brief contraction they suddenly expand and bend upward to 
line or form the mesial wall of the orbit, and again contract to articulate with the frontal 
at the superorbital fossa ; the mesial borders of the palatines articulate with the vomer 
and prefrontals ; and between the pterygoids and the vomer the palatines form the fore 
part of the lower half of the nasal passages. The orbital plate of the palatine sends ofl' 
an outer thin lamina, which terminates by a free margin at the back of the orbit. The 
palatine plates of the maxillaries (21) unite together for about an inch in front of the 
palatines, then slightly diverge to give place to the vomer (33), which, however, does 
not sink to their level ; in advance of the vomer the plates slightly diverge to their 

E 2 



28 PROFESSOR OWEN ON INDIAN CETACEA. 

anterior ends, giving place to the premaxillaries ( 22' ), which form the apex of the 
muzzle : the rest of the disposition of the maxillaries accords with Cuvier's account in 
Phoccena (jlobiceps; the superorbital plate (fig. 1, 2i») is divided by a notch from the 
rostral part ( 21 ) of the maxillary, and forms a tuberosity articulated with the under- 
lying malar ( 26' ). The premaxillaries ( 22 ) accord equally with those in P. glohiceps, 
save in their shorter proportions concomitantly with the shorter muzzle. They are 
perforated near the outer margin, between the posterior and middle third, the canal 
leading forward and inward. The three perforations (fig. 2, a, b, c) in the maxillary 
external to the nasal portions of the premaxillary ( 22'), are the upper outlets of canals 
which converge to open into an oblong fossa (fig. 3, 26) beneath the fore part of the roof 
of the orbit. 

The pterygoid (fig. .3, 24, 24' ) is a large sinuous plate folded upon itself from within, 
upward, outward, and backward; the thick fore part (24) articulates with the palatine, 
whence it continues the bony roof of the mouth backward for the extent of 1" 8'", 
with a convex surface, divided from its fellow by a vacancy of 8'" breadth, exposing the 
presphenoid and vomer ; the inner plate of the pterygoid forms the outer wall of the 
lower part of the nasal passage, and continues that passage obliquely backward, as an 
open canal (24'), beneath the base of the alisphenoid (6), as far as the otocranial 
plate of the basisphenoid (s' ). This posterior production of the pterygoid is three-sided ; 
the inner or narial one is concave ; the outer one is also concave, forming a channel 
leading upward and forward to the orbit ; the upper facet is sutural, and articulated 
with the basi-, pre-, ali-, and orbito-sphenoids. The anterior external lamina of the ptery- 
goid bends outward and upward to articulate with the corresponding free lamina of the 
palatine, bounding the narrow and deep sinuous fissure between the outer and inner 
portions of both bones. 

The malar, as in other DelpMnidcp, consists of the antorbital (PI. IX. fig. 1, 26' ) and 
styliform (26) portions. The former ( 26' ) is a narrow triangle, with the base thick, convex, 
turned forward, underpropping the fore part of the superorbital plate of the maxillary 
( 21* ), and articulating with the same part of the frontal ; the apex extends backward, 
and is wedged into the roof of the orbit between the frontal and maxillary. The 
styliform portion (26) is given off" by a process extending inward (mesiad), at right 
angles to the antorbital portion (fig. 3), and a few lines behind its fore part ; it sud- 
denly contracts and extends backward, with a slight bend, to the squamosal, articu- 
lating by a concave, oblique, terminal facet to a tubercle at the fore and imder part 
of the zygomatic process of the squamosal (fig. 1, 27). The length of this part of the 
malar is 3" ; its thickness throughout the greater extent is 1^'" by 1'" ; its squamosal 
articulation is 4'" across. The form of the orbit (ib. or) so defined below is longitu- 
dinally oblong, more arched above than below, 2" 2'" in fore-and-aft diameter, 1" 2'" 
in greatest vertical diameter ; the chamber communicates, of course, largely with the 
temporal fossa, and continues into the deep, ascending orbital fossa and the small 
antorbital fossa (d), external to which is the rough malomaxillary fossa [e). 



PROFESSOE OWEN ON INDIAN CETACEA. 29 

The squamosal consists chiefly of its articular or zygomatic part (PI. IX. figs. 1 & 3, 2-), 
which is deep in proportion to its length, truncate, and tluee-sided ; the outer side is 
slightly convex and rather rough, 1" 5'" in depth posteriorly ; the uiner side is divided 
between the articular cavity (fig. 3, g), rough for spidesmosis with the mandible, and 
the smoother surface internal to it, which extends mesiad in a triangular depressed form 
( 27' ) beneath the back part of the alisphenoid (6), but without joining it: the upper 
surface, of an inequilateral shape, contributes a lower wall to the temporal fossa. The 
squamous portion (fig. 1), continued upward from this facet, is triangular, with a rounded 
apex, about an inch in length, and rather more in height ; it is applied against the ali- 
sphenoid (e) and parietal (7). The rough posterior tract articulating with the parietal 
( 7 ) and exoccipital ( 2 ), and contributing to the outer wall of the otocrane (fig. 3, or), 
I consider to be the " mastoid" confluent with the squamosal, together forming the bone 
which should be termed " squamo-mastoid." The mastoid part termuiates below in a 
rough, flattened, triangular surface (fig. 3, s), 5" 7'" in diameter, which is divided from 
the zygomatic or articular process of the squamosal [g) by a deep fissure. On the inner 
side of the base or back part of the mastoid, m the Ime of its suture with the parietal, 
is the (stylomastoid!) fossa. The squamosal forms no part of the inner or proper 
wall of the cranial cavity. The glenoid or mandibul-articular surface {g) is longitu- 
dinally oblong, 1" 5'" by 8'" in diameter, moderately concave, least so transversely, and 
looking inward, do\vnward, and with a slight inclination forward. The mandible 



ofiers no notable peculiarity, save that which relates to shortness in proportion 
to the entu-e skull, concurrently with the same specific character of the upper ja\\-. 
The depth of the ramus at the coronoid process is relatively as great as in the longer- 
jawed species, and consequently bears a greater ratio to the length of the entire 
ramus : this in the present skull is 7", the gi-eatest vertical extent of the ramus being 
2" C" ; the shallowest part of the ramus is where it supports the teeth ; it deepens a 
little at the short symphysis. There are fourteen alveoli approximated in a conmion 
groove in each mandible, extending along 3" 3'" from the symphysis. The correspond- 
ing groove of the upper jaw (fig. 3) shows seventeen alveoli, along an extent of 3" 6'". The 
deeper part of the alveolus is distinct in the anterior teeth ; but, as they recede, the sockets 
are indicated by depressions merely in the common groove. The teeth are slender ones : 
the anterior ones in the upper jaw average a length of 8'", two-thu'ds of the UTegular 
cement-covered, thickened, and solid base being implanted, the exposed third forming a 
smooth, partially enamelled, pointed crown, with a circular transverse section and in 
most a slight inciu'vation ; the length of crown is from 3'" to 4'", the diameter of its 
base 1"', that of the inserted root 2'". 

As in other Belpkinidw, the bony palate is entfre, save at the slight median divarica- 
tion of the maxillaries and premaxUlaries, and the major part of this median fissure is 
closed by the vomer. A pair of small (neuro-vascular) foramina is situated near the 
maxillo-palatine suture, and one or two others obliquely groove and pierce the palatine 
plate of the maxillary. 



30 PEOrBSSOR OWEN ON INDIAN CETACBA. 

The optic foramen communicates or is blended with a larger vacuity or fissure 
between the orbitosphenoid, frontal and pterygoid, which might be termed the ^heno- 
frontal fissm-e. The foramen rotundum, in like manner, is blended with a larger vacuity 
between the ali- and orbito-sphenoids, answering to the "fissura lacera anterior" of 
anthropotomy, and which may be called the " mtersphenal fissure "^ 

The removal of the loosely attached petrotympanic exposes the wide otocranial 
vacuity (PI. IX. fig. 3, or) in the basal walls of the cranium, which is a characteristic 
feature of the Delphinoid as compared with the Physeteroid skuU (PI. XIII. fig. 2), 
where the otocranial is walled off from the cranial cavity. The otocrane, in both, is 
bounded by the paroccipital, basisphenoid, alisphenoid, and squamo-mastoid : in the 
pi'esent species of PJioccena it presents a subquadi-ate form, 1" 4'" in diameter, with the 
angles rounded off, notched anteriorly by the third division of the fifth, whereby the 
" foramen ovale" blends with this great vacuity. 

The entocarotid foramen pierces the outer and fore part of the base of the otocranial 
plate of the basisphenoid, close to, perhaps at, the line of confluence of the alisphenoid. 
There are neither olfactory nor lacrymal foramina. The absence of the rhinal capsules 
simplifies the condition of the prefrontals, and facilitates the comprehension of both the 
special and general homologies of these interesting bones. A pair of minute foramina 
lead from the cranial cavity to the narial ones piercing the prefontals ; but they do 
not give passage to olfactory nerves in the BeljiJirmdce. 

The departure from symmetry in the present Delphuioid skull is slight : it is seen 
in the gi-eater backward extension of the nasal plate of the right premaxillary (fig. 
2, 22" ), in the larger size of the prenarial plate of the right maxillary, and in a feeble 
inclination of the upper margin of the septum narium to the left. 

FamUy PHYSETERID^E (Cachalots or Sperm-Whales). 

Genus Euphysetes, MacJeay. 

PiiYSETER (Euphysetes) simus, Owen. 

The Snub-nosed Cachalot. (Plates X.-XIV.) 

The Cetacean which I have next to describe is represented by drawings of the adult 
male (side view, PI. XL to scale) and female (side view, PI. X. fig. 1 ; upper view, fig. 2 ; 
to scale). It is noted as " a kind of Porpoise " in Mr. Elliot's MS., and is known to 
the Telugu fishermen of the coast by the name of " Wonga." Tlie male, measuring 6 
feet 8 mches in length, was taken at Waltair, February 28, 1853. The female was taken 
on the 1st of March, 1853, at the same part of the coast ; she measured 6 feet in length. 

' It is noticed as " le trou spheno-orbitaire," by Cuvier, ' Oss. Boss.' torn. cit. p. 294. 



PROFESSOR OWEN ON INDIAN CETACEA. 31 

The resemblance to the Porpoise was suggested by the shortness of the snout ; but 
this is more obtuse, and is not marked off from the rest of the head by any sudden 
narrowing. More important differential characters suggest the affinity of the " Wonga" 
to a family of toothed Whales, distinct from the Uelphinidm. 

The first and most important of these is the mferior position of the mouth, beyond 
the smaU opening of which the blunt rostrum extends forward fi'om 4 to 6 inches. 
The blow-hole (PI. X. fig. 2) is single, but is not medial in position or symmetrical in 
shape ; it is hi advance of the eye, opens to the left of the mesial plane, is propor- 
tionally larger than in the Porpoise, and is crescentic, but curves obliquely from the 
mid line outward and backward, with the convexity turned forward and to the left, and 
tlie angles or " cresses" dii-ected backward and to the right. The anterior angle is 
5 inches from the end of the snout. The eye is small; the palpebral orifice is be- 
tween 7 and 8 inches from the end of the snout, and opens m the upper half of the 
head, seen in profile, near the boundary dividing it from the lower half. From the 
vertical Ime bisecting the eye to the end of the muzzle the head forms a cone with 
a blunt apex, less obtuse when viewed from above (fig. 1) than from the side (fig. 2), 
where the lower slope is interrupted by the small "rictus oris:" this is formed by a 
kind of semicu'cular excavation of the under part of the snout, into which the short 
dentigerous part of the lower jaw fits, like a box in its lid. The length of the " rictus ' 
in a side view, straight Ime, is 2^ inches in the male, 2 inches in the female. From the 
the parallel of the eye, the head, as it recedes, enlarges less rapidly; and the trunk 
continues gradually to expand to about midway between the end of the snout and the 
base of the tail. The widest part of the trunk is a little more forward in the male than 
in the female. 

According to the figures, the pectoral fin becomes free 1 foot 1 inch behind the 
snout in the male, and 1 foot 4 inches in the female ; but there may be some inaccuracy 
here. The length of the fin in both is 1 foot ; its extreme breadth is 4^ inches in the 
male, 4 inches in the female : its line of attachment is in the lower thu'd of the trunk, 
as seen in profile. The dorsal fin is well developed, subfalcate m sliape ; its anterior 
border is halfway between the snout and the base of the tail. The length of the base 
of the fin is 10 mches in the male, 9 inches in the female: the height of the fin. 
^■ertically at its back part, where the apex curves back a little beyond the basal attach- 
ment, is 7 inches m both. The anterior border of the fin is slightly convex ; its length, 
m a straight line, is 1 foot. 

The body, as has been said, gradually expands to near the origin of the dorsal fin, 
and thence contracts to the setting-on of the caudal fin : here the taU, or tail-end of the 
trunk, measures 3^ to 4 inches in vertical and nearly 2 inches in transverse diameter. 

The expansion of the trunk is pretty equal in every dfrection towards the dorsal fin, and 
the upper surface gives the appearance of the fore part being subdepressed : the diminu- 
tion beyond the dorsal is more rapid fi-om side to side than from above downward. 
The greatest vertical diameter of the trunk is, in the male, 1 foot 6^ inches, in the 



32 PKOFESSOR OWEN ON INDIAN CETACEA. 

female 1 foot 4^ inches: the greatest transverse diameter of the tmnk in the female is 
1 foot 2 inches. 

The caudal fin, the shape of which is given m fig. 2, PI. X., measures, in the female, 
1 foot 7 inches in extreme breadth, and 7 mches across the base of each lateral lobe. 
Between the dorsal and caudal fins, and nearer the latter, the mid line of tegument is 
raised into a longish, very low and obtuse ridge. The vent opens 1 foot 10 inches in 
advance of the posterior cleft of the tail-fin in the male, and 1 foot 7 inches fi-oni 
the same part in the female. It is 10 inches behind the vertical line di-opped from 
the back border of the dorsal fin, in the male, and 8 inches behind the same part 
in the female. The vulva is three mches in advance of the vent ; the prepuce of the 
male is 9 inches in advance. 

The note, as to colour, accompanying the drawings is — " Above shining black, 
smooth ; beneath paler, pinkish, but in one discoloiu-ed with blood." The dentition is 
t-J=20. (PI. XII. fig. 1, X, 32). 

The Physeterida (Cachalots or Sperm-Whales) are characterized by having the open- 
ing of the mouth inferior in position, not terminal. The largest known species {Physeter 
macrocephalus, Linn.) has a reduced or boss-like representative of the dorsal tcgu- 
mentary fin, and a dorsal longitudinal ridge has been attributed to it near the base of 
the tail. The soft parts of the head, which project in advance of the jaws or opening 
of the mouth, form a large obtuse tnmcate mass. Tlie external blow-hole is reduced 
by its operculum or flap to a single sigmoid fissure on the left side of the upper and fore 
part of the head, i. e. at or near to tlie summit of the truncate end of tlie snout. The 
functional teeth are limited to the lower jaw, and chiefly to the long symphysial part ; 
those of the upper jaw, when present, are minute and concealed in the thick gum, in 
fossae which receive the summits of the larger lower teeth when the mouth is closed. 
The maxillary bones are so developed as to bound a large concavity, or chamber, for 
the " spermaceti," at the upper part of the skull in advance of the short brain-case (PI. 
XIV. fig. 2, 21' ). 

The question put by Cuvicr', wliether any large Sperm-Whale may exist, characterized 
as above, but with a liigh dorsal fin, with the blow-hole near the forehead on the 
middle of the head, and with the mandibular rami not united at a long dcntigerous 
symphysis, still waits a reply from a du-ect and good observer of such problematic 
Cachalot. 

The Sperm- Whale towed ashore in the harbour of Port Jackson, New South Wales, 
December 1849, and referred by Macleay to the species " Catodon australis"'^, had 
the blow-hole situated at the upper termination of the snout, as in the true Sperm- 
Whale^ ; and the dentigerous symphysis of the mandible was more than half the entire 

' Ossemens Fossiles, 4to. vol. v. pt. i. p. 340. 

' ' History and Description of the Skeleton of a new Sperm- Whale, lately set up in the Australian Museum,' 
by Wm. S. Wall, Curator. 8vo. Sydney, 1851. ^ j},_ p n^ 



PROFESSOE OWEN ON INDIAN CBTACEA. 33 

length of the ramus (48 inches to 92 inches)'. The blubber-portion of the carcase 
having been removed previously to the articulator's arrival on the spot^, no obser\a 
on the condition of the dorsal fin or hump was made. 

Cuvier characterizes the "Cachalot macrocephale" [Catodon macrocephalus, Art., 
Physeter macrocephalus, Lrnn.) as having the back provided with a slightly raised 
prominence, which some have called " fin," others " longitudinal ridge", others " hump" 
or "tubercle" {loc. cit. p. 338): "II a une dorsale tres-peu saillante vers I'arriere 
dudos, quelquefois reduite a une protuberance, ou a deux ou trois" (ib. p. 339). In the 
' Regne Animal,' Cuvier says, " II n'a qu'une eminence calleuse au lieu de nageoire 
dorsale" (tom. i. p. 294, ed. 1829). In the judicious criticism on the alleged or nominal 
species of Sperm- Whales, in the ' Ossemens Fossiles,' Cuvier asks, " Existe-t-il en outre 
des Cachalots a haute dorsale I en existe-t-Q dont I'event soit perce pres du front sur le 
milieu de la tete 1 en existe-t-il oil les branches de la machohe inferieure ne soient pas 
reunies sur la plus grande partie de leur longueur en une symphyse cylindi-ique 1 Voila 
ce qui reste a chercher, ce qui restc a prouver autrement que par des figui-es tracees par 
des matelots. Ce n'est qu'apres que des hommes eclaires auront observe ces etres 
avec soin, et en aiu'ont depose les parties osseuses dans des collections oii elles puissent 
etre verifiees par des naturalistcs, qu'il sera possible a la critique de les admettre dans 
le catalogue des animaux" (torn. cit. p. 340). 

As regards large Cachalots these questions, as I have remarked, still wait their solu- 
tion. In the small Cetacean called " Wonga," of the seas washing the eastern coast of 
the Indian peninsula, we have, however, a satisfactory reply to them. 

In it we possess a member of the Physeteridw — a Cachalot in fact — though small, in 
which the dorsal is lofty, with the usual shape of such weU-developed fin in Cetacea, in 
which the blow-hole is not terminal but near the forehead, and in which, as will pre- 
sently be shown, the mandibular rami are united by a symphysis of less than half the 
length of the " rami." The inferior mouth, unsymmetrical blow-hole, and the second 
tegumentary production in form of the dorsal ridge, shown in the careful di'awings by 
the native artist, significantly indicated the fiimily affinities of the "Wonga:" the 
enlightened attention and care bestowed by Mr. Elliot on this seldom-studied branch of 
zoology has enabled me to place this conclusion on unequivocal grounds, through his 
transmissi(<n, with the di'awings, of the skuU of one of the individuals figui-ed. 

To the study and comparison of this precious evidence I have devoted full attention : 
it is figured, half the natural size, in Plates XII., XIII., & XIV. fig. 1. Its peculiarity 
of form is extreme : perhaps no other Cetacean skuU has yet been observed in which the 
cranial so greatly preponderates over the rostral part. In the degree in which this pro- 
portion prevails in the skuU first made known by De Blainville as of the Cachalot whidi 
he called Phjseter breviceps^, and in that subsequently described by Macleay* under 

' OiJ. cit. p. 9. ' Oj). cit. p. 4. 

' Aonales Fran^aises et Etrangcres d'Anatomie et de Phy.siologic, tom. ii. (8vo, 1838) p. 335 : " Sur les 
Cachalots." ■* Op. cit. 

TOI>. VI. — r.\RT I. F 



34 



PEOFESSOK OWEN ON IKDIAN CETACEA. 



the name Euphijsetes grayi, may be discerned at a glance the more immediate affinities 
of the present species, which I propose to call Physeter (Euphi/setes) shims, in reference 
to its peculiarly short obtuse muzzle. 

Bescrijition of the Skull. 
(Pis. XII., XIII., & XIV. %. 1.) 

Short as is the upper jaw in proportion to the skuU in Phoccena Iremrostris (PI. EX.), 
it is shorter in the subgenus or section of Phjseteridce represented by the Physeter 
breviceps, De Bl. (PI. XIV. fig. 3), and shortest of all in the present species (ib. fig. 1). 

In the foUowmg Table of admeasui-ements are given those of the Physeter {Euphysetes) 
grayi, Madeay (the larger species which was stranded on the Maroobrah beach, near 
Sydney, New South Wales, and the skeleton of which is now in the Australian 
Museum of that city), with the few admeasurements appended by De Blainville to his 
notice of Physeter hreviceps, fii-om the Cape of Good Hope^ 



P. simus. 



P. grai/i. 



P. hreviceps. 



Length from the back of occipital condyles to end of snout .... 

Breadth across postorbital processes 

Breadth across the beginning of malo-maxillary fissure 

From the back of occipital condyle to antorbital process of 

malar 

From the antorbital process of malar to end of snout 

From the back of occipital condyles to posterior wall of left 

nostril 

From the bottom of raalo-maxiUary fissm-e to end of snout . . . . 
From the beginning of malo-maxillary fissure to end of snout . . 
Breadth of snout between the fore part of the antorbital notches 

of maxillary 

Breadth of snout at its extremity 

Breadth of premaxillaries at the malo-maxiUary fissure 

Breadth between anterior ends of premaxillaries 

Antero-posterior diameter of left nostril 

Transverse diameter of left nostril 

Antero-posterior diameter of right nostril 

Transverse diameter of right nostril 

Length of interfrontal crest, straight line 

Width of occipital foramen 

Vertical diameter of foramen 

Between outer edges of occipital condyles 

Breadth between paroccipitals 

From the lower border of basioccipital to vertex 

Length of mandible, in a straight line 

Length of alveolar series 

Height of mandible at coronoid ridge 



inches. 

10 

9 



7 
4 

4 
5 

4 

4 
1 
2 

1 
1 


3 
1 
1 
2 
6 
5 
8 
2 
2 



lines, 

5 

5 
10 

7 
3 

10 




9 

4 

3 

4 

5 



3i 

4" 

6 

3 

8 

9 

2 

6 

6 

9 

1 



inches. 

16 

14 

9 



2 
3 
1 
2 
2 

1 
6 
1 
2 
4 
11 
11 
13 
5 
4 



lines. 

6 

6 





8 



9 
3 



H 

6 

2 
6 
4 



3 
3 
3 



inches. 
15 



lines. 
5 



13 10 



In the skull of the Physeter simus the occipital elements have coalesced with each 
other and with the surrounding bones. The vertical diameter of the basioccipital 

' Annales Fran^aises et Etrangeres d'Anatomie et de Physiologic, torn. ii. tab. x. (the admeasurements 
are given in French inches), viz. : — " Longueur du crane 14 pouces et demi," = 15" 5'", Engl. " Longueur de la 
machoire inferieure 13 pouces," = 13" 10'", Engl. Ecartement de ses condyles 12 pouces,"=12" 9"' Engl. 



PEOFESSOE OWEN ON INDIAN CETACEA. 85 

(Pis. XII., XIII., & XIV. fig. 2, 1 ) beneath the foramen magnum (ib. o) is 8 lines : it is 
here convex vertically, and concave transversely, showing a width between the lower 
end of the occipital condyles (to which it probably contributed) of only 4 lines. These 
(PI. XII. fig. 2, 2') are more sessile than in PJiocwna bremrostris, being raised only 
by a linear border from the contiguous bone, except at thek lower ends, which are 
rather more prominent : the long diameter of the condyle is 2" 2'", the greatest breadth 
1" : they are terminal, diverge as they ascend the sides of the foramen magnum, which 
is widest opposite their upper ends : the outer border of the condyle is more convex than 
the inner one. The foramen magnum is oval, with the larger end upward and not 
notched : the aspect of the plane of the aperture is backward and a little upward : in 
Physefer rnacrocephahis (PI. XIV. fig. 2, o) it is more upward than backward. The 
ex- ( 2 ) and superoccipital ( 3, 3' ) plate inclines from below, upward, outward, and 
forward, with a moderate convexity or indication of a pair of such. The exoc- 
cipital portion (PI. XII. 2) extends outward and slightly downward, expanding a 
little vertically, and thickenmg to form the paroccipital ( ^ ) ; this expanse is moderately 
concave transversely, convex vertically. The border of the paroccipital is thick and 
rugged: it is concave toward the otocrane (PL XII. fig. 1, and PI. XIII. fig. 2, e), of 
which it forms the posterior half of the upper, and part of the posterior wall : it is 
divided below by a fissure (Pis. XII. & XIII. fig. 2, I) from the otocranial plate of 
the basioccipito-sphenoid (PI. XII. fig. 1, and PI. XIII. fig. 2,5'): this plate arches out- 
ward and downward, with a slight obliquity backward, and is overlapped anteriorly by 
the pterygoid (ib. 21'), which seems to form an anterior continuation thereof, converging 
towards its fellow: but the free border of the basi sphenoidal otocranial plate (5') is 
more obtuse and thicker than that of its pterygoid prolongation (24). A trace of the 
suture between the exoccipital (PI. XII. fig. 1, 2) and squamosal (ib. 27) remains. The 
ridge across the vertex (Pis. XII. & XIII. fig. 1, 7,11,3) is obtuse, but well marked : 
the proportions contributed by the superoccipital (3), parietal (?), and interparietal (if 
any) cannot be determined; and the frontal (n), as it ascends, contracting from the 
superorbital roof, is also blended with those constituents of the ridge'. The instructive 
harmonia between basi- (Pis. XIII. & XIV. fig. 1, s) and presphenoid (ib. 9) remains. 
The alisphenoid (Pi. XIII. fig. 2,6), coalesced with the basisphenoid, where it is 
underlapped by the pterygoid ( 24' ), is horizontal ; it extends to the lower border of 

' To afford a comparison with Physeter macrocephalus, I propose to append, in the present note, descrip- 
tions of the homologous cranial boues of a fcetu,s of that species described, in my ' Catalogue of the Osteological 
Series in the Museum of the Koyal College of Surgeons,' 4to. 1853 : — " The elements of the occipital neural 
arch are unanchylosed. The lateral margins of the anterior half of the basioecipital are produced and bent 
obliquely downward. The exoecipitals are much produced and expanded laterally : they are deeply notched 
below. The superoccipital coutributes the upper ends of both condyles : it is in the form of a vertical plate, 
convex from side to side : a strong internal vertical crest is produced forwards ; it is overlapped at its lower 
and lateral angles by the exoecipitals, anterior to which it reaches the alisphenoids, and is notched externally 
for the reception of the upper angle of the squamosal" (op. cit. p. 442). 

r 2 



36 PEOFESSOR OWEN OX INDIAN CETACEA. 

the temporal fossa (ib. t), underlapping the squamosal (ib. 27), and thinning-off to its 
outer margin : its anterior border is notched by the intersphenal fossa (tr) : there is no 
distinct foramen ovale. It supports the natiform protuberance of the cerebrum, and is 
divided from the orbitosphenoid (ib. 10) by the intersphenal fissure (tr), from which 
two channels lead toward the back part of the orbital roof (or), blending together 
and widening as they grow shallow^. The temporal fossa (PI. XII. & PI. XIII. 
fig. 2, t) is 1" Y" in antero-posterior, and 2" in transverse extent, has its marginal 
boundary almost completed by the approximation of the postfi-ontal (ib. 12) to the zygo- 
matic part of the squamosal (ib. 27), the distance between their free ends being but 6'" ; 
but the zygoma terminates on a lower level (PI. XII. fig. 1, 27). 

The presphenoid (Pis. XIII. & XIV. fig. 2,9) retains its distinction from the basi- 
sphenoid (5), but has coalesced with the orbitosphenoids (10), as have these with the 
alisphcnoids (e). 

The orbitosphenoid (ib. 10) has its posterior boundary partially defined by the inter- 
sphenal fissure, at the fore part of which the optic canal is marked off' by an intercranial 
process arching over the same downward and backward (PI. XIV. fig. 2, w) : the orbito- 
sphenoids expand and ascend to form with the coalesced frontals the anterior wall of the 
cranial cavity ; the optic channel extends forward and outward from the intersphenal fissure, 
and, blending with the trigemmal one (PI. XIII. fig. 2, tr), is lost on the roof of the 
orbit (ib. or)-. The fossa (ib. d), into which the foramina on the frontal or nasal plate of the 
maxillary opens, is in advance of the optic chaimel (ib. 10). There is no intraorbital fossa 
answering to that in Phoccena brevirostris. The roof of the orbit is unbroken, gently con- 
cave from before backward, formed chiefly by thefi-ontal (PL XII. fig. 1, 11, u), which is 
notched near the middle of the superorbital ridge : this is tliick, obtuse, and produced 
backward and downward into a postfr'ontal or postorbital process (ib. 12). Above the ridge, 
the frontal (ib. 11') contracts; its surface is here free fi-om the maxiUary (21'), is slightly 
concave vertically, before it is reduced by the overlapping of the parietal ( 7 ) and superocci- 
pital (3) behind, and of the maxillaiy ( 21' ) in front, to the naiTow strip (n), which rises, 
bending convexly, to the vertex. The fore part of the superorbital ridge ( H' ) iS ob- 
tuse, and thickens to join the malar (26), from which it is partly divided by a notch^. 

' " The basisphenoid, or thick hexagonal bone, concave from side to side below, nearly flat above, is anchylosed 
to the alisphcnoids : these are perforated near the middle of their base by the foramina ovalia and rotunda, 
have a thick quadrate plate on their inner side, forming their eutocranial surface : they extend into a point 
anteriorly, and articulate both with the frontal and with the parietal angle of the superoccipital. The 
squamosal receives the alisphcnoid in a groove anteriorh"." — Phi/seter miicrocephalus, op. cit. p. 442. 

- " The presphenoid and the anchylosed orbitosphenoids form the anterior wall of the cranial cavity, and are 
perforated by the optic foramina ; they articulate anteriorly with the frontal, sending up a small process into 
the interspace at the beginning of the frontal suture, which process is impressed by a fossa in each of its 
sides : the posterior and lateral parts of the orbitosphenoids unite with the great alse ; the under and anterior 
part is overlapped by the vomer." — Physeter macrucephalus, op. cit. p. 447. 

' " The frontals are large triangular plates, concave externally, with the outer and fore angle produced into 



PROFESSOR OWES ON INDIAN CETACEA. 37 

The vomer (PI. XIII. figs. 1 & 2, i3, i3') has partially coalesced with the presphenoid (ib. 
fig. 2,9) and underlapsthe prefrontals (PI. XIV. fig. 1, 1 4): it appears upon the palate, 
about an inch in advance of the posterior fissure (PI. XIII. fig. 2, w), expands to a breadth 
of 6 lines (is), and is continued to the anterior end of the upper jaw, which it forms, 
contracting there to a breadth of 3 lines. Its under surface is flat ; its upper surface 
(fig. 1, 13), which is similarly exposed on that aspect of the muzzle, is smoothly and 
widely canaliculate : the groove lodges the cartilage in the fissure separating the prcmaxil- 
laries (ib. 22), which cartilage terminates anteriorly the series of vertebral centrums, of 
which the vomer is the mferior or cortical ossification. The fore margin of the confluent 
prefi-ontals (ib. u) is at 3 inches distance from the fore end of the vomer. The prefrontal, 
losing breadth and gaining depth, recedes with a slight bend to the left, forming the 
inner boundary of the large left nostril (ib. ol) and the corresponding wall of the small 
right nostril (PI. XIV. fig. 1, ol'). The nasal bones are confluent with that osseous mass 
(PI. XIII. fig. 1,15) which rises from the back of the septum narium and extends in a sinuous 
course, first convex to the left and then concave before subsiding at the vertex (is'): this 
ridge also sends off" a kind of " spur" (15) fi-om its right side, in the form of a short 
ridge, inclining to the right, with a convex border, thick and obtuse like that of the 
main ridge : the intervening space (ib. 1/) between these ridges expands as it extends 
forward, with a smooth sinuous surface concave across slightly contracting again as it 
ends behind the right nostril ' . 

A trace of the suture of the palatines (PI. XIII. fig. 2, 20) shows that they entered into the 
formation of the bony palate for half an inch at the postpalatal end of the vomer (i3'), 
almost meeting each other behind that part : as they extend outward, they expand to a 
fore-and-aft breadth of 10'", with a convex surface, most so in their- du-ection from 
within, outward and backward, contracting to terminate mesiad of the fossa (d) : they 
develope no outer or free lamella in Euphysetes. 

a loDg superorbital process, the channel on the under part of which contracts, as it approaches the cranium, into 
a long, deep, and narrow groove. The median anterior part of the bone unites with both orbito- and ali- 
sphenoid, and external to this is the broad sutural surface for the squamosal. The straight median margins 
of the frontals are thinned off and joined by a squamous fi-ontal suture, the right overlapping the left. The 
whole posterior and lateral border of the frontals, as far as the junction with the squamosal, presents a bi-oad 
oblique sutural surface, which joins, by overlapping, the contiguous border of the occipital. The smooth 
cerebral surface of the frontal is flat at the middle, arched at the sides, and not impressed by any convolutions." 
— Physeter macrocephaJiis, op. cit. p. 442. 

' M. de Blainville figures, but makes no mention of this bony ridge bisecting the "postnarial" cavity. 
Dr. Gray, in appending the term Kogia to the Physeter breviceps, De Blainv. (Zoology of the Erebus and 
Terror, " Cetacea," 4to. 1846, p. 22), is equally silent — indeed, adds nothing to De BlainvUle's mea<Te 
sketch of so remarkable a cranium, and quotes his admeasurements as in English inches and lines, without 
correction for the difference of the French " foot." Macleay was the first who pointed out the heavy ridge 
of bone that longitudinally divides the spermacetic cavity into two unequal parts (op. cit. p. 47) as sub- 
generically distinguishing his Euphijsetes from Physeter or Catodon. 



38 PROFESSOR OWEN ON INDIAN CETACEA. 

The maxiUary (PI. XII. fig. 1, PI. XIII. figs. 1 and 2, 21) forms the major part of the 
bony roof of the mouth : a small triangular strip of the premaxillary (PI. XIII. fig. 2, 22) 
is wedged into the short anterior interspace between the maxillary (x) and vomer (13'). 
The palatal surface (21*) is moderately convex transversely, straight lengthwise, and 
is impressed by an alveolar groove (al) retainuig one socket and tooth (PI. XII. 
fig. 1, x) at the fore end and contmued m a straight line backward for 3 mches 
(rather more on the left, rather less on the right side) without indications of 
alveoli, and ui a line not parallel with the outer margin of the bone, but receduig 
to a distance of 1 mch from it, posteriorly ; so that the teeth, if developed there, woidd 
be rather palatal than marginal in position. The outer border of the maxiUary thickens 
near the malo-maxiUary fissm-e (21, k), with a smooth convex exterior. That fissure dilates, 
as it sinks obliquely backward and inward, to a breadth of from 3 lines to 4 lines, its 
depth being 1 inch 6 lines (k). These fissures mark off the rostral portion of the .skuU, 
which is here an cquUateral triangle, including above (PI. XIII. fig. 1) parts of the 
vomer (i3), prefrontal (14), premaxillaries (22), and maxiUaries (21) : the surface so formed 
is concave transversely at its posterior three-fouiths, almost straight longitudmally. The 
maxiUary, expanding backward beyond the rostrum, bends (at A', fig. 1) round the upper 
and back part of the malo-maxiUary fissure ; and in close conjunction (here partial con- 
fluence) with the malar (26), it forms the large smooth tuberosity ( 21, 21;) external to 
the fissure: from the tuberosity the convex raised border of the posterior expanded 
plate of the maxillary comes into connexion with the frontal (11), whence it subsides 
to form a deep hoUow as it sweeps inward to rise again upon the bifurcate suiuous ridge 
(ib. 15,15") which divides this smgular postnarial tract, or spermacetic cavity, of the upper 
surface of the cranium. The total breadth of this cavity is 6 inches 4 lines, the posterior 
three-fourths of its cii'cumfercnce, so bounded by the maxiUaries and describing as much 
of a circle, being a little produced backward, subangulaiiy, at the hindmost part : the 
open anterior fourth is continued upon the more shaUow concavity of the triangular 
rostrum. The right maxiUary is verticaUy pierced by two foramina (PI. XIII. fig. 1, a, h), 
which converge to the common inferior outlet (ib. fig. 2, d). The upper fissure between 
the maxiUary and premaxiUary widens and deepens as it extends backward, and terminates 
in the canal (fig. 1, c), also conducting to the fossa (fig. 2, d), which, as it transmits 
maxUlary branches of the fifth pau- fi-om the orbit to the exterior of the skull, is 
homologous with the antorbital foramen of other mammals : the altered position of the 
outlet, as regards the orbit itself, is the result of the reflection, so to speak, of the facial 
surface and nasal plates of the maxiUaries upon the forehead above and behind the 
orbits. 

The pterygoids (PI. XIII. fig. 2, 24) meet at the midsurface of the roof of the mouth, 
and extend the palatine suture {pi) backward beyond the palatine bones (20). From this 
line each pterygoid extends outward and backward, and divides into an internal and 
external pterygoid plate : the former terminates in a short triedi-al process, representing 



PKOFESSOE, OWEN ON INDIAN CETACBA. 39 

the "hamular" one ; the outer portion, partly marked off by a ridge from the palatine 
plate of the imier portion, bends outward and backward with a convexity toward the 
palate, then slightly inward, as if twisted on itself, and, expanding at its upper attach- 
ments to the pre-, orbito-, ali-, and basi-sphenoids, terminates by developing the 
deep and broad plate (ib. 2r ) which appears to continue forward the otocranial 
plate (b') of the basioccipito-sphenoid. The inner surface of the outer part of the 
pterygoid is vertically concave to its posterior lamella, which is so bent as to make that 
sui-face somewhat convex: the concave channel prolongs backward the nasal passage 
(w) beyond the septum. A semiciixular emargmation divides the posterior subvertical 
plate from the palatme portion (24) of the pterygoid. The total length of the pterygoid 
is 4 inches 8 Imes ; the breadth of the pair of bones posteriorly is 5 inches ; the sutural 
union of the pterygoid with smTOunding bones persists'. 

The malar bone (PL XII. fig. 1, PI. XIII. figs. 1, 2, 26) is represented in the present skull 
by the portion of that in Belphinidce (PI. IX. figs. 1, 3,26') which is wedged like a 
lacrymaP between the frontal (ir) and maxillaiy (21") at the upper and fore part of the 
orbit {or) : it is here of a subtriedral conical shape, with its base notched for a wedged 
union with the maxillary above, and concave where it joins the frontal behind : the iimer 
angle of the base curves forward, with a slight twist, to unite again with the maxillary 
at the inner side of the malo-maxillary fissure {k). The outer facet of the malar is 
slightly concave vertically, convex trairsversely : the antero-internal facet is concave in 
both du-ections, except where it curves anteriorly round the obtuse angle between it and 
the outer surface : the internal or orbital surface is the nan-owest, and is conxex trans- 
versely, and straight vertically. The apex is subbifid, the outer part (PI. XII. fig. l,z) 
low and obtuse, the inner one longer, produced downward and rather backward, and 
terminating less obtusely ; but there is no sign of any slender zygomatic style having 
been continued from this part, as in Phocwna brevirostris (PI. IX. fig. 1, 26). It would 
seem, therefore, that the zygomatic processes of both malar and squamosal were short and 
free ; they are separated by an interval of more than 2 inches in the present skuU, which 
interval I found occupied by a ligament (" sclerous" state of malar) in a young Cachalot^. 

The squamosal forms an articular surface (PI. XIII. fig. 2, 27, g) for the mandible, look- 

' " The pterygoid, which is douhle the size of the palatine, extends backward to the basioccipital, articulating 
in that course by its expanded upper border with the pre-, basi-, and ali-sphenoids ; from this border the bone 
descends arching inward toward its fellow, which it joins along the anterior half of its extent : the remain- 
ing free border is divided from this bj a deep notch, and circumscribes the posterior bony aperture of the 
nostril." — Physeter macrocephalus, op. cit. p. 443. 

^ If this be the homologue of a lacrymal, it is not merely confluent, but connate with the malar. 

' " The malar is moderately long and slender, bent upon itself at an acute angle ; the upper portion, 
wedged between the maxillary and frontal, is the thickest ; the lower and more slender branch is bent down- 
ward and backward, circumscribing the orbit anteriorly and below, and is connected by ligament to the 
zygomatic process of the squamosal. There is no lacrymal bone." — Physeter macroeephalus, op. cit. p. 444. 



40 PKOFESSOll OWEN OX INDIAN CETACEA. 

ing downward and forward: the surface is rather convex at the anterior border from 
behind forward, and is very slightly concave in the rest of its extent ; it is smooth and 
with an ill-defined cii-cumference : the anterior boundary, which also forms the posterior 
one of the lower outlet of the temporal fossa, is concave : the wall (Pl.XlI. fig. 1, 27') which 
the squamosal contributes to the posterior and internal part of the temporal fossa (t) 
expands as it bends forward to join the parietal (7) and frontal (11): the suture with 
the superoccipital ( 3 ) is close to the upper boundary of the fossa ; that with the 
exoccipital (2) continues a short way beyond the squamosal, and indicates the extent of 
the exoccipital. On the outer part of the base of the zygomatic or articular process the 
bone is tuberous, and represents the mastoid ( 8 ) ; behind the articular surface it is 
roughly excavated (PI. XIII. fig. 2, 8'), where it contributes, with the paroccipital (1), 
to the otocranial cavity^ 

In the interior of the cranium (PI. XIV. fig. 1) the upper or epencephalic surface of 
the basioccipital is moderately concave, and is bounded laterally by a short, obtuse, longi- 
tudinal ridge, dii-ected mesiad, which may be where the exoccipital suture ran : the outer 
or lateral begiiming of the tentorium receives a short angular ossification, which forms 
the outer wall of the fossa (c), perforated by the vagal and acoustic foramina, both of 
which pass directly outward to that at the back part of the fundus of the otocranial 
cavity (PI. XII. fig. 1, PI. XIII. fig. 2, e). A small branch channel from the vagal one 
opens upon the outer surface of the exoccipital at the groove which runs to the cleft 
(PL XII. fig. 2, 1) between the otocranial plates of the basisphenoid (5) and jjaroccipitals 
(4). At the fore part of the tentorial process (PI. XIV. fig. I, v) is the foramen of 
a canal which opens outwardly upon the alisphenoid : it is too small for the carotid, and 
may have given exit to a vein. I cannot discover any distinct entocarotid canal, any 
more than a distmct foramen ovale, foramen rotundum, or foramen opticum : they all 
seem here to be confounded in the mtersphenal fissure (PI. XIII. fig. 2, fij. From the 
extreme shortness of the jaws, the nerves of sensation to the face must have been very 
small. The " sella" (PI. XIV. figs. 1 & 3), scarcely impresses the basisphenoid : its best 
antero-extemal boundaries are afforded by the superoptic processes of the orbitosphenoid 
(ib. «). There is no ossification of the falx-, no trace of olfactory foramina. The great- 
est diameter of the cranial cavity is in the direction of breadth. 

The lower jaw (PI. XII. fig. 1, 29-32) is 7 inches 4 lines in a straight line from the back 

' " The squamosal is a comparatively small, but strong and thick, triangular bone ; the upper end repre- 
sents the expanded squamous part in land mammals, and is articulated by broad, dcntated sutural margins to 
the frontal and exoccipital : its anterior border is grooved for the reception of the aHsjihcnoid : the lower angle 
is as it were truncated, and presents a rough surface for the attachment of the petro-tympnnic : a short, 
obtuse anterior angle bends forward and represents the zygomatic process : the under surface presents a 
smooth shallow cavity for the condyle of the lower jaw : the inner border of the glenoid surface is proaurcd 
downward into a slender process." — PhystUr macroccphahis, op. cit. p. 444. 

2 In the Great Cachalot "a strong medial crest is produced forward from the inner sniface uf the super- 
occipital" (he. cit. p. 442). 



PROFESSOR OWEN OX INDIAN CETACEA. 41 

of the condyle to the fore end of the symphysis. Each ramus has a convex, almost 
semicircular posterior margin, curving upward and backward from below (3o), where the 
angle normally exists in other mammals, and then forward to the seat of the coronoid 
process (29) : at the hindmost part of this curve the border is thickened to form the 
sessile condyle, adapted to the glenoid surface of the squamosal. Here the border bends 
outward: as the ramus advances, converging to its fellow, it is slightly bent witli the 
convexity outward, which again is changed to a concavity (lengthwise), where it joins 
the opposite ramus to form the elongate symphysis (32), which is continued straight 
forward to its termination. The symphysis here forms rather less than a thu-d of the 
entire length of the mandible, being 2 inches 4 lines in extent. The greatest vertical 
diameter of the ramus is 2 inches 2 lines ; that at the beginning of the symphysis is 
8 lines^ In the alveolar groove are partially excavated sockets for nine teeth ; the four 
middle intervals are severally equal to twice the basal diameter of the tooth : at the 
ends of the series, especially the anterior one, the alveolar intervals are less. The teetli 
(PL XII. fig. 1, and ^) are small, straight, conical, obtuse, not exceeding 8 lines in 
length, of which the cylindrical base has a diameter of 2 lines, that of the crown a 
diameter of 1^ line, with a length of 2^ lines, diminishing to a subrecurved apex. 

The loss of symmetry in this skuU. is hardly observable in the general contour, 
whether viewed from above (PI. XIII. fig. 1) or below (fig. 2) : it is chiefly, almost 
exclusively, confined to the nostrils and the bones concerned in the composition of those 
passages ; and this is only conspicuous in the upper surface of the skull. 

In Euphysetes breviceps, Bl., according to the figure of the side view of the skuU (copied 
in PI. XIV. fig. 3), the occipital condyle is more jn-ominent than m Eiiphysetes sinms(V\. 
XII. fig. 1) : the contour of the superoccipital is concave in Euphysetes breviceps, but is 
convex in Euphysetes simus — very feebly so, indeed, but as far as it departs from a straight 
line being in the direction of convexity. The most marked difference, however, is th<> 
greater proportional length of the rostral part of the, skuU — measured, viz., from the ma- 
lomaxiUary fissure (ib. & PI. XIII. k) to the end of the upper jaw (^2, x) : in Euphysetes 
breviceps it forms about two-fifths of the entire length of the skull, in Euphysetes simus 
about two-sevenths. The proportion of the maxillary, above the frontal and malar, on 

' " The condyle of the mandible projects from the posterior part of the ascending ramus, which is com- 
pressed and produced into a low obtuse coronoid process above, and into a similar angle below: a wide 
excavation, beginning at the inner side of the ascending ramus, deepens and contracts into the dental canal 
which enters the substance of the horizontal ramus : a fissure is continued along the inner side of the ramus 
from this canal, and is the sole indication of a compound structure of the jaw. The vessels and nerves emerge 
from several foramina at the outer side of the ramus, -where it is attached by its long symphysis to its fellow : 
the upper border of the syraphysial part of the ramus is excavated by a continuous dentigerous groove, some- 
what resembling, in the present foetal state, that in the upper jaw. The length of the symphysis in this 
skull is three-fourths that of the rest of the ramus. In the adult male the disproportionate growth of this 
part of the jaw leads to more excessive length of the symphysial part beyond the rest of the ramus." — 0/'. 
cit. p. 444, fa?tal Physeter macrQcephalus, 

VOL. VI PART I. G 



42 PROFESSOR OWEN OX INDIAN CETACEA. 

the exterior of the skull is much greater in Euphysetes breviceps than in Euphysetes simus, 
especially in vertical extent : in the upper ^'iew of the skull the porportion of the postnarial 
cavity, especially m breadth, to the extent of the rostrum is less in Euphysetes Ireviceps 
than m Euphysetes simus. To these differences must be added the difference in the number 
and shape of the teeth. In Euphysetes breviceps there are fourteen or fifteen teeth, or 
isockets for as many, in each mandibular ramus : the entire tooth, figured by De Blain- 
ville (copied in PI. XIV. fig 2. b), is 10 lines in length, and has a proportionally larger 
and more curved crown than in Euphysetes simus. De Blainville writes, " II me parait 
a pen pres certain qu'il n'y avait pas de dents a la machoire superieure" [l. c. p. 337) ; and 
these are equally absent in Euphysetes grayi : the first of the maxillary series remains 
exposed, as a functional tooth, in the quite adult skull of the smaller Indian species, 
Euphysetes simus. From Euphysetes grayi the present species differs not only in this 
dental character and its smaller size, but in its proportionally shorter muzzle, and in the 
minor number and wider disposition of the mandibular teeth. Thirteen teeth are found 
in each ramus of the lower jaw of the specimen of Euphysetes grayi in the Sydney 
Museum : they are divided by interspaces of less than theii- own basal diameter, and 
have relatively longer crowns than those of E. simus. There are twelve teeth in the 
right, and nine teeth in the left ramus of the mandible of Euphysetes breviceps, De 
Blainv. : they are as wide apart as in Euphysetes simus, but have crowns more slender 
and recurved. 

In the figures of the mandible given by De Blainville {loc. cit. pi. 10), and by 
Macleay {loc. cit. pi. 2. fig. 5), the breadth between the outer parts of the cond)les 
equals the length of the mandible m a straight line, that is, from the middle of the 
chord di-awn between the condyles to the end of the symphysis. In Euphysetes simus the 
breadth exceeds the length so taken. 

Among other differences between the present member of the Physeteiidce and the 
Belphinidoe (see Phoccena brevirostris, PI. IX. fig. 1) is the non-production of the upper 
or hinder expansion (naso-frontal plate) of the maxiUary (PI. XII. fig. l,2i», 2i") over 
the orbital process of the frontal ( u, ii' ) ; which, therefore, in Euphysetes simus as in 
Euphysetes breviceps, stands out free (PI. XII. fig. I, u' ) from the upper and lateral 
parts of the cranium behind the maxillary ( 21' 21"). 

Bones of the Trunk and Fins. (PI. XI. fig. 2.) 

Having been favoured with photographs of these bones in Euphysetes grayi by the 
present able Curator (]\Ir. Kreffts) of the Australian Museum, I have thought it might 
be useful to add the following notes : — 

Euphysetes (PI. XI. fig. 2) has fifty vertebrae, viz. seven cervical, fourteen dorsal, 
twenty-nine lumbari-sacro-caudal : in the latter series the hsemapophysial arch first 
appears between the sixth and seventh (or between the twenty-seventh and twenty- 
eighth vertebrae counting from the skuU) : the hsemapophyses cease to be developed at 



PEOFESSOR OWEN ON INDIAN CETACEA. 4a 

the twentieth (or forty-first from the skull), leaving ten, perhaps eleven, terminal 
vertebrae represented by depressed centrums, gradually diminishing to the last. The 
seven cerncals are anchylosed : the diapophyses distinguish the atlas and axis, the former 
of which vertebrae does not retain, as in Physeter, its separate condition ; the fifth, sixth, 
and seventh are lamelliform, from extreme anteroposterior compression. The dorsal spines 
progressively, but very gi'adually, gain in height to the last; beyond which they again, 
and more rapidly, shorten to the base of the tail, disappearing in the fortieth vertebra 
from the skull. The metapophysis begins to project above the prozygapophysis in the 
fifth dorsal, and supersedes that process in the articulation of the neural arches in the 
seventh or eighth dorsal. The four anterior pau's of ribs du-ectly join the sternum, 
which consists of three sternebers, each more or less completely divided at the middle 
line into two bones. The fii-st rib is broad, flat, and angularly bent, articulated by the 
tubercle to the first dorsal diapophysis, and by a ligament representing the head to the 
centrum of the seventh cervical : its connate sternal portion articulates with the antero- 
extemal angle of the manubrium. The second and six following ribs have both head 
and tubercle, the former abutting against the interspace of their owti and antecedent 
centrums ; the tubercle of the rib is attached to the diapophysis of its own vertebra : 
the second rib, less broad but one-fourth longer than the first, has a short, partly 
ossified cartilage, which joins the interspace between the first and second sternebers. 
The thii'd, gaining length, losing breadth, and with more regular curvature, is arti- 
culated by its short hsemapophysis to the interspace between the second and third 
sternebers. The fourth rib is joined to the end of the third stemeber. After the seventh the 
ribs lose their heads, become shorter, more slender, less curved- — gradually to the tenth, 
which is 9 inches in length — suddenly in the fourteenth, which is a straight style 
is hardly an inch long. There are two pairs of pelvic bone. The pectoral fins are 
relatively short and rather obtuse. The scapula is a flat triangular plate, with a con- 
vexly curved base, in extent equalling the fore-and-aft range of the five anterior dorsal 
spines. An obtuse rising near the anterior costa, at its humeral half, developes near 
the glenoid cavity a small coracoid directed forward. The acromion is much larger, and 
is produced from a greater extent of the anterior costa in the form of a parallelo- 
gram. The ulna developes scarcely any olecranon. There are five digits : the first and 
fifth are the shortest, each with a metacarpal and two phalanges; the second and 
third digits are the longest, with five and four phalanges respectively, besides the 
metacarpal ; the fourth digit, intermediate in length between the third and fifth, has a 
metacarpal and four phalanges. 

Conclusion. 

The first remark that I am led to make on a review of the cetacean characters above- 
defined in connexion with those pre\iously recorded is, that they are all gradational, 
and exemplify steps by which are gained the extreme modifications, especially in the 

skull and dentition. 

g2 



44 PEOFESSOE OWEN ON INDIAX CETACEA. 

Imperfect as may be the cetacean record, it yields several series of differential cha- 
racters, — as, e.g., in the proportion of the rostral to the cranial part of the skull, from 
Physeter simus to Physeter macrocephalus and Platanista — in the degree of expansion of 
the back part of the maxillaries, exemplified, step by step, in Balmna, Delphinus, PJio- 
cmna, Ziphius, Euphysetes, and Physeter, again culminating in Platanista — in the number 
of teeth, from zero (Balceiia and old Pelphinapteri), through Monodon, Zipkius, Euphy- 
setes, to the multitude of teeth in BelpMnus, Cuv. 

The formation of germs of teeth in parts of the jaws of foetal or young individuals of 
species which are edentulous in the full-grown mdividuals, the examples of which 
are too well known to need citation here, are, perhaps, amongst the most significant of 
the gradational modifications, above referred to, being due to deviations in offspring 
from the characters of parents. 

Such departures or variations may have been slight in the first instance, few and far 
between in the members of a contemporary generation, and rare exceptions to the rule of 
hereditary likeness ; but, occurring in the course of many generations, through long lapse 
of time, they might lead to " long-snouted" and " short-snouted" breeds, and to others 
exemplifying the various observed cranial and dental modifications of cetacean structures. 

In such conjectural mutations of specific characters may be discerned a fore-ordained 
law of deviation from primitive type, through the operance of which the ocean 
has at length become peopled with so many strange modifications of the cetacean 
structure. 

But such instances of exceptional freedom from the trammels of family likeness seem 
to be independent of external influences. The ocean has none of those diversities of 
condition which the dry land shows, and is exempt from the few which in fresh waters 
may be invoked to account for varieties in the species of fish. It is true that the trout 
{Salmofario) of the mountain-streamlets is small, while that of the wide river or wider 
lake is large ; but no such differences can be invoked to explain the origin of the 
dwarf Euphysetes or the giant Physeter : both have alike the unlimited seas for their 
range. 

But the same river may have the pike, the carp, the salmon, the eel, &c. ; these 
modifications of the piscine type exist in waters of the same temperature, same rate of 
riow, and same nature of bed. AVhere can we here discern selective influences equiva- 
lent to produce such changes of structure ? The hypothesis is still less conceivable in 
regard to the ocean. The various Cetacea of the Indian seas exist in a medium of the 
same nature, exempt from any influence of the earth beneath them, or of aught that 
may there live and grow. The external influence or power that could " select " the 
maxillary wall of the circumnarial basin, e. g., in Ilyperoodon, Ziphius, Euphysetes, 
Physeter, Platanista, is inconceivable. 

But the occasional departure from parental type, manifested by a so-called abnormal 
or monstrous proportion of the nasal or facial plate of the maxillary, may accord 



PROFESSOR OWEN ON INDIAN CETACEA. 45 

with the idea suggested by the obsei-ved steps in a gradation of such deviational 

developments. 

So far the species thereby characterized may be held as evidences of orderly succession 
and progression due to inherent organic force, operating according to a natural law or 
" secondary cause," of the precise nature of which we are yet in ignorance. But we 
may feel assured that the Power which called into bemg the first cetacean type fore- 
knew and planned, by predetermined degrees and kinds of departure from that type, 
all its subsequent modifications'. 

But much knowledge of the facts of organization is still needed for successfully 
gi-appling with these transcendent questions ; and the progress of zoology has been 
slower in regard to the Cetaceans than to most other orders of animals. 

This is due to their medium of existence, to the extreme latitudes at which some of 
the species have to be sought for, and to the vast bulk which certain species attain^. 
The latter characteristic precludes the preservation and exposition of the requisite spe- 
cimens in private collections or even in those of associations of the cultivators of 
natural history willing to carry on the work of advancement of the science at their 
own cost and to the extent of their means and usually limited incomes. 

The diversities of structure exemplifying specific characters in Balwna, Bal(enoptera, 
Physeter, Hyperoodon, Sec, and those which have suggested as many subgeneric divisions 
and names of the Cuvierian genera of those gigantic animals, are best exemplified in their 
skeletons, both by modifications of particular bones, and by proportions of the several 
regions of the skeleton ; but the framework of these animals, put together to exemplify 
their articulations and proportions, require for their exhibition the resources of a 
National Museum. There, and there only, can an intelligent public and the student of 
this branch of Mammalogy expect to find the means of contemplating and comparing 
the characters and structures of the strangest as well as hugest of animals— the most 
seldom seen, by reason of their ocean haunts— air-breathers, yet living in water— hot- 
blooded, tliough ever surrounded by a rapidly refrigerating medium— of man's own class 
by every essential of organization, but fishes in shape— a recent development of life- 
form on our planet, and the superseders of the great sea-lizards in their office in the 

ocean police. 

Hitherto the expectations of both student and sightseer have been disappointed. 
Space (the first essential towards fulfilling this exigency) has been found too costly ; 
at all events the guardians of the public purse have thought it not desirable, as yet, to 
vote the sums requisite for the galleries, however simple in structure, which are neede.l 
for the Cetaceous Department of a Zoological Museum^ 

• Owen, ' On the Nature of Limbs,' 1849, p. 86. 

' I may also add, from aggravating experience, the conflicting claims to the legal ownership of such monsters 
of the deep when they happen to be cast upon any part of the shores of Great Britain. 
= See Hansard, ' Debate on Museum of Natural History,' May 19th, 1862, p. 1928. 



46 PROi'ESSOE OAVEN ON INDIAN CETACEA. 

DESCRIPTION OF THE PLATES. 

PLATE HI. 

Delphinus (Steno) gadamu : diminished to scale. 
Fig. 1. Side view. 
Fig. 2. Upper view: b blow-hole. 

PLATE IV. 

Delphinus gadamu. 
Fig. 1. Side view of skull (wanting back part of cranium). 
Fig. 2. Side view of mandible. 
Fig. 3. Upper view of mandible : ss symphysis. 
Fig. 4. Symphysial end, inner view of mandible. 
Fig. 5. Bony palate. 

All the figures are nearly half the natural size. 

PLATE V. 

Fig. 1. Delphinus fusiformis, side view (diminished to scale). 
Fig. 2. Delphinus lentiginosus, side view (id.). 
Fig. 3. The same, upper view. 

PLATE VI. 

Fig. 1. Delphinus maculivenfer (to scale of Plate V.). 
Fig. 2. The same, upper view. 
Fig. 3. Delphinus pomeegra (id.). 

PLATE VII. 

Delphinus fusiformis. 
Fig. 1. Side view of cranium and upper jaw. 
Fig. 2. Side view of lower jaw. 
Fig. 3. Upper view of cranium and upper jaw. 
Fig. 4. Under view of ditto. 
Fig. 5. Upper view of symphysis of lower jaw. 

All the figures are nearly half the natural size. 

PLATE VIII. 

Fig. 1. Side view of cranium and upper jaw of Delphinus pomeegra. 
Fig. 2. Under view of upper jaw of ditto. 
Fig. 3. Side view of under jaw of ditto. 



PROFESSOR OWEN ON INDIAN CETACEA. 47 

Fig. 4. Upper view of symphysis of under jaw. 

Fig. 5. Under \'iew of upper jaw of Delphinus euphrosyne. 

All the figures are half the natural size. 

PLATE IX. 

Phocmna brevirostris. 
Fig. 1. Side view of cranium and upper jaw. 
Fig. 2. Upper view of ditto. 
Fig. 3. Under view of ditto. 

All the figures are nearly half the natural size. 

PLATE X. 

Euphysetes simus. 
Fig. 1. Side view of female. 
Fig. 2. Upper view of ditto (drawn to scale). 

PLATE XL 

Euphysetes simus. 
Fig. 1. Side view of male (to same scale as female, PI. X.). 
Fig. 2. Outline of ditto, with skeleton. 

PLATE XII. 

Evphysetes simus. 
Fig. 1. Side view of skull. 

Fig. 2. Back view of skull (rather more than half the natural size). 

PLATE XIII. 

Euphysetes simus. 
Fig. I. Upper view of skull. 
Fig. 2. Under view of ditto (half the natural size). 

PLATE XIV. 

Fig. 1. Section of cranium oi Euphysetes simus. 

Fig. 2. Section of cranium of Physeter macrocephalus. 

Fig. 3. Euphysetes hreviceps. 



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1 PHYSETER 5IMUS.2. PHYSETER MACRO CEPHALUS 3.PHYSETER BRBVICEPS 



[ 49 ] 



III. On the Osteology of the Dodo (Didus ineptus, Linn.). 
By Professor Owen, F.B.S., F.Z.S., dr. 

Read January 9th, 1S6G. 

[Plates XV. to XXIV.] 

§ 1. Introduction. 

IHE Dodo has long been one of the "curiosities of Natural History," through the 
rarity and paucity of the material evidences of the bird. 

The dried foot in the British Museum, the dried head and foot in the Ashmolean 
Museum at Oxford, the skull, lacking the lower jaw, and somewhat mutilated, in the 
Gottorf Museum at Copenhagen, were all the parts of the bird known to the authors 
of the admh'able monograph on the Dodo and its kindred at the date of its pub- 
lication'. 

Subsequently a portion of the bone of the upper beak has been discovered in the 
Museum of Natural History at Prague^. 

Such, until the present date, was the sum of the remains of this large, flightless, 
extuict bird which were knovm to have reached Europe. 

The happy perception, by the Danish Professor J. Reinhardt, in 1843^, of the 
resemblance of the beak of the Dodo to that of the tropical Doves, generically 
separated by Cuvier imder the name Vinago, on account of their proportionately larger, 
more strongly arched, and compressed beak than in other Pigeons, and the still closer 
resemblance, in miniature, of the beak of the Samoan Dove to that of the great Mau- 
ritian bird, which led Titian Peale to give to the former the generic name I/idun- 
cuius, du'ected the ornithologist and omithotomist to the family in which the most 
instructive comparisons might be made; and the results of these, so far as relates to 
to the head and foot and the bones of those parts, published by the authors of the 

' ' The Dodo and its kindred ; or, the History, Affinities, and Osteology of the Dodo, Solitaire, and other Ex- 
tinct Birds of the Islands Mauritius, Rodriguez, and Bourbon.' By H. E. Strickland, M.A., F.G.S., F.R.G.S., 
President of the Ashmolean Society, <fec., and A. G. Melville, M.D. Edin., M.E.C'.S. 4to, London, 1848. 

^ See Annals of Nat. Hist. ser. 2. vol. vi. p. 290 (18.50). 

' " Es war in 1843, dass ich auf den Gedanken kam, dass der Dodo eine anomale Taubenform sei ; ich 
Uberzeugte mich bald dass diese Auffassung die einzig lichtige sei, und iing an eine Arbeit iiber diesen Gegen- 
stand vorzubereiten. In 1845 wurde ich aber von meiner Kegierung beauftragt eine Eeise um die Welt mit 
einem diinischen Kriegsschiff mitzumachen ; meine Arbeit musste also vorlanfig bci Seite gelegt werden. Schon 
vor meine Abreise hat ich aber mehrere sowohl danische wie fi-emde Naturforscher niit meiner Ansicht bekannt 
gemacht, und der Beweis das es sich so verhiilt wird Owen finden konneu : — 

" 1. in den Forhandlingar de Scandinaviske Naturforskers Mode, i Kjobenhavn, 1847, p. 948 : und 

"2. in Sundevall, Arsberattelse om Framstegen i vertebrerade Djureus Naturalhistoria og Ethnographien, 
1845-50, p. 254." — Letter from Prof. J. Eeinhaedt to Dr. Albekt Gunihee. 

VOL. VI. PART II. H 



50 PROFESSOK OWEN ON THE 

above-cited work, left little doubt of the " striking affinity which exists between this 
extinct bird and the Pigeons"'. 

Whatever doubt, indeed, may have lingered in the minds of naturalists as to this 
affinity will probably be finally set at rest by the results of the comparison of the 
large proportion of the skeleton of the Didus ineptus which has at length been 
transmitted from the island of Mauritius to London, under the following circum- 
stances. 

In 1863, 1 was favoured by Miss A. Burdett Coutts with an introduction to the Bishop 
of Mauritius, then in this country, and I endeavoured to interest his lordship in aiding 
or promoting the acquisition, by the British Museum, of the zoological rarities of Mada. 
gascar, and especially of any remains of the Dodo which might be discovered in the 
island of Mauritius, to which his lordship was about to return. 

How speedily and successfully the Bishop has fulfilled my latter desire will be shown 
by the following letter, with which I was favoured in November, 1865. 

" St. James, Port Louis, 
'■' October 7, 1865. 
" My dear Sir, — when I had the pleasure of conversing with you for a short time 
in London two years ago, I promised to acquaint you with any facts or discoveries 
which might come to my knowledge, likely to interest you in connexion with Mada- 
gascar. I have not anything as yet to communicate definitely respecting that island 
in the way of natm-al history, but I have strong reasons to believe that a discovery 
has been made here recently which will gratify you very much. Mr. George Clark, 
who has for many years devoted himself to the work of teaching in this island with 
great success, is an ardent student of natural history, and has explored many parts of 
the island in search of information on the subject. From careful observation he was 
led to conclude that no remains of the Dodo were likely to be found in any of our 
watercourses, because of their steep descent and the immense rush of water which 
sweeps down them at times. But he had also frequently expressed his opinion that 
in certain marshes, with high banks of sand between them and the sea, such remains 
would probably be found. In one of these places he has found several of the bones 
of the Dodo (as he believes), and is now forwarding them home for your in- 
spection ^. 

At his request, I write these lines to ask for your kind care of his interests in 
securing any reward which may accrue to him. It would be a great pleasure to me 
to find that his discovery was really important, and likely to be useful to himself; 
for he has pursued these and similar investigations with an amount of intelligence, 
skill, and dihgence, in his vacation -times (by no means extensive), which deserves much 
credit and encouragement. 

' Eeinhardt, quoted by Strickland, op. cit. p. 41 (see also p. 70). 
. ' This Collection was purchased by the Trustees of Ihe British Museum for the sum of ^100. 



OSTEOLOGY OF THE DODO. 51 

" The book which you kindly sent me on the Aye-Aye has been read by many, and 
especially by medical men, with much interest. I entrusted the other copy to Mr. 
John Douglas for the Society here. 

" I remain, my dear Sir, 

" Your very faithful Servant, 

(Signed) " Vincent N. Mauritius." 

" Professor Owen." 

This letter was accompanied with the following " Statement " by Mr. George Clark, 
Master of the Government School at Maheboiu-g, Island of Mauritius : — 

" On the estate called ' Plaisance,' about three miles from Mahebourg, in the island 
of Mamitius, there is a ravine of no great depth or steepness, which, apparently, once 
conveyed to the sea the drainings of a considerable extent of circumjacent land, but 
which has been stopped to seaward, most likely for ages, by an accumulation of sand 
extending all along the shore. The outlet from this ravine having been thus impeded, 
a sort of bog has been formed, called ' La Mare aux Songes,' in which is a deposit of 
alluvium, varying in depth, on account of the inequalities of the bottom, which is formed 
of large masses of basalt, from three to ten or twelve feet. The proprietor of the 
estate a few weeks ago conceived the idea of employing this alluvium as manure ; and 
shortly after, the men began digging in it ; when they had got to a depth of three or 
four feet they found numerous bones of large tortoises, among which were a carapace 
and a plastron pretty nearly entire, as also several crania.^ 

" When I heard of this, it immediately struck me that the spot was one of the most 
likely possible to contain bones of the Dodo, and I gave directions to the men working 
there to look out for any bones they might find. Nothing, however, was turned up but 
a fragment of what I supposed to be the humerus of a large bird. This encouraged 
me to look further ; and my search was rewarded by the discovery of several tibia, more 
or less perfect, two tarsi, one nearly perfect pelvis, and fragments of three others. 

"These were found imbedded in a black vegetable mould, the lighter-coloured 
specimens being near the springs.. My reasons for believing these to be remains of the 
Dodo are : — the certainty that that bird once existed in Mauritius ; the similarity of 
these bones to what the representations of the Dodo which I have seen would lead one 
to expect, particularly the breadth of the pelvis, the stoutness of the tibiae and tarsi, 
and the shortness of the latter ; the favourable nature of the spot in which they were 
found for the haunts of such birds when liAing— a sheltered hollow with two springs 
in it ; the non-existence, actual or traditional, in Mauritius of any bird to which bones 
such as these could have belonged ; the indubitable antiquity of these bones, proved by 
the deposit of alluvium which covered them. 

" During neariy thirty years that I have mhabited this colony, I have made frequent 
inquiries of old people as to the finchng of the bones of large bu'ds, and have offered liberal 



52 PEOFESSOR OWEN ON THE 

rewards for such ; and I have consulted with the late Dr. Ayres as to the spots most 
likely to contain them. We agreed that the floods which sweep the hill-sides and the 
ravines in the rainy season would be most likely to carry any remains into the sea ; 
and this would doubtless have been the case here, but for the stoppage occasioned by 
the sand-do^vn. (Signed) " George Claek. 1865." 

The above " Statement " was authenticated by the following testimony : — 
" Having visited the place with Mr. Clark, 1 can vouch for the truth of the facts 
herein mentioned. (Signed) " William Thomas Banks, 

" Civil Chaplain, Mauritius." 
"The Eev. W. T. Banks, Civil Chaplain at Mahebourg, in this diocese, and Mr. 
George Clark, Master of the Government School at Mahebourg, are well known to me, 
and deserving implicit credit for their statements as to matters of fact. 

(Signed) "Vincent N. Mauritius. Oct. 6, 1865." 

§ 2. Description of the Bones. 
The bones of the Dodo [Bidus ineptus, Linn.) discovered by Mr. Clark, under the 

above circumstances, which have reached me up to the present date (December 20th, 

1865) are the following: — 

Name. Number of bones or parts. 

Cranium and lower jaw, in parts 14 

Vertebrae and pelvis 30 

Ribs 22 

Sternum 2 

Scapular arch, in parts 7 

Humerus, ulna, radius 6 

Femora 5 

Tibise 6 

Fibulae 4 

Metatarsals 4 

Total number of parts of skeleton of the Dodo 100 

The known characters of the skull and metatarsus of the Bidus inejjtus sei-ved to 
identify those bones as belonging to that species : the agreement in relative size, colour, 
condition, and locality left no room for hesitation in referring the other bones in the 
above list to the same species'. They belong, however, to four or five individuals 

' So determined, subsequent sets of bones transmitted from the Mauritius, and from which I was privileged to 
select the most perfect specimens for the present memoir, got into the market and were sold by auction since the 
present memoir wa.s in type, as bones of the Dodo. I hare to express my sincere and grateful acknowledgements 
to those gentlemen into whose hands these lots have fallen, who have forborne their own advantage and refrained 
from rushing into print with figures from inferior specimens to anticipate the appearance of a memoir notified 



OSTEOLOGY OF THE DODO. 53 

varying somewhat in size. Vt'ith the bones of the Dodo were the end of the lower jaw 
of a broad-billed Parrot, two bones (radius) of a small Mammal, and part of the skull of 
a large Tortoise'. 

To the description of the Dodo's bones I now proceed. 

§ 2. Vertebrce. 
The dorsal vertebrae are chiefly represented, in this series of bones, by three which are 
anchylosed together by their bodies and neural arches (PI. XVII. figs. 1-5) : the posterior 
articular surface of the body of the last of these vertebrae (ib., fig. 4, c) is subquadrate, 
longer in the vertical than the transverse direction, concave vertically, convex trans- 
versely, almost fitting, but being rather too small for, the anterior articular surface of 
the body of the first of the sacral series (PI. XIX. fig. 1, c). The difference is such as 
to indicate that only one dorsal vertebra may have intervened ; and I conclude that 
the last of the three coalesced vertebrae is the penultimate dorsal. The anterior arti- 
cular surface of the foremost of the three (PI. XVI. fig. 1, c) is 11 lines in transverse, 
and 4 to 5 lines in vertical diameter : it is concave transversely for the middle three- 
fifths, and convex transversely at the two outer fifths of its extent : it is more or less 
convex vertically throughout its extent. The bodies of these vertebrae are compressed 
and wedged-shaped, slightly expanded at their coalesced ends, produced below into 
subquadrate hypapophyses in the first and second (PI. XVII. fig. 1, hy) ; while this 
process is restricted to the fore part (ib. hy 3), or may be represented only by a slight 
anterior production of the lower edge of the wedge, in the thii-d (ib. fig. 5, hy 3). 



(in the ' Proceedings of the Zoological Society,' January 9th, 1866) as destined " to be published entire in the 
Society's Transactions," and therefore necessarily awaiting the lithographing of " illustrations," which every true 
promoter of science for its own sake must have desired to see as complete as the best-selected materials would 
permit to be given. — K. 0., June 1866. 

' In the quaint print, in foUo 3, of the " Narration Historique du Voiage faict par les huict Navires cV Am- 
sterdam an mois de Mars I'An 1598. soubs la conduitte de 1" admiral Jaqnes CorueiUe Necq,'' &c., the first- 
named object, No 1, " Sont Tortues qui se tiennent siir I'haut pays, frustez d' aisles pour nage, de telle grandeur, 
qu'ils chargent ung homme et rampent encore fort roidement, prennent aussi des Ecriuisses de la grandeur d'un 
pied qu'ils mcngent. 2. Est ung oiseau, par nous nomme Oiserni de Nmisee, h, I'instar d'une Eigne, ont le cul 
rond, convert de deux ou trois plumettes crespues, carent des aisles, mais en lieu d'icelles ont Uz trois ou 
quatre plumettes noires, des susdiets oiseaux avons nous prins une certaine quantite, accompaigne d'aucunes 
tourtureUes et autres oiseaux, qui par noz compaignous furet prins, la premiere fois qu'U arrivoyent au pays, 
pour chercher la plus profonde et plus fraische Kiviere, et si les navires y pourroyent estre sauvez, et retour- 
nerent d'une grande joye, distribuant chasque navire, de leur Vcnoison prins, dont nous partismes le lendemain 
vers le port, fournismes chasque navire d'un PUote de ceux qui au paravant y avoyent este, avons cuict cest 
oiseau, estoit si coriace que ne le povions asses boviUer, mais I'avons menge a demy cm. Si tost qu'arrivames 
au port, envoya le Vice-Admiral nous, avecq une certaine troupe au pays, pour trouver aucun penple, mais 
n'ont trouve personne, que des TourtureUes et autres en grande abondance, lesquels nous prismes et tuames, 
car veu qu'U n'y eust personne qui les effraia, n'avoient Uz de nous nuUe crainte, tindret lieu, se laisserent 
assomer. En some c'est un pays abodant en poisso et oiseaux, voire tellemet qu'il excella tons les autres 
audit voyage." — Le Second Livre de la Navigation des Indes Orientales, fol., 1601, 



54 PEOFESSOE OWEN ON THE 

The hypapophysis of the first of the three expands at its termination (PI. XVI. fig. 1, 
hy), with the hinder angle bent back to coalesce with the front one of the next hypapo- 
physis, which is somewhat longer, and bent forward with a similar terminal expansion : 
a full elliptical space is intercepted by this terminal confluence of these hypapophyses 
(PI. XVII. figs. 1 & 5, hy). Each vertebra shows an elliptical articular cavity (ib. figs. 
] Si. b, p,j)^) for the head of the rib, near to the anterior articular surface ; the long 
axis of this costal surface is directed from above obliquely downward and forward. The 
surface of the rib's tubercle cuts obliquely the lower part of the free end of the dia- 
pophysis (PI. XVI. fig. 1, d). 

The neural arch circumscribes a canal the anterior outlet of which (ib. fig. I, n) is 
oval with the small end downward, 5 lines in vertical, and 3 J in transverse diameter: 
the sides of the neural canal slightly project inward above the lower third : the posterior 
outlet (PI. XVII. fig. 4, n) is more regularly elliptical in form, and rather narrower in 
proportion to its vertical diameter. The neurapophysis sends off from the outer and 
fore part of its base a stout process, which expands and divides into zygapophyses (PL 
XVI. fig. 1, z) and diapophyses (ib. d); the articular surface of the former is of a 
full oval shape, flat, looking obliquely upward and inward; the diapophyses extend 
outward and a little backward : the articular surface for the tubercle of the rib is 
transversely elliptical and nearly flat. The hinder part of the neurapophysis exjiands 
into the postzygapophyses : these have coalesced with the preezygapoj^hyses in the suc- 
ceeding vertebra (PI. XVII. fig. 2, z), as has happened also between this and the tliii'd 
vertebra. In the last of the three vertebrae the postzygapophyses are entire (ib. z 3), 
and show very slightly concave, oval articular surfaces, looking obliquely downward and 
outward (ib. fig. 4, ;:). The conjugational foramina, continuously surrounded by bone, 
are a full ellipse, and large, the anterior one (ib. figs. 1 & &,_/") being 5^ lines in vertical 
diameter ; the second (ib. /'') is somewhat less : these foramina are also rather larger in 
one of the specimens than in the other. The length of the three coalesced dorsals is the 
same in both, viz. 2 inches 3 lines. The neural spines have run together into a con- 
tinuous ridge in fig. 1, ns ; in fig. 5 the summit is broken off in both, leaving only the 
anterior angle of the foremost entire ; in both this inclines forward ; the hinder border 
of the third vertebra (fig. 1, «s) has the same vertical parallel as the back part of the 
centrum. The anterior margin of the base of the spine shows a rough surface for the 
attachment of ligament (PI. XVI. fig. I, ns). A small foramen behind the base of each 
of the coalesced zygapophyses (PI. XVII. fig. 2, z z) leads to a canal descending to 
the neural one, and indicates superiorly the limits of the othervrise continuously ossified 
neural arches. 

In the series of detached vertebrse, one (PI. XVII. fig. 6 & 7) indicates by its neural 
spine and hypapophysis a position at the base of the neck. The centrum is barely an 
inch in length ; its anterior surface (ib. fig. 7, c) is narrow vertically, broad transversely ; 
both fore and hind surfaces indicate freedom and extent of flexure. The hypapophysis 



OSTEOLOGY OF THE DODO. 55 

has a broad, bituberculate base (ib. hij), but is limited in fore and aft extent to the 
middle third of the under surface of the centrum : its length is shown in fig. 6, hy. The 
parapophysis (fig. 7, p) is slender, and expands at both attachments, with an indication 
of a terminal surface. The diapophysis {(I) has a larger costal surface : it sends for- 
ward a convex ridge midway between the di- and zygapophysis [z). The neural canal 
(fig. 7, n) has wider and more fully elliptical outlets than the hinder dorsal vertebra, 
in relation to the greater extent of motion at the fore part of the series. I conclude 
that a free pleurapophysis {pi) existed, indicating the present to be the first of the dorsal 
series, as shown in PI. XV. Tiie neural spine is short, broad, obtusely pointed, with a 
vertically oblong syndesmotic surface (fig. 7) before and behind. Each postzygapophysis 
(fig. 6, z') supports an anapophysial tubercle [a). 

A cervical vertebra from a position just in advance of the above has lost the neural 
spine, but retains the hypapophysis. This process (ib. figs. 8 & 9, hy) is compressed 
and directed obliquely downward and forward for an extent of G lines ; the extremity is 
rounded : the length of the centrum of this vertebra is 1 inch -3 lines ; the anterior 
articular surface is longest transversely, and concave in that direction, convex vertically ; 
the proportions and curvatures are transposed in the posterior surface (fig. 9, c). The 
parapophysis (ib. p) is continued from the anterior border of the centrum to the 
middle; it is a depressed plate, confluent with the rib (ib. d). The diapophysis 
forms a short, obtuse projection above its anchylosis with the rib (ib. pi): this 
projects backward 7 lines in length, terminating obtusely, and cuxumscribing a ver- 
tebrarterial foramen (ib. v) of a full elliptic shape, 5^ lines in long diameter. The 
surfaces of the praezygapophyses {z) are larger, and look more upward and less inward, 
than in the preceding and the dorsal vertebra; : they are veiy slightly concave. Those 
of the postzygapophyses (fig. 8, z), with a downward and slightly outward aspect, are 
in a similar degree convex. The neural canal, as usual in the cei-vical series, expands 
at its outlets, most so posteriorly (fig. 9, n) ; the middle of the upper surface of the 
neural arch is impressed by an elliptical, rough, ligamentous surface, which slightly 
rising in the middle is the sole indication of a neural spine. The upper surface of 
each postzygapophysis developes a tuberous anapophysis (figs. 8 & 9, a). 

The three cervicals that succeed the axis show progressively sinking neural spines, 
which subside in the six following vertebrae (PI. XV.). The third cervical has also the 
hypapophysis (PI. XXIII. fig. 3, hy). 

In all the other cervicals of the present series the hypapophysis is wanting, but each 
parapophysis developes a plate (PI. XVII. figs. 10 & 11, PI. XX. fig. 1, p) to form the 
sides of the haemal canal through which the carotids ran ; and the position of such ver- 
tebrae in the cervical series is indicated, respectively, by the degree of convergence of 
these processes, in none of which, where entire, have they met so as to circumscribe the 
canal : in some of these vertebrae, however, they are mutilated. They differ chiefly in 
the position and shape of the anapophyses (fig. 10, a), which advance from above the 



56 PEOFESSOK OWEN ON THE 

postzyo-apophyses (z'), converging towards the middle of the upper surface of the neural 
arch, bemo- aiTested, save in one instance, at the sides of the ligamentous surface 
occupying the common position of the base of the neiu-al spme. 

In the axis vertebra (PI. XVII. figs. 12 & 13) the posterior articular surface, concave 
vertically, and 3 lines in that extent at its middle part, is very convex transversely, being 
continued upon the sides of the posterior part of the centrum ; a thick obtuse hyp- 
apophysis (fig. 13,/?^) descends below this surface: the anterior or odontoid surface 
presents the usual form in bu-ds; the odontoid process (ib. x) has a pit at its apex. 
The prezyo-apophyses (fig. 12, z), of very small size, project from the outer and fore 
border of the nem-al arch, with their articular surface looking outward and slightly 
upward; a ridge is continued from their back part to the base of the postzygapo- 
physes: the surface (fig. 13, z') in these, 4^ lines in long diameter, is three times the 
size of the anterior one ; it is concave transversely, and looks downward and a little out- 
ward. The anapophyses (ib. fig. 12, a) are large tubercles rising above the articular 
surfaces. The base of the neural spine, 9 lines in length (ib. ns), is coextensive with 
the neural arch ; the spine rises posteriorly to a height of 6 lines, with a thickness of 2 
lines, having a convex upper margin (PI. XV.). 

The relative size and position of the cervical vertebrae, as coadjusted in the position 
and degree of flexure of the neck represented in Sir Hans Sloane's life-size painting of 
the Dodo, in the British Museum, are given in Plate XV. with the varying proportions 
pf the pleurapophyses and other processes. 

§ 3. Bibs. (Plates XV. & XVI.) 
The specimens of ribs include both vertebral and sternal portions ; that which appears 
to be the second or thu'd on the right side (PI. XVI. figs. 7, 7 a) is 4 inches 4 lines 
in length (following the outer curve), and expands to a breadth of 7 Imes at its lower 
part; the interval between the articular surfaces of the head and tubercle is 6 lines, 
'i'he appendage (ib. a) has coalesced with the middle of the hind margin of the shaft. 
The neck js compressed, with a thin upper margm ; the lower one is continued with 
a curve upon a strong internal buttress-like ridge (ib. b), which runs to near the fore 
part of the flattened body of the rib, where it meets the ridge continued from the 
tubercle, about 2 inches down the rib : there is a shallow chaimel between these 
ridges, contracting to their confluence. The inner surface of the rib is impressed by 
a deeper and broader channel behind the buttress : the posterior border expands in 
in the form of a triangular plate, with a base of about an inch in extent, due to the 
complete confluence there of tjie epipleural process. The anterior border is thicker, 
and is almost straight. Towards the sternal end the plem-apophysis contracts and thickens, 
terminating in a rough syndesmotic elliptical surface, 3 lines by 2 (fig. 7,/"), for the at- 
tachment of the hsemapophysis or sternal rib. 

A vertebral rib (ib. fig, 2) which is entire, measi^res 9 inphps in length (following the 



OSTEOLOGY OF THE DODO. 57 

outer curve). The head and tubercle are at the same distance as in the preceding, but 
the tubercle is broader. The characters of the body of the rib are very similar ; but it 
is narrower,' not attaining a breadth of 5^ lines at its lower end; the narrowing and 
thickening to the articular surface for the sternal rib is more gradual. 

A last vertebral rib is adapted, by tlie longitudinal extent and partial division of the 
tubercle, to the vertebra which forms the first of the coalesced series of sacrals ; and the 
body of the rib, instead of preserving the regular outward curve of the antecedent 
ones, is more suddenly bent soon after it emerges beyond the margin of the ilium ; the 
lamelliform part thence continued is straighter, and, moreover, shows upon its outer 
surface a flattened facet, indicative of pressure or friction by the movements to and fro 
of the thigh over a rib in such position. Beyond this surface the rib curves in a way 
not shown in the other specimens ; the distal end has the flat syndesmotic articular 
surface to which had been attached a haemapophysis not reaching the sternum. In 
this last (eighth) free rib there is no epipleural process, nor any definitely marked liga- 
mental surface on the posterior mai-gin indicative of the attachment of such process. 

The body of a posterior vertebral rib (PI. XVI. fig. 10) shows a fracture which has 
been healed, with some irregular ossific deposit on the inner surface. AU the ribs have 
a pneumatic foramen (ib. figs. 2, 7, 8 p) at the fore part of the neck, near the base of 
the tubercle. 

The eight left vertebral ribs (PI. XV.) and the five right ones do not, either of them, 
constitute a consecutive series, but have come from different individuals, of diff'erent 
sizes, as exemplified in the third rib figured in Plates XV. and XVI. 

The sternal ribs (PI. XVI. figs. 3 & 12) are characterized by the two facets, nearly 
or quite meeting at an open angle, into which their .sternal end expands (ib. fig. 3, c). 
One of these ribs, which is entire, shows the single, elliptic syndesmotic surface at the 
opposite end (ib. h) ; it is ?>\ inches in length, with a greatest breadth of 5 lines, and 
is straight. Another and longer specimen (ib. 12) shows a moderate degree of curvature. 
A third specimen is 6 inches in length : the proximal end has a breadth of nearly 
half an inch (the penultimate rib in PI. XV.). 

Five successive sternal ribs are indicated by gradational size and curvature, and a 
sixth, which does not reach the sternum. Before describing this bone I shall proceed 
with the account of the sacral vertebrae, and the expanded haemal arches of such as 
complete the pelvis. 

§ 4. Pelvis. (Plates XV. & XIX.) 

The pelvis of the Dodo is chiefly remarkable for the flatness and great breadth of the 
posterior half, corresponding with the characteristic proportions of that part of the body 
in the old woodcuts of the Dutch " Dodaersen'"'. It includes sixteen coalesced sacral 
vertebiae, with which the iliac bones are continuously confluent. 

' See, especially, Bontekoe's figure, copied by Strickland, in tlie title-page and at p. 03 of the above-cited work. 

VOL. VI. P.^RT II. I 



58 PBOPESSOB OWEN ON THE 

The first sacral shows the transversely extended and concave articular surface of the 
centrum (PI. XIX. lig. 1, c) ; the subcircular pit (ib. 2') for the head of the rib is behind 
the middle of the side of the centrum, at its upper part ; the inferior surface is ridged 
lengthwise ; and a transverse low but sharp ridge defines the posterior boundary, the 
depressions in front of which indicate the hindmost origins of the subvertebral muscle 
(longus coUi 1). The anterior outlet of the neural canal (ib. n) is subcircular in one spe- 
cimen, vertically elliptic in others, and 3 lines or less m transverse diameter. From the 
sides of the neui-apophyses stretch out the strong buttresses of bone which blend with 
the under part of the ilia, giving off from the fore part of theu' base the prsezygapophyses 
(ib. z), and from the back part of their apex the surface (ib. d), or part of it, for the 
tubercle of the last moveable rib, the ilium in the latter variety affording the rest of 
that surface. The fore part of the strong neural spine (ib. ns) is roughened by a syn- 
desmotic surface ; it rises to a height of 14 Imes, ciu'ving forward, and is confluent at its 
summit with the approximated anterior margins of the ilia. A continuous track of bone, 
forming a smoothly obtuse longitudinal ridge, represents the summits of the succeeding 
sacral spines (ib. fig. 2, nts) to the hindmost vertebra of the series, without any trace of 
their primiti\'e division ; but this track rises, posteriorly, above the shallow channel on 
each side, in which are the foramina (ib. o), indicating most of the constituent vertebrae. 

The second sacral vertebra abuts against the ilium by a pleurapophysis (ib. fig. 1, 
j)l 2), as well as a diapophysis (ib. d 2) ; but the former is a slender, straight filament, or 
narrow plate of bone, confluent at both ends. 

In the next two vertebrae the pleurapophysis (ib. ^j/3&4) assumes more breadth and 
robustness, but is short and straight, abutting against the inner surface of the ilium an 
inch in advance of the acetabulum. The flrst of these rib-buttresses inclines forward, 
and is completely confluent with tlie ilium ; the thicker one (ib. jjl 4 ) has retained part 
of its primitive ligamentous attachment to the ilium : the proportions of both are sub- 
ject to some variety. 

These are succeeded by three or four vertebrae in whicli the pleurapophysis is not de- 
veloped, the attachment to the ilia being by diapophyses only (ib. d d), which are short 
slender lamellae, directed upward and backward ; below and between them are the double 
orifices for the separate motory and sensory roots of the sacro-spinal nerves. In the 
next vertebra the pleurapophysis (ib. 2^1 ^ ) reappears, longer but more .slender than in 
the fourth sacral, extending obliquely backward, and expanding at its extremity to abut 
against a prominence on the underside of the ilium, opposite the hind part of the 
acetabulum, with which prominence the rib has completely coalesced by an expanded 
end. The under part of all these vertebrae is traversed by a sharp median longitudinal 
ridge, which is more feebly and interruptedly continued to near the end of the sacral 
series. 

Eight vertebrae, abutting by diapophyses only (ib. d d) against the ilia, succeed the 
one last described ; their coalesced bodies are less than half the breadth of those of the 



OSTEOLOGY OF THE DODO. 59 

preceding vertebrae : they gradually diminish in depth to the last, without loss of breadth. 
The diapophyses proceed obliquely outward and backward, are lamelliform, about 9 
lines in length, and intercept oblong cavities of the same extent and direction, into which 
open the orifices (ib. fig. 2, o) noticed on the upper surface of that part of the pelvis. 
The articular surface of the body of the last sacral is transversely elliptic, 4 lines by 
2 lines, and very slightly convex. The outlet of the neural canal, above it, is circular, 
and about a line in diameter, the whole vertical extent of the last sacral being 5 lines, 
while that of the first sacral is 2 inches 2 lines. 

The ilium is divided, as usual, into two parts by the ridge on its upper or outer 
surface (ib. fig. 2, r), extending obliquely backward to behind the acetabulum — the an- 
terior di^'ision being narrower and concave, the posterior broader and convex but m a 
minor degree. The anterior (slightly thickened) border of the ilium is curved with the 
convexity forward, extending 8 or 9 lines in advance of the fore part of the neural spine 
of the first sacral vertebra. The ilia almost meet above that of the second and third 
sacrals, with which they coalesce, and then diverge to the oblique boundary ridge, 
which is thence continued, in some with an angular bend, more directly outward. 
At this angle the bone is so confluent with the sacrum that the orifices leading 
to the ileoneural canals' are almost or quite obliterated. These canals are, here 
(ib. i i), the longitudinally extended cavities intercepted between the fore parts of 
the ilia and the continuous coalesced sacral spines and diapophyses, widening to 
their anterior outlets. The extent of that part of the ilium in advance of the 
acetabulum is 3 inches 8 lines ; the breadth at its middle part is 2 inches. As 
the ilium approaches the acetabulum it increases in thickness, and is grooved at 
the outer margin by a vessel which leaves impressions of its ramifications upon the 
upper concave surface of the bone (ib. fig. 2, 62). The acetabulum (ib. a a) is cir- 
cular, 11 lines in the diameter of its outlet, 9 or 10 lines in that of its inner circum- 
ference, being widely open, as usual in birds, towards the cavity of the pelvis ; the tro- 
chanterian surface (ib. 1 1) above the acetabulum is elliptic, with the long axis length- 
wise, 9 lines by 6 in its diameter, with its upper border sharp and produced ; the 
anterior border (ib. h) of the acetabulum is slightly produced ; the position of this ar- 
ticular cavity is about midway between the fore and hind ends of the pelvis. The 
oblique external ridge of the ilium terminates in the outer margin of the broader part 
of the bone (ib. j"*), 7 lines above the sharp and prominent margin of the trochanterian 
surface (ib. t). The ilia have diverged from each other for the extent of an inch and a 
half behind the beginning of the boundary line (ib. r), which interval is occupied ex- 
teriorly by lateral ossification from the neural spines to the diapophyses of that part of 
the sacrum: the mesial borders of the ilia (ib. fig. 2, 62') slightly converge to the 
fifteenth sacral vertebra, where they are separated by an interspace of 1 inch, and tlien 
again diverge to the last sacral ; they coalesce with the diapophyses (ib. fig. 2 d d). 
' Owen, 'Anatomy of Vertebrates,' 1866, vol. ii. p. 3i!. 

I 2 



60 PEOFESSOE OAVEN ON THE 

The inner or under surface of the iUum is thickened into a kind of buttress (ib. fig. 1, e), 
termuiating behind the ischiadic foramen. The breadth of the iliac bones and inter- 
venmg sacrals, 1 inch behind the acetabulum, is 5 inches; at the back part of the 
pelvis it is 4 inches. The outer border of the posterior part of the ilium (ib. fig. 2, g) 
projects as an obtuse ridge above the ischiadic foramen and the succeeding expanded 
and confluent part of the ischium (ib. 63), which is vertically concave externally : the 
ilium, ischium, and pubis (ib. fig. 1, m) have completely coalesced around the aceta- 
bulum. The pubis, vchich in this part is 7 lines thick, contracts as it becomes free 
to a diameter of 4 lines ; it is smooth and convex below, and has been broken off near 
the acetabulum on both sides ; the fracture shows its pneumatic structure. The ischium, 
as it recedes from the acetabulum, contracts to a trihedral column, with a vertical dia- 
meter of 4 lines; it is conca\e outwardly, convex inwardly, and suddenly expands 
below, about an inch from the acetabulum, to form part of the posterior boundary of the 
obturator foramen (ib. fig. 1, /'), which is 9 lines in length, and is situated one lialf in 
advance of, and the other half beneath, the ischiadic foramen (ib. in). Tliis latter 
is oval, with the large end forwards, 1 inch 3 lines by 10 lines in its principal dia- 
meters. Behind this foramen the ischium is confluent with the ilium for an extent of 
2 inches, or perhaps rather more, as the posterior margin of the pelvis is not entire in 
any of my specimens. The inner surface of the ischium forms a low, obtuse longitu- 
dinal ridge towards the pelvic cavity, losing thickness as it recedes from the acetabulum. 
The chief pneumatic foramina in the pehis are on the inner surface, above tlie aceta- 
bulum, behind the trochanterian articulation, and behind the iliac confluence of the last 
sacral pleurapophyses, — also at the hinder part of the ilium, on each side of the transverse 
buttress (ib. e) near its posterior junction with the ischium. The prcerenal fossa (be- 
tween^? 4 & j)Z 8, fig. 1) is deep and subdivided by the diapophysial plates : the post- 
renal fossa is wide and shallow. 

§ 5. Sternum. 

Of this instructive and determinative bone there are two specimens, the one most 
entire (Pis. XV., XVI. fig. 4 4 , & XVIII.) measuring in a straight line, from the costal 
process to the hind border, 7 inches. The extreme breadth between the lateral pro- 
cesses (PI. XVI. h) is 4^ inches; from this diameter the bone contracts anteriorly to a 
breadth of ?)\ inches at the costal processes (ib. d), and posteriorly it contracts more 
rapidly to an obtuse, horizontally flattened apex (PI. XVIII. fig. 3). The anterior 
border of the sternum (PI. XVI. fig. 4.) is widely and rather deeply emarginate at 
the middle (e), less deeply so on each side : the breadth of the mid notch [h e h) is 
1 inch 9 lines, that of each side notch {b d) is 1 inch 2 lines. The sternum is deeply 
hollowed above (PI. XXIII. fig. 4), correspondingly convex beneath (ib.); the keel (s) 
is low and thick, commencing by a pair of broad obtuse ridges (Pis. XVI. fig. 4, & 
XVIII. fig. 1, r r) from the mesial ends of the outer walls of the coracoid grooves 



OSTEOLOGY OF THE DODO. 61 

(ib. b' ), which gradually rise from the surface of the bone as they extend backward, con- 
verging to form the beginning of the keel about 2 inches from the anterior emargina- 
tion {(') : the keel gains a depth of f of an inch at the middle of the sternum, then 
gradually sinks to the level of the bone, as it extends backward, at li inch from the 
hind end (PI. XVIII. fig. 3), a little increasing in thickness as it subsides : its free 
border describes a pretty regular convex curve (PI. XV.) ; it is thick, flat, partially 
canaliculate : the sides of the base of the keel expand, to be continued gradually into 
the body of the sternum (PI. XXIII. fig. 4). Behind the costal surface (PI. XVIII. r), 
on each side, extends a lamelliform process (Pis. XV. & XVIII. h), ^ an incli in breadth, 
upward and a little outward, slightly expanding to its free termination, which, however, 
is not entire in either specimen : the longitudinal extent of this characteristic process, 
where it is best preserved, is 1 inch ; it is conjecturally restored in Plate XV. ; it answers 
to the ectolateral process (h) of the gallinaceous sternum (Pis. X. & XIV. fig. 3): 
there is no trace of an entolateral process (ib. /). The thin margin of the Dodo's breast- 
bone, behind the ectolateral process (Pis. XV. & XVIII. Ji), is entire and uninterrupted 
to the obtuse apex, and the body of the sternum is imperforate : the notch (f) behind 
the process (/*) represents the ectolateral notch of the gallinaceous sternum (PI. XXIV. 
figs. 1 & o,f). The costal border (PI. XVIII. fig. 2, c) is 1 inch 9 lines in extent, and 
6 lines across its broadest part ; it shows articular surfaces for five sternal ribs, of which 
the four posterior (2—5) are bdobcd, the anterior one (c 1) simple, and limited to the 
outer half of the border ; the second sternum shows some variety in this respect : the 
deep interspaces, in both, are perforated by pneumatic foramina. The costal process 
{(l)^ in advance of these surfaces expands, as it rises upward and a little outward and 
forward, to the extent of nearly an inch ; the hinder and outer side is impressed by a 
concavity, continued fi'om the costal border ; the inner side is smooth and convex : it is 
not quite entire on either side. The coracoid grooves (PI. XVI. fig. 4, b b') are small 
in proportion to the sternum, and are divided from eacli other by an interspace of about 
an inch ; the outer wall of the groove (i'), 9 lines in extent, is moderately produced and 
convex ; it appears to be a continuation of one of the initial ridges (/•) of the keel : the 
inner wall of the groove (h) is deeper, and is formed by the obtuse angle of the anterior 
border of the sternum, between tlie medial and lateral emarginations. External to each 
coracoid groove is a large elliptical pneumatic foramen {p) or depression. Tliere is no epi- 
sternal process. On the convex outer surface of the body of the sternum the '• j^ectoral " 
ridge (PI. XVIII. fig. 1, Ji)~ is feebly indicated, extending from the outer end of the 
coracoid groove backward and inward to near the posterior third of the keel. The con- 
cave surface of the sternum (ib. fig. 2) shows a number of small pneumatic foramina, 
chiefly along the middle line to near the posterior third. Behind the costal border the 

' Called "hyosternal '" in the Geoffiroyan determination of parts of the bird's sternum. 

^ The intermuscular ridges (' pectoral,' ' subcostal,' carinal ') are, with other parts of the bird's sternum, . 
here named as defined in my ' Anatomy of Vertebrates,' vol. ii. pp. 10-23. 



62 PROFESSOR OWEK ON THE 

substance of the sternum gradually increases in thickness from the sharp lateral margins 
to the middle, above the base of the keel, and shows there a fine pneumocancellous 
texture (PI. XXIII. fig. 4). 

§ 6. Scapular Arch. (Plates XV. & XX.) 
This consists of the scapula (PI. XX. figs. 6, 7, 8 & 9, si), coracoid (ib. figs. 4 & 5, s'j), and 
clavicle (ib. ss), the latter ending in a point and here tied by ligament to its fellow, to form 
a furculum. I have received the elements of this arch in three conditions : — one in which 
the bones, though of full size, are separate ; a second, in which the scapula and coracoid 
are confluent, but the clavicle distinct ; a third, in which the three bones are confluent 
at the ends converging to the humeral ai'ticulation. The scapula (ib. figs. 6 & 7, 8 & 9, 
5i), 3 inches 7 or 8 lines in length, has the usual sabre-shaped body, slightly expanding 
and decurved at its free extremity, the breadth of which is 7 lines : it terminates ob- 
tusely : varieties of shape are shown in figures 6 & 8. The outer surface of the bone, 
at the two posterior thirds of its extent, is slightly concave and marked by muscular 
attachments ; the inner surface of that part is smooth and slightly convex : the bone 
increases in breadth, with some diminution of thickness, towards the articular end, and 
is remarkable for sending off from the lower border, at 7 or 8 lines from that end, 
a sliort process (ib. 5i) ; between this process and the articulation the breadth of the 
bone is little more than •" linos ; the breadth of the articular end is 9 lines. Nearly 
une-half of it is occupied by the almost fiat, subcircular humeral surface (fig. 8, a), with 
a diameter of A\ lines, and directed upward, outward, and a little forward. From this is 
continued an oblong, much narrower coracoidal surface, beyond which the acromial pro- 
cess (fig. 6, c) extends forward, curving toward the coracoid, and terminating obtusely. 

The coracoid (ib. figs. 4, 5, 8 & 9, 62), averaging a length of 3 inches 7 lines, expands 
to a breadth of 1 inch 3 lines at its sternal end (52), of which the articular surface {e) 
occupies an inch ; the non-articular part forms the outer angle [in), and extends in advance 
of the pneumatic foramen (PI. XVI. fig. 4, ^>) at that part of the breast-bone: the outer 
border which extends from this free angle to the body of the bone, into which it sub.sides, 
at one-third of the extent of the bone, is sharp ; the inner border is obtuse to near the , 
inner angle (PI. XX. figs. 4 & 5, n). The outer surface of the expanded sternal end 
is smooth and convex; the inner surface is flatter and more irregular, perforated by 
pneumatic foramina ; the diameter of the subcylindrical part of the shaft is 4 lines : the 
extremes of difference in the distal expansion of the coracoid are shown in figs. 4 & 8, 
.■)2, PI. XX. A muscular ridge and rough surface (ib. fig. 9, r) mark the back part 
below the middle of the shaft. The bone then expands to its upper articular end, 
which is obliquely truncate from within outward : it shows, fii'st, the oblong surface 
for the scapula, which is extended upon the inner prominence of that end ; next, the 
larger and full oval surface for the humerus (A), from which the thick, obtuse, inner 
'continuation of the scapular end projects inward, forward, with a slightly upAvard curve. 



OSTEOLOGY OF THE DODO. 63 

and shows the narrow oblong surface for the articulation and ultimate confluence of the 
clavicle (os). The coracoid unites with the scapula at an angle of 100°. 

The clavicle (ib. figs. 4 & 5, os), at its scapular end, is slightly expanded, compressed, 
with an obtuse recurved termination articulating with the above-named surface of the 
coracoid, and in one instance coalescing therewith, and by extended ossification with the 
" acromion scapulae" (ib. figs. 8 & 9). As the clavicle descends it curves slightly and 
contracts to a point. The angle at which the pair meet is shown in figs. 4 & 5. 

§ 7. Bones of the Wing. (Pis. XV. & XX. figs. 12-17.) 
Of the humerus the series contains two specimens, both measuring 4 inches 3 lines in 
length, one right, and the other left (PI. XX. figs. 12-14), but difi'ering slightly in their 
proportions and in colour — one being of the olive-brown tint with which most of the 
bones are stained, the other black. The articular head (ib. a) is an elongate oval con- 
vexity, with the larger end toward the radial side, prominent toward the back and rather 
flattened toward the front of the bone, which there swells out beyond the base of the 
articular surface. The radial tubercle is small, and descends from the radial end of the 
head for about 5 lines; the pectoral process (ib. h) is triangular, obtuse, short, and 
bent, or directed toward the front side of the bone : the ulnar tuberosity (ib. c) is more 
produced in that direction ; it is oblong, obtuse, with its base impressed by a large pit 
both above (fig. 12, h) and below — the lower one (ib. fj) being the deepest, and perforated 
by a pneumatic foramen ; the convex, broad, ulnar border of this tuberosity has two 
slightly produced processes, an upper or posterior (ib. fig. 12, c) and a lower and internal 
(ib. //), which is the smallest. The breadth of the proximal end of the humerus, across 
the tuberosities, is 1 inch 5 lines, beyond them the bone contracts to a smooth subcylin- 
drical shaft, showing at the back part of the proximal third a longitudinal ridge (fig. 12, »•), 
half an inch in length ; it gradually expands at the distal third to a breadth of 10 lines, 
where the articulations off'er the usual avian characteristics of the elbow-joint. The 
head of the humerus is occupied by a fine cancellous structure : into the large vacuity 
below this, crossed in the section figured (PI. XXIII. fig. 5) by a transverse slender bar 
of bone, the small pneumatic foramina at the bottom of the wide and deep fossa for the 
axillary au--cell open. The part of the hollow proximal end giving off" the pectoral 
and other processes for the attachment of muscles is strengthened by similar abutments. 
The pneumatic cavity of the main part of the shaft of the humerus is simple, with a 
compact wall thicker than at the ends of the humerus, but not exceeding that which is 
characteristic of the long air-bones in birds. The portion of the distal end chiefly 
serving for muscular attachments and the antibrachial articulation are also cancellous. 

The radius (Pis. XV. & XX. fig. 15) is a straight and slender bone, 3 inches 1 line in 
in length, and 2 lines in chief diameter of the shaft. The proximal articular surface is 
subcircular, 3 lines in diameter, moderately concave ; the distal end expands to the same 
extent, but is compressed, as usual. 



64 PEOFESSOR OWEN ON THE 

The ulna (Pis. XV. & XX. figs. 16 & 17) is 3 inches 1 line in length, of the usual 
ornithic character, with a well-defined, narrow, elliptic, rough muscular depression, 8 lines 
in leno-th (fig. 16, c), extending upon the shaft from below the anterior or palmar angle 
of the proximal articular surface. This bone has no pneumatic foramen ; the orifice for 
the medullary artery is above the middle of the same palmar surface, the canal inclining 
distad. The shaft of the bone is nearly straight ; the back or anconal surface, which is 
slio-htly convex, shows feeble impressions of the attaching ligaments of the alar plumes, 
which are represented in all the figures of the entire or living bird. A second ulna is 3 
inches 3 lines in length. 

There was no carpal or pmion bone in the collection of remains submitted to me : this 
pari of the wing is conjecturaUy restored in dotted outline in Plate XV. 

§ 8. Bones of the Leg. (Pis. XV., XXL, XXII. & XXIII.). 

Of the &\e femora in the above defined series of remains of the Dodo, two measure 
6 inches 3 lines in length; one (PI. XXI.) is 6 inches 4^ lines ; the shortest is a little 
under 6 inches, with proportionate difierences in the diameter of the shaft. All of them 
show a small pneumatic foramen (PI. XXI. figs. 1 & 2,^j) on the inner side of the anterior 
ridge of the great trochanter (ib. c), and on the same transverse line with the head of the 
bone. This part shows an oblong depression (ib. figs. 2 & 3, a) for the " ligamentum 
teres" at the upper and back part. The articular surface on the same aspect of the neck 
(ib. fig. 3, b), adapted to the trochantcrian prominence of the pelvis (PI. XIX. t), is 
well-defined. The trochanter (PI. XXI. fig. 1, c) rises, ridge-Uke, above the level of the 
head, and is continued from behind the middle of the articular surface on the neck, 
forward, with a convex outline upon the fore and outer part of the shaft, where it gra- 
dually subsides; a narrow intermuscular ridge (ib. fig. 1, r), inclining to the middle of 
the fore part of the shaft, is continued from the trochantcrian one. The small tro- 
chanter (ib. fig. 3, d) is a small subcircular tuberosity, in some specimens a ridge, 3 to 
4 lines in length, on the inner side of the shaft, about an inch below the head. The 
muscular impressions on the fore part of the bone are well defined. A minute medul- 
lary canal (ib. fig. 8, m) perforates the middle of the back part of the shaft; the 
popliteal fossa (ib. fig. 3, o) shows a few small pneumatic orifices ; a triangular rough 
fiat surface divides the fossa from the outer condyle. Above the fibular depression 
(ib. fig. 3, y) there is a well-defined, slightly raised, rough surface (ib. k) for the head 
of the ectogastrocnemius muscle. The ridge (ib. n) extending to the back part of the 
inner condyle is not sharp ; the rotular groove (ib. fig. 1,^) is deep and moderately wide, 
with the inner boundary, formed by the narrow anterior part of the inner condyle (ib. 
fig. 5, e^), most produced. The breadth of this end of the longer femora is 1 inch 9 lines ; 
the character of the distal articular surface is shown in PI. XXI. fig. 5. 

The head, neck, and great trochanter (PI. XXIII. fig. 6) are occupied by a pneumatic 
cancellous structure, with a thin compact wall on the upper part and sides : this begins 



OSTEOLOGY OF TKB DODO. 65 

to gain thickness at the under part of the neck and at the lower and back part of the 
trochanter, the compact wall acquiring a thickness of a line at the beginning of the 
shaft, where the cancellous structure is confined to the outer side of the pneumatic 
cavity ; this structure gives way to a few delicate filaments of bone crossing the cavity 
of the major part of the shaft, and is not resumed until the bone expands to form the 
distal condyles (ib. fig. 7). 

The five tihice of Didus in the same collection range in length from 8 inches 8 lines 
to 9 inches. The procnemial ridge (PL XXII. figs. 1, 2, 4, f) is a triangular plate, 
with the base longest and the apex rounded off: it inclines outwardly, and does not 
extend much more than half an inch from the level of the proximal end of the bone : 
the length of its base rather exceeds an inch : on its inner side a triangular muscular 
sui-face is well defined by an irregular inferior line or ridge (ib. fig. 2, n). The ectocnemial 
process (ib. figs. 1, 3, 4, e) is thicker, shorter, and terminates roughly and obtusely. 
There is a low, narrow ridge (ib. fig. 2, g), about half an hich in length, on the inner 
side of the proximal end of the shaft, begimiing about 9 lines below the articular sur- 
face at that end. The fibular ridge (ib. figs. 1 & 3, h), beginning 1 inch 8 Imes from 
the proximal end, extends about 2 inches down the outer side of the shaft. The epi- 
cnemial ridge (ib. figs. 1 & 4, k) is obtuse, and but little produced above the upper 
articular surfaces or condyles {t d) of the tibia : the breadth of that end of the bone, 
in the longest specimen, is 2 inches 3 lines. The tendinal canal at the fore part of 
the distal end is bridged by bone (ib. fig. 1, I), and is situated on the inner half of that 
aspect of the shaft ; the lower opening is subcircular and close to the anterior end of the 
inner lower condyle (ib. «), which is more produced forward than the outer one (ib. h). 
Theii- hind ends project very little beyond the level of that aspect of the shaft of the 
tibia. An intermuscular ridge (ib. fig. 1, r) strengthens into a tuberosity (/•') at the inner 
side of the tendinal groove. 

The cancellous structure in the tibia is limited to an extent of about half an inch 
below the proximal articular surfaces (PI. XXIII. fig. 8), and to about an incli and a 
half from the distal end of the line (ib. fig. 9) : the shaft is occupied by a large air-cavity, 
with a compact wall of half a line in thickness at the upper third, gradually increasing 
to about a line at the lower fourth, until the cancellous structure is reestablished ; the 
transverse direction of a plate of this structm-e indicates the extent of the original 
distal epiphysis of the tibia (fig. 8). 

The fibula (PI. XXII. figs. 6-8) presents the usual ornithic characters of the bone : 
it varies from 4 inches 4 lines to 4 inches 6 lines in length, with a greatest j^roximal 
breadth of 8 lines. No adequate gain would result from a detailed description or com- 
parison of this bone ; and the rest of the bones of the foot have received every requisite 
attention in this way in the excellent work on the Dodo and its kindred, already 
quoted. A longitudinal section of the metatarsus, taken in the direction from side to side 
(PI. XXIII. fig. 10), shows the loose cancellous textiu'e of the common epiphysis of 

VOL. VI. — PAET II. K 



66 PHOFESSOB OWEN OX THE 

the three long metatarsals, and the remnant of their contiguous coalesced walls reduced 
to a thin lamella of bone. As the moiety of the bone figured is the posterior one (of 
the left metatarsus), the usual oblique position of the middle metatarsal {in), with its 
proximal end nearer the back part and its distal end nearer the fore part of the coalesced 
series, produces a corresponding direction of the section, with narrowing and termination 
of the exposed part of the medullary canal about one-third from the distal end of that 
metatarsal. The medullary canal of the outer metatarsal [iv) is wider and descends 
lower before the breaking up of the inner surface into decussating lamellfe or filaments, 
than that of the inner metatarsal (/?) : the peripheral compact wall of the inner is twice 
the thickness of that of the outer metatarsal. I may remark that the more posterior 
position of the middle metatarsal at its proximal end, from which and the corresponding 
part of the common epiphysis the calcaneal process is developed, is related to the greater 
share taken by the middle toe in the act of walking and scratching. I will only remark 
that of the four metatarsals of as many Dodos in the present series, one exceeds by a 
line the length of that figured in plate xi. op. cit., and one falls short thereof to the 
same trifling amount. 

§ 9. Skull. (Plates XV. & XXIII. fig. 1.) 

Of the skull of the Dodo, the series of bones transmitted to me include the cranial 
part with the detached upper mandibular bone (more or less mutilated) of two mature 
birds, and the lower mandible of three individuals. In the latter the dentary elements 
(PI. XXIII. fig. 1,32), confluent at the " gonys," are distinct from the liinder halves 
of the rami formed by the confluent, or perhaps connate, articular, surangular and an- 
gular elements (ib. 3i) : if the " splenial " were ever distinct, it has coalesced with the 
dentary, where its upper boundary is indicated by a linear groove or series of small 
foramina. 

In size, shape, and all other cliaracters of these important evidences of the specific 
character of the remains from the Mahebourg morass', they agree Avith those of Bidus 
ineptus detailed in the 'Proceedings of the Zoological Society' for January 11th, 1848 
(part xvi. pp. 2-8), and in the work entitled " The Dodo and its Kindred," pp. 76-96. 

The occipital condyle (ib. i) presents the same hemispheroid or rcniform shape, with 
the median vertical notch or depression above. The upper margin of the foramen mag- 
num is broad, as it were excised, with the sides slightly prominent. The supcroccipital 
foramen is present in both specimens, as in the one originally described (Proc. Zool. 
Soc. part xvi. p. 2). This foramen also exists in Owls and Parrots, but not in all Pigeons ; 
the Bidimculus (PI. XV. fig. 2) shows no trace of it ; I have also failed to find it in the 
skull of a Crown-pigeon (Goura coronata). The supcroccipital ridge is defined by the 
subsidence of the surface beneatlr it being continued directly from the upper, almost 
flat, smooth surface of the cranium : the middle part of the ridge is more produced than 

' " La Marc aux Songes." 



OSTEOLOGY OF THE DODO. 67 

the angles. In the great breadth of the occipital surface compared with its depth, in 
its flatness from side to side, and its aspect backward and a little upward, Didics most 
resembles Binornis. The basioccipital curves downward, and unites with the basi- 
sphenoid in developing the pair of larger tuberosities (PL XXIII. fig. 1, o), which ter- 
minate about \ an inch below the occipital condyle. There is nothing of this structure 
in the Columbine cranium. In one of my Dodo's skulls there is a pair of small 
tubercles between the larger basioccipital ones ; these are not developed in the other 
cranium. The basisphenoid is subquadrate, and fiattish below, impressed by a shallow 
median longitudinal channel. 

The hypoglossal nerve escapes by two small foramina on each side of the base of the 
condyle ; external to these is the vagal foramen ; still more external is the depression 
(ib. a) perforated below by the entocarotid, glossopharyngeal, and sympathetic, above 
by the tympanic vein. The entocarotid canal opens into the hind part of the sella or 
pituitary fossa : the vagal canal begins within the skull, above the hypoglossal foramina. 
The paroccipital carries the posterior surface of the skull downward and outward to a 
much greater degree than in any Dove, but to a less degree than in Dinornis. The 
Eustachian tubes impress the outer and fore part of the basisphenoid. 

The temporal fossse (PI. XV.), in the present specimens, show the same contracti(3n in 
proportion to their depth by which the original skull of the Dodo, compared with that 
of the Dinornis, ' Proc. Zool. Soc' (1848, p. -3), differed from the larger extinct wingless 
bird. In the approximation of the postorbital process to the mastoid, IJidunculus shows a 
closer resemblance to Bidus than does Goura, in which the temporal fossa, besides being 
narrow, is shallow. The temporal muscle appears to spread its origin above the fossa 
upon the sides of the cranium, forward half an inch in advance of the postfrontal 
process, and backward to the outer angle of the superoccipital ridge. 

The parietal region is broad, flat, and short, as in Dinornis, not convex as in Doves ; 
it is also impressed at its middle part by a shallow transverse groove, continued out- 
ward and forward of less depth and definition, so as to mark off" the convex interorbital 
part of the swollen frontals. 

The outer side of the mastoid is convex, smooth, overhanging the tympanic ca%dty, 
and sending off' a short process, the base of which is defined in one cranium by a trans- 
verse ridge in front of the anterior articular cup for the tympanic bone. A similar 
process is developed in Didunculus, not in Goura, where it is barely indicated. 

The presphenoid is compressed, but thickened and rounded below, where the pala- 
tines and pterygoids at their junction with each other abut against it: the pterygoid 
sends off" a short process from the middle of its hinder border ; but this is not met by 
a corresponding " pterygoid process" of the basisphenoid as in Didunculus. 

The frontals are broad and convex, rising abruptly (as in Didunculus) above the 
coalesced cranial ends of the nasals and premaxillary (PI. XV.) ; in Didus the breadth 
greatly exceeds the length of the interorbital frontal convexity, as compared with 



68 PEOFBSSOE OWEN ON THE 

Diduncuhs, and the convexity reigns in the transverse as well as the antero-posterior 
direction ; in Didunculus, however, it is less concave ti-ansversely than in Goura. In 
the breadth or thickness of the interorbital septum Didus resembles Apteryx and Palap- 
teryx and shows the same pneumatic cancellous structui'e. The posterior olfactory 
chambers are partially divided, as in Binornis, by an upper median septum ; each 
compaitment, which is 7 lines across and an inch in length, is perforated postei-iorly by 
an olfactory foramen more than a line in diameter, from which grooved impressions of 
ramifications of the nerve diverge upon the hind and upper wall of the chamber : ex- 
ternal to the olfactory foramen is a longer one for the passage of a vein into the fore 
and inner part of the orbit. 

The cranial ends of the nasals and nasal process of the premaxillary (PI. XXIII. 
fig. 1, 23) are flat, depressed, thin plates; the latter at its junction with the frontal is 6 
lines broad, partially divided by a median groove above and a ridge below, and by short 
linear fissures from the nasals : the forward extension of these bones is feebly indicated 
by linear grooves terminating at the outer margins of the nasal branch of the premaxil- 
lary, about 4 inches from its vertical end. The proportion of the base of the upper 
mandible attached to the frontal contributed by the nasals is the same as that indicated 
in the ' Proc. Zool. Soc' I. c. The nasal branch of the premaxillary presents a full 
elliptical transverse section where it quits the maxillary processes, losing both depth 
and breadth as it recedes to join the nasals; here it retains its breadth, viz. 6 lines, but 
continues to be thinned off vertically to the plate above named joining the frontal. The 
under surface of the narrower part of the stem is angular, the upper one being gently 
convex. 

" Where the nasal and maxillary processes diverge, there is a deep groove externally, 
terminating in a canal directed forwards into the rostral part or body of the premaxillary"'. 
This part is subdecurved, pointed, roughened by irregular vascular peiforations and 
grooves, with a sharp alveolar border, which desci-ibes a sigmoid curve lengthuase, and 
with a deeper conca\ity of the palatal surface than in Dinornis or Didunculus. Moreover 
the concavity is partially dinded lengthwise by a median ridge. The palatal surfaces of 
the maxillary processes and maxillaries are narrow and very convex transversely, inter- 
cepting a long narrow palato-nasal fissure. The outer side of the maxillary process is deep 
vertically and sHghtly concave lengthwise — a structure not known in Didunculus or any 
Dove, and related, like most other deviations from the Columbine cranial characteristics, 
to the provision of unwonted strength of beak in the Dodo. The maxillary branches 
of the premaxillary have completely coalesced \vith the maxillaries, as these have with 
the palatines ; and the halves of the upper mandible here swell out laterally and more 
so vertically, the maxillaries rising to combine with the outer divisions of the nasals, and 
sending back a short process from their lower and lateral part to join the malar. The 
inner surface of the maxillary process (PI. XXIII. fig. 1, 22») is smooth and slightly 
convex vertically ; both upper and lower borders are obtuse and thick. 

' Proc. Zool. Soc. I. c. p. 5. 



OSTEOLOGY OP THE DODO. 60 

The palatines arch outward from their posterior attachments, are broad and smooth 
mesially ; the margin here is angular, with a slightly produced obtuse apex, divided by 
a channel on the under surface of the palatine from the outer convex border ; the upper 
and outer ridge extends forward to the maxillary ; the inner one subsides before 
reaching that bone. " The palatines form the posterior boundaries of the naso- 
palatine aperture, and approximate each other at both ends, but more closely posteriorly, 
yet here without meeting; whilst in Didunculus they coalesce before receiving the 
abutment of the pterygoids. 

" The tympanic bone is subquadrate, with the four angles produced, and the upper 
and hinder are bifurcate, forming the double condyle for the mastoid articulation" '. 
There is a larger pneumatic foramen, communicating with the tympanic cavity, between 
the articulating cavities for these condyles. 

The brain is singularly small in the present species of Didus : and if it be viewed as 
an index of intelligence of the bird, the latter may well be termed ineptus. The length 
of the cranial cavity (PI. XXIII. fig. 1, t; c) is 1 inch 8 liaes, its extreme breadth 1 
inch 6 lines, its greatest height 1 inch (and this is at the cerebellar fossa). The most 
remarkable feature in the cranial structure of Didus is the disproportionate size of the 
brain-case to the important part of the neural axis it contained and protected : some 
approximation to this condition is made by Dinornis'^, the Owls, and a few large Cocka- 
toos, e. (J. Microglossum aterrimum ; but it is fully paralleled only by the Elephant 
among air-breathing vertebrates, as may be seen by comparing the section PI. XXIII. 
fig. 1 with the figures of a similar section quoted below^ 

Not only was the brain of very small proportional size in the present large extinct 
bird, but the division of the cranial cavity appropriate to the cerebrum proper is less in 
proportion to that for the cerebellum and optic lobes, at least in vertical and longitu- 
dinal diameters, than in any other known bird. 

In the Elephant the thickness of the pneumatic diploe between the fore part of the 
cerebral cavity and that of the outer cranial wall equals the longitudinal diameter of the 
cavity containing the cerebral hemispheres : in Didus it exceeds that diameter. The thick- 
ness of the pneumatic diploe above the cerebral cavity equals the vertical diameter of 
that cavity in Didus : the diploe gradually decreases in thickness as it approaches the 
foramen magnum. The disposition of the osseous lamellse forming the cells or ca\ities 
of the diploe is very difierent in the Elephant and Dodo : they extend for the most part 
vertically between the outer and inner tables of the skull in the proboscidian mammal, 
leaving long and narrow interspaces ; in the heavy ground-bird they form a congeries 
of small subequal and subspherical air-cells, and this structure obtains in the basal and 
lateral walls as well as in the superior or " roofing" wall of the cranial ca\'ity. The 

' Proc, Zool. Soo. I. c. p. 6. 

> Zool. Trans, vol. iv. pi. 24. fig. 4. 

' Odontography, pi. 146. fig. 1 ; Anat. of Vertebrates, vol. ii. p. 439. fig. 296. 



70 PROFESSOR OWEN ON THE 

extent of this cancellous structure at the sides of the cranial cavity may be known by 
the ratio of the breadth of that cavity to the breadth of the cranium, which is 3 inches 
and 8 lines at the broadest part of the brain, viz. the prosencephalon. It would seem, 
at first sight, as if the poorly developed brain of the Dodo had needed, on some account, 
unusual protection ; but the true explanation rests on the size, weight, and power of the 
bill, and the concomitant necessity for adequate extent of attachment of the facial to the 
cranial part of the skuU, and of the muscles from the trunk destined to sustain and wield 
the long and heavy-beaked head. The cerebrum of the Dodo does not greatly, and by no 
means proportionally, exceed the size of that part of the brain in the Crown-pigeons 
(Goura). If the great Ground-dove of the Mauritius gradually gained bulk in the 
long course of successive generations in that uninhabited thickly-wooded island, and, 
exempt from the attacks of any enemy, with food enough scattered over the ground, 
ceased to exert the wings to raise the heavy trunk, then, on Lamarck's principle, the 
disused members would atrophy, while the hind limbs, through the increased exercise 
by habitual motion on land, with increasing weight to support, would hypertrophy. 

In the long course of generations subject to this slow rate of change, there would be 
nothing in the contemporaneous condition of tlie Mauritian fauna to alarm or in any 
way to put the Dodo to its wits ; being, like other Pigeons, monogamous, the excite- 
ment, even, of a seasonal or prenuptial combat, might, as in them, be wanting : we may 
well suppose the bird to go on feeding and breeding in a lazy, stupid fashion, without 
call or stimulus to any growth of cerebrum proportionate to the gradually accruing in- 
crement of the bulk of the body. Whatever part of the brain was concerned in regu- 
lating or controlling muscular actions, might, indeed, be expected to show some concur- 
rent rate of increase with the growing mass of the voluntary contractile fibres ; and the 
size of the cerebellar division (PI. XXIII. fig. 1, w o) of the cranial cavity accords with 
tlie generally accepted physiology of the superincumbent mass of the epencephalon. 
The lateral depression at the fore and under part of the side of the postcerebral division 
of the cranial cavity indicates that the optic lobes, like the eyes, remained almost 
stationary during the progressive acquisition of the bulk that distinguishes the Dodo 
from the largest existing Doves. 

The proportions of Didus, Pezophaps, Casuanus, Rhea, Dromaius, Struthio, Aptornis, 
Cnemioniis, Palapteryx, ^pyornis, Dinornis, &c. among terrestrial birds, of Notornis 
among the lake-haunting Coots, and of Aptmodytes and AIca impennis among sea- 
birds, point to the disuse of wings in flight as the main condition of increase of size in 
species of birds — the next condition being absence of lethal enemies during the years 
requisite for such course and rate of growth. 

Let foes arise from whom a power of flight is the main condition of escape, and the 
wingless giants of the feathered class soon succumb. Among the genera above-cited, 
Aptornis, Cnemiorms, jEpyornis, Palapteryx, Dinornis, Pidus, and Pezophaps, mth the 
largest of the Auks, have thus passed away, while Notornis and Apteryx are on the 



OSTEOLOGY OF THE DODO. 71 

verge of extinction through the rapid increase of population in the small island to 
which they are restricted In sparsely peopled continents, such as Africa, South 
America, and Australia, brevipennate giants may still range the deserts, pampas, and 
unfrequented wilds. The ascertained recent advent of ilan in New Zealand, New 
Britain, Ceram, Banda, Salwattie, Mauritius, Rodriguez, significantly points to the 
conditions under which have come to pass, in lapse of time, so strange an anomaly as a 
bird with the specially modified instruments of flight reduced below the power of 
exerting that mode of locomotion, yet, as a bird, retaining the conditions of the 
respiratory and tegumentary systems of the volant class, of which it has become a 
degenerate member. With the cessation of the chief of those conditions, viz. the 
absence of enemies, such birds necessarily perish. 

Refraining, however, from further indulgence in an easy and seductive vein of specu- 
lation, I would recall attention to the notable protuberance in the cranial cavity of the 
Dodo (PI. XXIII. fig. 1, o) developed towards the upper part of the vertical tentorium, 
contracting at its lower part into the ridge dividing the prosencephalic from the mesen- 
cephalic chamber. In the latter are the orifices for the issue of the trigeminal nerve, the 
larger and posterior (ib. t)'} giving passage to the third and second divisions, and answering 
to the combined foramen ovale and rotundum of mammals, and the smaller and anterior 
foramen dismissing the first or orbital division of the fifth nerve. At the upper part of 
the mesencephalic fossa the narrow groove for the lateral venous sinus impresses and 
defines the back part of the tentorial protuberance, above which it bifurcates, the lower 
branch bounding or defining the wall of the superior semicircular canal and the upper 
part of the primitive acoustic capsule. Below this arch is an oblong cerebellar fossa (ib. ?;) 
which appears to have received veins from the cranial diploe. Beneath this fossa, and 
just behind the mesencephalic chamber, is the multiperforate internal auditory depres- 
sion. Next behind this is the outlet for the vagal nerve and entojugular vein. Below 
this are the small precondyloid foramina. There is a falcial ridge, low and thick, indi- 
cating the division of the prosencephalic chamber into lateral compartments for lienii- 
spheres ; and this ridge shows a narrow groove as for a small longitudinal sinus. A 
transverse linear groove abruptly defines the fore part of the ridge. 

The vertically expanded anterior part of the premaxillary (ib. fig. I, 22) has a large 
pneumatic cavity communicating by a reticidate wall with the cells of a cancellous struc- 
ture, larger than those of the cranial diploe. The maxillary branch of the premaxiUary 
(ib. 22*) consists of a light open-work air-diploe, with a very thin outer case of bone. 
The short symphysis mandibulfe shows a small cavity, suiTOunded by more minutely can- 
cellous structure and thicker compact walls, especially at the upper and hinder parts. 

Although some characters have been too much insisted on (e. g. the " superoccipital 
foramen ") as exemplifying the affinity of the Dodo, the more essential characters of the 
skull relate to its true Columbine character, while the deviations from that part of the 
skeleton of volant Doves are explicable in the adaptive developments needed for the 



72 PKOFESSOE O^VEN ON THE 

wicldino- of long, powerful, massive mandibles, serving most probably to enable the bird 
to subsist on some proportion of animal diet, in addition to such vegetable food as it 
mio'ht gain from the ground. Such indiscriminate feeding doubtless rendered its flesh 
less palatable than that of the winged Pigeons of the Mauritius to the Dutch navigators 
of the sixteenth and seventeenth centuries. 

But the affinities of Didus will be more fully and decisively brought out in the com- 
parison of the, in this respect, moi-e instructive and light-giving parts of the skeleton. 

§ 10. Comparison of the Skeleton. 

The dorsal region of the vertebral column shows, in some birds, a confluence of 
certain vertebrae : I have observed four to be so welded together by both centrums and 
neural spines in Phoenicopterus, viz. the second to the fifth dorsal inclusive, leaving the 
sixth free, which articulates with the first costigerous sacral vertebra. In Platalea 
three dorsals coalesce in advance of the antepenultimate free vertebra. In the smaller 
diurnal bkds of prey five dorsal vertebrae are usually confluent, leaving one free vertebra 
for the lateral movements of the trunk between such dorsal " sacrum " and the pelvic one. 
In Vultures, Plovers, Bustards, Ci-anes, Psophia, C'ariamn, Palamedea, the Penguins, and 
in all flightless land-bii-ds save the Dodo, no such anchylosis takes place. The Colum- 
bidm are the species in which the dorsal vertebrse, homologous and the same in number 
with those of Didus, undergo the process of confluence into one mass of bone : they are 
the three which immediately precede the last (moveable) dorsal vertebra ; and of these 
the two anterior devclope, in Goiira and Didunculus, hypapophyses closely corresponding 
in shape and proportion with those in the Dodo. 

The chief difference which Didus offers in the present region of the vertebral column 
from that of Colunibidw is in the greater number of the vertebrse or segments which 
are typically completed by bony haemapophyses articulating with pleurapophyses and 
directly with their mass of coalesced and expanded haemal spines constituting the ster- 
num. Of these typical thoracic segments there were five in Didus (PL XV.) ; Didun- 
culus (ib.) shows four ; Goura three. In both existing genera these segments are suc- 
ceeded by a single one, anchylosed to the fore part of the sacrum, but with the pleura- 
pophysis long and moveable, with its hsemapophysis terminating in a point before 
reaching the sternum, and extensi^'ely connected with the antecedent haemapophysis or 
sternal rib : in both genera two dorsal vertebrae in advance of the typically complete one 
have moveable pleurapophyses terminatuig freely in a point, with no haemapophyses 
other than the costal processes of the sternum may represent. In Qoura, which has six 
pairs of moveable or thoracic ribs, the second pair belong to the first of the three anchy- 
losed dorsal vertebrae : in Didunculus, which has seven pairs of thoracic ribs, the second 
pair belongs to the free dorsal immediately in advance of the anchylosed mass. Sup- 
posing Didus to have had one pair of ribs behind, and two pairs in front of those that 
directly articulate with the sternum, as the vertebra PI. XVII. fig. 7 indicates, it 



OSTEOLOGY OF THE DODO. 73 

would have had eight pairs of thoracic ribs; and I think this excess of one pair beyond 
the formula in Didunculus to be very probable in the large-bodied, small-winged extinct 
Ground-dove. 

As far as the series of Dodo's neck-vertebrse under my observation exhibit such 
characters, the proportion of those with neural spines, or with hypapophyses, or both, 
is the same as in the Columbidce. In this family, as in most birds, the greater part of 
the series want both processes. The cervical parapophyses, descending to form the 
sides of the carotid canal, do not meet, coalesce, and circumscribe it in any cervical 
vertebra of Goura or Didunculus ; and not any of the vertebrae of Bidus, which I have 
yet received, shows such circumscription of the hsemal canal. The majority of the 
cervicals in Bidus (those, viz., that lack both neural .spines and hypapophyses) are 
broader and more massive in proportion to their length than in the winged Doves. The 
third cervical in Bidus has both the above processes, as in Columhidw : the characters 
of the axis vertebra in the same family are closely repeated in that of the Dodo. In 
the Raptores the axis vertebra is shorter in proportion to its length, and a greater pro 
portion of the cervical vertebrEe at both ends of the series have both neural spines and 
hypapophyses. 

The ribs of the Dodo are as broad, in proportion to their length, as in Doves, but are 
relatively longer in proportion to the dorsal region, encompassing a more capacious 
thoracic-abdominal cavity. The ribs of the Vulture are more expanded than in Bidus, 
especially where they afford the extensive attachment to the epipleurals. But I shall 
not dwell further on the comparative characters of this part of the skeleton, as more 
decisive ones of the affinity of Bidus are afforded by other parts. 

In comparing the sternum of the Dodo with that of Doves of flight, the first well- 
marked difference is in the adaptive development of the keel in the last (PI. XV. fig. 2, 
Bidunculus), and in the provision for the concomitantly broader coracoids, the grooves for 
which meet and run into each other across the fore part of the bone in existing Coluin- 
hidm (PI. XXIV. fig. 2, h) ; consequently the inner or upper wall of the confluent grooves 
forms a median prominence (ib. e) at the front margin of the sternum, contrasting with 
the wide notch at that part of the bone in the Dodo (PI. XVI. fig. 4). The next differ- 
ence, as compared with Goura and most Pigeons, is the absence of the entolateral 
processes (PI. XXIV. fig. 3, i) in the Dodo's sternum : but Bidunmlus singulariy exem- 
plifies its nearer affinity to Bidus by a like absence of those processes ; only the sternal 
margins behind the ectolateral processes (ib. fig. 1, A), instead of converging with a 
slight convexity to an obtuse apex, as in PI. X\TII., describe a concavity, through an 
expansion of the posterior truncate end of the breast-bone. The sternum oi Bidunculus 
may be said to show one pair of posterior notches (PI. XXIV. fig. 1,/), that of other 
Pigeons two paii'S (ib. fig. 3,//') ; but the sternum oi Bidus, which is relatively broader, 
shows no other trace of the anterior notch (PI. XVIII. /) than is afforded by the 
rounded angle at which the ectolateral process {h) rises from the bone. Although the 

VOL. VI. PART II. " ^ 



74 PROFESSOR OWEN ON THE 

costal margin is relatively shorter in Doves of flight than in the Dodo, again an inter- 
mediate condition is manifested by Diduncidus as compared with Goura, in whicli latter 
Dove there are articular surfaces for three sternal ribs (PI. XXIV. fig. 3, c i, 2, 3), whilst 
in Bidunculus there are four (ib. fig. 1, c). Didunculus also exhibits, more strongly 
than Goiira, the obtuse ridges (ib. fig. 2, r) converging like buttresses from the outer 
wall of the coracoid groove to the fore part of the keel, where they subside. In Didun- 
culus there is a pneumatic foramen exterior to the coracoid groove, corresponding with 
p, fig. 4, PI. XVI., which I do not find in the sternum of Goura; but in the Crown- 
pigeons the pneumatic foramina along the middle line of the upper surface of the 
sternum are conspicuous ; they are confined to the fore part of that surface in Didun- 
culus (PI. XXIV. fig. 1). 

In the direction of the ectolateral processes Goura (ib. fig. 3, Jt) is intermediate be- 
tween Didunculus and Didus. The pectoral ridge on the outer surface of the sternum, 
continued backward from the outer end of the coracoid groove, is adaptively better marked 
in Pigeons of flight than in the Dodo ; and the pair- of ridges are more nearly parallel 
in theii' backward course, not so convergent as in Didus. In Goura the subcostal ridge 
is better marked than in Bidunculus. In no Dove of flight is the body of the sternum 
so broad and hollow as in Didus (PI. XXIII. fig. 4) ; in this respect the Vulture more 
nearly resembles the Dodo, as it does also in the more convex anterior contour of the 
keel : but the vulturine sternum does not lose breadth as it extends backward : it is a 
square-shaped shield in birds of prey, shorter in proportion to its breadth, with a greater 
extent of costal process and margin, and with the ectolateral processes, when they exist, 
extending backward as far as the hinder border of the bone. In the thorough quest of 
resemblances to the Dodo's sternum which I have made through the class of Birds, I 
came upon an unexpected superficial likeness to it m the sternum of a Night-jar [Po- 
daryus humeralis). The ectolateral processes (PI. XXIV. fig. 4, h) rise behind the 
moderately extended costal borders, c ; and beyond them the body of the sternum con- 
verges to an obtuse end, witlr a contour similar to that in Didus. Moreover the cora- 
coid grooves are divided from each other by a free concave border, less deep and exten- 
sive, indeed, than in Didus, but as free from any trace of episternal projection. The 
ectolateral processes, however, are extended backward to beyond the sternal body ; and 
this part usually shows a pair of small entolateral notches, f, of which one was present 
on one side in the specimen figured. 

Through the reduction of the coracoids in all flightless bu'ds, there is an interval 
between their sternal articulations: this is long and concave in the Dodo, but is 
longest and most deeply concave in Apteryx ; it is long but almost straight in Mhea ; in 
Casuarius and Dromaius it is narrow but deeply notched ; in Struthio it developes a short 
episternal process. In no Grallatorial sternum with both ecto- and ento-lateral pro- 
cesses (as e.g. Otis, (Edicnemus, Charadrius) do the former project, as in Didus and the 
Rasores, immediately behind the costal margin, but they are continued, parallel with 



OSTEOLOGY OF THE DODO. 75 

the keel, from the outer and posterior angle of the sternum, distant from the costal 
margin. In old Plovers the entolateral process joins the contiguous angle of the sternal 
body, and converts the inner notch into a foramen. 

In the breast-bone of the Dodo we plainly discern the Columbine modification of the 
Gallinaceous type, simplified in the minor development of those parts relating adaptively 
to the povFer of flight, and expanded and excavated for the support of the larger gizzard 
with its heavier grindstones'. 

In comparing the pelvis of Dtdunadus and Goiim (PI. XXIV. fig. 5) with that of 
Didus (PI. XIX. fig. 1), the correspondences are : — in the general shape, proportions and 
disposition of the ilia ; in the articulation therewith of the last pair- of moveable ribs, and 
of the .short straight confluent pleurapophyses of the three succeeding sacral vertebree ; 
then follow, as in Didus, three vertebrae without pleurapophyses, these reappearing in 
the next two with theii- extremities converging to abut against a prominence of the inner 
surface of the ilium in the same relative position. The difference here is in the two 
equal and more slender rib-buttresses, in place of the single stronger one, which is the 
more common structure in Didus ; but in Goura I have noted an instance in which it 
agreed with the Didunculus on the left side, and with Didus on the right, in the last- 
specified character. In the Crown-pigeons, also, there is an indication of the transverse 
ridge marking off' the under part of the centrum of the first sacral from the rest, and those 
that follow are less expanded than in the Uodlets ; moreover in Didunculus they show 
a median canal instead of a ridge, while the ridge is feebly indicated here and there and 
there is no canal in Goura. In neither Didunculus nor Goura do the sacral centrums 
behind the last rib-abutments diminish in breadth so suddenly as in Didus : in both the 
winged Pigeons the hinder part of the pelvic cavity is relatively deeper and narrower 
than in -D/(Zi<s ; in both, also, the upper and anterior concave tracks of the ilia are deeper; 
and in Didunculus the mesial borders do not attain the neural crest, but leave a pair of 
open longitudinal canals at that part of the pelvis; in Goura those margins reach the 
neural crest, but do not overtop it at any part. In Goura the acetabula are more in 
advance of a median position than in Didunculus, Columha magnifica, or Didus. Although 
the ischiadic foramina are completed by terminal confluence of the ilium and ischium in 

' The habit of the Dodo to avail itself of extraneous crushers to a gallinaceous or struthious degree, is attested 
by the following fruit of the extensive research of the learned and conscientious author of the Article Dodo, in 
the ' Penny Cyclopedia : ' — 

" About 1638, as I walked London streets, I saw the picture of a strange fowle hong out upon a cloth ; and 
myselfe with one or two more then in company went in to see it. It was kept in a chamber, and was a great 
fowle, somewhat bigger than the largest Turkey-cock, and so legged and footed, but stouter and thicker and of 
a more erect shape, coloured before Hke the breast of a young cock feasan, and on the back of a dunn or deere 
coulour. The keeper called it a Dodo, and in the end of a chymney in the chamber there lay a heap of large 
pebble-stones, whereof hee gave it many in our sight, some as big as nutmegs, and the keeper told us shee eats 
them (conducing to digestion)." Sir Thos. Brown's Works (WUkin's Edition, 4 vols. : London, 1830), vol. i, 
p. 369; vol. ii. p. 173. 

l2 



76 PROFESSOR OWEN ON THE 

Uromaius and Casuarius, yet the length of those foramina (which are unclosed) in 
Struthio and Aptenjx, concomitant with the greater relative length of the pelvis, shows 
the difference of Bidus from the cursorial Brevipennates in this part of the skeleton. 
The ischia of the winged Pigeons resemble those of the Dodo; but the inner longitu- 
dinal ridge is more strongly marked in Bidunculus : in the Goura it is less developed 
than in Didus ; the bone is longer also in proportion to its breadth, and the ischiadic 
foramen is lono-er and narrower : the proportions of that in Bidunculus are more like 
those in Didus. In Bidunculus the pubis coalesces with the ischium behind the small 
obtm-ator foramen, but leaves a second or posterior elongate ischio-pubic vacuity. The 
greatest amount of resemblances with the pehis of the Dodo is found in that of difierent 
members of the Dove-tribe. 

In comparmg the pelvis of the Dodo with that of the Vulture (PI. XXIV. fig. 6), we 
find in the latter that the first two confluent sacral vertebrae supporting moveable ribs 
are succeeded by several with short abutting ribs, the extent of this part of the sacrum 
bein" nearly one-half of tlie whole, instead of one-fourth as in ZHdus and the Doves. 
The reappearance of rib-abutments after four ribless sacrals is in the posterior thu-d of 
the sacrum, and they are continued to the end of that bone from the last four vertebra 
of the series, constituting a very marked diiference, both as to number and the character 
of the vertebra; in the sacral part of the pelvis. 

With regard to the iliac bones, the anterior concave track occupies two-thii-ds of the 
extent of the bone in VuHiir, not one-half as in Bidus and most Doves ; the breadth of the 
posterior parts of the ilia with the intervening sacrum in the Vulture is relatively less 
than in the winged Doves, and differs in a greater degree from that characteristic part in 
the sacrum of Bidus. In Ciconia the antacetabular part of the pelvis is relatively 
lon"-er, and the iliac bones are more expanded anteriorly. In Platalea the proportions 
are more nearly those in Bidus. In Otis the ilia touch the fore part of the sacro-spinal 
ridge, but leave both posterior and anterior apertures of the ilio-neural canals widely 
open. In (Edicnemus and Charadrius they are grooves, the ilia not reaching the sacral 
spines. The external concavity of the Uium is longer, narrower, and deeper, in most 
waders, than in Bidus. In Eudyptes and A])tenodytes the ilia are more expanded ante- 
riorly, but the whole pelvis is narrower and longer than in Bidus. The Gar-fowl 
{Alca im])ennis)\ Uria, Podicejjs, and Colymhus, all show still longer and narrower pro- 
portions of the pelvis. 

In the Doves of flight the proportions and relative position of the three compart- 
ments of the cranial cavity differ fi-om those in the Dodo. Both the pros- and mesen- 
cephalic ones are proportionally larger than the epencephalic ; and the mesencephalic 
compartment lies more directly below the prosencephalic one. A very thin stratum of 
finely cellular diploe divides the two tables of the skull along the medial line of the 
upper surface : it is thicker between the orbits. The falcial ridge at the inner surface 

' Trans. Zool. Soc. vol. v. pL 51. 



OSTEOLOaT OF THE DODO. 77 

of the prosencephalic roof resembles that in Bidus. The tentorial ridge bifurcates half- 
way down, the front portion dividing, almost horizontally, the pros- from the mesen- 
cephalic compartment, the hinder and more obtuse ridge dividing, almost vertically, 
the mes- from the epencephalic compartment. The angle of bifurcation is slightly 
produced and obtuse, but rei^resents very feebly the tentorial tuberosity (PI. XXIII. 
fig. 1, o) in the Dodo : from it, in Goura, is continued backward the arch of bone formed 
by the superior semicircular canal, above which is the groove for the venous sinus, as 
in Bidus. The internal auditory fossa is less deep than in Bidus : above it is a similarly 
vertically oblong cerebellar pit. The nerve-foramina correspond with those in Bidus : 
the entocarotid canal opens into a rather deeper sella in Columba palumbus. 

On comparing the cranial cavity, as exposed by a vertical longitudinal section in the 
Dodo (PI. XXIII. fig. 1), with that of a Dinornis similarly exposed', the first difference 
is the smaller proportional depth of the diploe in the larger wingless bird, which is not 
greater over the prosencephalic than over the epencephalic compartment ; next may be 
noticed the larger relative size of the former compartment, indicating the larger cere- 
brum of the Dinornis, then the absence of the tentorial tuberosity, the sharper and 
more produced superior part of the tentorial ridge arching transversely between the 
cerebrum and cerebellum, the smaller mtemal auditory fossa, and the deeper sella : the 
mesencephalic compartment, or cavity for the optic lobe, is less in proportion to the 
prosencephalic compartment than in Bidus; it holds, however, a similar relative posi- 
tion: finally, the cerebellar pit, above the internal auditory fossa, is wanting in the 
Dinornis. 

The Dodo agrees with the Doves in possessing a slender furculum, forming an acute 
angle : it resembles Columba galeata, more especially, in the halves of that bone being 
united by ligament below, and forming separate styles or " clavicles." 

The humerus of the Goura closely repeats most of the characters described in that of 
the Dodo ; but its length is proportionally greater, being 3 inches 9 lines, nearly equal to 
that of the sternum or pelvis, whereas the humerus of the Dodo is little more than half 
the length of either sternum or pelvis. The processes for the attachment of the muscles 
are, nevertheless, fully as strongly developed in Bidus (PI. XX. figs. 12 & 14) as in the 
volant Doves (PI. XXIV. figs. 8 & 9, Goura) ; tliat, indeed, which is a ridge (r) on the 
back part of the shaft in Bidus, is a mere rough surface in Goura, and does not sliow 
in Bidunculus. The pneumatic fossa, which varies in depth in the two humeri of the 
Dodo, is in both relatively larger and shallower than in Goura. The pectoral process 
is thmner, but relatively rather more produced, in Bidunculus. The humerus in (Edi- 
cnemus, Otis, and Charadrius has a more longitudinally extended, thinner, and more 
produced pectoral ridge than in Bidus and the Columhidce ; there is a more marked 
ectocondyloid tuberosity, which in Charadrius becomes a pointed process. 

There is nothing to be gained by giving the details of the more striking diflFerences 

' Trans. Zool. .Soc. vol. iv. pi. 24. fig. 4. 



78 PEOFESSOE OAVEX ON THE 

which the humerus presents in Penguins, Auks, and birds of prey, as compared with 
that bone in the Dodo ; but a few words may be recorded of the comparison of the 
humerus of the Dodo with that of the flightless bird of New Zealand, so nearly ap- 
proaching to it in size, which bird is described in the 5th volume of the ' Transactions' 
of the Society under the name of Cnemiornis (p. 395, pi. 6G. figs. 7-10). In that extinct 
species, although the humerus is 5^ inches in length, the parts mdicative of the forces by 
which it was worked are comparatively feebly developed. The ulnar tuberosity is nar- 
rower, thicker, more obtuse, and its base has neither the upper nor lower excavation ; 
it rises above the articular head, which is less prominent and narrower than in Didus ; 
the pectoral ridge is shorter and situated lower down upon the shaft, not on the same 
level with the radial tuberosity as it is in Didus ; the distal articulation is of the same 
size as in Didus, but neither the radial nor the ulnar convexity is so prominent or well- 
defined. 

The ulna of the Dodo is shorter absolutely, and much more so proportionally, than in 
the Goura and most other volant Doves. In these it exceeds the humerus by about one- 
fourth its own length ; in Didunculus (PL XV.) it is a little longer than the humerus ; 
in the Dodo (ib.) it is shorter than the humerus. The length of the ulna in Goura 
coronata is 4 inches 6 lines; it is more bent than in the Dodo ; the quill-tubercles, seven 
cr eight in number, are more prominent ; nevertheless the rough depression for the 
insertion of the chief flexor is less deep and less defined. The plumed winglet of the 
Dodo would seem, therefore, to have been frequently and forcibly moved. 

In comparing the femur of the Dodo with that of the largest Dove, the bone appears 
gigantic. The length of the femur in Goura coronata (PI. XXIV. fig. 11) is but 3 
inches 3 lines, and it is more slender in proportion to its length than in the Dodo ; it, 
however, repeats the few characteristics, if they may be so termed, of the Dodo's femur. 
It has the pneumatic foramen in the same position, perhaps proportionally larger ; it 
has the same large oblong surface for the ligament at the head of the bone ; the great 
trochanter has the same form and disposition, but is not quite so much produced an- 
teriorly ; there is a slight depression instead of a ridge for the trochanter minor ; the 
fore part of the inner condyle is relatively thicker and less produced. The femur in 
Otis and (Edicnemus has a thicker and shorter trochanter major, a more narrow and 
shallow rotular channel ; it is shorter in comparison with the tibia, and more especially 
with the metatarsus, than in Didus and the Doves. 

The femur of Aptornis otidiformis^ is of the same size as that of the Dodo; but it has 
no pneumatic foramen, the head is more hemispheroid and inclined forward, the liga- 
mentous pit is deeper and more circular, the supracervical articular surface is not 
defined from that of the head, there is a wider and deeper depression at the fore 
part of the proximal end of the femur, and a more prominent tuberosity on the back 
part; the ridge continued from the back part of the shaft to that of the inner con- 

' Trans. Zool. Soc. vol. v. pi. 65. fig. 3. 



OSTEOLOGY OF THE DODO. 79 

dyle is more produced and sharper in Ajitornis, the fore part of the same condyle is 
less produced. 

The femur in Cnemiornis^ and Linornis^ is much thicker, in proportion to its length, 
than in either Aptornis or Didiis. In Pezophaps the great trochanterian ridge rises 
higher above the neck, and the shaft has a more uniform thickness, with the inner 
contour less concave, than in Didus. 

The characters which have been noted at the proximal and distal ends of the tibia of 
Didus are repeated in those of the tibia of the Goura. The difference in size is more 
marked than in the femur ; the length of the tibia of Goura coronata is 4 inches 7 lines, 
and its shaft is more slender, in proportion to its length (PI. XXIV. fig. 13), than in 
Didus (PI. XXII.). The tendency to a trihedral form of the shaft is less marked in 
Goura ; the anterior prominences of the distal condyles are thicker in proportion to the 
intervening fossa. 

In the Vulture the fibular ridge is more parallel with the long axis of the shaft than 
in Didus ; the tendinal canal is less cylindrical, has an oblique course from the middle 
of the anterior surface towards the inner condyle ; the fore parts of both distal condyles 
are less produced and less convex ; the distal end is narrower from before backwards in 
proportion to its breadth ; both extremities of the bone are less expanded in proportion 
to the shaft than in the Dodo. 

In the great Plover [CEdicnenms crepitans) the tibia, as in other Grallse, is longer in 
proportion to its thickness than in Didus ; the epicnemial process rises higher above and 
projects further in front of the condylar sui-faces before it divides into the pro- and 
ectocnemial plates ; and these are relatively more produced. The fibular ridge is 
shorter in proportion to the length of the tibia, is more prominent, and more parallel 
with the axis of the shaft. The distal condyles project further backward than in Didus. 
The tibia in Charadrius, Otis, Tantalus, Grus, Cicoma, Mycteria, Porphyrio, opposes 
similar or equivalent differences to those in (Edicnemus, against the aflanity of Didus to 
any of those Grallse. 

In the comparison of the tibia of this extinct flightless bird with that of the Cneniiornis, 
the wonderful development of the plates and processes at the proximal end of the bones 
in the New Zealand bird is strikingly manifested. In Cnemiornis the fibular ridge runs 
in a line with the shaft, and does not incline from above obliquely forward as in Didus 
and the Doves ; the ridge on the outer side of the distal fourth of the bone is stronger 
and sharper in Cnemiornis ; the tendinal canal is transversely elliptical, medial in 
position, with a slight inward inclination ; the intercondyloid fossa is much wider in 
Cnemiornis. The differences, indeed, in all the characters of the tibia, as compared with 
Didus, in the Vultures, Plovers, Penguins, and terrestrial flightless birds tend to render 
more instructive and convincing the resemblances which Pigeons present in the same 
characters to the extinct Mauritian bird. 

' Trans. Zool. Soc. vol. v. pi. 65. fig. 1. ' Ibid. fig. .5. 



80 PEOFESSOE OWEN ON THE 

§ 10. Conclusion. 

The affinities or place in nature of the Dodo being thus determined by the characters 
of its skeleton, but few words remain to be said on the bearings of present knowledge 
of this species upon other zoological generalizations. 

The researches and observations of naturalists have been carried out to such an extent 
as to support the conclusion that the Didus ineptus does not now live in any part of the 
world, and that it never existed save in that part of which the island of Mauritius 
may be a remnant. Consequently the species there originated ; and the most intelli- 
gible conception of its mode of origin is that to which I have alluded in the description 
of the brain-case (p. 70). 

The Dodo exemplifies Buffon's idea' of the origin of species through departure from 
a more perfect original type by degeneration ; and the known consequences of the 
disuse of one locomotive organ and extra use of another indicate the nature of the 
secondary causes that may have operated in the creation of this species of bird, agree- 
ably with Lamarck's philosophical conception of the influence of such physiological con- 
ditions of atrophy and hypertrophy 2. The young of all Doves are hatched with wings 
as small as in the Dodo : that species retained the immature character. The main con- 
dition making possible the production and contmuance of such a species in the island 
of Mauritius was the absence of any animal that could kill a great bird incapable of 
flight. The introduction of such a destroyer became fatal to the species which had lost 
such means of escape^. The IMauritian Doves [Cohnnba nitidissima and C. meyeri) that 
retained their powers of flight continue to exist there. 

As I have no reason to ofier why one kind of Pigeon should have retained and another 
lost its powers of flight, nor am able to adduce a particle of evidence of the hypothetical 
degrees of diminution of the wing-bones to their stunted proportions in Didus, any 
more than in Dinornis, I feel that in the foregoing remarks I lay myself open to the 
rebuke of fellow-labourers who may think with the able authors who last treated of 
the present subject. 

They warn their readers to " beware of attributing anything like imperfection to these 
anomalous organisms, however deficient they may be in those complicated structures 
which we so much admire in other creatures. Each animal and plant has received its 
peculiar organization for the purpose, not of exciting the admiration of other beings, 
but of sustaining its own existence. Its perfection, therefore, consists, not in the 
number or complication of its organs, but in the adaptation of its whole structure to 
the external circumstances in which it is destined to live. And, in this point of view, 
we shall find that every department of the oi'ganic creation is equally perfect, the 

' Histoire Naturolle, &c., 4to, torn. xiv. "Degeneration des Animaux: " 1760. 
' Philosophie Zoologique, 8vo, 1809, torn. i. chaps. 3, 6, & 7. 

' Agreeably with the principle of the " contest for existence " by which I explained the extinction of the 
species of Dinornis, Trans. Zool. Soc. toI. iv. p. 14, 1851. 



OSTEOLOGY OF THE DODO. 81 

humblest animalcule or the simplest conferva being as completely organized with 
reference to its appropriate habitat and its destined functions as Man himself, who 
claims to be lord of all. Such a view of the creation is surely more philosophical than 
the crude and profane ideas entertained by Buffon and his disciples" '. 

Nevertheless the truth, as we have or feel it, should be told. In the end it may prove 
to be the more acceptable service. The Didus ine])tus, L., through its degenerate or 
imperfect structure, howsoever acquired, has perished. What have the stigmatizers of 
Buflbn to offer in lieu of his theory as applied to the origin of this species of bird ] 
They begin by asking, "Why does the whale possess the germs of teeth which are never 
used for mastication \ and why was the Dodo endowed with mngs at all, when those 
wings were useless for locomotion'? This question," they own, "is too wide and too deep 
to plunge into at present." They nevertheless proceed to remark, " These apparently 
anomalous facts are really the indications of laws which the Creator has been pleased 
to follow in the construction of organized beings ; they are inscriptions in an unknown 
hieroglyphic, which we are quite sure mean something, but of which we have scarcely 
begun to master the alphabet. There appear, however, reasonable gi-ounds for believing 
that the Creator has assigned to each class of animals a definite type or structure, from 
which He has never departed, even in the most exceptional or eccentric modifications of 
form. Thus, if we suj^pose, for instance, that the abstract idea of a Mammal implied 
the presence of teeth, and the idea of a Bird the presence of wings, we may then 
comprehend why in the Whale and the Dodo these organs are merely suppressed, not 
wholly annihilated"^. 

This notion of type-forms or centres, unfortunately, has not merely relation to abstract 
biological speculations or theories, but to practical questions on which the true progress 
of Natural History vitaUy depends. If such types do exist, the National Museum, it is 
argued, may be restricted to their exhibition : and so our legislators and the public were 
assured by the Professor of Natural History in the Government School of Mines ^, when 
the question was before the " House" four years ago. I have let slip no suitable occa- 
sion* to combat and expose what has seemed to me to be both an erroneous and mis- 
chievous view, most obstructive to the best interests of the science ; and, standing alone 

' Strickland and MelviUe, ' The Dodo and its Kindred,' 4to, 1848, p. 34. 

" Op. cit. p. 34. 

' See letter in ' The Times' of May 21st, 1862, advocating the limitation of the National Museum of Natural 
History to " six rooms," signed Thomas H. Huxley, F.B.S. 

* Eeply to the above in ' The Times ' of May 2nd, 1866, and in both editions (1861, 1862) of my ' Discourse 
on the Extent and Aims of a National Museum of Natural History.' " Some naturalists urge that it is only 
necessary to exhibit the type-form of each genus or family. But they do not tell us what is such ' type- 
form.' It is a metaphysical term, which implies that the Creative Force had a guiding pattern for the con- 
struction of all the varying or divergent forms in each genus or family. The idea is devoid of proof; and those 
who are loudest in advocating the restriction of exhibited specimens to ' types ' have contributed least to lighten 
the difficulties of the practical curator in making the selection." (Ed. 1862, p. 24 ; see also pp. 26-34.) 
VOL. VI. — PART II. M 



82 PEOFBSSOE OWEN ON THE 

as I seemed to do on this point in the aiTay of evidence before the "Parliamentary 
Committee on the British Museum," I was glad to find my views on type-forms adopted 
and paraphrased by the President in his Addi-ess to the British Association at the 
meeting at Nottingham', in the present year. 

DESCRIPTION OF THE PLATES. 

PLATE XV. 

Fig. 1. Side view of the skeleton of the Dodo {Bidus ineptus, L.), with an outline of 
the bird as represented in the oil-painting presented to the British Museum 
by Edwards, Naturalist and Librarian of the Royal Society, to whom it had 
been given by Sir Hans Sloane, P.R.S., with the statement that the painting 
had been made, of the natural size, from a living specimen of the Dodo, in 
Holland. The bones represented in profile, of the natural size^, testify to the 
accuracy of the form and proportions of the Dodo given in the painting. 

Fio-. 2. An outline of the Samoan Dove or Dodlet (Didunculus strigirostris, Peale; 
Gnathodon strigirostris, Jardine^) of the natural size, from a specimen living 
in 1865 in the Gardens of the Zoological Society of London, with a view of 
the skeleton corresponding with that of the Dodo. 

PLATE XVI. 

Fig. 1. Front view of the fourth (or first of the three confluent) dorsal vertebrae (centrum 

and neural arch). 
Fig. 2. Vertebral rib, or pleurapophysis, of the same vertebra, front view. 
Fig. 3. Sternal rib, or ha?mapophysis, of the same vertebra : a, outer side ; b, upper or 

pleural end ; c, lower or sternal end ; d, front margin ; e, inner surface. 
Fig. 4. Front view of sternum, or connate mass of hsemal spines, including that of the 

same (fourth dorsal) vertebra. 
Fig. 5. Inner surface of an anterior pleurapophysis, with coalesced appendage, a. 
Fig. 6. Oblique view of ditto, ditto. 

' " The doctrine of typical nuclei seems only a mode of evading the difSculty. Experience does not give us 
the types of theory ; and, after all, what are these types ? It must he admitted there are none in reality. 
How are we led to the theory of them ? Simply by a process of abstraction from classified existences. Having 
grouped from natural simihtudes certain natural forms into a class, we select attributes common to each 
member of the class, and call the assemblage of such attributes a type of the class. This process gives us an 
abstract idea ; and we then transfer this idea to the Creator, and make Him start with that which our own 
imperfect generalization has derived." (Address, &c., by William R. Gbove, Esq., Q.C., M.A. 8vo, London, 
1866: p. 31.) 

- The scapular arch is rotated in advance of the ribs to show the character of the anterior dorsal vertebrae. 

■■> See also Goidd, ' Birds of Australia,' part 22 (March, 1846). 



OSTEOLOGY OF THE DODO. 83 

Fig. 7. Anterior pleui-apophysis with appendage, a, front view: c, capitular end; t/, 
tubercular end ; /, haemal end ; 7 a, outer surface ; 7 b, inner surface. 

Fig. 8. An anterior pleurapophysis, front \-iew. 

Fig. 9. Posterior surface of the upper end of a posterior pleurapophysis : 9 «, body and 
loAver end of ditto. 

Fig. 10. Part of a pleurapophysis which has been broken and healed. 

Fig. 11. I>ower end of a posterior dorsal pleurapophysis, with connate rudiment of 
appendage, a. 

Fig. 12. Hsemapophysis. 

PLATE XVII. 
Fig. 1. Fourth, fifth, and sixth dorsal vertebrse, anchylosed, side riew. 
Fig. 2. Ditto, ditto, upper view. 
Fig. 3. Ditto, ditto, under view. 
Fig. 4. Ditto, ditto, back view. 
Fig. 5. Ditto, ditto, mutilated, of another Dodo. 
Fig. 6. Anterior dorsal vertebra, side view. 
Fig. 7. Ditto, front view; pl^ outline of heads of floating rib. 
Fig. 8. Penultimate cervical vertebra, side riew. 
Fig. 9. Ditto, back view. 
Fig. 10. Middle cervical vertebra, upper view. 
Fig. 11. Ditto, under view. 

Fig. 12. Axis, or second cervical vertebra, upper view. 
Fig. 13. Ditto, under view. 

PLATE XVIII. 

Fig. 1. Under view of sternum. 
Fig. 2. LTpper or inner view. 
Fig. 3. Back view. 

PLATE XIX. 

Fig. 1. Under or inner view of pelvis. 
Fig. 2. Upper or outer view of pelvis. 

PLATE XX. 



Fig. 1. Middle cervical vertebra, upper view. 
Fig. 2. Fifth cervical vertebra, upper view. 
Fig. 3. Fourth cervical vertebra, under view. 
Fig. 4. Right coracoid and clavicle. 



84 PEOFESSOE OWEN ON THE 

Fig. 5. Left coracoid and clavicle. 

Fig. 6. Right scapula, outer view. 

Fig. 7, Right scapula, inner view. 

Fig. 8. Left moiety of scapular arch, outer view. 

Fig. 9. Ditto, inner view. 

Fig. 10. Upper articular end of right coracoid. 

Fig. 11. Lower ditto. 

Fig. 12. Left humenis, anconal or back surface. 

Fig. 13. Left ditto, ulnar or inner surface. 

Fig. 14. Left ditto, palmar or front surface. 

A. Ditto, proximal or upper end. 

B. Ditto, radial side of upper half. 

C. Ditto, distal end. 
Fig. 15. Eight radius. 

Fig. 16. Right ulna, inner or radial side. 
Fig. 17. Right ulna, outer or ulnar side. 



PLATE XXL 



Fig. 1. Left femur, front view. 

Fig. 2. Ditto, inner view. 

Fig. 3. Ditto, back Aiew. 

Fig. 4. Ditto, upper end. 

Fig. 5. Ditto, lower end. 



Fig. 1. Left tibia, front view. 
Fig. 2. Ditto, inner view. 
Fig. 3. Ditto, back view. 
Fig. 4. Ditto, upper end. 
Fig. 5. Ditto, lower end. 
Fig. 6. Left fibula, outer view. 
Fig. 7. Ditto, inner view. 
Fig. 8. Ditto, upper view. 



PLATE XXII. 



PLATE XXIII. 



Fig. 1. Longitudinal vertical section of mutilated skull. 

Fig. 2. Ditto of third cervical vertebra. 

Fig. 3. Ditto of lower cervical vertebra. 

Fig. 4. Transverse vertical section of sternum. 



OSTEOLOGY OF THE DODO. 85 



Fig. 5. Longitudinal section of humerus. 
Fig. G. Ditto of upper end of femur. 
Fig. 7. Ditto of lower end of femur. 
Fig. 8. Ditto of upper end of tibia. 
Fig. 9. Ditto of lower end of tibia. 
Fig. 10. Ditto of metatarsus. 



PLATE XXIV. 

Fig. 1. Sternum oi Didunculus, upper view. 

Fig. 2. Ditto, front \iew. 

Fig. 3. Sternum of Goura, ufiper view. 

Fig. 4. Sternum oi Podargus Immeralis, under view. 

Fig. 5. Pelvis of Goura, under or inner view, half natural size. 

Fig. 6. Pelvis of Gi/ps (Vulture), under or inner view, half natural size. 

Fig. 7. Left moiety of scapular arch, Goura. 

Fig. 8. Left humerus of Goura, anconal surface. 

Fig. 9. Left humerus of Goura, palmar surface of upper end. 

Fig. 10. Left humerus of Goura, palmar surface of lower end. 

Fig. 11. Right femur of Goura, front view. 

Fig. 12. Right femur of Goura, back view of upper end, and back view of lower end. 

Fig. 13. Right tibia and fibula of Goura, front view. 

All the figures are of the natural size, save when otherwise expressed. The letters 
are explained in the text. 



VOL. VI. — PART II. 



PLATE XV 





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DIDUNCDLU; 



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Sii-lV; INEPTUS./ 



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irrut 



imp 



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KrofTi jirtJb on Stone by J Erxleben . 



MA: If Harthart.imp 



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Vio 1 



fin 5 








:0 




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S Smjt Ji;.h 



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[ 87 ] 



IV. DeHcription of the Skeleton of Inia geoffrensis and of the Skull of Pontopoiia 
blainvillii, ivith Eemarks on the Systematic Position of these Animals in the Order 
Cetacea. By AVilliam Henry Flower, F.B.S., F.B.C.S., F.Z.S., &c.. Conservator 
of the Museum of the Royal College of Surgeons of England. 

Read November 22nd, 1866. 

[Plates XXV. to XXYIIL] 

I. On the Skeleton of Inia geoffrensis. 

Of the several species of Cetaceans which are inhabitants of the waters of the 
Amazon and its great tributaiy streams, one has particularly attracted the attention of 
zoologists on account of certain peculiarities of its external conformation and of its 
skull and teeth, the only parts of its structure hitherto described. 

The Inia, so called by M. Alcide d'Orbigny, from the name by which the animal is 
known to one of the Indian tribes of Bolivia, is chiefly characterized by the long, 
harrow, and almost cylindrical rostrum, furnished with scattered, stout and crisp hairs, 
by the broad, long, and obtuse pectoral fins, by the dorsal fin reduced to a mere ridge, 
and especially by the development of a large lobe on the inner side of all the posterior 
teeth. 

The species is mentioned by Spix and Martius* as DelpUnus amazonicus; but for 
the most complete account of its external characters, habits, and geographical distribu- 
tion we are indebted to d'Orbigny, who described it under the name of Inia holiviensis\. 
He also gives a figure of the animal, and a side view of a skull which he brought home 
and deposited in the Museum at the Jardin des Plantes, with some details of the teeth. 
I will quote from this memoir two observations — the first referring to the habits, the 
second to the structure of this singular Cetacean : — " Toutes ces observations nous font 
regarder cette espece comme ayant des mceurs beaucoup plus terrestres qu'aucune des 
especes connues." — " Tons ces caracteres reunis a uTie dorsale peu apparente, nous font 
proposer la formation d'un nouveau genre, qui etablerait le passage entre les sousous 
[Platanistci^ et les stelleres " [Sirenia']. 

* Keise in Brasil. t. iii. pp. 1119 & 1133 (1831). Von Martius States that his Delphinus amazonieus agrees 
very closely with Desmarest's description of D. geoffroyi, and even suggests that it may possibly belong to the 
same species. His description of the teeth is suificient to detennine the animal spoken of; but he says 
" pinna dorsalis distincta, elata." Perhaps he has here confounded it with some of the other species of fresh- 
water dolphins of the Amazon, the existence of which he did not suspect. The rude Httle figure he gives 
(fig. 34) more resembles Delphinus fluviatilis (Gervais) of Castelnau's Voyage than the Inia. 

t Nouv. Ann. Mus. Paris, torn. iii. p. 23 (1834). 
VOL. VI. — PART in. O 



88 ME. "W. H. FLOWEE ON THE OSTEOLOGY OF 

In the Zoology of d'Orbigny's ' Voyage en Amerique meridionale,' " Mammiferes," 
by d'Orbigny and Gervais (1847), more careful figures both of the upper and lateral sur- 
face, and of the teeth, of the same skull are given (pi. 22), but unaccompanied by any 
further description. It is, however, suggested that the animal belongs to the same 
species as a stuffed and painted specimen received at the Paris Museum from the Musee 
d'Ajuda at Lisbon among the spoils of Napoleon's Peninsular campaign, and described 
by de Blainville in the Article " Dauphin " in the ' Nouveau Dictionnaire d'Histoire 
Naturelle,' t. ix. p. 151 (1817), as Belphhms geoffrensis, and subsequently by Desmarest* 
as D.geoffroyi. 

In a later notice by Professor Gervais, in the Zoology of Castelnau's ' Expedition 
dans les parties centrales de 1' Amerique du Sud,' "Mammiferes," p. 90(1855), this sup- 
sition is confirmed, and the name Inia geoffrensis definitively adopted. In this notice 
some further details are given respecting the original skull brought home by d'Orbigny ; 
and a new figure of the external appearance of the animal is added, differing chiefly 
from that of d'Orbigny in the position of the pectoral limb. 

A few years ago that enterprising naturalist Mr. H. W. Bates obtained at Ega two 
skulls, which are now in the British Museum. Of one of these. Dr. Gray has given ihe 
dimensions f. 

According to information received from my friend Dr. Peters, there is in the Ana- 
tomical Museum at Berlin a skull brought home by Natterer. No description of this, 
however, has been published. 

In the early part of the present year Mr. Edward Bartlett, while collecting zoological 
specimens on the upper Amazon, above Nauta, succeeded, after encountering many diffi- 
culties, in obtaining a complete animal, the carefully prepared skeleton of which has 
now been purchased for our National Collection. For the opportunity of examining 
and describing this rare and interesting specimen, before it was deposited in the 
Museum, I am indebted to the kindness of Dr. Gray. 

The skeleton is that of a young animal, the epiphyses being not united to the bodies 
of the vertebrae from the axis to the tenth caudal ; but the arches have completely 
coalesced with the bodies throughout the spinal column. The head of the humerus 
stiU retains its epiphysial condition. The total length of the living animal, judging 
from the skull and vertebree, and allowing for the intervertebral spaces, would be but 
little more than 5', the skull being 16-4". The specimen obtained by d'Orbigny is 
stated to have measured 2'"-4 = 6' 8" Eng., and its skull is 0'"-48 or 19". The skulls 
collected by Mr. Bates indicate animals of still larger size, the one being 19-4", the 
other 20-7" long. The skull at Berlin, as Professor Peters has informed me, is 19^'' 
Eng. in length. Martius states the length of the animal to be from 7 to 8 feet. Finally, 
Castelnau gives 2'"-80 or 8' 4" as the length of an individual taken at Nauta. 

* Mammalogie, p. 512 (1822). 

t Catalogue of Seals and Whales in the British Museum, p. 227 (1866). 



INIA GEOPFEENSIS AKD PONTOPORIA BLAIFSaLLII. 



89 



As Inia has always been supposed to have certain affinities with Platanista, I 
have in the follomng description compared the different bones with those of that sin- 
gularly modified Cetacean on the one hand, and of several of the ordinary Deljihinidm 
on tlie other. Fortunately the Museum of the College of Surgeons contains a 
skeleton of the Gangetic Dolphin of nearly corresponding age with the subject of 
the present communication, and I have also had frequently to refer to Eschricht's 
valuable memoir upon the species*. 

The leading features of the skull have already been made known by d'Orbigny and 
Gervais ; but I am able to add some further details regarding its structure. 

A comparison of the two skulls at the British Museum sent by Mr. Bates, with the 
present example, shows only such differences as might be expected from the greater 
age of the former, such as a more marked development of the ridges and prominences 
in proportion to the size of the brain-case. The postnarial prominence especially is 
more elevated and angular in the older specimens. The teeth differ somewhat in 
number, as will be mentioned further on. 

The principal dimensions of the three skulls are as follows : — 



Extreme length 

Length of rostrum (from anterior end of premaxiilary to bottom 

of antorbital notch of maxillary) 

From anterior end of premaxiilary to lower edge of nasal bones" .' 
From anterior end of premaxiilary to hinder edge of palate 
Greatest breadth, across zygomatic processes of squamosals 

Breadth of foramen magnum 

Breadth of the occipital condyles '.'.'..'.'... 

Breadth across antorbital processes of frontals 

Breadth of rostrum at base (bottom of antorbital notches of 

maxillaries) 

Breadth of rostrum at middle 

Mandible, length 

Length of symphysis 

Greatest breadth across the posterior ends of the rami 

Height of ramus at coronoid process 



Collected by 



Mr. Bates, ' Mr. Bates. Mr. Bartlett. 

a. \ b. \ 



20-7" 

13-5 
16-7 
15-7 
9-4 
1-4 
3-2 
6-1 

4-2 
1-4 
18-2 
9-8 
8-7 
3-9 



:9-4" 

12-7 

15-7 

t 

8-1 
1-2 
30 
5-3 

3-5 

1-2 
17-0 
9-4 
7-3 
3-4 



lC-7" 

11-0 
13-6 
12-6 
7-0 
1-3 
2-9 
4-5 

31 
11 
14-2 
73 
6-5 
2-9 



The want of symmetry so prevalent in the skulls of Dolphins is but slightly marked. 
It can, however, be detected in a slight twist to the right of the hinder part of the 
narrow median space between the premaxillaiy bones, and in the greater elevation on 
the same side of the postnarial prominence of the frontal bones. Both maxillary and 
premaxiilary bones extend backwards to an equal extent on the two sides. 

In the cranium of the young specimen which forms the subject of the present commu- 

• " Cm Oangesdelphinen," Trans. Roy. Dan. Acad. 1851. Translated in Ann. &, Mag. Nat. Hist, for March 
1852. f Broken. 

o2 



9Q ME, W. H. TLOWEE ON THE OSTEOLOGY OP 

nication (see Plates XXV. & XXVI.), the elements of the occipital bone have completely 
coalesced with each other and with the basisphenoid, and partially with the parietals. 
The foramen magnum is subcircular, the greatest vertical and transverse diameters being 
exactly equal ; but it is rather broader above than below. Its plane is nearly vertical 
when the skull is held horizontally. The condyles are large and prominent ; they do not 
meet below by a space of -7". In the middle line on the supraoccipital, just above the 
margin of the foramen magnum, is a deep triangular depression, continuous with a 
broad and shallow median groove which ascends nearly to the vertex, and with lateral 
grooves which pass outwards above the upper edge of the condyles to the concave surface 
of the exoccipitals. In the lower part of the median groove the surface of the bone 
is very rough, being channeUed out for a plexus of blood-vessels ; and there are several 
rounded perforations, one of them as much as -1" in diameter, by which these vessels 
would apparently communicate with the interior of the cranial cavity. Corresponding to 
this groove, on the inner side, is a median bony ridge, but there is no transverse tentorial 
ossification. The lateral boundaries of the supraoccipitals are raised into strong narrow 
ridges, on the summit of which the occipito-parietal suture is situated. These are nearly 
parallel until they come opposite to the posterior angle of the maxillaries ; then they 
rapidly converge, enclosing a triangle with a truncated apex which projects forward 
into the high postnarial eminence of the frontals. 

The temporal fossa, as noticed by d'Orhigny, ia tery much larger in proportion to the 
size of the cranium than in any other Dolphin, excei^t Phi fanista, not only occupying a 
larger space on the lateral surface of the skull, but being prolonged forward at the 
expense of the orbit. Its form is that of a long oval, with the small end turned forwards. 
Its posterior nearly semicircular boundary is formed by the ridge, before spoken of, at the 
junction of the occipital with the squamosal and parietal. The superior border, continued 
forwards fi-om the latter, is a nearly straight, sharp, thin crest, projecting outwards and 
upwards, 3" long, and averaging more than half an inch in height, formed by the 
maxillary uniting with the edge of the frontal, and posteriorly with the parietal. The 
inferior border is formed by a long and strong zygomatic process of the squamosal, 
approaching, but not equalling, that of Platanista in size, and a triangular pointed 
postorbital process of the frontal, -7" in length, and directed backwards and downwards, 
but which does not meet the process of the squamosal, by a space equal to its own 
length. In Platanista there is no space or postorbital process, the anterior end of the 
prodigiously developed zygomatic process of the squamosal reaching so far forward as 
even to be lodged in a hollow in that part of the orbital plate of the frontal from 
which such a process is usually developed. 

The bones whicli enter into the formation ol the temporal fossa resemble in their 
number and arrangement those of the true Dolphins rather than of Platanista. The 
parietal appears in the shape of a wide arch, receiving in its concavity the squamosal, 
and articulating for a space of '&' with the well-developed alisphenoid, thus completely 



INIA GEOFFEENSIS JlND PONTOPOHIA BLAINVILLII. 91 

shutting off the squamosal from the frontal ; whereas in Plntanista the last-named 
bones unite for a considerable distance below the pointed anterior end of the somewhat 
triangular parietal, and the alisphenoid does not appear in the fossa at all. 

The orbit, in its structure, as well as its size, is intermediate between that of Pla- 
tanista and Belpkinus. Its antero-posterior diameter is 1". The malar bone is shorter 
and more thick and tuberous than in the Dolphins generally, and contributes chiefly to 
the formation of the prominent rounded antorbital eminence. The ends of the styli- 
form processes are unfortunately broken off; but the portions that remain adhere to 
the form prevalent among the Belphinidce. In the larger skull in the British Museum 
this process on one side is 1" long, and appears to have a free, natural, rounded termi- 
nation, not uniting, by .a very considerable interval, with the zygomatic process of the 
squamosal. If this is constantly the case, Inia presents, in this respect, a remarkable 
exception to all other Dolphins. There is no distinct lachrymal bone. 

The upper surface of the facial portion of the skull behind the rostrum is longer 
and narrower than in the Belphinidm generally. It is distinctly bounded on each side 
by the sharp, straight, and nearly parallel crest before spoken of as forming the upper 
margin of the temporal fossa. Within these crests, on each side, the narrow upward 
prolongations of the maxillaries are deeply hollowed. Their hinder edge extends an 
inch further back than the anterior apex of the supraoccipital, and they curve inwards 
round the top of the premaxillaries to articulate with the nasals, and enter for a small 
space, between these bones and the premaxillaries, into the formation of the lateral 
boundaries of the narial opening. It is the narrowness and excavation, combined with 
the straightness and elevation of the outer borders, of the maxillariea, which gives 
the peculiar character to the upper surface of the skull of Inia as compared with that of 
Belphinus. The difference is only one of arrangement of the same parts; there is 
nothing superadded like the extraordinary outgrowths upon the maxillae of Platanista. 

Immediately behind the narial opening is a somewhat square-shaped elevation, rising 
vertically in front, sloping behind, and hollowed out and overhanging at the sides, 
formed chiefly of the frontal bones, and suggestive of the peculiar elevation of this 
part so characteristic of the Ziphioids. The nasal bones are applied to the front wall of 
this elevation, but do not reach the top of it. In general form they are irregularly 
quadrilateral, prominent and thick near their longest, straight, inner border, where 
they meet each other in the middle line, and deeply hollowed and notched in their 
upper and lower margins. Their shorter, but straight and thick, outer border articulates 
with the maxiUaiy. Above and below they are bounded by the frontal, on which they 
rest. The greatest length of each bone is -9", the greatest breadth -7". They present 
no marked deviation from bilateral symmetry. Attached to the upper outer angle of 
each, and lodged in the groove between the frontal and maxillary, is a minute oval 
bone, -25" long, apparently originally distinct, though now partially united with the 
nasal ; and their inferior internal angles rest upon a median single triangular piece, -3" 



92 MR. W. H. PLOAVER ON THE OSTEOLOGY OF 

broad and -25" high, distinctly separated by a suture from the frontals. It will be seen 
from the above description that the nasals are extremely different from those of most of 
the Deljihi)iidw, in which they are generally reduced to irregular, oval, unsymmetrical 
nodules. PhoccBna, however, differs from its allies in this respect, and closely approxi- 
mates to Inia. In Flatanista also the nasal bones are well-developed flattened plates ; 
but they partake of the great elongation, naiTowness, and lateral distortion which per- 
vades this region of the skull. 

The opening formed by the junction of the anterior nares is 1" long, and the same 
width posteriorly. It is bounded laterally and in front by two very prominent, rounded, 
longitudinal elevations, formed by a thickening of the premaxillaries, like that seen in 
this region in Phoccena and Beluga, but considerably more marked. No part of the 
maxillaries comes to the surface in the middle line in front of the narial aperture as in 
many of the Delfhinidce (e. g. Olohiocephalus). 

The rostrum is exceedingly long and narrow, and, except at its base, much compressed. 
The diminution of its breadth takes place rapidly for the first fourth of its length, but 
for the remaining portion only very gradually. The bone of which it is composed is of 
dense texture ; and, even in this young subject, the sutures between the premaxillaries 
and maxillaries are almost obliterated. The width of the premaxillaries scarcely alters 
through their entire length, their outer boundaries being parallel, and the general 
diminution in the breadth of the rostrum taking place solely at the expense of the 
maxillaries. There is a narrow interval throughout in the middle line between the 
premaxillaries, and the subjacent cavity for the median ethmoid cartilage is not filled up 
with bone as in many of the Zijahiinw. 

On each side of the inferior surface of the rostrum (Plate XXVI. fig. 1) the alveolar 
tract, marked by the row of deep and distinctly separated tooth-sockets, extends from 
the apex to 1^" from the bottom of the antorbital notch. Between these tracts the 
palatine surface is quite flat, and in the anterior three-fourths slightly raised above 
their level. At the middle of the rostrum it is only "4" wide, but gradually expands 
posteriorly. Between the two maxillary bones, in the median line is a narrow fissure, 
in which, 1" behind the middle of the rostrum, a thin strip of the vomer appears, and 
continues visible as far as the posterior edge of the palate. 

The remarkable conformation of the bones of the hinder part of the palatial region 
in the Gangetic Dolphin has been well described by Eschriclit, who pointed out tliat the 
great lamella of bone which continues backwards the palatine portion of the maxillaries, 
and passes outwards and upwards to articulate with the squamosals and frontals, is 
really the pterygoid, and not the palatine as Cuvier supposed *. Tlie easily separable 
condition of the bones of the young Flatanista skull in the Museum of the Royal 
College of Surgeons has enabled me to confirm Eschi'icht's view ; for on removing this 
plate the true palatine is seen, forming as usual the greater part of the anterior and 
* Ossemens Eossiles, 4™° edit. (1836) tome viii. p. 130. 



INIA GEOFFEENSIS AJSTD PONTOPOEIA BLAINVILLII. 93 

outer wall of the nasal passage, but not entering in the slightest degree into the com- 
position of the free surface of the bony palate. 

In this disposition of the palatine and pterygoid bones Flatanista stands alone 
among Cetaceans ; even Inia presents no approximation to it. There are, however, in 
the latter genus some peculiarities in this region by which it may be distinguished 
from the ordinary Dolphins. 

Behind the posterior pair of teeth the palate loses its flatness, and begins to rise to a 
ridge in the middle line and slope away at the sides towards the roof of the orbits. 
The summit of the ridge is formed by the vomer, which is quite uncovered in the 
middle line by the palate-bones. The inner edges of these bones, applied to the 
surface of the vomer, are distinctly marked, and posteriorly are -4" distant from each 
other. The suture between them and the maxillaries is completely obliterated, so that 
then- limits forwards and outwards cannot be definitely stated. As in the ordinary 
Dolphins, the palatines have each an outstanding, nearly vertical, plate running outwards 
and backwards, unattached posteriorly, and forming the upper part (in the natural 
position of the skull) of the outer waU of the chamber which lodges the great post- 
palatine air-sinus. This plate is slightly developed and very thin, perforated by 
numerous large lacunae, and, owing to the non-development of the outer reflected portion 
of the pterygoids, is completely free along its inferior edge. 

The pterj'goids are comparatively simple, and also very thin and lacunated. As usual, 
the upper or attached portion forms a ridge along the side of the cranium, continuous 
posteriorly with the ridge on the side of the basisphenoid, which forms the inner 
wall of the cavity for the lodgment of the ear-bones. This portion articulates by 
nearly the whole of its inner edge with the hinder expanded part of the vomer, and 
externally with the alisphenoid and orbitosphenoid. From its anterior part springs 
the recurved descending plate which bounds externally the posterior nares, and, then 
turning inwards and backwards, forms the anterior wall of these passages below the 
palatines. This last-named plate of the pterygoid forms the hinder part of the bony 
palate ; anteriorly it lies on the hinder free edge of the inferior sm-face of the vomer, 
but does not quite cover it to the middle line ; behind the vomer it diverges rather 
more from its fellow, leaving a gap of from -1" to -2" in breadth. Posteriorly each 
terminates by a concave free margin. The third portion of the pterygoid, which 
exists in all ordinary Dolphins (excluding the Physeteridw), and which when 
present completely conceals that last described, being reflected from its hinder and 
inner edge outwards and upwards to meet the edge of the projecting plate of the 
palatine, and so close in the postpalatine sinus below, is wanting in Inia, or only 
represented in the osseous cranium by some small irregular body-excrescences. The 
result is that the cavity for the sinus is widely open below. It might be conjectured 
that this plate, being thin, brittle, and much exposed to injury during the process of 
cleaning the skull, had been broken away. It is certainly possible that such is the case ; 



H ■ ME. W. H. FLOWER OX THE OSTEOLOGY OF 

but as the adult and apparently perfect skull from Ega, in the British Museum, shows a 
precisely similar condition to that above described*, it is probable that, if ossification 
takes place at all, it is of a very imperfect character. 

Both petro-tympanic bones are unfortunately absent from the skull. The fossa 
at the base of the cranium for tlieir lodgment is shallow, and the aperture left 
in the cranial wall by their removal large, compared with that in an ordinary Dol- 
phin. It is ii-regular, circular, and averages 1" in diameter. In the largest skull 
in the British Museum these bones are present, and enter considerably into the forma- 
tion of the cranial wall, the inner and upper surface of the petrosal being seen in 
the interior of the cerebral cavity, on a level with the internal surface of the other 
bonesf. 

One circumstance in which the petro-tympanic bones of Inta differ from those of 
Platanista is their loose connexion with the rest of the cranium ; for they are only 
attached by ligament, as in Belphinus, and not locked in their place by a process of 
the mastoid. In general form the tympanic bullae resemble those of Deljihinus, though 
they are larger than in a member of that genus of corresponding size, and have their 
anterior (Eustachian) extremity rather more prolonged and pointed, though to a far 
less degree than in Platanista. Their antero-posterior length in the adult skull is 
1-65", their greatest breadth Tl". 

The mandible presents a remarkable miniature resemblance to that of a Cachalot. It 
differs from the mandible of all the true Deljihinidce by the great length, narrowness, 
and shallowness of the symphysial portion, which includes three-fourths of the tooth- 
bearing part of the rami. The consequence is that the hinder parts of the rami diverge 
much more rapidly from each other tlian in the true Dolphins. The coronoid process 
is unusually elevated. The lower jaw of Platanista, as is well known, presents all these 
characters, but in a much more exaggerated degree. 

The characteristics of the teeth have been well described by d'Orbigny and GeiTais. 
They are distinguished from those of all other Cetaceans by the peculiar and very 

* In the smaller skull in the same Collection nearly the whole of the pterygoids have heen destroyed. 

t After noticing that in certain Dclphinoids the aperture left hetween the hinder edge of the alisphe- 
noid, the exoceipital, basioccipital, and basisphenoid is eicceedingly Bmall, so that the tympano-periotic is 
stiU more shut out from the cranial cavity than in Baliena, Professor Huxley remarks that " in Platanista 
the aperture is large, and the periotic appears in the interior of the cranial canity in the ordinary way " (Ele- 
ments of Comparative Anatomy, 1864, p. 276). This is certainly the case in the two small Platanista skulls 
in the Museum of the College of Surgeons, upon which the observation was founded ; but it is worthy of not* 
that in a large and apparently aged skull of an individual of the same genus in the British Museum the 
periotic bones are completely shut out of the cerebral cavity by the excessive development of the proper cranial 
bones, and communicate with it only by a narrow passage fuHy an inch in length. MTiethcr this difference 
depends on age or on species I cannot at present determine ; but it shows that the relative position of these bones 
to the rest of the cranium may vary, even in most closely allied forms. 



miA GEOFFEENSIS AND PONTOPOEIA BLAIXVILLII. 95 

marked rugosity of the sui-face of their crowns*, and especially by the broad, rounded 
lobe, developed on the inner side of the base of the crown of those situated in the pos- 
terior part of both upper and lower jaws (see Plate XXVI. figs 1, 2 & 3). In the 
anterior two-thirds they are simple, conical, and slightly incurved. They giadually 
increase in size from the front of the jaws until the fourth from the posterior end of 
the series, after which they diminish again. Unlike those of most Dolphins, the teeth 
are imjDlanted by large and generally somewhat tveisted and flattened fangs (in the hinder 
teeth very wide transversely), which fit so tightly into deep alveoli that it is almost im- 
possible to extract them, even in the dried skull, without injury to the bone. When 
the mouth is shut they fit closely into the interspaces of the opposite series ; but there 
is little sign of attrition to be seen anywhere on their sui'face. 

The number of the teeth in the different specimens of Inia examined shows a conside- 
rable range of variation, presuming that they all belong to one species. In the one now 
described there are grigiTif =104. The larger specimen in the British Museum from 

OQ '■70 

Ega has 2gr:^=109, and also two minute rudimentary teeth in the gum behind the 
last in the left maxiUa. In the smaller skull from the same place there are 
281^=110. In the skiill in the Paris Museum, brought by d'Orbigny, there are» 

OQ OO 

according to Gervais, 33:^33=132 ; but in the type specimen in the same museum, taken 

OA Ofi 

from Lisbon, the number is given by de Blainville as ggii^e^^l^'^- ^^ ^^^ Berlin skull the 
teeth are g3:^,=131f. Von Martins in his diagnosis of the species gives 291159= 114. 

The bones of the hyoid apparatus scarcely differ from those of the ordinary Dol- 
phins. Their general form is shown in the figures (PI. XXVI. figs. 4 & 5) at half theu- 
natural size. The basihyal and thyrohyals are not yet united by continuous ossifica- 
tion. The stylohyals are thick, subcylindi-ical, slightly curved, and somewhat flattened 
towards the ends. 

" 
Antero-posterior diameter of the basihyal 1 -0 

Transverse diameter I'o 

Length of thyrohyal 2'0 

Greatest breadth 06 

Distance between the outer extremities of the thyrohyals... 3-4 

Length of stylohyal 2-7 

Greatest thickness 0'4 

The spinal column (PI. XXV. figs. 1 & 2) appears complete to the end of the tail, and 
' onsists of but 41 vertebrae, the smallest number known in any CetaceanJ. Of these, 

* Some Dolphins of the genus Steno of Gray present a similar though far less marked rugosity; and indi- 
cations of it are seen in young specimens of Orea and Fseudorca. t Peters, in a letter. 

J As the bones had been separated from each other adnclcaned at the time that they came into my hands lor 
VOL. VI. PART III. P 



96 MB. W. H. FLOWER ON THE OSTEOLOGY OF 

7 belong to the cervical, 13 to the thoracic, and 21 to the lumbo-caudal region. When 
the vertebrae are placed in order, with their bodies in contact, the whole column mea- 
sures 38-8". 

The cervical region, as in Platanista, occupies a larger proportional space than in 
most other Cetaceans, being 3-3" long, or t§Jo of the whole column. In a common Por- 
poise, measured for the purpose of comparison, it is but yggiy. All the vertebrae are 
distinct, as in Platanista, Beluga, and Monodon alone among toothed Whales. 

The atlas (PL XXVII. fig. 1), very large for the size of the animal, greatly resembles 
that of Platanista, but is higher in proportion to its breadth. Its neural arch is strong, 
and has on its upper surface a slight longitudinal ridge representing tlie spine. The 
base of the arch is not perforated as in many Cetaceans, and the groove for the sub- 
occipital nerve is but slightly marked. On each side, between the anterior and pos- 
terior articular surfaces, are two rounded eminences, the rudiments of an upper and 
lower transverse process. In Platanista there is only a single intermediate process (which 
Eschricht considers to represent the lower process), but it is developed to a much greater 
length. In Beluga both processes are present as in Inia, and upon corresponding parts 
of the surface of the bone. As in the other Odontoceti having a free atlas, there is a 
strong process developed from the hinder edge of the lower arch of the bone, which 
passes under and articulates with the inferior surface of the axis (see PL XXV. fig. 2). 
This is bifid at the extremity, and much more powerfully developed than in the young 
Platanista wliich served for comparison. 

The axis has a massive body, and a high neural arch. There is no distinct odontoid 
process, but only a general (though strongly marked) prominence of the anterior surface of 
the body, especially towards its lower margin. On the under surface of this there is a 
large rounded articular facet for the inferior process of the atlas. This is continuous at 
the sides with the anterior articular facets, and would indicate a tolerably free motion 
between the first two bones of the neck. In Platanista this anterior projection of the 
body of the axis is still more strongly marked, forming a process quite comparable with 
the " odontoid " of other Mammalia. In Beluga it is almost wanting. The other pro- 
cesses of this vertebra differ somewhat in detail from those of Platanista. The spinous 
process is broad and bifid ; the posterior zygapophyses are much less prominent, and their 
surfaces look more backwards. A proper transverse process can scarcely be said to exist. 
There are, however, instead of the single, conical, backward-directed process of Plata- 
nista, slight rudiments of an upper and a lower process, with a groove between them, on 
the hinder surface of the lateral wings of the bone which support the great articular 
facets for the atlas. The posterior epiphysis of the body was not ankylosed. 



description, I must admit the possibility of some of them being lost ; but the circumstances under wliioh the 
skeleton was prepared render this, at the least, extremely improbable. When it arrived in this country the 
vertebra; were all united by their natural ligaments. Unfortunately they were not counted when in this state. 



INIA GEOFFEEXSIS AXD PONTOPORIA BLAINVILLII. 97 

The remaining five cei-vical vertebrae are compressed in the antero-posterior direction, 
but less so than in most Cetaceans. They do not present the peculiar depression and 
transverse extension characteristic of the cer\ical vertebrae of Platanista, but their 
bodies are nearly circular in outline, and the height of the neural canal bears a more 
considerable proportion to its breadth. The bodies increase but very slightly in 
thickness from before backveards. The arches are wide and low, their sides meeting 
above at very obtuse angle, and so narrow in the antero-posterior direction as to 
leave spaces between them about equal to their ovra breadth. They increase but very 
slightly in height from the third to the seventh, and possess but a mere rudiment of 
a spine, scarcely recognizable in the third, and but -2" in height in the seventh. The 
anterior and posterior articular facets of the arches are well developed in all, and ha^f' 
their usual relations. 

The transverse processes are, as usual, two on each side, upper and lower ; the upper 
springs from the arch, the lower from the body of the vertebra. In the third vertebra 
these two are very near together, and approximate at their ends so as to enclose an 
oval foramen or canal -2" in its greatest diameter. On the left side this canal is com- 
pletely surrounded by bone; on the right side it is not quite completely inclosed. 
In Belufja similar rings are formed by the transverse processes of this vertebra, also in 
the Platanista described by Eschiicht, though in the College specimen there is but a 
single broad imperforate transverse process. In the fourth vertebra the processes are wider 
apart, short, and obtuse, and of about equal length ; a small elevation rises from the side 
of the body of the bone, midway between tliem. In the fifth vertebra they are still 
wider apart, owing to the upper one, which is short and conical, rising higher on the 
side of the arch. The lower process is much larger, stouter, rounded at the end, and 
directed backwards. Although upwards of -|" long, it was evidently not fully deve- 
loped in this immature individual, being tipped with cartilage. The prominence of 
this process, contrasting with the almost rudimentary condition of all the others, is a 
marked characteristic of the cervical region. In Platanista and Beluga, as in most 
other Mammalia, it is the sixth vertebra which has the most largely developed inferior 
transverse process, in the former very remarkably so. It is worthy of note, however, 
that the Dugong (ffalicore) agrees Avith Inia in this respect, as well as in many other of 
the characters of the neck-vertebrse. 

In the sixth vertebra, both upper and lower processes are small and conical. In the 
seventh vertebra the upper process is more developed ; the lower one still exists, but 
in quite a rudimentary state ; behind it is a shallow excavation for the head of the first 
rib. The laminae of the arch of this vertebra are wider than in the others ; its spine, 
as before said, is slightly higher; and the posterior surface of its body is transversely 
extended. 

The thirteen thoracic vertebrae measure in length when placed in close contact 12-5". 
Their bodies increase at first rapidly, then more giadually in length — the fu-st mea- 

p2 



98 ME. W. H. FLOWER ON THE OSTEOLOGY OP 

siiring '5", the sixth -9", and the last 1-2". Their arches are surmounted by rather 
long, erect, and (especially in the hinder part of the region) very broad spines trun- 
cated at the top. The antero-posterior breadth of these processes presents a constant 
relation to the length of the body, being always nearly equal with it, and forms rather 
a remarkable feature in the general aspect of the vertebral column. The height of the 
spine of the first thoracic vertebra is scarcely inferior to that of the others, which are 
almost precisely equal. In the sixth, from the inferior edge of the body to the junc- 
tion of the laminae of the arch measures 1'6"; the spine above this point is 2'2". 
Distinct articular facets or zygapophyses are developed on both the anterior and pos- 
tei'ior edges of the arches as far as the ninth vertebra, and on the anterior edge only of 
the tenth and eleventh. These, as usual, are broad and wide apart at the commencement 
of the series, and gradually become narrow and approximated as they shift from the 
sides to the summit of the progressively diminishing neural arch. 

The so-called oblique processes (metapophyses of Owen) begin to separate them- 
selves from the transverse processes at the fifth or sixth vertebra, and gradually 
pass upwards and inwards on the anterior edge of the arch towards the prozygapo- 
physes, which they supersede on the twelfth vertebra. Owing to the comparatively 
slight development both of these processes and the zygapophyses, the thoracic vertebrae 
of Inia are not locked together in the manner which distinguishes those of Platanista. 

It remains only to speak of the processes for the articulation of the ribs, which offer 
some interesting peculiarities. In all the ordinary Delphinido' the anterior ribs are 
articulated by their tubercle to a well-developed trans^'erse process standing out from the 
side of the arch, and by a long neck to the hinder edge of the body or root of the arch 
of the antecedent vertebra. There is usually no indication of any articular surface for 
the head of its own rib on the front edge of the body of the vertebra. At about the 
middle of the series the heads suddenly cease to be developed, and the rib is only attached 
by its tubercle to the end of the transverse process, still arising from the arch, but 
gradually lengthening and becoming lower in its pomt of origin, till at the end of the 
series it springs rather from the body of the vertebra than from the arch, and is in a line 
with the transverse processes of the lumbar vertebrae. This arrangement, departing con- 
siderably from .that found in the ordinary mammal, occurs in Delphinus, Phoccena, Orca, 
GIohiocepJialKs, Beluga, Monodon, and their immediate allies — in fact, in all the Del- 
])Mnidce which have ossified costal ribs. In the remarkably aberrant Hyperoodon and 
Physeter a totally different arrangement takes place in the hinder part of the dorsal 
region, which, however, is equally peculiar among the Mammalia. The upper transverse 
processes springing from the arch (diapophyses, Owen) suddenly cease, and the rib retains 
its connexion with the body only : the articular surfaces of the latter push out a process 
(which, on Owen's system, would be called a parapophysis), at the end of which the rib 
is attached, and which becomes the transverse process, being continuous serially with the 
transverse processes of the lumber region. In the first case, the transverse process on the 



INIA GEOFFEEXSIS AND PONTOPORIA BLAINVILLII. 99 

body of the last dorsal vertebra is arrived at by a gradual lowering of the transverse process 
of the arch of the first ; in the second it is a new process, first appearing on the body 
rather abruptly, as the process on the arch ceases, but for the space of two or three verte- 
brse coexisting with it, as in the cemcal region : or, to explain the case in other words, 
the anterior ribs in both have an upper and a lower connexion with the vertebrae ; in the 
first instance they lose their lower connexion by the non-development of their neck and 
head, but the gradual lowering of the transverse process brings the headless rib again 
in connexion with the body, by the intervention of a long straight process; in the 
second instance they always retain their lower connexion, but the development of 
a process out of the articular surface of the body, with concurrent shortening of the 
neck of the rib, and disappearance of the upper process of the vertebra, produces an 
exactly similar result. 

In Inia the mode of attachment of the ribs is, as far as I know, peculiar among 
Cetaceans, being intermediate between the two distinct forms above described, and far 
more resembling that which obtains in the Sirenia and the terrestrial mammals. The 
anterior vertebrae have as usual a tolerably well-developed, thick and rounded transverse 
process, springing from the arch at the junction of the pedicle with the lamma, and 
pointing upwards and forwards, with a large articular facet at its extremity ; this process 
gradually becomes shorter, till in the seventh vertebra little more than the articular 
facet remains on the side of the arch. On each side of the body of the first vertebra 
are two distinct articular facets, each receiving part of the head of the first and second 
ribs respectively. The same occurs in the two following vertebrae, though the facets are 
less distinctly marked, the head of the rib apparently articulating chiefly to the inter- 
vertebral substance in front of its own vertebra. In the fourth, and more distinctly 
in the fifth and succeeding vertebrae, there is a strongly-marked articular facet on the 
anterior edge of the body, while that on the posterior edge has entirely disappeared (a 
condition, it will be observed, never found in the true JDeljihinidw). Hereafter each 
rib is solely articulated to its own vertebra, and its lower attachment becomes moved 
by degrees from the anterior edge to the middle of the body. As far as the seventh 
vertebra the rib has a double attachment ; but in the eighth the upper and lower arti- 
cular surfaces (that on the arch and that on the body) have coalesced, though the part 
that originally belonged to the transverse process and that on the body are distinctly 
recognizable. This coalescence, however, becomes more complete ; and, by the diminu- 
tion of its upper part, the articular facet, at first elongated vertically, becomes oval in the 
opposite du-ection in the eleventh vertebra, and also begins to rise out from the body as 
a short thick process. This process is somewhat elongated and flattened in the twelfth, 
and notably so in the thirteenth vertebra ; and at the same time the articular surface be- 
comes gradually reduced in size, corresponding with that of the head of the rib. We have 
thus among the toothed Whales a third method by which the transformation fi-ora the 
first thoracic vertebra with its doubly attached rib, to the last with its singly attached 



100 ME. W. H. FLOWES ON THE OSTEOLOGY OF 

rib, is effected, not in this case by the disappearance of either the lower or the upper 
attachment, but by their gradual coalescence. 

In Platanista the attachment of the ribs is again different in detail, being something 
between that found in the true Delphinidm and in Inia. Each of the first seven ribs 
is attached to the transverse process of its own vertebra and to the body chiefly of 
the preceding vertebra; but the transverse processes differ from those of the iJel- 
iMnidoe in being very short, and in being more rapidly transferred down to the 
bodies; indeed this takes place as early as the sixth vertebra, and before the disap- 
pearance of the articular facet for the head of the rib, leading to a blending of the two 
articulations in one as in Inia. 

The remaining vertebrse (lumbo-caudal) are twenty-one in number. In accordance 
with the usual (and most correct custom) of reckoning the caudal region of the Cetacea 
as commencing with the first vertebra which bears a chevron bone*, there are but three, 
or at most four, vertcbrce, which can properly be called lumbar. The uncertainty rests 
upon the difficulty of determining, in a skeleton of which the bones are all separated, 
and in which, owing to its immaturity, the articular surfaces and processes are not very 
distinctly marked, to which of the vertebrae the first (always very small) pair of hsema- 
pophyses was attached. I think, however, that there can be little doubt that the 
fourth of the vertebrae behind the thoracic region did bear such bones, not only from 
indications on its own surface, but also because the facets on the hinder edge of the 
under surface of the fifth are too strongly pronounced to be the attachments of the 
small first pair. Taking, then, the true lumbar vertebrae at only three, Inia presents 

* As a unifoi-m system of nomenclature in enumerating the vertebrtc of Cetacea is veiy desirable, it is to be 
regretted that Esohricht and Reinhardt, in their most recent works on Cetology, should have given the weight 
of their high authority to reckoning as the last of the lumbar vertebraB the one immediately preceding the first 
chevro]! bone, and which has commonly been regarded as the first caudal. The only reason given for this 
change is, that " the anus, which may justly be said to mark externally the limits between the abdomen and the 
tail, is situated directly beneath the first chevron bone " '. This, however, does not prove the case ; for if we 
look at the skeleton of any terrestrial mammal in which the distinction between the different regions of the 
vertebral column is definitely marked, we may see that the commencement of the caudal region is situated some 
way in front of the position of the anus. We ought rather, according to this criterion, to reckon two or three 
of the vertebrK in the Cetacea commonly called lumbar to the region of the taU, — a view further strengthened 
by the fact that, iu the ordinary mammals, the chevron bones, when present, begin generally not on the fii'st, but 
on the second or third caudal vertebra. Such a division would, however, be quite impracticable. 

Each chevron bone belongs essentially to the vertebra in front of it. This is most clearly seen when they are 
small, as in the commencement of the series. In the skeleton of a Physeter that I lately examined, the first is 
even ankylosed to the posterior edge of the body of its proper vertebra, and has no connexion with that behind 
it. It is quite certain that any vertebra bearing a chevron bone cannot consistently be regarded as one of the 
lumbar series. We may therefore conveniently reckon the first vertebra which is so distinguished as the com- 
mencement of the caudal region. 



' Recent Memoirs on the Cetacea, published by the Ray Society, 1866 : Eschricht and Reinhardt on the 
Greenland Whale ; p. 105 ; and Reinhardt on Pseudorca cmssklens, p. 204. 



rS'IA GEOFFKENSIS AND PONTOPOEIA BLAINVILLII, 101 

an extraordinary deviation from all other Cetaceans, among which the number, though 
certainly very variable, is usually considerable, ranging from eight in PJatanista and 
Physeter to twenty-four in some of the Belphini and Lagenorhynchi. On the other 
hand, in the Sirenia, the lumbar region of the vertebral column is, as in Liia, extremely 
restricted. 

The three lumbar vertebrae are very remarkable for the great antero-posterior 
breadth of their processes, both spinous and transverse. The bodies are large, being 
respectively 1'3", 1'4", and 1-5" in length; their extremities are subcircular, and, as 
usual in the Cetacea, the middle of the side below the origin of the transverse process 
is much contracted, so that the median line of the under surface forms a sharp ridge, 
from which a strongly marked arterial groove runs outwards and backwards to the 
hinder edge of the root of the transverse process. The spinous processes resemble 
those of the posterior dorsal region ; the first two are slightly curved forwards, the last 
is nearly vertical and somewhat smaller. The oblique processes (metapophyses) are 
short, flat, rounded projections from the upper part of the laminae of the arch, very 
closely approximated to each other. The transverse processes rise from the whole 
length of the side of the body ; they are of nearly equal length, but increase in breadth, 
especially by the development of a considerable angular process on the middle of their 
anterior border, most conspicuous in the third vertebra; beyond this process the 
anterior border is sharply cut off, so that the extremity appears to point backwards. 
The hinder border is nearly straight, with a notch close to its origin from the body, 
continuous with the groove before spoken of on the inferior surface of the bone. 

The vertebra here reckoned as the first caudal closely resembles the last lumbar. 
Its body is of the same length, but its transverse process is even broader. The suc- 
ceeding tail-vertebrae keep up the same general character, having large heavy bodies 
and broad processes. The projecting surfaces on the hinder edges for the attachment 
of the chevi-on bones are very strongly marked as far as the ninth, after which they 
become obscure ; they are not seen on the anterior edge until the fifth. It is difficult 
to determine exactly how many chevron bones there were, but probably not more 
than eleven. The spinous processes, broad and rounded at their summits, become 
gradually lower, until in the tenth the greatly reduced vertebral canal is scarcely closed 
in by the laminae of the neural arch, and there is no longer a true spine. In the 
eleventh, the canal is altogether open above. The metapophyses continue in much the 
same relative development and situation as far backward as the spinous processes 
extend. The transverse processes gradually diminish in length, and lose their charac- 
teristic form. Already in the second that cutting away of the anterior edge noticed in 
the lumbar region is lost ; and in the third and succeeding vertebrae the anterior edge is 
straight, and the hinder one sloping, so that they appear to point forwards. In the 
eighth they form but a slight prominence on the anterior part of the body, and in the 
ninth they have altogether disappeared. The vertical perforations for the lateral 



102 MR. W. H. FLOWER ON THE OSTEOLOGY OF 

ascending branches of the caudal artery, so characteristic of a certain region of the tail- 
vertebrtE of the Cetacea, occur first iii the fifth vertebra, but only on the left side ; in the 
sixth they are seen on both sides, perforating the body of the bone, not the root of the 
transverse process. 

As in all Cetacea, the caudal vertebrae suddenly change theii- characters at the point 
where they enter the laterally expanded part of the tail aiid where the chevron bones 
cease to be developed. They now lose their cylindrical form, and become broad, de- 
pressed, and angular. There are seven such vertebrae in the present specimen ; and the 
eighth from the end of the series, or the eleventh caudal, reckoning from the be- 
ginning, is what may be called the transitional vertebra, being intermediate in form 
and size between its two exceedingly diflerent neighbours. The last two show a 
rapid diminution in width. The terminal one is triangular in outline when seen from 
above. 

Nothing can well be more dissimilar than the lumbo-caudal region of the spinal 
column in Inia and Platanista. In the latter the short bodies, the long narrow trans- 
verse processes, and high spines curving forwards and bearing immense laterally deve- 
loped oblique processes with (throughout the lumbar region) well-marked anterior and 
posterior articular surfaces, form most striking distinguishing characters. 

The chevron bones sent with the skeleton are ten in number. It is probable that 
the first is wanting, as there is none corresponding with the form this usually has in 
the Cetacea. I have therefore indicated its situation with a dotted outline in the figure 
of the vertebral column (PI. XXV. fig. 2). These bones agree in general characters 
with the processes of the vertebrae with which they are connected, being of moderate 
length, very broad and rounded at their free extremity. The lateral halves of the last 
three are not united in the middle line. 

There are thirteen pairs of ribs (PI. XXVII. fig. 2), the last being well developed 
and articulating with the transverse processes of the corresponding vertebrae. They 
are stout and heavy for their length, more so than in the ordinary Dolphins. In their 
comparatively cylindrical form they present a marked contrast to the broad flat ribs of 
Platanista. The last two or three are, however, much more compressed than the 
others. The curve, very strong and angular in the first, gradually diminishes and 
becomes more regular. The last has a slight turn outwards at the lower end, giving a 
gentle sigmoid curve to the whole bone. 

The anterior ribs have long and broad, somewhat compressed capitular processes, 
with distinct articular surfaces at the extremity and at the tubercle. In the fifth the 
length of this process is sensibly diminished. In the sixth, seventh, and eighth it 
shortens rapidly, the two articular surfaces being already confluent in the seventh. In 
the ninth a rounded projection of the lower border of the vertebral end indicates the 
rudimentary process ; in the tenth it has disappeared altogether, and henceforward the 
upper end of the rib ends in a somewhat dilated, oval, convex, articular surface, gradu- 



First 


3-7 


Second 


5-7 


Third 


6-9 


Fourth 


7-3 


Fifth 


7-3 


Sixth 


7-2 


Seventh 


7-1 



miA GEOFFEENSIS AND PONTOPORIA BLAINVILLII. 103 

ally diminishing in size. The mode of attachment of the ribs to the vertebral column 
has been noticed in the description of the thoracic vertebrae. 

The extreme length of the ribs of the right side in a straight line is as follows : — 

Eighth 6-9 

Ninth 6-7 

Tenth 6-5 

Eleventh 6-4 

Twelfth 5-9 

Thirteenth 5-3 

The costal cartilages, as in Platanista and all the Physeteridae, are not ossified. How 
many may have reached the sternum it is, in the present state of the skeleton, impos- 
sible to determine ; but indications of the attachment of only two pairs aie to be seen 
on this bone, wliich, if confirmed, would be most exceptional among Cetacea, and be 
another feature of resemblance with the Sirenia. 

The sternum (Fl. XXVII. figs. 3, 4 & 5) is very peculiar in shape, quite unlike that 
of any other Cetacean with which I am acquainted, and in its shortness, breadth, and the 
deep notch on the anterior border somewhat recalling that of the Manatee. It difl'ers 
from this, however, in its gi'eater solidity, especially towards the anterior part, and in 
possessing two strong triangular processes {I) projecting downwards and outwards from 
the fore part of the external surface. 

It consists of a single bone, which is at present but incompletely developed, all the 
prominences and the whole hinder margin terminating in cartilage. 

The extreme length of the ossified portion of this singular bone is 4"'2 ; its greatest 
breadth, near the middle, is 3". Its general form is irregularly oval. In the anterior 
border is a notch 1" in depth, vAth. smooth, rounded edges. On each side of this are 
two thick conical processes («), projecting directly forwards, •1" apart at their ends. As 
these have dried cartilage both on their tips and inner surfaces, it is possible that in 
the adult animal their ossification might extend so far as to convert the notch into a 
foramen. On each side of the hinder half of the notch the bone becomes very thick, 
running out on the external or inferior surface into the triangular process before no- 
ticed (5), and backwards and upwards into a thick in-egular edge {c), apparently for 
the attachment of the cartilage of the first rib. The hinder half of the bone is flat, 
and gradually becomes thinner towards its rounded and incomplete posterior edge, which 
is divided into two lobes by a narrow cleft, situated slightly to tlie right of the median 
line. About the middle of the left lateral margin is a small transverse notch, re- 
presented on the right side by an oblique j^erforation, apparently for the passage of a 
blood-vessel. Immediately behind this the margin is thickened and excavated for the 
attachment of the cartilage of the second pair of ribs (</). There are no other indica- 

VOL. VI. — PART III. Q 



104 MR. W. H. FLOWER ON TILE OSTEOLOGY OF 

tions of such attachments, though it is possible that the cartilaginous hinder margin 
may have been connected with another pair. 

In Plcdanista, according to Eschricht, four pairs of ribs are attached directly by their 
cartilages to the sternum, and the form of this bone has nothing in common with that 
of Inia. The manubrium is flat and triangular, very broad in front, with a straight 
anterior edge, and without either of the processes so prominent in Inia. This is 
succeeded by a distinct body, ossified from two lateral centres, and a xiphoid process 
wholly cartilaginous in the young specimen described. Many of the true Dolphins have 
two conspicuous pairs of processes on the manubrium sterni, evidently for the attach- 
ment of muscles— one projecting forwards and outwards, in front of and within the sur- 
face for the attachment of the first pair of sternal ribs, the other rising from the lateral 
border between the surfaces for the articulation of the first and second sternal ribs, and 
dh-ected somewhat backwards. These are especially developed in Monodon. It is to 
these that the processes of the sternum of Inia appear to correspond, though much 
modified in direction. The sternum of Phoccena entirely wants these processes; 
otherwise it presents some resemblance to that of Inia in its breadth, flatness, and in 
consisting of a single piece. 

The pectoral limbs of Inia are described by d'Orbigny as "larges, longues, et 
obtuses ;" and the present skeleton fully corroborates this account. 

The scapula (Plate XXV. fig. .3) does not present that -singularly aberrant character 
which is one of the most peculiar features of the skeleton of Flatanista, but conforms 
more to the ordinary type of the Dolphin-family. Its superior costa is long, and with 
a tolerably regular arch ; the anterior and posterior costse (of which the former is 
slightly the longer) are much hollowed out, so that the lower half of the bone is 
naiTower from side to side than in most Dolphins. Both the acromion process and 
coracoid are very long, flat, and expanding and truncated at their extremities. The 
glenoid fossa is large. 

The principal dimensions are : — 

Extreme height, from glenoid fossa to middle of superior 

costa 37 

Extreme breadth 4-8 

Breadth of body at root of acromion process 1 "2 

Length of acromion 1'7 

Length of coracoid process 1'3 

Length of glenoid fossa 1"2 

Breadth of glenoid fossa <J"9 

Tlie humerus is unusually long in proportion to the other segments of the limb, and 
very simple in its character. The tuberosity is very small ; but it is probably not com- 
pletely ossified. The neck is but slightly marked. The distal end of the bone is 



INI A GEOFFEEiSrSIS AND PONTOPOEIA BLAINVILLII. 105 

flattened, and not much expanded in width. The inner surface is quite smooth and 
slightly concave longitudinally. The outer surface is rougher, and has a rather deep 
pit a little way below the neck. 

The radius and ulna are considerably shorter than the humerus, contrary to what 
obtains in most Cetacea. They are vciy simple, broad and flat bones, but have a con- 
siderable space between them, owing to the concavity of the contiguous borders of the 
ulna and radius. The ulna presents the great peculiarity of possessing no rudiment 
of an olecranon process. 

Length of humerus 3'2 

Width at middle 1-1 

Width at lower end 1-6 

Length of radius 2-5 

Width at middle 1-2 

Width at lower end 1-4 

, Length of ulna 1-9 

Width at middle 1-0 

Width at lower end 1-6 

The carpal region is large, and composed in the present specimen in great measiu'e 
of cartilage. There are five principal ossifications. Intending to discuss fully the 
homologies of the carpal bones of the Cetacea with those of the terrestrial mammals 
in my Osteography of the genus Physeter, I will only say here that these appear to 
represent : — 1 the scapho-trapezium, 2 the lunar, 3 the cuneiform, 4 the unciform, 
and 5 the magno-trapezoid. They have probably been somewhat disturbed from their 
natural position by unequal shrinking of the surrounding cartilage in drying. In 
addition to these five, an oval bone (6) projects from the ulnar border of the carpus, 
which must represent the pisiform bone, although considerably displaced from its 
normal situation. The bone which appears to belong to the second row of the carpus 
near the radial border, and which might well be taken for a trapezium, is probably 
the first metacarpal, as already determined in other Cetaceans by Cuvier, Gegen- 
baur, and Van Bambeke. 

The digital portion of the hand consists of five fingers of moderate length, and 
spreading somewhat from each other. The second digit is the longest, the third nearly 
approaches it, the fourth and fifth are much shorter. It is possible that the terminal pha- 
langes of the digits are not present in every case, especially as they do not always ossify 
before the animal has attained a considerable age ; but the following are the numbers of 
the phalanges present, exclusive of the metacarpals: — I. 1, II. 5, III. 4, IV. 2, V. 2. 
The indindual phalanges are thus not numerous ; but they are long in proportion to 
theii" breadth. 

From the humerus downwards the pectoral limb of Inia presents considerable re- 
semblance to that of Platanista, both agreeing in the great length of the humerus as 

Q 



9 



106 ME. W. H. FLOWER O^N THE OSTEOLOGY OF 

compared with the forearm, and in the absence of the olecranon process. In the 
carpus, to judge by Eschriclit's figure, some differences of detail may be found. They 
agree in the comparative length and slenderness of the phalanges and spread of the 
fingers; but Platanista differs from Liia and all the other Dolphins in the nearly 
equal development of the four outer digits, giving the remarkable truncated form to 
the termination of the extremity. 

The pelvic bones have unfortunately not been preserved with the skeleton. They 
are also unknown in Platanista. 

II. On the Skull of Pontoporia blainvillii. 

In the Museum at the Jardin des Plantes, Paris, is the skull of a small Dolphin 
brought by M. de Freminville, an officer in the French navy, from the neighbourhood 
of Monte Video, at the mouth of the Rio de la Plata. This was first described by 
Professor P. Gervais, in the ' Bullet, de la Soc. Philomathique de Paris,' 1844, (27 Avril) 
p. 38, as Belphinus Blainvillei ; also in ' ITnstitut,' of the same year. 

In the part of the ' Zoology of the Voyage of the Erebus and Terror ' devoted to 
the Cetacea, published in 1846, Dr. Gray gave a figure and brief description of this 
skull, and constituted the genus Pontoporia for the reception of the animal to which it 
belonged. 

Professor Gervais, in the description of the " Mammiferes " of d'Orbigny's ' Voyage 
en Amerique Mcridionale,' published in 1847, but the introduction to which bears the 
date of December 1846, redescribcd and figured the skuU (plate 23), pointing out 
that its peculiarities were sufficient to entitle it to rank as a subgenus, for which the 
name of Stenodel2)Ms was proposed. In the same plate a figure is given of a long- 
beaked Dolphin, observed by d'Orbigny off the coast of Patagonia, but of which no 
portion was brought home ; and a conjecture is thrown out that this Dolphin belonged 
to the same species as the skull presented to the Museum by M. de Freminville. 
Although this is a mere assumption, and not a very well founded one, as even the 
colour does not correspond with the brief description given by M. de Freminville*, it 
has unfortunately been treated as a certainty in most systematic worksf, and thus Pon- 
tojwria, the skull of which shows such near affinities with those of the river-Dolphins 
Inia and Platanista, and which from its only known habitat may be wholly or partially 
fluviatile, and of which the external form is entirely unknown, is now regularly installed 
in zoological literature as an oceanic Dolphin with a high falcate dorsal fin ! 

A few weeks ago, and after the whole of the foregoing description of the skeleton of 

* " D'apres un renseiguement favori par M. de Freminville, le Dauphin dont provient ce crane, est long de 
quatre pieds, et U est blanc, avec uno bande doi-sale noire." 

t See Gervais, Hist. Nat. des Mammiferes (1855), vol. ii. p. 322; Gray, Cat. Seals and Whales, Brit. Mus. 
(1866) p. 231. 



INIA GEOPPEENSIS AND PONTOPOEIA BLAINVILLII. 107 

Inia was written, a second skull of Ponfoporia, also from the mouth of the Eio de la 
Plata, was received at the British Museum, as a present from Dr. Hermann Burmeister, 
of Buenos Ayres. With his wonted liberality, Dr. Gray immediately informed me of 
its arrival, and has permitted me to add to the description of the skuU of Inia a com- 
parison with this nearly allied form. 

The skuU (PI. XXVIII.) is that of a perfectly adult animal. The sutures are par- 
tially obliterated, and the bones are compact and heavy. Many of the teeth are broken, 
some having been lost during life and the alveoli filled up ; the remainder are considerably 
worn at the pomts. The rostrum is curved downwards towards the extremity, much 
more so than in the Paris specimen ; this is probably the eifect of age, as a similar 
change takes place in Inia and some other Dolphins. The mandible partakes also 
of this curve. The small, roimded and depressed cranium, and very long, narrow and 
compressed beak, give a remarkable appearance to this skuU, reminding one, as Gervais 
remarks, of the head of a scolopacine bird. 

The principal dimensions are : — 

II 
Extreme length 15'8 

Length of rostrum (from anterior end of premaxillary to 

bottom of antorbital notch of maxillary) 11'2 

From anterior end of premaxillary to lower edge of nasal 

bones 1 3"5 

Greatest breadth (across zygomatic process of squamosals) 4-8 

Breadth of foramen magnum IT 

Breadth of occipital condyles 2-4 

Breadth across antorbital processes of frontals 2-6 

Breadth of rostrum at base 1'8 

Breadth of rostrum at middle 0'6 

Mandible, length 13"7 

Mandible, length of symphysis S'O 

Greatest breadth posteriorly 4-5 

Height at the coronoid process 2-3 

The supraoccipital is broader and shorter than in Liia, terminating in front by a much 
more open angle, and on each side ui a low ridge, coming in close contact with the broad 
posterior extremities of the suprafrontal plates of the maxiUaries. In the ankylosed 
condition of the bones it is impossible to say whether any of the frontal intervenes 
between them. The temporal fossa resembles that of Inia in its extent and form. 
The zygomatic process of the squamosal is proportionally longer, and meets the post- 
orbital process of the frontal. The relative forms of the parietal, squamosal, and frontal 
bones, as they appear in the temporal fossa, more resemble those of Platanista than of 
Inia ; but a narrow piece of the parietal prevents the union of the frontal and squamosal 



108 ME. W. H. FLOWER ON THE OSTEOLOGY OF 

below. The alisphenoid is concealed by a plate of the pterygoid, which articulates with 
all three bones just mentioned. 

The orbit is slightly larger in proportion to the length of the cranium than in Inia, 
and therefore considerably more so than in Platanista. The upper margin forms a 
wider arch than in the former ; the postorbital process is broader and shorter ; the 
antorbital tuberosity much smaller, but still chiefly formed by the malar bone. The 
styliform processes are unfortunately broken oflT. 

The upper surface of the skull is remarkably flat, showing scarcely a trace of the 
postnarial elevation. On this surface the frontal bones appear in a narrow, slightly 
raised median piece behind the nasal bones, "T" long, and '5" wide, bounded laterally 
by the posterior extensions of the maxillaries — and on each side in the supraorbital 
plates, of which a much broader piece is left uncovered by the maxillaries than in 
Inia. The nasals are flattened, irregularly quadrate plates, as in Inia, but, in con- 
sequence of the direction of the frontals, lying nearly horizontally instead of vertically. 

The narial aperture is broader, but shorter, than in Inia, being encroached upon by 
the largely developed antenarial tuberosities of the premaxillaries, which are broader and 
flatter on the surface than in Inia. The upper obtusely pointed ends of the pre- 
maxillaries extend to a level with the inferior border of the nasals, but do not arti- 
culate with them, as a strip of the maxillary comes between. The hmder ends of 
the maxillaries are broader and flatter than in Liia ; but in front of the nostrils they 
are much more contracted, and above the orbits have, a small but distinct longitudinal 
crest, •?>" high at the middle and gradually subsiding at the ends. This is not a mere 
elevation of the edge of the bone, as in Inia, but a distinct ridge placed some way within 
the suture between the maxillary and the orbital plate of the frontal, and of which there 
is no trace in Inia. Between this crest and the elevated portion of the premaxillary there 
is a very deep and narrow fossa, continuous in front with an extremely narrow but deep 
groove, which lies between the maxillary and premaxillary along the entire length of the 
rostrum, and which is only faintly indicated in Inia. The rostrum is considerably 
longer and narrower in proportion to the size of the cranium than in Inia. 

The palate-bones resemble those of Inia in not covering the vomer in the middle 
line. They have a small free external plate. Unfortunately the greater part of the 
pterygoids is broken away ; but enough remains to show that these bones do not conform 
to the type of the ordinary Dolphins, but are arranged in a peculiar manner, apparently 
intermediate between those of Inia and Platanista. A broad outer lamella, resembling 
that so characteristic of Platanista, remains on each side, and, though not covering the 
palatine anteriorly as in that genus, passes upwards and outwards to the temporal 
fossa, overlying the alisphenoid and articulating with the squamosal, parietal, and 
frontal 

The petrotympanic bones are wanting on both sides, showing that their mode of 
attachment resembles that of Inia rather than that of Platanista. 



IXIA GEOFFRENSIS AND PONTOPOEIA BLAINVILLII. 109 

The mandible resembles that of both Inia and Flatanisfa, and is intermediate 
between the two in narrowness and comparative length of the symphysis. Its osseous 
substance is very dense, and the two rami are completely ankylosed at the symphysis. 
Running along each side of the symphysial portion is a deep and narrow groove, corre- 
sponding to that on the rostrum between the maxillary and premaxillary. 

The teeth are implanted in distinct alveoli. As many have been lost from tlie 
anterior part of the lower jaw during life, and the sockets completely filled up, their 
number cannot be estimated with perfect accuracy, but it may be estimated as follows : 
54?— &4? ^"'''- -^1^ h.2i\e broad fangs, much compressed laterally, surmounted by a 
crowTi, the base of which, when seen from aboye, is of a quadrilateral form, with the 
angles rounded off, longer from before backwards than from side to side ; this suddenly 
contracts into a slender subconical apical portion, much compressed in the opposite 
du-ection, and slightly incurved at the apex, which is worn off in nearly all the teeth of 
this old specimen. The enlarged base of the crown, M'hich forms a sort of cingulum, is 
slightly granulated on the surface, and in the natural state is entirely concealed within 
the gum. The projecting contracted portion has a smooth glossy surface. The teeth 
vary but little in size or form throughout the whole series of both jaws. The dimen- 
sions of one taken from the middle of the lower jaw are : — 

Length of fang -15 

Length of crown -24 

Antero-posterior breadth of cingulum -17 

Transverse breadth of cingulum •!! 

Antero-posterior breadth of apical part at middle ... "05 

Transverse breadth '10 

This peculiar form of the teeth, which distinguishes Pontopcnna from all the ordinary 
Dolphins, and affords another evidence of its afiinity with Inia, has not been observed 
in the Paris specimen. Gervais's description is as follows : — " Les dents * * * * sont 
petites, longues de 5 ou 6 millimetres au plus, toutes plus ou moins aigues, et au 
nombre de 53 ou 54 superieurement, ainsi quinferieurement. Les posterieures sont un 
pen moins aigues que les autres, et leur partie terminale est un peu recourbce." 

The Paris skull, moreover, according to the figures, has a less elongated and slender 
rostrum than the present specimen — a difference which may certainly depend on age, 
presuming that the two animals belong to the same species. 

• I II. On the Systematic Position of Inia and Pontoporia in the order Cetacea. 

Tlie foregoing sketch of the principal osteological features of Inia shows that this 
Cetacean presents peculiarities sufficient to constitute it a well-mai-ked genus among 
the Dolphins. Its natural position in the order, and its affinities, howeVer, can only be 



110 • ME. W. H. FLOWEB ON THE OSTEOLOGY OF 

determined when a complete and satisfactory classification of the entire group can be 
arrived at. The requisite materials for accomplishing this are at present wanting. 
The anatomy of many distinct forms is still but imperfectly known ; and moreover it is 
probable that there are many others existing as yet undiscovered. We know enough, 
however, to arrive at certain general conclusions. The larger natural divisions may be 
indicated with tolerable certainty ; and when the extent and limits of these become 
generally recognized, much will have been done towards clearing the ground for future 
observation. We shall at least be spared from the irrelevant compai-isons, between 
objects essentially dissimilar, with wliich anatomical treatises on the Cetacea are too 
often encumbered. 

Before proceeding further with this part of the subject, I would remark, in passing, that 
sevei'al resemblances pointed out above between the skeleton of this Cetacean and 
that of the Sirenia, accordmg singidarly with d'Orbigny's observations upon its external 
foim and habits, can scarcely be regarded as evidences of affinity ; they only add 
somewhat to the numerous morphological analogies between the members of these 
essentially distinct orders. 

The interval which separates the Whalebone-Whales frbm all the Whales with teeth, 
in almost every point of their structure, is far greater than can be found between the 
most widely divergent forms of the latter. Hence the division of the Cetacea into 
several primary groups or families, of which the Whalebone-Whales constitute one, and 
are therefore treated as equivalent to some of the minor groups of the Toothed Whales, 
is quite inadmissible. The recognition of two great and distinct groups (suborders) is 
the first requisite to a right appreciation of the classification of the Cetacea. 

The principal distinctive characters of these two groups were defined in a former 
paper*. Increased knowledge of their structure, especially of the Odontoceti, has 
rendered some slight modifications of these characters necessary. They may at present 
stand thus : — 

1. Mystacoceti or Bal^noidea. Teeth never functionally developed, but always disap- 
pearing before the close of intra-uterine life. Upper jaw provided with plates of 
baleen. Olfactory organ distmctly developed. External respu-atory aperture 
double. Skull symmetrical. Maxilla produced in front of, but not over, the 
orbital process of the frontal. Lachrymal bones small and distinct from the jugal. 
Rami of mandible arched outwards, their anterior ends meeting at an angle, and 
connected by fibrous tissue, without any true symphysis. Sternum composed of a 
single piece, generally broader than long, and connected only with the first pair- of 
ribs. No costo-sternal bones. All the ribs at their upper extremity articulating 
only with the transverse processes of the vertebrae ; their capitular .processes, when 
present, not articulating immediately with the bodies of the vertebrae. 

* Proc. Zool. Soc. 1864, p. 388. 



INIA GBOFFREXSIS AND PONTOPOEIA BLAINVILLII. Ill 

2. Odontoceti or Uelpuinoidea. Teeth always developed after birth, and generally 
numerous, sometimes few and early deciduous. No baleen. Olfactory organ rudi- 
mentary or absent. External respiratory aperture single. Upper surface of the 
skull generally, if not always, unsymmetrical. Hinder end of the maxilla ex- 
panded, and covering the greater part of the orbital plate of the frontal bone. 
Lachrymal bone either inseparable from the jugal or, when distinct, very large 
and forming part of the roof of the orbit. Rami of mandible nearly straight, 
much expanded in height posteriorly, and coming into contact in front by a surface 
of variable length, but always constituting a true symphysis. Sternum almost 
always composed of several pieces placed one behind the other, and always con- 
nected with several pairs of ribs, either by cartilage or by distinct costo-stcmal 
bones. Many of the ribs with capitular processes developed, and articulating 
with the bodies of the vertebrae. 
It is not necessary to pursue further the arrangement of the Mystacoceti, as it has no 
direct bearing upon the subject of this memoir, and as moreover I have no reason to 
make any alteration in the divisions into families and genera sketched out in the paper 
above referred to. 

The subdivision of the Odontoceti, according to their structural affinities, presents at 
first sight considerable difficulty. To relate all the various attempts, more or less 
successful, that have been made to unravel this problem would be out of place here. 
I will only add one more to the number, founded chiefly on an examination of the 
osteological characters of the principal members of the group*. 

In seeking for some starting-point ft-om which to commence the formation of a 
natural division of the Toothed AVhales, one has occurred to me which I have not found 
hitherto noticed. The strong and well-defined bones which connect the ribs with the 
sternum, ossified even at birth, common to the Porpoise, true Dolphins, and their 
nearest allies, are represented even in the adult Hyperoodon by an entii-ely unossificd 
cartilage. In the four skeletons of Physeter macrocephalus that I have had the oppor- 
tunity of examining, I have looked in vain for sterno-costal bones, some of which would 
certainly have been preserved if they approached in relative magnitude and density 
those of the true Dolphins. In answer to my inquiries on the subject. Dr. George 
Bennett has kindly informed me that, in both the skeletons of the genus Kogia, now 
mounted in the Sydney Museum, the cartilages are unossified ; and I am indebted to 
Professor Van Beneden for similar information respecting the skeleton of the ziphioid 
Micropteron preserved iu the Zoological Museum at Brussels. From these facts, I think 
that we may safely infer that the absence of ossified sternal ribs is a character common 
to the large natural group which includes Physeter, Hyperoodon, and the Ziphioids. To 
* The arrangement here proposed nearly coincides with that arrived at by Professor Huxley and myself, 
when discussing this subject together before the delivery of the course of Hunterian Lectures at the Koyal 
College of Surgeons for the present year (see 'Lancet,' 18C6, vol. i. p. 381). 

VOL. VI. PART III. R 



112 ME. W. H. FLOWER ON THE OSTEOLOGY OF 

these may also be added Plafanisfa and Inia. Here, then, is a character derived from a 
part of the organization apparently less liable to adaptive modification than the teeth or 
fins, which may be taken as the basis of a primary division. It must now be seen whe- 
ther the remaining essential structural modifications are in accordance with it. Still 
confining our attention to the axial skeleton, there are certain tolerably obvious pecu- 
liarities about the vertebral column, more especially in the thoracic region, that will 
afford considerable assistance. As before indicated (p. 98), a peculiar mode of attach- 
ment of the ribs to the vertebrae is constantly found associated with the sterno-costal 
bones. The genera thus characterized may therefore be separated at once as a distinct 
natural group. They have also several minor characters in common, which will be 
pointed out presently. 

Should the whole of the genera with cartilaginous sternal ribs be united into a single 
group, equivalent to that just marked off'? I am inclined to think that they should 
not. To revert to the same point of structure just mentioned, it was shown before that 
Physeter and Hyperoodon agree in a very peculiar condition of thoracic vertebrte and 
rib-attachments. Whether Kogia and the Ziphioids conform with their nearest allies 
in this respect I am not at present able to say ; but we may assume with tolerable 
certainty that they do. But here, as well as in many more trivial characters, including 
the teeth and pectoral limbs, Inia and Platanista differ- — and differ, as it appears to me, 
more than any of the true Dolphins do, inter se. I would therefore raise the Cacha- 
lots and Ziphioids on the one hand, and Platanista and Liia on the- other, to the rank 
of primary divisions of the Toothed Whales. With the latter it is in the highest degree 
probable that the genus Pontoporia should be associated. This group is not so com- 
pact and easily defined by positive characters as the other two, between which it 
naturally stands. The two genera whose structure is most completely known vaiy 
widely from each other, one diverging towards the Physeteridce, the other towards the 
DelphinidsB, yet distinctly marked off from either. The validity of the group as a 
natural one will be greatly strengthened if the skeleton of Pontoporia should be found 
to possess the characters common to Platanista and Inia*. It would be interesting, 
moreover, if it should be discovered that this Dolphin is, like the members of the other 
two genera, habitually fluviatilef. 

* Dr. Gray in the " Zoology of the Voyage of the Erebus and Terror" placed Inia and Ponto2yona in one 
section at the end of the family DeljAinida;, following immediately upon Platanista. In his recently published 
Catalogue, Platanista constitutes the fourth family (Platanistidae) of the Cetacoa, foUowiug the Catodoutidoe ; 
Inia forms a separate (the fifth) family, Iiiiida; ; and Pontoporia commences the sixth family (Delphinidae), 
comprising idl the remaining Dolphins except the Globiocephalidae and the Ziphiidio. 

Gervais (Hist. Nat. des Mammiferes, 1855) unites Platanista, Inia, and Stenodelphis {Pontoporia) to form 
one of the five tribes {Platanistim, Delphim7is, Orcins, Monoclontins, and Phodnins) into which the family 
DelphiniJes is divided. The primary divisions of the order or families arc : — Phjsdcrides, ZiphiiJes, Delphinides, 
and BaUnides. 

t It is to be hoped that Dr. Burmeister may be able to obtain information on this point. I should mention 



INIA GEOFFRENSIS AJSTD PONTOPOEIA BLAINVILLII. 113 

I will now endeavour to formularize the distinctive characters of these three jn-imary 
groups of the Odontoceti, giving them the rank of families. 

I. PHTSETERiDJi:. Costal Cartilages not ossified. The hinder ribs losing their tuber- 

cular and retaining their capitular articulation with the vertebrse. The greater 
number of the cervical vertebrae ankylosed together. Pterj'goid bones thick, 
produced backwards, meeting in the middle line, and not involuted to form 
the outer wall of the postpalatine au--sinus. Symphysis of mandible of moderate 
or excessive length. Xo functional teeth in upper jaw. Mandibular teeth various, 
often much reduced in number. Lachrymal bones usually large and distinct. 
Bones of the skull raised so as to form an elevated prominence or crest behind the 
anterior nares. Orbit of small or moderate size. Pectoral limbs small. Dorsal 
fin usually present. 

II. Platanistidje. Costal cartilages not ossified. The tubercular and capitular articu- 

lations of the ribs blending together posteriorly. Cervical vertebroe all free. Ptery- 
goid bones thin, not conforming in their mode of arrangement with either of the 
other sections. Jaws very long and narrow ; both with numerous teeth having 
compressed fangs. Symphysis of mandible very long, exceeding half the length of 
the entu-e ramus. Orbit very small. Lachrymal bones not distinct from the 
jugal. Pectoral limbs large. Dorsal fin rudimentary*. 

III. Delphinid^. Costal cartilages firmly ossified. Posterior ribs losing their capi- 
tular articulation, and only uniting with the transverse processes of the vertebrae 
by the tubercle. Anterior (2-6) cervical, in most, ankylosed together. Ptciy- 
goid bones short, thin, involuted to form, with a process of the palatine bone, the 
outer wall of the postpalatine air-sinus. Numerous teeth in both jaws [Monodon 
excepted), sometimes deciduous. Sj'mphysis of mandible short or moderate, never 
exceeding one-thu-d the length of the ramus. Bones of the skuU not raised into 
a distinct crest behind the anterior nares. Orbit of moderate size. Lachrymal 
bone not distinct from the jugal. Pectoral limbs vaiying much in form and size. 
Dorsal fin usually present. 

I. The Physeferido' appear to constitute a very natural groupf . This may, however, 
be divided into two weU-marked subfamilies : — 

that Mr. Danvin has informed me that he met with no evidence of the existence of a freshwater Dolphin in the 
La Plata system of rivers, and that no mention is made by Azara of any such animal. 

* These characters are subject to modification when more is known of the structure of Pontopona. 

t Van Beneden insists strongly upon the close aifinity of Phifseter with the Ziphioids ; he says, " Comme on le 
voit, les Cachalots sont pour nous dta Ziphioides veritables, portant une rangce de dents fortes ct espacees sur 
chaque branche de maxillaij'e " (Mem, sur mie Xouv. Espeee de p^iphius, Mem. de I'Acad. Eoyale de Belgique, 
t. xvi. 1863). 

s2 



114 IVfB. W. H. FLOWER ON THE OSTEOLOGY OF 

1. Physeterince, characterized by the numerous teeth in the lower jaw, and having 

no distinct lachrymal bone, including the genera Pkyseter and Kogia (Gray)*. 

2. Zij)hiince, with only one or two pairs of teeth in the lower jaw (besides the ru- 

dimentary concealed teeth), and a distinct lachrymal bone. This includes 
Hyperoodon, Berardius, Ziphius, Micropferon, Dioplodon, and several extinct 
forms. 

II. The two best-known genera of the Platanistidce must each be placed in a distinct 
subfamily, characterized thus : — 

1. Platanistinm. Maxillary bones supporting large bony incurved crests. No 

cingulum or tubercle at the base of the crown of the teeth. Pectoral fins 
truncated. Visual organs rudimentary. External respiratory aperture longi- 
tudinal, linear. 

2. IniincB. Maxillary crests absent, or very slightly developed. Many of the 

teeth with a complete cingulum or a distinct tubercle at the base of the 
crown. Pectoral fin ovate, obtusely pointed. 

The position of Pontoporia cannot be definitely determined until more is known of 
its general structui-e ; but as its cranial and dental characters accord most nearly with 
those of Inia, it may be placed provisionally in the same subfamily. 

III. Although the DeJpkinidce present considerable diversity in the characters of their 
dentition, in the relative length of the rostral part of the skull, in the form and struc- 
ture of the pectoral limb, and in the form and size of the dorsal fin, it is by no means 
easy to subdivide them into natural gi-oups. It is even difficult to define neatly the 
distinguishing characters of the genera, so much do they blend one into the other. 

_^ The Narwhal and the Beluga appear to separate themselves from all the rest, by 
certain well-marked structural conditions, especially the characters of the cervical ver- 
tebra. As these two animals are in almost every part of their skeleton nearly identical, 
even to the number of the vertebrae and phalanges, I am disposed to look upon the ex- 
ceptional dentition of the former as an aberration of secondary importance, and to unite 
the two genera into a distinct subfamily, placing it next to the Platanistidce. Among 
the remaining genera, none stand out in equal prominence. We must either group 
them together in one subfamily or make almost as many subfamilies as there are genera. 
For the present I prefer adopting the former course. Phocoena and Neomeris stand by 
themselves in the form of their teeth and certain cranial characters. Orca is distin- 
guished from all the others by its excessively broad manus, and GloMocephalus by the 
extreme length and narrowness of the same member. Belphinus and its allies are charac- 
terized by the long narrow rostrum and numerous teeth. Each of these genera might 
* A genus quite distinct from Physeter. It has also been called Euphysetes (Wall. Descr. New Sperm Whale, 
&.Q., 1851) ; but Gray's name (Zool. Erebus and Terror, 1846) clearly has the priority. 



INIA GEOFTEENSIS AND PONTOPOEIA BLAINVILLII. 



115 



easily be made the type of a distinct subfamily, were it not for the difficulty of placing 
the numerous osculant forms, Pseudorca, Grampus, Lagenorhynckus, &c. 

In the following tabular view of the arrangement of the Cetacca, many of the genera 
lately formed, chiefly by subdivision of the old genus JDelphinvs, are not introduced. 
It must not be be inferred from this that I question their validity, though such as are 
founded on skulls alone may require revision when the entire skeleton is known. But 
as the present object is to determine the position of Inia and Pontoporia in the order, 
it is only necessary to mention the well-established and generally recognized generic 
divisions. 



Order CETACEA. 



Suhorderi. 

I. MrslACOCETI* 

or 
Balsnoidea. 



Families. Subfamilies. 

f BatenidsE Balaemnae . . . 



i_ Balaenopteridae 



Genera. 

f Baltena. 
\ EubalsEna. 



iJIegapterinffi Megaptera. 
f Physalus. 
Balisnopterina; i Sibbaldiua. 

[ Baloenoptera. 



"" PhyseteridsB 



II. ODONTOCETlt 

or < 

Delphinoidca. 



Platanistidae 



Delphinidse 



■i 



Phj-setennffi i -^g, 



Physeter. 



I 



rHyperoodon. 
Bcrardius. 

Ziphiinffi ■< Ziphius. 

j Dioplodon. 
1^ Micropteron. 



{Platanistinffi Platanista, 
J Pontoporii 
Inuns I Inia. 

f Monodon. 
' BelugmiE I Beluga. 



Phocsena. 

• ^ Neomcris. 

Grampus. 
Orea. 

. DelphinmsB ? <! Pseudorca. 

Lagenorhynchus. 
Delphinus. 
Dcliihinaptcrus. 
(^ Globiocephalus. 



* fivariii, KiJTOs ; equivalent to the German " Barten-Walle." 



t OCOIIS, KtJTOS. 



116 OSTEOLOGY OP INIA GfEOITEENSIS AND PONTOPOEIA BLAINVILLIl. 

DESCRIPTION OF THE PLATES. 

The figures in Plates XXV., XXVI., and XXVII. are drawn from the skeleton of the 
young Inia geoffrensis described above. 

PLATE XXV. 

Fig. 1. Upper surface of the cranium and vertebral column of Inia geoffrensis. One- 
fourth the natural size. 
Fig. 2. Side view of the skull and vertebral column. One-fourth the natural size. 
Fig. 3. Bones of the right pectoral limb. Half the natural size. 

PLATE XXVI. 

Fig. 1. Inferior surface of the cranium of Inia geoffrensis. Half the natural size. 

Fig. 2. Superior siu'face of the mandible. Half the natural size. 

Fig. 3. A maxillary tooth from the left side, the fourth from the posterior end of the 

series. Natural size. 
Fig. 4. The basi- and thyro-hyals. Half the natural size. 
Fig. 5. One of the stylo-hyals. Half the natural size. 

PLATE XXVII. 

Details of the osteology of Inia geoffrensis. All the figures half the natural size. 

Fig. 1. Anterior surfaces of the seven cervical vertebra;. 
Fig. 2. The thirteen ribs of the right side. 
Fig. 3. Side view of the sternum. 

a. Anterior process. 

b. Lateral process. 

c. Surface for attachment of cartilage of first rib. 

d. Sui-face for attachment of cartilage of second rib. 
Fig. 4. Internal surface of sternum. 

Fig. 5. External surface of stenium. 

PLATE XXVIII. 

Skull of adult Pontoporia hlainvillii. All the figures (except fig. 5) half tlie 
natural size. 

Fig. 1. Side view of cranium. 
Fig. 2. Side view of mandible. 
Fig. 3. Upper surface of cranium. 
Fig. 4. Inferior surface of mandible. 

Fig. 5. A maxillary tooth from the left side, the fourth from the posterior end of the 
series. Twice the natural size. 



Fi<j I 




vl 



-J Sm\t . lith . 



Fi^.3 




Fl^ 2. 



I z 



^ i i tS 7 g 



± 

4- 




INIA GEOFFR 






%^2M^dM^6. 



9 10 II 12 



'3 It /.J /^ I J ig 




C.P 



A& 




fg 



/J 




'° " '2 « /+ ;j (6 (7 "^ 



M &Nfra2ihart inj* 



'■■llj . ■/ 



jM^3JMm^m<^6'. 




MA-JIHantiorbimp . 



INIA GEOFFRENSIS 



v/^Z/nJ.. 



fl/J l 




FuJf- 3 




jrio A' 



Frn 5 





r<i 










MAitilAmartiii^ 



INIA GEOFFREl^SIS T 



?^' 




f 



Faj 3. 






J.Snat lith.. 



M *N Hajitiarl imp 



PONTOPORIA ■ BLAINVILLII , 2- 



[ 117 ] 



V. On a Baptorial Bird transmitted by Mr. Andersson /ro)« Bamara Land. 

By J. H. Gurnet, F.Z.S. 

Read November Uth, 1865. 

[Plate XXIX.] 

i HE raptorial bird now exhibited has been recently sent to me, with some other birds 
collected in Damara Land, by my friend Mr. Charles J. Andersson, to whose exertions 
we have already been frequently indebted for valuable contributions to oiu- knowledge 
of the ornithology of that part of South-western Africa. 

Mr. Andersson remarks, with reference to the present specimen, which was procured 
at Objimbmque, Damara Land, on the 10th of March last, " I have only obtamed this 
individual, a female, shot by my servant, who observed another, which was probably 
the male. I imagine I have once or twice observed this species near my place (Objim 
binque) just before dusk. I strongly suspect that it is a nocturnal or seminocturnal 
bird. I found only a Bat in the stomach of the specimen sent, of which the description 
and measurements are as follows : — 

" Irides bright lemon-yellow ; extremities of mandibles black ; basal parts and gape 
bluish lead-colour ; tarsi and toes bluish white ; claws bluish black. 
"Entu-e length 1 ft. 6^0 in.; length of wings when folded 1 ft. 1\\ in.; length of 
tarsus 2^in. ; length of middle toe 2-ji-2- in. ; length of tail 7^^^ in. ; length of 
bill from comer of gape to the tip of the mandible, straight, 1-^ in." 

To the above remarks of Mr. Andersson I have to add the following : — The colours 
of the plumage are dark brown mingled with pure white, the tint of the brown being 
very similar to that of a dark specimen of Buteo vulgaris ; a very few feathers of a still 
darker tinge, however, are apparent on the occiput and back. With the exception of a 
Ime of white above and below the eye, the feathers on the upper part of the head are 
brown : this colouring extends slightly below the gape, and also over the whole of the 
upper surface of the bii-d, mcluding the wings and tail ; but the basal parts of the 
feathers on the upper part of the head, the nape, and back are white, though this is not 
apparent except when a feather is displaced ; but this white becomes somewhat more 
visible where it is mingled with the brown, in the form of bars and spots, on all the 
feathers of the wings, both above and below, and includhig the upper and under wuig- 
coverts, as also on the upper and under tail-coverts ; the upper surface of the tail bears 
five transverse bars of a pale browia, which on the lower surface of the tail-feathers are 
white, and the tail is also very slightly tipped with dirty white. The throat is white, 
but is bisected for the upper three-fouiths of its length by a brown medial line, starting 



118 HIE. J. H. GUENEI ON A EAPTOEIAL BIRD. 

from the angle of the lower mandible, and extending for about 3 inches in a straight line 
towards the sternum. The feathers of the breast and sides are of a mingled brown and 
white, the latter predominating m the vicinity of the throat. 

The abdomen and inner sides of the thighs are white, the outer sides of the thighs 
are brown, the plumage of the thighs also extending over about one-fifth of the upper 
portion of the tarsus. 

The occipital feathers are lanceolate and slightly darker (some of them being also a little 
longer) than the feathers of the adjacent plumage, thus presenting an appearance similar 
to that which is frequently to be obsen'ed in adult specimens of Pernis cristatus. 

Of the primary feathers the third is the longest, the second next, then the fourth, the 
fifth, and the first successively ; the points of the primaries, when closed, reach to within 
three-quarters of an inch of the tip of the tail. 

The tail, which consists of twelve feathers, is very slightly forked, the centre feathers 
being the shortest, and the pair next to the outside pair the longest. 

The bill is singularly small for the size of the bkd; but the gape s very large, 
extending backwards till it reaches a point directly below the centre of the eye. Be- 
tween the eye and the upper mandible a row of small bristles takes its rise, pointing 
towards and extending over the upper edge of the mandible as far as the nostrils, which 
are uncovered and of a narrow oval form. As in the case of the American Vultures, 
there appears Jo be no septum between the nostrils. The ridge of the upper mandible 
is remarkably keel-shaped, and there is a very noticeable depression intervening between 
it and the cutting-edge of the mandible, which latter is entirely destitute of anything in 
the nature of a tooth, a notch, or a festoon. 

The tarsi and toes are slender in theu- character, and the scales with which they are 
covered are (with the exception of those covering the last joint of each toe) remarkably 
small. The middle toe, which is considerably elongated, has a promment roughened 
pad below each end of the last joint; the inner toe is similarly provided, but with the 
hinder pad thrown further back ; the outer toe has two of these appendages situated 
as those on the middle toe, and two others placed further back ; the hinder toe has one 
large pad only, seated immediately behind the root of the claw. 

The inner edge of the middle claw projects laterally, and appears to me to present a 
rudimentary pectination resembling that which is found in the Owls, a tribe to which 
the present species seems also to offer some resemblance in the form of its bill and the 
extent of its gape. 

P.S. I had intended proposing the name of Stringonyx anderssoni for this singular 
form, supposing it to be undescribed ; but, as has been pointed out by Mr. Bartlett 
since my paper was read*, it is no doubt identical with the Machcerhamphus alcinus of 
Westermanf , the type of which is in the Museum at Leyden. The present specimen 
has been added to the collection in the Nonvich Museum. 

* Proc. Zool. Soc. 1866, p. S2-i. t "Westenn. Bijd. t. d. Dierk. i. p. 29. 



^^a%^M. ?I/f^/29. 




Mk 



-V 




^. 



^ 






J-WaKdeletlitti, 



MACHEIRHAMPHUS ALCINUS, 



J«f (JrlfBiniiirlimj . 



[ 119 ] 



YI. On some Fossil Birds from the Zehhurj Cave, Malta. 
By W. K. Paeker, F.B.S., F.Z.S., d-c. 

Bead and received for publication Deo. 12th, 1865. 

[Plate XXX.] 

i* IVE years have elapsed since I first examined numerous bony remaius from the Zobbug 
Ca^■e, the " lamellu'ostral" nature of which was apparent to Dr. Falconer and myself 
from the first. I transmitted a list of them to that lamented palaeontologist for his and 
Captain Spratt's inspection, the latter gentleman having taken an active part in exhuming 
these treasures. A fresh examination of them has not changed my views as to their 
nature ; and I can now refer to figures of the most important, drawn side by side with 
theu' counterparts in the common Swan [Cygnus olor). The specimen of this species, the 
bones of which I have used for comparison, was a fine old female, 5 feet long from tlie 
tip of the beak to the end of the taU, not so large as the male, but a large bird notwith- 
standing. As half or more of the fossil bones evidently belonged to a Swan about one- 
thu'd larger than my specimen of the tame kiad, it must have been a noble creature, 
and its extinction is to be deplored as much as that of the Dinornis and the Dodo. 

Many of the bones belonged to a smaller kind than even the common mute species : 
it was about the size of a male Bewick's Swan, or the female of the Common Hooper 
(C. musicus); some, however, belonged to a bird as large as the male Hooper. There 
were also some bones of much smaller dimensions ; these appear to have belonged to a 
small Bemicle, such as the Bernicla brenta. 

On June the 10th, 1861 (the next summer), I received, through Professor Eupert 
Jones, another parcel of these bones ; and last autumn Mr. Busk put into my hands the 
hinder part of the skull of the largest kind, which, with a few thigh-bones of the same 
species, he had received from Dr. Leith Adams, of Malta. 

Altogether there are in my hands about three pounds' weight of fragments, amounting 
to several dozen in number. About one-fifth of these are indeterminable, on account of 
their worn and comminuted condition. The only bones quite perfect are phalanges ; and, 
with the exception of the lower part of a til)ia of the largest kind, which is 6^ inches 
long, the pieces are from 1 to 4 inches in length. Mr. Erxleben suggests that they are 
the remains of feasts held by foxes — a very good suggestion, as far as I can see. 

The specimens of bones belonging to the largest kind of Swan, which I propose to 
call Cygnus falconeri, in honour of the great palaeontologist whose loss we have so lately 
suffered, are as follows : — 

VOL. YI. — PAKT in. s 



120 



ME. PABKEE ON SOJIE FOSSIL BIEDS 



Skull (posterior fragment) 2 specimens. 

Ribs (upper part) 3 „ 

Ulna (middle) 1 specimen. 

Femur (various parts) 12 specimens. 

Tibia (various parts) -20 „ 

Tarso-metatarse (various parts) .... 20 „ 

Phalanges (perfect) 3 „ 

Of the smaller kind of Swan {Cygnus 7nusicus?) there axe — 

Cervical vertebra (2nd or 3rd) .... 2 „ 

Sternum (anterior part) 1 specimen. 

Scapula (proximal part) 7 specimens. 

Humeri (various parts) 18 „ 

Ulna (various parts) 7 „ 

Radius (various parts) 7 or 8 „ 

Metacarpus (various parts) 5 „ 

Phalanx (proximal, perfect) . . . . ^ 1 specimen. 

Phalanx (distal, perfect) 2 specimens. 

Sacrum (various) 3 

Femur (shaft-part) 2 

Tibia (various) 3 

Tarso-metatarse (various parts ) . . . . 4 
Phalanges (perfect) 5 



Of the small Goose-bones (Bernicla 
Coracoid (head) . . . 



■ ?) there are — 



Radius (distal and middle portions) 

Ulna (middle) 

Metacarpus (almost perfect) . . 
Femur (nearly perfect) . . . , 
Tibia (lower end) .... 



1 specimen. 

2 specimens. 
1 specimen. 
1 

1 „ 

1 



Some of these bones are of a beautiful ferruginous dark brown ; others are of a light 
colour, like the clay in which they were imbedded. 

Cygnus falconeri, Parker. Skull. 
Dr. Leith Adams's specimen of this part of the great Swan came to hand too late to 
ba figured ; I was able to make out that it belonged to a Swan nearly one-third larger 
tahn Cygnus olor, and to see the occipital plane, foramen, and condyle, as well as part 
of the parietal and temporal regions. With this specimen of the skull there were two or 
three fine " ossa femoris," which coiToborated the conclusion I came to as to the skull be- 



FEOM THE ZEBBUG CAVE, MALTA. 121 

longing to C. falconeri ; for the thigh-bones were the exact counterpart of those which 
had come earlier under my notice. 

I annex a Table of measurements of the bones of C. falconen, as compared with 
those of the large female C. olor : — 

Tvj-jji i-u • -u ^- ^^0''- ^- falconeri. 

Middle thoracic rib — .... . r ,. 

inches, lines. inches, hnes. 

a. Across the neck ....0 5...0 6 
I. Width of outer edge ... 3 ... 4 

Uka— 

Diameter of shaft 5...0 7 



Radius — 

Diameter of shaft 





3 


Femur — 

Across head and trochanter 
Width of middle of shaft . . 
Width across lower condyles . 


1 


1 


1 



Tibia — 

Fore-and-aft width of head 
Thickness of head .... 
Width of shaft 


1 




5 
5* 



Width across lower condyles , 11 

Tarso-metatarse — 

Extreme length 4 3 



Width across head . . . 
Width across shaft . . . 
Width across condyles . . 

Phalanx (proximal, middle) — 

Length 

Thickness of proximal end . 
Thickness of shaft .... 

Mr. Erxleben's figures show very faithfully the perfect agreement, in everything but 
size, between the great extinct Swan and Cynus olor. The largest bones of C. falconeri 
are not, however, displayed in the Plate, for this reason, that the most perfect bones 
for figuring were apparently those oi females; but there are bones still larger in tlio col- 
lection, most likely those of male birds. 

s2 



1 








5 


1 





2 


^ 





H 





3 



1 


5 





8 


1 


5i 


1 


9 


1 


2 





8 


1 


4i 


5 


3 


1 


2i 





6 


1 


4 


1 


H 





71 





4i 



122 ME. PABKEE OX SOBIE FOSSIL BIKDS 

The coarseness of these bones is well shown when the diploe is displayed (see PI. XXX. 
figs. 10, 11 & 12), and the walls of the tibial diaphyses are a line and a half (one-eighth 
of an inch) thick in the stoutest specimens. 

The Ribs. — The coarseness of the bones, and their great size as compared with those 
of the tame species, are well seen in the three fragments of ribs ; they are, altogether, 
one-fourth larger than their counterparts in the living Swans. 

The Ulna (PI. XXX. figs. 4 & 5). — The diameter of the uhia, as seen in fig 5, is as 7 
to 5 compared with fig. 5 a ; and the strength of the shaft is well sho\vn in fig. 4. The 
oblong quill-knobs, confluent by means of an elevated ridge, are well shown to be precisely 
alike in the extinct and the tame species. 

The Femur (PI. XXX. figs. 6, 7, 8, 9, 10 & 11).— These figures of the left femur, 
although not of the most massive specimens, give a good idea of the stoutness of this lost 
bu'd : its head, trochanter, shaft, and lower condyles are seen to be most exactly like those 
of the tame kind, save and except such intensification of the ridges and general surface- 
marking as is due to the origin and insertion of the muscles of a much mightier bird. 

Tibia (PI. XXX. figs. 12, 13, 14 & 15).— This, again, is evidently the bone of a fe- 
male, as there are considerably larger specimens, although not so perfect, in the collec- 
tion. Fig. 12 shows the strengtii of the shaft ; fig. 13 is an anterior view of the distal end 
of the right tibia, showing the broad tendon-bridge and groove, the space for attachment 
of the fibula, and two depressions in the space for the precalcaneal knob, which are but 
faint in the tame kind. Fig. 14 shows the extent of the lower condyle as seen laterally 
on the inside, and fig. 15 its division into an inner and outer lobe. 

Tarso-metatarse (PI. XXX. figs. 16, 17, 18 & 19).— The length of this right shank 
is seen to be gi'eater in proportion to its thickness than in the tame Swan; but their 
general agreement is most accurate. 

The low precalcaneal knob, the postcalcaneal ridges, grooves, and bridge, and the form 
and relative proportions of the lower bifid condyles are well seen. There is, however, a 
passage, shown in the head of the shank of the tame Swan (fig. 19 a), which does not ap- 
pear in fig. 19 : this Taisiaken foramen escaped me when examining the proof-plates ; it 
was made by me in the tame Swan's bone for the purpose of syringing out the marrow. 

The bony bridge uniting the outer and middle condyles (figs. 16, 17, 18) is seen to 
correspond beautifully in the two birds ; the pei"fection of the figui'es exonerates me 
from detailed description. 

Phalanges (PI. XXX. figs. 20, 21 & 22). — There are only three phalanges which 1 
can safely refer to the largest Swan ; but they are very remarkable, being quite unlike 
what we see in the species of Swans still living ; for fig. 20, as compared with fig. 20«, is 
seen to be full one-third thicker, and but little more than two-thirds the length. This 
is the case with the proximal phalanx of the great or middle toe ; and the other two are 
quite similar in shortness and robustness. 

If this shortness of the toes be remembered, along with the fact that the shank is 



FKOM THE ZEBBUG CAVE, MALTA. 123 

longer in proportion than in the recent kinds, we shall see that the great extinct Swan 
was rather generalized in character, being somewhat of a Goose, possessing, as he did, 
longer legs and shorter toes than the typical Swans. 

It would appear, however, that, like the gigantic Adjutant among the Storks, this bii'd 
had its wings of the full relative size : the immense ulna shows this (see PI. XXX. figs. 1, 
4&5). 

As tlie feet were shorter, it is probable that the extinct bird was not so expert at 
rowing as the smaller but more elegant kinds ; on land he may have shown better ; and 
perhaps he was altogether more terrestrial. 

It is worthy of remark, that the most generalized type of all the " LameUirostres," 
viz. the Palamedea — that in which the lamellae of the beak are arrested in their gi'owth, 
and which has no webs to connect the toes — has the digits longer even than the Swans. 
This bird, however, is not unrelated to the Grallatorial " Macrodactyli." 

Cygnus musicus(\). 

The most important bone of those belonging to the smaller Swan, which, as the fore- 
going list shows, are very numerous, is the front part of the sternum. This fine fi'ag- 
ment is well shown in PI. XXX. figs. 1, 2, 3 ; and, besides exhibiting the separated 
coracoid grooves, anterior part of keel, costal process, condyles for sternal ribs, ridge for 
middle pectoral, &c., is especially interesting because of the well-displayed anterior part 
of the cavity for the wind-pipe. Fig. 8 shows the smooth, rounded cavity; fig. 2 part of 
its left wall ; and fig. 1 the eminence caused by it on the midline of the sternum : the 
two rows of wind-passages are also well seen. 

This, then, is the sternum of one of the Wild Swans, perhaps the greater species 
{C. mxisicus), perhaps C. hewickii, or, it may be, some species nearly allied to these. At any 
rate it is interestmg to find that C. musicus is still to be found in lands bordering the 
Mediterranean, the Eev. H. B. Tristram having, in his last travels, received it from 
Solomon's Pool, near Jerusalem (see Proc. Zool. Soc, 1864, p. 453). 

The similarity of the bones m the species of Swans is so great that I feel it to be 
unnecessary to desci-ibe the rest of the bones of the smaller kind ; they are nearly all 
fragmentary, like those of C.falconeri, and the fragments are in the same good condition. 
The birds which owned these bones varied in size from that of a small female tame 
Swan to that of a medium-sized Black Swan ; yet the difference is scarcely more than 
varietal and sexual. There may have been more than two species buried in the Zebbug 
Cave ; but we lack positive evidence. 

The smallest " lamellirostral " bones are intermediate in size between those of the 
Wild Goose {Anser cinereus) and those of the Mallard {Anas boschas) ; so that they may 
have belonged to a small female Bernicle, such as the black-faced kind {Bernicla 
brenta). 

But, few as these are, they probably belonged to two kinds ; for the femur and tibia 



124 ME. PARKER ON SOME FOSSIL BIRDS FROM THE ZEBBUG CAVE, MALTA. 

arc relatively larger than the coracoid and metacarpus : these latter bones are not larger 
than those of a good-sized tame Duck (A. hoschas). 

DESCRIPTION OF PLATE XXX. 

(N.B. — ^The figures are all of the natural size.) 

Fig. 1. Anterior fragment of sternum of Cygnus musicus (l) ; upper view. 

Fig. 2. Anterior fragment of sternum of C. musiciis (?) ; side view. 

Fig. 3. Anterior fragment of sternum of C. mimcus (1) ; front view. 

Fig. 4. Ulna of C. falconeri ; end view of fragment. 

Fig. 5. Ulna of C. falconeri; side view of fragment. 

Fig. 5 a. Ulna of C. olor ; side view of fragment. 

Fig. 6. Femur (left) of C. falconeri ; front view. 

Fig. 7. Femur (left) of C. falconeri ; lower view. 

Fig. 8. Femur (left) of C. falconeri ; hinder riew. 

Fig. 9. Femur (left) of C. falconeri; upper -view. 

Figs. 6 a- 9 a. Femur (left) of C. olor. 

Figs. 10, 11. Femur of C. falconeri; fragments. 

Fig. 12. Tibia (right) of C. falconeri ; end view of fragment. 

Fig. 13. Tibia (right) of C. falconeri; front view of distal end. 

Fig. 14. Tibia (right) of C. falconeri; side view of distal end. 

Fig. 15. Tibia (right) of C. falconeri ; end riew of distal end. 

Fig. 13«. Tibia (right) of C. olor; front view of distal end. 

Fig. 16. Tarso-metatarse (right) of C. falconeri; hinder view. 

Fig. 16a. Tarso-metatarse (right) of C. olor; hinder view. 

Fig. 17. Tarso-metatarse (right) of C. falconeri ; lower view. 

Fig. 17a. Tarso-metatarse (right) of C. olor; lower view. 

Fig. 18. Tarso-metatarse (right) of C. falconeri; front view. 

Fig. 18a. Tarso-metatarse (right) of C. olor; front view. 

Fig. 19. Tarso-metatarse (right) of C. falconeri; upper view. 

Fig. 19a. Tarso-metatarse (right) of C. olor; upper view*. 

Fig. 20. Phalanx (proximal, of middle toe) of C. falconeri; upper view. 

Fig. 20 a. Phalanx (proximal, of middle toe) of C. olor; upper view. 

Fig. 21. Phalanx of C. /aZcowen ; side riew. 

Fig. 21a. Phalanx of C. olor; side view. 

Figs. 22, 23. Phalanx of C. falconeri ; end riews. 

Figs. 22a, 23a. Phalanx of C. olor; end views. 

* The circular hole in this view is of artificial ori'riii. 



j^^j.^jt.dc^'^M^MJO. 




J.Erxle"ben.cLel et kth. . 



1_3 CYGHUS MUSICUS ? 4_ 23 .C . FALCOEERI, P. 4a,.23a-_C OLOR. 



M&N Hinharl, Imp? 



[ 125 ] 



VII. SynojJsis of the species of recent Crocodilians or Emydosaurians, cAif/?y_/b?<wrfe<Z on 
the specimens in the British Museum and the Royal College of Surgeons. By Dr. 
John Edward Gray, F.R.S., V.P.Z.S., F.L.S., &c. 

Eead December 9th, 1862. ' ■■ *-'»-• ^^' ^^ ^ f - 

[Plates XXXI. to XXXIV.] 

1 HE distinction of the species of Crocodiles has hitherto been one of the difficult 
problems in systematic zoology ; and therefore I believe that it may be of some slight 
use to lay before the Society the result of my examination of the very large collection 
of Crocodiles, of all ages and from various localities, which are contained in the British 
Museum. Knowing the difficulty that suiTounds the subject, I have made great 
exertions to obtain specimens from different countries ; and the examination of these 
specimens has shown that the characters of the species, when allowance is made for the 
changes that take place in the growth of the animal, are quite as permanent as in any 
other group of Reptiles, and not more difficult to define. 

An outline of the synopsis of the Crocodilidse or Alligatoridee was published in the 
'Annals and Magazine of Natural History' for 1861 (3rd series, vol. \\\\.). Since that period 
I have examined the additional specimens which have been received in the British Museum, 
and also those in other collections, especially the skulls in the Museum of the Iloyal 
College of Surgeons, the specimens in. the two museums at Liverpool, and in other local 
collections within my reach. Among the specimens recently received by the British 
Museum are some typical skulls from the Dutch possessions in the East, obtained 
from Leyden, which enable me to determine with certainty the species described by the 
Dutch zoologists. 

The determination of the species of the Crocodilians has always been attended with 
considerable uncertainty ; and if we may judge by the manner in which the specimens and 
the skulls of them are named in Museums, or sent about by the more scientific dealers, 
it would appear that as yet they are not properly understood. 

I do not mean as to the precise limit of a species — that is to say, whether the specimens 
from different districts of the same zoological or geographical province are mere local 
varieties of the same species, or are distinct ; for that is a question which I admit must, 
with the materials at our command, for the present remain unsolved and open to discus- 
sion. But it is not unusual to find most distinct species confused under the same name, 
and specimens of the same species, only different in age, separated under two or more 
names. 

In this paper I have endeavoured to condense into a short synopsis the principal 
leading characters, especially those furnished by the examination of the skull and the 

VOL. VI. PART IV. T 



126 DE. J. E. GHAT'S SYNOPSIS OF THE SPECIES 

nuchal and dorsal plates, by which the different species of Crocodiles and Alligators may 
be most easily determined. 

My object in this paper is to furnish the zoologist with the best character to distin- 
guish the different species of Crocodile and AUigator, without any pretence of giving 
an account of the comparative anatomy or osteology of the species. I make this state- 
ment, as confusion arises in the student's mind between the object of the studies of the 
two branches of the science, both equally important ; but the one ought to be based on 
the examination and comparison of the largest possible number of specimens and 
species, while the most important papers on comparative anatomy are often those that - 
arise from the examination of a single example of the animal. 

I am well aware that there is a prejudice against such short papers, and that they 
incur the reproach of certain continental and native naturalists ; but after considering 
their objection and their practice, I am still of the opinion that papers of the kind are 
far more usefid to the working natui-alist than the long descriptions of species which it 
is the custom of these naturalists to prepare, when their descriptions, instead of merely 
presenting the peculiar character of the species under consideration, give in full detail 
under each species (so as to hide in a bushel of words the characters which you are 
looking for) the character of the genus, or even often of the family or order to which 
the species belongs. Macleay well observes, " The modem art of describing is too long, 
often insufferably long, while human life remains as short as ever " (Illust. Zool. S. 
Afi-ica, p. 5-i). 

I know by experience that synoptical papers take far more mental and bodily labour to 
prepare than the description of a single specimen, often taken at haphazard and regarded 
as the type of a species because it presents some striking peculiarities of appearance. 

This paper, short as it is, is the result of the examination and repeated reexamination, 
at different periods, of more than two hundred specimens of Crocodiles, — a series of the 
most characteristic specimens of each species having been laid out so that they could be 
viewed and studied together and at leisure, and their peculiarities and likenesses noted 
down. 

If all the notes made during these comparisons were printed, as is the custom with 
many naturalists, they would fill many pages, and thus make a long paper. Many 
papers and books are estimated by their size, rather than by the extent of labour that 
has been bestowed upon them ; while the results of much labour and careful study, 
condensed into a few pages, are often spoken of by critics, who never undertook such 
researches, or who dislike the labour of condensing their observations into systematic 
order, as merely the short notes of a hasty examination : at least that is the way in 
which some papers, which were the results of equally extensive examinations, have been 
regarded by naturalists who should have known better. 

I may further observe that, even after so much study, when new specimens have been 
accumulated and with additional experience, one frequently finds peculiarities overlooked 



or RECENT CKOCODILIANS. • 127 

and facts requiring verification, when the old and the newly acquired specimens are 
submitted to a reexamination and study. It is this experience that makes me inclined to 
place less reliance than other naturalists upon essays prepared by persons who come and 
look at a series of specimens for the first time, and describe them oflFhand. Yet such 
works are often regarded as of authority, very often on account of their length, or the 
beautiful manner in which they are printed or illustrated. 

The references to the catalogue of the osteological specimens in the College of 
Surgeons are based on the examination of the specimens in that collection ; and I have 
to thank the Council of the College for their permission to examine them, and Mr. 
Flower, the energetic Curator of the collection, for his kindness and assistance in 
determining them. 

If any evidence were required of the difficulties of determining the species of this 
family, I need only refer to the nomenclatui'e of the skull in the catalogue above 
referred to, which was prepared by the late Curator of the collection. Professor Owen. 

In this collection, for example, I found what I consider to be three distinct species 
in one case, and two distinct species in another, confounded under the same name ; and 
on the other hand, I found what I regard as skulls of the same species inserted under 
three different names. 

The skull of a Crocodile which is found in the internal rivers of India, is named 
Crocodilus rhombifer, Cuvier (which is an American species), though the specimen in the 
College ^Museum was received from Bengal. 

I do not by any means regard my determination of these skulls as infallible ; but I 
have taken every care to make it correct by repeated examination. I first arranged 
the skulls as they appeared to be alike, according to the characters here assigned to 
them, without paying any attention to the names given, placing them in order according 
as the size showed the change in the growth; and Mr. Flower, Mr. Gerrard, and some 
other zoologists who are used to the examination of bones, agree with me in m) 
determination, and were much interested in observing how gradually the skulls of 
different ages glided into each other ^ 

I must observe, if there is this difference of opinion in the determination of skulls of 
recent Crocodiles, where the series of skulls for different-aged animals can be compared, 
and where the skulls are in a perfect state, how much more difficult it must be to have 
confidence in the determination of the skull of the fossil, or some fossil species where 
the skulls are generally more or less imperfect, and perhaps only single specimens 
(often very imperfect specimens) have been examined ! 

' The following is the result of my examinations of the specimens of Crocodiles in the Museum of the College 
of Surgeons (the numbers refer to the numbers in the catalogue) : — 
682—707. Guvialis (jawjetkus=Guvi(tlis. 

710. C'rocodihts catuphractxis^Mecistops mtaphractvs, the t^'pe specimen. 

711, 712, 714, 716. Crocodilus acidus=Moliahi ame ricaim , from America. 



128 DR. J. B. GEAT'S SYNOPSIS OF THE SPECIES 

The chief difficulty in distinguishing tlie species has originated from the very great 
change of forms that takes place in the shape and proportions of the head of the animal 
in its different stages of growth ; but the changes seem nearly similar in all the species, 
and therefore when once observed they can be easily allowed for. The difference may 
be divided into thi'ee stages, exemplified in the young, the nearly full-grown, and the 
adult or aged specimens. The head and beak of the young are generally depressed, with 
more or less distinctly marked symmetrical ridge and depressions ; and these characters 
are gradually modified until the animal assumes its nearly full size, — the skull becoming 
thicker and more solid, but yet retaining most of the characters that distinguish its 
young state. After this period, as the animal increases in age, the skull becomes more 
and more convex and swollen and heavy, and assumes a very different external form. 

It is to be observed that in all these changes in the external form of the skull, the 
bones themselves of which it is composed preserve their general form and relation to 
each other ; and the sutures between these bones appear to me to offer some of the best 
characters to separate the species into groups. In many instances, when I have been in 
doubt, the sight of the intermaxillary suture has at once solved the difficulty, which has 
been verified by the examination of the locality of the specimen. 

These changes in the form of the head have been among the causes that liave made the 
study of the species of Crocodiles so difficult. If this is the case with the recent species, 
how much more caution is requisite to determine the fossil remains of the animal ! 
Cuvier set a very good example in that respect : he commenced the study of each group 
of animals with an examination of the osteology and external characters of the living 
species, and then applied the knowledge he thus acquired, to the distinction of the fossil 
remains ; but now we often find palaeontologists, as they call themselves, neglecting, or, 
at most, only taking tiie outline of the osteological and zoological characters of the living 
species at second hand, and describing the fossil, and often forming genera and species 
on a small fragment, thus encumbering the science with a multitude of names. 

At one time I proposed to give accurate measurements of the different parts of the 

713. Crocodihts acutus^Oopholis porosus of India. 
71.'>. Crocodilus acutus= Crocwlilus vulgaris of Africa. 

717. CrocoiUlus vuhjaris, much distorted. 

718. CrocodUns vulgaris=Bomhifrons, perhaps B. simnemis. 
719-724, 727, 728. Crocodfbis hiporcatvis^=Oopholis porosus. 

725. Crocodiliis biponatiis^Croeodihis vulgtiris. 

726. Crocodilus biporcati(S = Bontbifrons indicus. 

750, 751. Crocodilus rhombifer, from Hvngal^^Bombifrons imlicus. 
752. Crocodilus palustris'!=Bombifrons indicus. 
760-762. Alligator lurius— Alligator mississippieiisis. 
764. Alligator niger=.Triciirc' nigra. 
' Dr. J. E. Gray " On the Change of Form of the Heads of Crocodiles," Transactions of the Sections in ' Report 
of tlie British Association of Science,' Cambridge, 1862, p. 109. 



OF RECENT CEOCODILIANS. 129 

skull of each of the specimens of the different species in the British Museum Collection ; 
but I am satisfied that the importance of such tables of measurement is over-estimated : 
no doubt it has a very imposing appearance ; but a good figure is more useful than any 
amount of measurement. Every species has its normal measurements ; but these are 
liable to vary in the different individuals; and any difference sufficient to show a 
distinction of species is easily appreciated by the eye, as it must alter the general 
proportions of the different parts of the head. 

It has been suggested that I ought to give the description of each separate bone of 
which the skull is composed. This may be of use to the student of comparative anatomy, 
but is not of so much importance to the zoologist ; for though each bone has a normal 
form in each species of Crocodile, yet they are each liable to considerable variation 
within certain limits in the different individuals of the species. 

The bones of the different genera have been described in several works on osteology, 
and they are well figured by De Blainville and others. 

De Blainville, in his ' Osteographie,' devotes five folio plates to the osteology and 
dentition of recent Crocodiles, giving details of Crocodilus hqwrcatus, C. lucius, C. 
vulgaris, C. schlegelii, C. longirostris, C. rhomhifer, and C. sderops. These plates 
were prepared to accompany an essay that M. de Blainville was preparing for the 
' Memoires de I'Academie des vSciences de France ' when he died. 

Professor Carl Bernhard Briihl, of the Universities of Cracow and Pesth, has published 
twenty quarto etchings of the skeletons of Crocodiles and Alligators, giving details of 
three or four species. The plates are exceedingly accurate, and full of details, being 
drawn and etched by the Professor and his wife direct from the specimens. They were 
published at Vienna in 1862. There is a continuation of the work, containing three 
additional plates, published in 1865, principally devoted to the canals of the ear-bone. 

I must here refer to a paper by Professor Huxley, entitled " On the Dermal Armour of 
oiJacare and Caiman, with notes on the Specific and Generic Characters of recent Croco- 
dilia," Journ. Proc. Linn. Soc. Zool. iv. p. 1. As this paper contains an excellent account 
of the ostcological differences between the different genera of Crocodilia, 1 have not 
considered it desirable to repeat them here, more especially as they were chiefly drawn 
up from specimens in the British Museum. 

Order EMYDOSAURI (Emydosaurians). 

Emydosauri, Blainville, Gray, Ann. Phil.x.l95, 1835; Cat. Tortoises & Crocodiles Brit. Mus. 38,1844. 
Crocodilia, Huxley, Journ. Proc. Linn. Soc. Zool. iv. p. 1. 

The Emydosaurians or Crocodilians may be divided into three families : — 
A. TJie cervical and dorsal plates forming one dorsal shield. 

I. Gavialid^. The large front teeth and the canines in the lower jaw fit into notches 
in the margin of the upper jaw. 



130 



DE. J. E. GEAT'S SYNOPSIS OF THE SPECIES 



B. The cervical shield forms a small group, ivhich is separate from the dorsal shield. 
II. Ceocodilid^. The canines fit into notches in the upper jaw, and the large front 

teeth fit into pits or perforations in the front of the upper jaw. 
III. ALLiGATOKiDiE. The large front teeth and the canines fit into pits or perforations 
in the edge of the upper jaw. 

The large front teeth of the Garials fit into a notch in the front of the upper jaw, 
and the canines into a notch also. In the Crocodiles the canines fit into a notch, as in 
the Garials, but the large front teeth fit into a pit or perforation in the front of the 
upper jaw; and in the Alligators both the canines and the large front teeth fit into pits 
or perforations in the edge of the upper jaw. 

The geographical distribution of the genera may be thus exhibited : — 



Africa. 


Asia asb ArsxRALAsiA. 
Fam. GfwialideB. 
Gavialis. 
Tomistonia. 

Fam. Crocodilidie. 


America. 


Crocodilus. 


Oopholis. 






Bombifrons. 


Palinia. 


Halcrosia. 




Molinia. 


Mecistops. 




Fam. Alligatoridis. 
Alligator. 
Caiman. 
J acare. 



In Africa there are three species of Crocodiles. They seem all to have been known 
to Adanson. They are, 1. The common Crocodile (called the Olive Crocodile by 
Adanson), Crocodilus vulgaris, which is spread over the whole of Africa, from north to 
south and from east to west ; 2. The Black Crocodile of Adanson {Halcrosia nigra) ; 
and, 3. The False Gavial of Adanson, the Mecistops catapihractus. The two latter are 
confined to the rivers on the west coast of Africa. 

In India' there are also three species of Crocodiles: — 1. The Oopholis j)orosus (or 
Crocodilus biporcatus of Cuvier), which is found only in the estuaries at the mouths of 
the large rivers ; 2. The Muggar" {Bomhifrons indicus) ; and 3. The Garial (or Ghurrial), 

' See Dr. J. E. Gray " On the Cjocodiles of India and Africa," Transactions of the Sections in ' Report of the 
British Association of Science,' Cambridge, 1862, p. 107. 

- Dr. Falconer says, the proper name of the CrocodOe is Coomheer. The Eapacious Shark is called the Mii(/gtir ; 
iinil by reflection this name is also sometimes given to the Crocodile, because it is a rapacious animal. 



OF EECEXT CROCODILIANS. 131 

which is confined to the rivers in the interior of the country. The Coombeer or Muggar 
ascends the rivers to the mountains, where the water is often frozen. The Ghurrial, on 
the contrary, is confined to the lower level, where the climate is warm. 

In stating that there are three species of Crocodiles in India, I only intend to state 
there are three distinct forms ; for I will not undertake to say for certain that the 
Muggar of Ceylon, of Siam, and of India are not distinct species. 

Mr. Blyth observes, " Both the Gangetic species of Crocodiles have been received by 
the Asiatic Society, Calcutta, from Java. The Crocodiles are known to abound in 
Timor, from which island they may well have passed to iiustralia. Governor Grey met 
with them in the north-west." — Blyth, Eep. Austral. Vert, in Mus. A. S. C. 

If by " both the Gangetic species of Crocodile " Mr. Blyth means the estuarine Croco- 
dile (OojifioHs jwrosus) and the Coombeer or Muggar {Bomhifrons milieus), no example 
of the latter animals from either Java, Timor, or Australia has occurred to me, and the 
animal figured as Crocodilus raninus by Dr. Salomon Muller is certainly Oo])holts ^orosus ; 
and there is in the British Museum a fine adult skull of that species sent by the Leyden 
Museum from Java. 

The observations of MM. Dumeril and Bibron (Erp. Gen. 25, 47), that Crocodiles are 
not. found in Australia, and that the American Crocodiles are confined to the islands of 
that continent, are no longer consistent with facts ; indeed, long before the publication 
of their work, various travellers had recorded the occui'rence of Crocodiles on the north 
coast of Australia. 

The estuarine Oopholis jJOrosus was observed by Governor Grey on the north-west 
coast of Australia. There is in the British Museum a skull of the species sent thence, 
and also a full-grown specimen which was killed and preserved in that country. 

The Island of Borneo is inhabited by a false Garial, named Tomistoma schlegelii. 
I am not aware that it has been found in any of the other islands of the archipelago. 
It is intermediate in character between the true Garial and the Crocodiles. 

The Crocodiles and Alligators are widely distributed in America. There are four 
American Crocodiles, and nine Alligators. One of the Crocodiles, Palinia rhomhifer, 
is peculiar to the island of Cuba. The other species of Crocodiles and the Alligators 
are found on the mainland. The Alligator mississippensis is found far north, where the 
waters are often frozen ; all the other Alligators and American Crocodiles are confined 
to the tropical and subtropical parts of the continent. Molinia americana is found in 
Cuba and St. Domingo, as well as in the rivers of the east and west side of the conti- 
nent, showing the incoiTectness of the assertion of MM. Dumeril and Bibron that the 
Crocodiles of America are confined to the islands of that continent (Erp. Gen. 25, 47}'. 

• ' In the ' Gentleman's Magazine ' for August 1866 appears an article, entitled " Notes on a Yoi^ng Crocodile 
found in a Farmyard at Over Norton, Oxfordshire," by George R. Wright, F.S.A. Mr. Wright observed the 
specimen in a case of birds and animals, preserved by Mr. WiUiam PhilUps, who said that it was foimd lying 
dead in a gutter m his farmyard, evidently but lately killed ; its bowels protruded from a wound in the beUy. 



132 DR. J. E. GRAY'S SYNOPSIS OF THE SPECIES 

FamUyl. GAVIALIDiE. 

The cervical and dorsal plates formed into a single continuous shield. Teeth nearly 
of uniform size, all fitting into notches on the edge of the upper jaw. The fi-ont large 
teeth fitting into a notch m the front, the canines into a notch on the sides of the 
fi'ont of the upper jaw. The jaws elongate, slender. 

Crocodilida (part.). Gray, Ann. Philos. x. 195, 1825. 
Crocodilida § *, Gray, Cat. Tortoises & Crocod. B.M. 36. 
Gavialida, Huxley, Joiiru. Proc. Linn. Soc. Zool. iv. p. 16, 1859. 



Synopsis of Genera. 
Gavialis. Beak elongate, linear, end swollen. The lateral teeth oblique, not received 

into pits. 
ToiiiSTOMA. Beak conical, thick at the back, the lateral teeth erect, received into pits 

between the teeth. 

1. Gavialis. 

OT 07 oa f>A 

Beak of skull linear, end dilated from the enlarged nostrils. Teeth 2o-^2b i °^' 26—36 - 
The mandibular symphysis extends to the twenty-third or twenty-fourth tooth. Most 
of the lateral teeth of both jaws are directed obliquely, and not received into inter- 
dental pits. The front margin of the orbit is much raised. 

Gavial, Oppel. Le gavial, Cuvier. 

Gavialis, Merrem, Gray, Ann. PhU. x. 195, 1825 ; Cat. Tortoises, &c., B. M. 36, 57, 1844. Geoff. 

Mem. Mus. xii. Huxley, Proc. Linn. Soc. Zool. iv. p. 20, 1859. 
Gavialia, Fleming, Phil. Zool. 
Ramphostoma, Wagler, Syst. Amph. 441. Rhamphognatkus, Yogt, Zool. Brief, ii. 289. 

1. Gavialis gangeticus. (The Garial or Nakoo.) 

Narrow-beaked Crocodile, Edw. Phil. Trans, xlix. 639, t. 19. 

Le gavial, Lacep. Q. O. 1235, t. 15. Faugas, Mont. S. P. 235, t. 8. f. 46, 47. 

Lacerta gangetica, Gmelin, S.N. i. 1057. Shaw, Zool. iii. 197, t. CO. 

The men said it ran out of the stack of wood, they killed it, hut they could easily get him another ; he offered 
n ifuinca for auother specimen, dead or aUve ; but the reward was never claimed. 

An account of the discovery appeared in the ' Field Newspaper ' for 1861 or 1862 ; aud another, with a 
figure of the specimen, was published in Hardwicke's 'Science Gossip,' Jan. 1, 1867, p. 7, figs. 1 it 2. Dr. 
Yesalius Pettigrew and Jlr. Frank Buckland thought it was a very young Crocodile that had escaped from some 
travelling show. I should suspect that it was much more likely to be a just-hatched specimen that had 
been preserved in spirit and thrown away. The wound in the belly was probably the umbih'cim. The figure 
shows too long and slender a beak fcr a young specimen of any Crocodile I have seen. 



OF EECEJST CKOCODILIANS. 133 

Crococlilus longirostris, Schneid. Amph. 160. Daudin, Kept. 4-293. Blainv. Osteog. Crocod. t. 2. f. 4, 

t. 3. f. 6, t. 4. f. C, t. 5. f. 5. 
Crocodilus ardirostris, Daiid. Rept. ii. 393. 
Crococlilus tenuirostris, Cuvier, Ann. Mus. x. t. 1. Tiedem. Amph. 1. 15. Wagler, Syst. t. 7. f. 111. 

Merrem, Tent. 38. 
Gavialis ganyeticus, Geoff. Mem. Mus. sii. Gray, Syn. Rept. 36; Cat. Tortoises &c. B. M. 57. 

Dnm. & Bib. Erp. Gen. iii. 135, t. 26. f. 2. Huxley, Journ. Proc. Linn. Soc. Zool. iv. 

p. 20, 1859. Briihl, Skelet. Krokod. t. 8, 9, 10, 11, & 17. 
Crocodilus gangeticus, Tied. Oppel, & Libosch., Naturg. Amph. 81, t. 14. Geoif. Mem. Mus. H. N. 

xii. 118. 
Gavialis longirostris, Merrem, Amph. 37. 

Gavialis tenuirostris, Merrem, Amph. 38. Guerin, Icon. R. Anim. t. 2. f. 3. 
Ramphostoma teimirostre, Wagler, Nat. Syst. Amph. 141, t. 8. f. 3. 
Le gavial, Lacep. H. N. Q. Ovip. i. 235, t. 15. 
Gavial, Owen, Monogr. Fossil Reptilia of the London Clay, t. 11. 1849 (skeleton). 

JIab. Indian rivers. Bengal, Nepal, Malabar. 

2. TOMISTOMA. 

Beak of the head conical, thick at the base. Teeth fgnfg. The mandibular sym- 
physis extends to the fifteenth tooth ; the hinder tooth of the upper jaw, and most of 
those of the lower jaw received into interdental pits. Premaxillary hardly expanded, 
orbital mai'gins not raised. 

Gavialis, sp., Miiller ; Owen. 

Tomistoma, S. Miiller, Wiegm. Arch. 1846, i. 122. 

Rhynchosuchus, Huxley, Journ. Proc. Linn. Soc. Zool. iv. p. 16, 1859. 

The upper edge of the intermaxillary bone extends back as far as the second canine 
tooth ; and in this character it differs from the skull of the slender-nose Crocodiles, as 
Croc, gravesii and 3Iecisto]}s cataphractus. 

Dr. Falconer, when describing the skull of Crocodilus cataphr actus, in Ann. and Mag. 
Nat. Hist. 1866, xviii. 362, observes, " Crocodilus schlegelii constitutes the passage 
from the true Crocodiles into the Gavials," and he shows how the skull agrees with the 
Crocodiles' in the position of the nasal bones. 

Professor Owen, in the first ' Essay on the fossil reptiles of the London Clay,' Crocodiles, 
p. 15, observes, " The Bomean species, Crocodilus schlegelii, was in fact originally de- 
scribed as a new species of Gavial ; but tlie nasal bones, as in the fossil from Sheppey 
(figured in t. 2. f. 5), extend to the hinder border of the external nostrils." This does 
not agree with oui- skull, nor with the figures of the skull in Blainville's ' Osteographie.' 
See also Huxley, Journ. Proc. Linn. Soc. Zool. iv. p. 18. 



VOL. VI. — PAET IV. 



134 DR. J. E. GEAT'S SYNOPSIS OF THE SPECIES 

1. ToMiSTOMA sc'HLEGELii. (Bornean Gavial.) 

Crocodilus gavialis schlcgeUi, Miillcr, Naturgesch. Ost. Ind. t. 123. f. 1-5. 

Crucodilus schleyelii, Blainv. Osteog. Crocod. t. 2. f. 3; t. 5. f. 4. Briihl, Skelet. Krok. t. 8. f. G. 

Owen, Fossils of the London Clay, p. 15. 
Rhynchosuchus schleyelii, Huxley, Proc. Linn. Sec. iv. (1859) p. 17; Ann. & Mag. Nat. Hist. 1859. 
Mecistopsjournei, Gray, Cat. Tortoises &c. B. i\I. 38, not synonyma. 

Hah. Australasia, Borneo (Muller, Brit. Mus.). 

The two figures of the skull in Mi'tUer and Schlegel, t. 3. f. 1 and 2, show the difference 
that occurs in the form of the skull of the same species. 

In the British Museum tliere is a young sjjecimen in spirits, and an adult skull 
received from the Leyden Collection, and a very fine adult skull from Borneo, obtained 
from Mr. Mitten. 

Family IL CEOCODILID^. 

The cervical plates forming a distinct shield, separate from the doi'sal shield. Teeth 
strong, very unequal in size, hinder larger. The 9th upper and the 11th lower teeth 
larger, like canines, the large teeth of the lower fitting into pits or perforations, and 
the canines fitting into notches on the edge of the upper jaws. Nose of both sexes 
simple. 

Tlic upperside of the intermaxillary is slightly expanded behind, and its hinder end 
is divided, and separated into two parts by the front end of the nasal bone. 

Crocodilida §**, Gray, Cat. Tortoises &c. B.M. 36, 1844. 
Crocodilida, Huxley, Proc. Linn. Soc. Zool. iv. 5. 
Crocodilus, Cmder; Gray, Aim. Phil. 1825, x. 195. 
Champse, Merrem, Tent. 

Professor Huxley divides this family into two genera, Crocodilus and Mecistops. See 
Journ. Proc. Linn. Soc. Zool. iv. 6. 

The Crocodiles when they are first hatched have a very short beak to the head. This 
is even the case with the long-beaked Meci.ftops cataphractus, which in its very young 
state is hardly to be distinguished in the form of its beak from the young of the com- 
mon Crocodile, Crocodilus vulgaris. 

As the young obtain strength the beak developes itself more or less rapidly according 
to the species, until its normal character is attained. 

The head seems to continue of nearly the same form, merely increasing in size, for 
some time, perhaps years ; for we know little of the duration of the life of the Crocodiles ; 
and they are probably long-lived animals. As they reach maturity, and as old age creeps 
on, the skull thickens considerably, and the jaws dilate and thicken on the sides. The 
growth of tlie teeth, which are produced in succession, and greatly enlarge in diameter, 
and the enlargement of the jaws proceed pari passu : the latter is also influenced by the 
development of these teeth and the larger alveoli required to support them. 



OF RECENT CROCODILIAKS. 135 

The head of the Crocodile first mcreases in length compared with its width, and then, 
having arrived at a certain form, increases in width, thickness, and solidity. 

The same change takes place in the head and skull of the Bornean Garial, Tomi- 
stomn schlegelii, as is found in Mtiller and Schlegel's figures of the half-grown and 
adult skulls in their work. 

It is to be observed that each of the Crocodiles of India and Africa (and it may also 
be the case with those of America) seems to present two varieties — one with a broad and 
the other with a narrower face ; this variation occurring in each species appears to me 
to show that it is more probably a local, or perhaps even sexual variation than a specific 
distinction. 

If it were a sexual distinction, it might be soon settled by observers in the country 
where they abound ; but the sex of the skin and the skull sent to Europe is rarely, if 
ever, marked on the specimens. 

The broad-nosed variety is much more abundant in the Museum than the narrow-nosed 
one.; and this is against the form of the face being a sexual distinction, as one would 
suppose that they would be nearly equal in number, unless the narrow-nosed specimens 
are the males and they are more wary and not so frequently cauglit. 

Some naturalists might be inclined to regard them as distinct species ; but in the 
Museum series, large as it is, we have not sufficient materials to decide the question 
with any confidence. Perhaps, if the skulls of specimens from each locality could be 
compared, other characters might be found ; but this must be left for my successors in 
tliis field of research. 

In the short-nosed species the upperside of the intermaxillary bones is short, and the 
nasal bones are produced between their edges to the edge of the nostril ; and in the genus 
Halcrosia tliey are produced beyond it, and form a bony septum between the nostrils. 
In the long arid slender-nosed species the intermaxillary bones are rather produced 
behind and the nasal bone does not reach the edge as does the long nostril in the genus 
Mecistops ; they are considerably short of them ; but still the nasal bones come between 
the hinder ends of the intermaxillaries, and this character at once separates the skull of 
that genus from the two genera of Garials which have short nasal bones. 

The skulls of Crocodiles may be separated thus : — 

1. Nasal bone produced, and separating the nostril into two parts. Halcrosia. 

2. Nasal bone produced, and dividing the edges of the nostril. Oopholis, Crocodilu.s, 
Molinia (americana), Bombifrons, Palinia. 

3. Nasal bone not reaching the nostril. Molinia {intermedia), Mecistops. 

The intermaxillary bone in Bomlifrons and Falinia is short and truncated behind. 
In Halcrosia it is rather produced behind, the straight sides converging to a point. In 
all the other genera it is produced behind, with the hinder edges converging on the sides 
and truncated at the end. 

r2 



136 DB. J. B. GEAT'S SYNOPSIS OF TUB SPBCIBS 

The palatal bone in all the genera is truncated or rounded in front, except in 
3Iecisto2)s, where it is narrow, short, and acute in front. 

The skulls of the genera Bomhifrom, Oojjholis, and Molinia are easily distinguished in 
the young state, — the face of Oopholis being much longer and narrower than that of 
Bomhifrons, and that of Molinia is longer and narrower than that of Oojjholis. The 
measurements following are for three skulls which appear to be from animals nearly of the 
state of growth, same in inches and lines : — 

Bomhifro)is. Oojjholis. Molinia. 

in. lines. in, lines. in. lines. 

length of the skull, entire .-t 8 5 8 6 9 

Length of face to front of orbit 2 8 3 6 4 4 

Length of forehead to front of orbit 2 2 1 2 4 

Length of palate from condyle to front end o^lo-ii '^ a SIO 

palatine J 

Length of middle suture of maxilla 1 2 1 1| 1 7 

Length of middle suture of iutermaxiUa 9 1 3 1 6 

Width at occiput 2 6 2 5 2 10^ 

Width at hinder contraction of beak 1 6 1 4 1 4^^ 

Width at notch 9 9 9 

The dorsal scales present considerable variations in diflferent specimens from the same 
locality ; but, allowing for such variations, the genera Uiay be arranged thus : — 

1. The dorsal scales nearly uniformly keeled, in four or six longitudinal series; the 
outer series ovate-elongate. Oopholis. 

2. The dorsal scales nearly uniformly keeled, quadrilateral, as broad as long. Croco- 
dilus, Palinia, Molinia, and ilecistops. 

3. The dorsal scales quadrilateral, as broad as long; the vertebral series scarcely 
keeled, the lateral series irregular and keeled. IMcrosia and Molinia. 

The eyelid of the genus Halcrosia is thickened with hard bony plates, as in 
some of the Alligators, with which it also agrees in the external form of the head and 
the disposition of the nuchal shield. In all the other genera it is thin and mem- 
branaceous. 

Synopsis of Genera. 
I. Cervical disk rhombic, separated from the dorsal shield. Normal Crocodiles. 

A. Nuchal scutella none. Dorsal plates ovate-elongate, in four or six longitudinal 

series. Estuarine Crocodiles. 

1. Oopholis. Asia and North Australia. 

B. Nuchal plates four, in a transverse series. Dorsal plates as broad as long, 

square. FluviatUe Crocodiles. 



OF RECENT CROCODILIANS. 137 

a. Intermasillary bone truncated behind, with a nearly straight hinder 

edge. Face broad, oblong. 

2. BoMBiFKONS. Toes webbed. Legs distinctly fringed. Asia. 

3. Palinia. Toes short, free. Legs with only an indistinct fringe- 

America. 

b. IntermaxiUary bone elongate, produced, and truncated behind; 

sutures slojnng backwards and converging, then transverse or 
sinuous. Toes webbed. Legs fringed. 

4. Crocodilus. Face oblong, without any ridge from front of orbit, 

forehead flat. Africa. 

5. MoLiNiA. Face elongate, forehead convex, smooth, without any ridge 

fi-om orbits. America. 

II. Cervical disk strongly keeled on each side, and nearly continuous with the dorsal 
shield. Aberrant Crocodiles. 

* Face broad, nasal bone produced into the nostrils. AUigatoroid Crocodiles. 

6. Halcrosia. Africa. 

** Face very long, slender, nasal bones not reaching the nostrils. Ga-\aaloid 
Crocodiles. 

7. Mecistops. Africa. 

1. The nape nnth a rhombic disk formed of six plates, ivhich is well separated 
from the dorsal shield. Normal Crocodiles. 

A. Nuchal scutella none. Dorsal scales in four or six longitudinal series; the outer 
series ovate-elongate. Toes webbed. Legs fringed. The intermaxillary bon^ 
produced, truncated, and converging on the sides. Estuarine or brackish-water 
Crocodiles. 

1. OOPHOLIS. 

Face oblong ; orbits with an elongated, longitudinal, more or less sinuous ridge in 
front. Nuchal scutella none, or rudimentary. Cervical disk rhombic, of six plates. 
Dorsal plates uniformly keeled, in four or six longitudinal series ; the vertebral series 
with straight internal edges, the outer ovate-elongate. Legs acutely fringed. Toes 
broadly webbed. Intermaxillary bone produced, and truncated behind, the sutures 
sloping backwards and converging, and then transverse or sinuous. 
Oopholis, Gray, Cat. Tortoises & Crocodiles in B. M. 1844 ; Ann. & Mag. Nat. Hist. 3rd series, x. 267. 



138 DR. J. E. GRAY'S SYNOPSIS OF THE SPECIES 

a. The dorsal scales in six longitudinal series ; the vertebral ones elongated 

like the others. 

1. OoPHOLis POROSUS. (The Saltwater Crocodile.) 

Crocodiliis jiorosiis, Sclin. Araph. 159. Gray, Cat. Tort. & Croc. &c. Brit. Mus. 58; P. Z. S. 1861, UO. 

Crocodilus ooptwlis, Sclm. Amph. ii. 165. , 

CrocodUus biporcaius, Cuvier, Oss. Foss. v. 65, t. 1. f. 4, 18, 19 (young skulls) ; t. 2. f. 8. Miiller 
and Schlcgel, Verh. t. 3. f. 6 (middle-aged skull). Owen, Cat. Osteol. Mus. Col. Surg. 159, 
iios. 719, 723, 724, 727, 738. Huxley, Journ. Proc. Linn. See. Zool. iv. 11. Blainv. Osteogr. 
Crocod. t. 1, t. 3. f. 1, t. 4. f. , t. 52. 

Crocodilus acutus, Owen, Cat. Osteol. Mus. Col. Surg. 157, no. 713. 

Chumpse fissipes, Wagler, Amph. t. 17. 

Crocodilus biporcaUts raiiinus, Miiller and Schlegel, Verh. t. 3. f. 7 (aged skull) ! 

Oopholis porosus, Gray, Ann. & Mag. Nat. Hist. 3rd series, x. 267, 1862. 

Ilab. Asia and Australia ; India, Bengal, and Venang (Ilardwicke) ; China. (Lindsa//); 
Trincomalce ; Borneo {Belcher); Tenasserim coast {Packman); Siam, Cambogia {Mouhot). 

^'ar. avstralis, Giinther. 

Crocodile, Landsljorough, Explor. of Australia, i. 70. 

Hab. North Australia {Elseij & Kraig). 

Dr. Giinther has pointed out to me that all the Australian specimens which we have 
examined have one cross band of the shield less than the Indian specimens; that is to 
say, they have sixteen, and the Indian specimens seventeen bands of shields from the 
neck to the base of the tail. That is the case both in the small specimen in spirits and 
the large specimen, 17^ feet long, which was procured by Islv. Kraig. 

In the British Museum there is the skin of an adult from N.E. Australia, another, 
13 feet long, received from the Zoological Society, and several (two-thirds half-grown) 
young specimens, stuffed, and several young specimens in spirits. 

The largest skull in the British Museum is 29 inches long ; the adult skulls vary from 
29 to 31 inches in length ; a half-grown species is 19 inches long. The skull 26 inches 
long, is said to be from an animal caught in Bengal that was 33 feet long. 

Cuvier figures the skulls of young and half-grown specimens. S. Miiller and Schlcgel 
figure two skulls, one under the name of C. liporcatvs (f C), and the other C. hipor- 
catus raninus (f. 7) : the latter seems to be from an adult or aged animal ; the former 
(f. 6) fiom a full-grown one before the skull is thickened and spread out. Another 
specimen, figured as ('. liijxircatus raninvs (f. 8), appears to be from a specimen of 
Crocodilus or Bomhifrons siamensis. It certainly is not an Oopholis, from the foini of 
the dorsal scales and the presence of the nuchal ones. 

There is a good series of skulls of this species in the Museum of the College of 
Surgeons; but No. 725, named C. hipurcatns in the Catalogue, is the skull of an adult 
Crocodilus vulgaris; and No. 713, called Crocodilus acutus in the Catalogue, is Oopholis 
porosus. 



OF RECENT CKOCODILIANS. 139 

The British Museum received from the Leyden Museum au adult skull of the 
Crocodibos {hiporcatus) ranimis from Borneo ; it is 22 inches long, and agrees in every 
respect with the Oojjhoh's porosus from India. 

Mr. Landsborough observes, " harmless as this animal is in Australia, we were not 
anxious for his company in his native element." — Exploration of Australia, p. 70. 

b. The dorsal scales in four series ; the vertebral series broader than long, the oxter 

series elongate-ovate. 

2. OoPHOUS POXDiCHERiANUS. (Pondicherry Crocodile.) 

Oopholis pondicheriarms. Gray, Ami. & Mag. N. H. Srd series, x. 268. 
Crocodilus pondicerianus, Gunther, Kept. B. I. t. 7. 

The specimen of this species in the British Museum is small, and only just hatched, 
but it is quite distinct from all the others. The vertebral series of plates are nearly 
twice as broad as those in 0. porosus ; the others are also rather wider in comparison ; 
all the dorsal scales are more keeled, and the keels on the scales on the side of the 
base of the tail are higher, and more prominent. The black spots are larger and 
further apart. 

The specimen was purchased of M. Parzudaki of Paris, it having formed part of a 
collection which he received from the French Museum. 

B. Nuchal pjlates four, or rarely two or five, in a cross series. The dorsal plates as 
broad as long, in four or six series. Flu^datile or River Crocodiles. 

a. The intermaxillary bones trnncated behind, with a nearly straight premaxillary 

suture. Face broad, oblong. 

To observe the form of the premaxillary suture in the preserved specimens, it is only 
necessary to elevate the skin of the front of the palate, and lay the bones bare. 

* Toes loebbed. Legs distinctly fringed. Asiatic Crocodiles. 

2. BOMBIFRONS. 

The premaxillary suture straight, or rather convex forwards. The face oblong; 
forehead mth nodules in front of the orbits, but no distinct preorbital ridges. Nuchal 
plates four, in a curved line. Cervical plates six, in the form of a rhombic shield, 
distinct from the dorsal one. Dorsal plates oblong, rather elongate, all keeled, in six 
longitudinal series, and with two short lateral series of keeled scales. The legs fringed 
with a series of triangular elongated scales. Toes webbed. 
Bombifrons, Gray, Ann. & Mag. N. H. 3 series, x. 269. • 

Skull with the nostril separate, the internal nostril as broad as wide, with a 
deep pit on each side in front of it, and rather bent down, so as to open neai'ly 
horizontally. 



140 BR. J. E. GEAT'S SYNOPSIS OP THE SPECIES 

1. BoMBiFRONS INDICES. (The Muggar.) (Plate XXXI., figs. 1, 2, 3.) 
The intermaxillary short, nearly semicircular. 
Crocodilus vulgaris, var. indicus, Gray, Syn. Rept. 58, 1831 ! 
Crocodilus dubius, (jcofF. Ann. du Mus. xii. 122? 
Crocodilus suchus, var. D., Dum. Enc. Meth. Rept. 27. 
Crocodilus palustris, Lesson, Belanger, Voy. 305. Gray, Cat. Tort. & Croc. B. M. 62 (young). 

Owen, Cat. Osteol. Mus. Coll. Surg. 164 & 752 ! Gunther, Rept. B. lud. t. 8. f. a. 
Crocodilus bombifrons, Gray, Cat. Tortoises & Crocodiles &c. B. M. 59, 184'4 (adult) ! 
Crocodilus bombifrons (palustris 'f) , Huxley, Proc. Linn. Soc. Zool. iv. 13! 1859. 
Crocodilus biporcatus, Cautley, Asiat. Research, xix. t. 3. f. 1. p. 3 ! (not Cuvier). 
Crocodilus trigonops, Gray, Cat. Tort. & Croc. B. M. 62, 1844 (young) ! 
Bombifrons trigonops, Gray, Ann. & Mag. N. H. 3rd series, x. 269 ! 
Crocodilus vulgaris, var. B. Dumer. & Bibron, Erp. Gen. iv. 108. 
Crocodilus rhombifer, Owen, Cat. Osteol. Mus. Coll. Surg. 164, n. 752 ! (not Cuiier) . 
Crocodilus V Owen, Cat. Osteol. Mus. Col. Surg. 159, n. 726! 

Hab. India: Gdcages [Dr. Sm/er); Madras (Ji»;-fZo«) ; Cey\oi[i {Kelaart). 

The dorsal shields in four series, all equally keeled, with two irregular series of 
plates on the sides. The shields are often nearly of the same form and size; hut 
sometimes there are larger and broader shields intermixed in and deranging the series, 
and at otlier times the whole vertebral series is formed of wider shields. 

This species has generally been confounded with Oopholis liporcatus and Crocodilus 
vulgaris. 

The face of the younger specimen is rugulose and depressed, with a deep pit on the 
sides over the eighth and ninth teeth ; there are two arched ridges on each side behind 
the nostril, and some rugosities in front of the orbits. In the older skull the face is 
very convex and rounded, rugose, with some more or less distinct rugosities in front 
of the orbits, but not the distinct longitudinal ridge so characteristic of Oojj/wlis po7-osus. 

Professor Owen described the peculiar form of the premaxillary in a skull in the 
College of Surgeons Museum, sent from Bengal by Dr. Wallich ; but he refers the skull 
to Crododilus rhombifer of Cuvier, which is an American species. 

The smallest specimen in the British Museum is 19 inches, and the largest nearly 
10 feet long; there are skulls showing that it grows to a much larger size. The 
specimen I described as C. trigonalis is 24^ inches long. 

In my Catalogue of the Tortoises and Crocodiles in the British Museum, published 
in 1844, 1 described it, from two adult skulls from India of 18 and 20 inches long, as a 
new species, which I called Crocodilus bombifrons, pointing out the straightness of the 
suture between the intermaxillary and the maxillary bones. I observed that I had 
seen in the Paris Museum a large specimen which had been described by Dumeril and 
Bibron as an adult of Crocodilus biporcatus, which appeared to belong to this species, 
stating that it was immediately known from C. porosus by the breadth and convexity of 
the face. 



OF EECENT CROCODILIANS. 



141 



In the same work 1 separated the Indian specimen from the common African Croco- 
dihis, under the name of Crocodilus palustris of Lesson, and pointed out that it seemed 
to be the same as the Crocodilus biporcatus raniniis of Miiller and Schegel ; and I 
described two other very young specimens under the name of Crocodilus trigonojjs, on 
account of the shortness and width of the head. 

The examination of the specimens on which these species were founded, and the com- 
parison of them one with another when ranged in a series, with the other specimens 
since obtained interlocatcd in their places according to their size, have convinced me 
that they are referable to mere variations of growth of a single species, which is generall)- 
spread over the Indian peninsula. 

Var. Nose narrow, the intermaxillary bones rather longer and narrower. 

Hah. Ceylon (skull, Kelaart). 




Fig. 1. and h. Skull of adult C homb'ifrons, Gray, 1S47. Presented by C'apt. Oriel. 

There may be two species of Ceylon Muggars, as in one of the heads the intermax- 
illaries appear to be longer and nariower than in the others from the same country. I 



VOL. VI. PART. IV. 



142 



DR. J. E. GRAY'S STXOPSIS OF THE SPECIES 



have not sufficient materials to satisfy myself as to the distinctness of this species and 
the permanence of the forms. 




Fig. 2. Skull of B. huUcus, nearly adult. 

Fig. 1. Fig. 2. Fig. 3. Fig. 4. 

in. lines. in. lines. in. lines, in. lines. 

Leugth of skull 20 17 3 9 10 4 8 

Length from occiput to front of orbit ..60 59 37 28 

Length of face 13 3 11 6 6 3 2 

Length of lower jaw 27 23 none. 5 5 

Width at occiput 13 5 Id 6 5 11 2 (5 

Width at hinder notch i) 2 (1 9 3 9 1 

Width at notch 5 4 5 U 2 4 (J 9 

The face becomes shorter, compared with the width of the middle of the face, as the 
animal becomes older. 

In the young, fig. 4, the length of the head is rather more than three times the width 
of the swollen part behind the notch. In fig. 3 it is just three times, and in fig. 2 it is 



OP EECENT CROCODILIANS. 



143 



twice and a half the length of the width at the same part; and in the old skull, fig. 1, 
It IS only a little more than twice the width of the face in length. 

Fig. .3. Skull of B. indkug, half-grown. India, Sir John Boileau. 



sW'i" 

Fig. 4. Skull of young, of natural size (C. ^nV/ojio^w, Gray). 

As a good illustration of the difference in the appearance of the skulls of the individuals 
of the species, I may gi^•e the measurement of two skulls of " Muggars " from India, of 
the same size, in the British Museum Collection :— 

Broad variety. Narrow variety. 

T iv p ii_ 1 11 ■ , inches. inches, 

i^cngtu oi the skull along the forehead 9i 91 

Length of side of the skull IQi 10^ 

■Width of back of skuU r^i =! 

Width in front of orbits 41 4" 

Width over largest tooth 33 31 

Width at notch 9i 9 „„ 1 1 .i 

~2 -"^ '■It 

The broad-nose variety (fig. 3) was presented by Sii- John Boileau, and the narrow one 
by Capt. Boys. 

22 




^ 



144 DB. J. E. GEAY'S SYNOPSIS OF THE SPECIES 

When the two skulls are placed side by side, the large teeth are just the same 
distance apart, and the different teeth in the two skulls exactly agree in size, position 
and distance from each other. 

2. BoMBiFRONS siAMENSis. (Siamese Muggar.) 

The face depressed, elongate, nearly smooth, with a slight nodule in front of the 
orbits. Intermaxillaries rather elongate, half oblong. 

Crocodilus niloticus, Latr. Kept. i. 206, t. — . From Faujas St. Fond, Mont. St. Pierre, t. 43. 
Crocodilus siamensis, Schn. Amph. 157. Gray, Syn. 60 ; Cat. Tort. & Croc. B. M. 63 (monstrosity) ? 

From Perrault, Hist. Acad. Sci. iii. 355, t. 54. Gunther, Eept. B. I. t. 18. f. 3. 
Crocodi/us ffaleatus,C\ivier,Oss.Foas. v. 52, 1. 1, f. 9 (from Perrault). Dum. & Bibr. Erp. Gen.iii.113. 
Crocodilus pahtstris (part.), Dum. & Bibr. Erp. Gen. iii. 113. 
Crocodilus vulgaris (part.). Gray, Syn. 58. Dum. & Bibr. Erp. Gen. ii. 108 ? ^liiller & Schlegel, 

Verb. t. 3. f. 9 (head?). 
Crocodilus vulgaris, Owen, Cat. Osteol. Mus. Col. Surg. 107. n. 718 ? 
Bombifrons siamensis, Gray, Ann. & Mag. N. H. 3rd series, .\. 269. 

Hab. Siam, Cambogia (M. Mouhot). 

There is a well-preserved half-grown specimen of this species in the British Museum.* 
It differs from all the specimens of Bombifrons indicus in the collection in the face 
being much longer, and not so tubercular and pitted. 

It has four series of nearly equal-sized, uniformly shaped, and keeled shields, with 
three interrupted series of unequal-sized smaller shields on each of the sides ; those of 
the outer series are the longest. 

As the head agrees with the figure of the head from which Schneider named his 
species, I have retained it; and I have little doubt that the two keels which are 
present in that specimen are either an individual peculiarity, or perhaps a character 
that developed itself as the animal approached old age. 

The skull of the young animal in the Museum of the College of Surgeons, no. 718, 
appears to belong to this species ; but it requires more comparison. It is clearly a 
Bombifrons, and it is much smoother and longer than the skull of B. indicus of the 
same size and age. Professor Owen observes, "The palatine suture between the 
premaxillary and maxillary bones passes obliquely backwards a little way at its 
commencement, and then extends truncated across; but the premaxillary bones are 
larger than in the second Gangetic Crocodile." There is a small palpebrary ossicle 
above the anterior angle of the eyes. — Owen, I. c. p. 157. n. 718. 

There is a young specimen of a Crocodile, received from Singapore, which somewhat 
resembles the one from Siam in the form of the liead, and lias six series of strongly 
keeled shields on the back ; but the four middle ones, of nearly equal size and form, and 
those of the outer series, are narrower, and there is a series of much smaller ones below 
on each of the sides. I am by no means convinced that this will form a distinct species, 
it is probably only an accidental or a local variety. 



OP EECBNT CEOCODILIANS. 145 

** The legs with an indented fringe of short, narrow scales. Toes short, 
nearly free. American Crocodiles. 

3. Palinia. 

The face oblong ; forehead very convex, with a ridge in front of each orbit, con- 
verging in front and forming a lozenge-shaped space. Nuchal plates two or four, 
unequul. Cervical disk rhombic, of six large plates. Dorsal plates lai-ge, broad, in 
six series; the vertebral series nearly smooth, the lateral one strongly keeled. The 
intermaxillary short, truncated behind the premaxillary ; suture straight, transverse. — 
See Cuvier, Oss. Foss. iii. 72, t. 3. f. 1-5. 
Palinia, Gray, Cat. Tortoises & Crocodiles, B. M. 1844 ; Ann. & Mag. Nat. Hist. .3rd series, x. 270. 

1. Palinia rhombipera. (Cuban Palinia.) 

The upper surface of the forearms and thighs covered with convex keeled scales ; 
the outer edge of the legs and feet with a series of very elongate scarcely raised scales, 
forming only a slight fringe. The toes short, scarcely webbed. 

Aquez palin, Hernand. Nov. Mexic. ii. 2. 

Crocodilus rhomhifer, Cuvier, Ann. Mus. H. N. x. 51 ; Oss. Foss. v. 51, t. 3. f. 1-4. Tiedem., Oppel, & 
Leboscli,Nat.Ampli.75,t. 10. Gray, Syn. Kept. 59. Dum. &Bibr.Erp.Gen.iii.97. Sagra,Cuba, 
t. 4! Huxley, Proc. Linn. See. iv. 10. Blainv. Osteog. Croc. t. 5. f. 3 (head?) (not Owen) . 

Crocodilus [Palinia) rhomhifer, Gray, Cat. Tort. Croc. B. M. 63; Ann. & Mag. Nat. Hist. x. 270. 

Crocodilus planirostris. Graves, Ann. Gen. des Sei. Phys. de Bordeaux, ii. 348. Gray, Syn. Rept. 59. 

Crocodilus gravesii, Bory de St. Vincent, Diet. Class. H. N. iii. 109, t. Dum. & Bibr. Erp. Gen. iii. 101 . 

Eab. South America, Cuba {W. S. Macleay, Eamon de la Sagra). 

In the British Museum there is a well-grown specimen, 5 feet 4 inches long, of this 
species, collected in Cuba by M. Ramon de la Sagra, and sent from the French Museum. 
Two young specimens in spirits, sent from Cuba by Mr. W. S. Macleay, are almost 
2 feet long, are pale brown, with small dots on the head, and a dark spot on the middle of 
many of the dorsal scutella ; the face is irregularly tessellated with square brown spots. 

Cuvier described the Crocodilus rhomhifer from two specimens : — one in the Cabinet of 
the Academy of Sciences, in a nearly entii-e state ; and the other, a very mutilated skin, 
in the Museum, which also furnished him with the skull figured in t. 3. f 1, 2, 3, 4, 5 
of his work on Fossil Bones, pp. 51-70. The original habitats of these specimens 
were not marked. But M. Ramon de la Sagra sent a young living .specimen to the 
Jardin des Plantes, proving that this is an American species ; and it is probable that 
the Crocodile which Hernandez describeSvand figures as coming from New Spain, under 
the name of Aquez palin, belongs to this species. 

M. Graves, in the ' Annales Generales des Sciences Physiques de Bordeaux,' describes 
a Crocodile under the name of C. planirostris, from a specimen which was formerly in 
the Collection of the Academy of Bordeaux, but is now in the Museum of that town. It 
was procured from M. Journee, a surgeon of a ship that for some time traded with the 
negroes of the coast of Congo. M. Bory de St. Vincent for these reasons thought it niight 



146 DB. J. E. GBAT'a SYNOPSIS OP THE SPECIES 

have come from Africa ; and he figured and described it under the name of Crocodilus 
qravesii in the Dictionnaire Classique d'Hist. Nat. vol. iii. p. 109, t. 

MM. Dumeril and Bibron observe that, when they asked for a new account of the 
specimen, it was in such a bad condition that they could only reproduce the description 
given by M. Graves. The study of the description and figure, which are the only 
material now left for the purpose, lead to the idea that it was not distinct from 
Crocodilus rliomhifer, and was most probably brought from the island of Cuba; and 
the ships which are engaged in trade with the negroes on the coast of Congo 
frequently visit Cuba, as that island is furnished with slaves from the Congo coast ; so 
that it is not at all unlikely that the specimen was brought from tliat island. 

2. Palixia? MORELETii. (Yucatan Palinia.) 
Crocodilus moreletii, Dum. Arcli. du ]Mus. vi. 255, t. 20 ; Cat. Rei3t. 28, n. 5*. 
Palinia? moreletii, Gray, Ann. & Mag. N. H. 3rd series, x. 271. 

Dorsal scales keeled, nearly square; scales of the sides and limbs smooth, without 
tubercles. 

Ilab. Yucatan, Lac Flores (M. Morelet). 

This species is described from a specimen in the Museum of Paris, which is very 
badly figured and indistinctly described in the memoir above cited. 

There are two young specimens of Crocodiles, in spirit, without habitats, in the 
British Museum, which are peculiar in the large size of the nuchal shield, the strength 
of the keels of the dorsal shields, and the large keeled scales of the forearms and thighs, 
in which they agree with Palinia rhombifera ; but there is so much diflierence between 
the two, and between each of them and the specimens of that species from Cuba, that I 
think they must be left in doubt for further elucidation. There are also two small 
stuffed specimens in the collection (purchased of dealers, without any locality attached), 
which are peculiar in having six series of imiform, squarish, very strongly keeled dorsal 
scales ; they are very unlike any other specimen in the collection, and may be new ; 
but I do not like to describe them in the present imperfect state of our knowledge. 

b. The intermaxiUaryhone elongate, produced and truncated hehind ; the sutures sloping 
backwards and converging, and then transverse or sinuous. Toes webbed. Legs 
with a fringe of elongated triangular scales. 

4. Crocodilus. 

Face oblong, depressed, without any ridge in front of the orbits. Nuchal shields 
four, in an arched series. Cervical disk rhombic, of six shields. Dorsal plates quadri- 
lateral, as broad as long; the vertebral series rather the widest and most keeled. 
Intermaxillary produced behind. 
Crocodilus, Gray, Ann. & Mag. N. H. 3rd scries, x. 271. 

" The crocodiles live on the mud-banks or swimming about the rivers " of Africa. 

Dr. Balfour Baikie observes : — " The ninth upper tooth of crocodiles is said to be 



OF RECENT CEOCODILIANS. 



147 



enlarged like a canine ; but this is not correct. I have examined the dentition of eighteen 
skulls of various species; in the lower jaw there are always nineteen teeth, but in the 
upper jaw the number in the adult is seventeen on either side, while in the young it 
is eighteen. This is owing to the second incisor being deciduous ; and in old skulls the 
socket is completely obliterated by the enlargement of foramen for the two anterior 
teeth. Thus in old animals there are only four teeth in each intermaxillary bone, while 
in the younger individuals there are always five. So, more strictly, it is the tenth, and 
not the ninth, upper tooth which is enlarged." — P. Z. !S. 1857, p. 50. 

Crocodilus vulgaris. (OMve African Crocodile.) 
Crocodihts niloticus (part.), Daud. Kept. ii. 267. Wagler, Syst. Amph. t. 7. f. 11. 1, 2. 
Crocodilus vulgaris, Cu^der, Oss. Foss. v. 42, t. 1. f. 5 & 12, t. 2. f. 7. Blainv. Osteogi-. Crocod. 12(). 

Gray, Ann. & Mag. N. H. 3rd series, x. 271. Huxley, Proc. Linn. Soc. iv. 6. 
C. suchus, Geoff. Ann. Mus. x. 84, t. 3. f. 2-4. 
C. chamses, Bory, Diet. Class. H. N. v. 105. 
C. lacunosus, Geoff. Croc. d'Egypte, 167. 

C. marginatus, Geoff. Desc. d'Egypte, 365. Gray, Cat. Tortois. 61. 
Crocodilus cataphractus, Riippell, MS. Gray, Syn. Kept. 78. Mus. Frankfort. 
Crocodilus verd de Senegal, Adanson, Seueg. Cuvier Oss. Foss. v. 4. 
Crocodilus acutus, Owen, Cat. Osteol. Mus. Coll. Surg. p. 157. n. 715, not Cuvier. 
Crocodilus binuensis, Balfour Baikie, Proe. Zool. Soc. 1857, xxv. 484. Skull described. 
Green crocodile, Gray, Rep. of Brit. Assoc. 1862, Sections, p. 107. 

Hah. African rivers. Living on the mud-banks : North Africa, Egypt; West Africa, 
Senegal {Adanson) ; Gaboon {Murray, Cope) ; South Africa, Cape of Good Hope ; Central 
Africa, Kwora and Binui {Baikie) ; Madagascar {Ilavet, fide Cuvier, Oss. Foss. 44). 



Fis. 5. 



Fiff. ' 




Fis. 6. 





!«|fli/'/Wli^a 



Figs. 5, 6, 7, 8. Head and nuchal and cervical shields of Crotodihis vult/ai-is. 



148 DE. J. E. GEAT'S SYNOPSIS OF THE SPECIES 

The largest specimen in the British Museum is nearly 15 feet long. There is a very 
tine skull received from old Calabar, whose greatest width behind is 13 inches, length 
above upper surface from end of nose to back of occiput 22 inches, width at the larger 
lateral tooth 7f inches, at the notch 4f inches. The intermaxillary bones are produced 
backwards between the ends of the maxilla. The hinder nasal opening is transverse, 
inferior, and ascending nearly perpendicularly. The nose has two large oblong diver- 
ging prominences on the sides — one over the hinder edge of the notch, and the other 
o\'er the hinder part of the root of the largest tooth, behind the notch. 

There is a second skull from Western Africa in the Museum, of nearly the same 
length, which is considerably narrower in all its parts. Length along the upper surface 
from end of nose to back edge of occiput 20^ inches; greatest width behind, 12 
inches, at largest lateral tooth 6| inches, at the notch 3f inches. 

These two skulls rather differ in the direction of the suture behind the maxillary 
bones ; in the wider specimen it is much more produced behind tlian in the other. 

I have examined and compared with care specimens of different ages from North 
Africa near the Nile, from West Africa at Senegal and Gaboon, South Africa at the 
Cape of Good Hope and Natal, and a specimen brought from Central Africa by Dr. 
Baikie ; and thougli they each exhibited certain peculiarities, yet I believe, as far as the 
specimens at my command enable me to form a judgment, that they all belong to a 
single species which is generally distributed over the African continent. 

At the same time, from the slight differences which the specimens from the different 
localities do exhibit, I should not be surprised, if we had a complete series of perfect 
specimens and of skulls of different ages from each locality, to find that there were 
sufficient differences between them to show that each locality has a special local 
variety or, perhaps, species; but unfortunately there is not in the British Museum, or in 
the other museums and collections to which I ha^■e access, such a series ; all the speci- 
mens from the cape of Good Hope and West Africa seem to be either in the adult or 
very young state, while those from the other localities are either ^ery young, or of an 
intermediate age. 

On the other hand the series of specimens- from the same locality, as from South 
Africa for example, whence we have most specimens, exhibit variations among them- 
selves, quite as great as between the specimens from \ arious parts of Africa. 

It is therefore more safe to regard them all as one species. 

These species grow to a large size ; we have a specimen from the Nile and some from 
the Cape of Good Hope in the British Museum which are nearly 15 feet long. 

The skulls which seem to belong to larger specimens often reacli tlie length of 
24 or 25 inches. 

The history of the Nile Crocodile is given in great detail in the fifth volume of 
Cuvier's ' Recherches sur les Ossemens Fossiles,' v. 43. 

Geoffrey St. Hilaire, in his ' Essay on the Crocodiles of Egypt,' separated the Egyptian 



or EECENT CEOCODLLIANS. 14!) 

specimens into two species under the name of Crocodilm lacunosus and C. marffinatus. 
In the "Annalesdu Museum," vol. x. p. 83, he described a third, under the name 
of C. suchus. 

Professor Owen has figured the skull of a crocodile, from an Egyptian mummy, imder 
the name of C'rocodilus suchus, Geoff., in the ' Monograph on the Fossil Reptilia of 
the London Clay,' published by the Palgeontographical Society, 1850, t. 1. f. 2. I do 
not see how it diifers from the crocodiles at present found in the Nile. See also Huxley, 
Journ. Proc. Linn. Soc. iv. 15. 

In the ' Catalogue of Tortoises and Crocodiles,' p. Gl, I separated the adult Cape 
crocodiles from the North-African specimens, under the name of C. mare/inatus, because 
the head is not so narrow ; but it is to be observed that most of the North-African speci- 
mens with which I had compared them were of small size, and consequently had the 
head less developed. 

Dr. Baikie described the crocodile of Central Africa, found in the river Kwora 
and Binue (or Niger and Twedda), under the name of Crocodilus binuensis ; it is of a 
dark green colour, and lives on the mud-banks or swimming in the rivers. 

Mr. Cope, ' Proceedings of the Academy of Natural Sciences of Philadelphia ' for 
1859, p. 296, regards the crocodile of Equatorial Western Africa (Ogobai) as the Croco- 
dilus margiiiatus of Geofii-oy. 

Dr. A. Smith, referring the Cape specimens to Crocodilus marginatus, observes, " they 
are occasionally found in the rivers west of Port Natal, but more abundantly in those to 
the eastward and northward, and occur in such numbers in the rivers in a district north 
of Kurrichane, between 24° and 22° south latitude, that the natives who used to reside 
there were known by the appellation Iiaquana:=th.e people of the crocodile." — Zool. 
South Africa, Appendix 2, 1845. 

MM. Dumeril and Bibron in their ' Erpetologie Generale,' iv. 104, divided their 
Crocodilus vulgaris into four varieties, thus : — 
Var. a. The Crocodilus vulgaris of Geoffroy, from North Africa, Egypt, and the 

Nile. 
Var. b. Crocodilus palustris. Lesson, described from a specimen sent from the Ganges 

by M. Duvaucel, and from the coast of Malabar by M. Dussumier. 
Var. c. the Crocodilus marginatus, 1. Geoffi-oy, from North Egypt and the Cape of 

Good Hope. 
Var. d. the Crocodile verd of Adanson, fi-om the Nile, the Niger, and Senegal. 

There is no doubt that vars. a, c, and d are true Crocodiles, and are what is considered, 
in this essay to be the Crocodilus vulgaris of Africa. 

Var. b on the other hand does not belong to the same genus. I have not the sHghtest 
doubt this variety is founded on young and half-grown specimens of Bombifrons indicus, 
most distinct from Crocodilus vulgaris by the form of the head and the structure of the 
skull, as MM. Dumeril and Bibron would have found, if they had examined any of 

VOL. VI. P.\RT IV. Y 



150 DE. J. E. GEAT'S SYNOPSIS OF THE SPECIES 

the twelve specimens which they say they procured. They have named the adult 
specimen in the Paris Museum Crocodihis hijiorcattis. 

In the ' Annals and Magazine of Natural History,' vol. xviii. t. 7, Dr. Falconer figures 
the skull of a Crocodile under the name of C. marginatws, which is in the Belfast Museum. 
It is said to have been brought from Sierra Leone ; but I think that this must be a 
mistake: it is not like the .skull of any Crocodile I have seen from West Africa, and it 
is not a bad representation of the skull of a half-grown Bombifrons mdints from India. 
Can the habitat be a mistake ? perhaps the habitat was only intended for the first-de- 
scribed species, Cataphractits meristops, for which it is the true locality. 

A skull of Crocodihis vulgaris is described in Professor Owen's ' Catalogue of Os- 
teological Specimens in the Museum of the College of Surgeons ' under the name of 
Crocodilus acutus, p. 157. n. 715. 

5. MOLINIA. 

Face elongate ; forehead swollen, convex, especially in the adult; orbits without any 
anterior ridge. Nuchal plates two or four, small. Cervical disk rhombic, of six 
plates, the side plates generally small.' The legs fringed with a series of triangular 
elongate scales. Toes webbed. Scales of the forearm and thigh thin, smooth. 

Muzzle oblong, elongate, slender, with a swollen convexity on the middle of the 
face before the eyes. Nostril not separated by a long ridge : the internal nostril pos- 
terior, with an oblong sloping opening; the intermaxillary suture produced behind 
between the ends of the maxillae. 
Molinia, Gray. Ann. & Mag. N. H. 3rd series, x. 272. 

* Face slender. Dorsal plates irregular ; the central series small, keeled ; lateral 
scattered, strongly keeled. Nasal hones produced to the nostrils. Molinia. 

1. Molinia Americana (American Crocodile). 
Crocodilus americanus (Plumieri), Schn. Ampli. ii. 23. Gray, Cat. Tort. & Croc. &c. B. M. 60. 
Crocodihis acutus, Geoff. Ann. Mus. ii. 53, t. 57. f. 1. Cuvier, Oss. Foss. v. t. 1. f. 3 & 14, t. 2. f. 5. 

Gray, Syn. 60. Dum. & Bib. Erp. Gen. iii. 120. Owen, Cat. Osteol. Spec. Mus. Col. Surg. 

1 57. n. 711, 712, 714, 716 ; Reptiles of the London Clay, t. 25. f. 10. Briilil, Skclct. Krokod. 

t. 8 & 9, t. 10, t. 17. 
Crocodilus americanus {acutus, Cuv.), Huxley, Journ. Proc. Linn. Soc. iv. 11, 1859. 
Molinia umericana, Gray, Ann. & Mag. N. H. 3 ser. x. 272. 

Tt Crocodilus hiscutatus (part.), Cuvier, Oss. Foss. x. t. 2. f. 6. Tiedem. Anipli. t. 12. 
Crocodilus de St. Dominyue, Geoff. Ann. du j\Ius. ii. 53, t. 27. f. 1 . 

Hah. Tropical America. Cuba {W. S. Macleay); Jamaica {B.M.)\ West Ecuador 
Nicaragua (Fra5<;r ; Michardso7i)\ West coast of America (.S^'/c/^e;-); St Domingo (CVwV/-); 
Guatemala [Salvin). 

The specimens in the British Museum vary in length from 19 to 103 inches ; and the 
skulls show that they grow to a larger size. 



OF EECENT CEOCODILIANS. 151 

Var. with two additional small cervical scutella behind the others. B.M. 

Crocodilus americanus, var. ? Gray, Cat. Tort. & Croc. B. M. 60. 
Crocodilus acutus, var., A. Dum. Cat. Rept. 28; Arch, du Mus. vi. 256. 
Molinia americana, var.. Gray, Ann. & Mag. N. H. x. 2~2. 

Hab. West coast of America {Belcher); Mexico (Wanvick). 

Cuvier in his essay gives the history of this species under the name of Le Crocodile a 
museau effile, ou de Saint Domingue (Crocodilus acutus, nob.), Oss. Foss. v. 458, and 
figures the skull at t. 1. f. 3 & 14, and the nuchal shield at t. 2. f. 5. 

Professor Briihl described and figured the skeleton of this species in his work. There 
is the skeleton of a well-grown specimen in the British Museum, and several skulls. 
The central prominence of the hinder part of the muzzle is sometimes much less 
developed than in the typical skulls. 

** Face very slender. Dorsal plates nearly uniform. Nasal bones not produced quite 

to the nostrils. Temsacus. 

2. Molinia intermedia (Orinoco Crocodile). (Plate XXXIl. figs. 4-6.) 
Dorsal plates in six rows, all slightly and nearly equally elevated ; the keels of the two 
vertebral series rather larger than the others, quadrilateral, rather broader than long ; the 
lateral ones oval, with five or six large plates forming an interrupted line on the sides. 

Crocodilus intermedins, Graves, Ann. Sci. Phys. ii. 344. Gray, Syn. 59. 

Crocodilus journei, Bory, Diet. d'H. N. v. iii. Dum. & Bib. Erp. Gen. iii. 129. A. Dum. Arch, du 

Mus. X. 172, t. 14. f. 3 (head). Huxley, Proc. Linn. Soc. iv. 11. 
Crocodile de I'Orenoque, Parzudaki, MS. 

Mecistops journei (part.). Gray, Cat. Tort. & Croc. B. M. 58, from Bory. 
Molinia intermedia, Gray, Ann. & Mag. N. H. 3rd series, x. 272. 
"il Mecistops bathyrhynchus. Cope, Proc. Acad. N. S. Philad. 1860, xii. 550 (skull). 

Hab. America: Orinoco. 

There is a young specimen in spirits in the British Museum, sent by M. Brandt, of 
Hamburg, as Crocodilus acutus, and an adult skull, 20 inches long, received from Paris 
as Crocodile de VOrenoque, and a second very large skull purchased in London. 

In my Catalogue of Tortoises and Crocodiles in the British Museum Collection, from 
all I could then learn, I was induced to believe that the Crocodilus intermedius of 
Graves was the same as the Crocodilus schlegclii of Borneo, and therefore called the 
Bornean animal Mecistops journei. M. Dumcril, in his paper in the Archives du 
Museum, not seeing the mistake, says that I refer the true Crocodilus intermedius to 
the genus Mecisto])s, and suggests that Crocodilus acutus ought also to belong to it. 

M. Auguste Dumeril, for the purpose of comparing the head of this Crocodile with 
that of Crocodilus leptorhynckus of West Africa, gave a figure of the head and front part 
of the back of the Crocodile de Journee, Archiv. du Mus. x. 173, t. 14. f. 3 ; but it does 
not appear whether it is from a specimen, or only an enlarged copy of the figure of 

t2 



152 



DH. J. E. GEAT'S SYNOPSIS OP THE SPECIES 



M. Bory de St. Vincent. If the latter, it is so embellished that one is unable to recognize 



its origin. 



Pig. 9 




Pigs, y & 1(1. Skull oi Molinia intermedia : adult. 

II. Xape with a broad flat-topped shield formed of two or three pairs of keeled plates, 
strongly keeled on each side, and nearly continuous with the dorsal shield. Legs 
fringed. Toes welled. Abnormal Crocodiles. 
* Face Iroad ; nasal bone produced into the nostril. Alligatorian Crocodiles. 

6. Halcrosia. 

The premaxillary suture transverse, rather convex backwards. Nasal bones produced 
beyond .the intermaxillar)-, and forming a bony septum between the nostrils. The 
palatine bone produced to the same level as the lateral opening — that is, to the lateral 
inflection of the skull. The face oblong, broad, without any ridge in front of the orbit. 
Eyelids with two bony plates. Nuchal plates four, in a cross row, strongly keeled. 
Cervical plates three or four pairs, forming a ridge ou each side, the hinder one smaller. 
Dorsal plates in foui- series ; the central broad, slightly keeled, the outer narrow, dis- 
tinctly keeled ; sides with large convex scales. 
Halcrosia, Gray, Ann. & Mag. N. H. 3rd series, x. 273. 
Osteolamus, Cope, Proc. Acad. N. S. Pliilad. xii. 550. " 

It has the square head and the elongated cervical shield formed of single pairs of 
scutella, and the bony eyelids, of the Alligators with bony eyelids ; but it is a Crocodile, 
and there arc two bones in the eyelid instead of one as in Caiman pmlpelrosus. 



OF EECENT CROCODILIAKS. 



153 



The skull of the AUigato)' ])a^p('i)'Osus is easily known from that of this species even 
in the young by the cheeks of the former being flattened and nearly erect, and of the 
latter spread out, and in the supratemporal fossae being open, while in the Alligator they 
are closed even in the young specimens. 

Most probably it was from the examination of a skull of this Crocodile that the 
statement has arisen that in some Alligators the canine teeth sometimes fit into a notch 
in the upper jaw, and not into a \i\t as they normally do in that genus. 1 will not under- 
take to say that such an abnormal state does not exist in the genus Alligator; but 
I have not observed it. 



Halcrosia nigra (Black African Crocodile). (Plate XXXI. figs. 4, 5, 6.) 
Krokodile noir du Niger, Adanson, MS., Mus. Paris. See Cuvicr, Oss. Toss. iii. 41. 
Crocoddus niger, Latr. H. N. Rept. i. .510, from Adanson. 
Crocodilus palpebrosus, var. 3, Cuvier, Oss. Foss. iii. -11, t. 3. f. 6 (part.). 
Crocoddus trigonatus (part.), Cuvier, Oss. Foss. iii. 65. 
African Black Crocodile, Gray, Rep. Brit. Assoc. 1863, Sections, 107. 
Osteolamus tetraspes, Cope, Proc. Acad. N. S. Philad. xii. 550. 
Crocodilus frontatm,A.'Sl\3Jcra.j, Proc. Zool.Soc. 1863, pp. 1.39, 21.3,fig. head, t. 30. by Ford. Strauoh, 

Syu. Croc. t. 1 (head, young) . 
Halcrosia frontata, Gray, Ann. & Mag. Nat. Hist. 3rd series, x. 377. 

Hab. West Africa : Senegal [Adanson) ; Gaboon ; Old Calabar ; Ogobai River (Cope). 
Fiff. 11. iiwi Fis. 13. 





Fig. 12. 




Fis. 14. 




Figs. 11-14. Head and cervical and nuchal plates of young IlaLrositi niijrn 



154 DE. J. E. GEAT'S SYNOPSIS OF THE SPECIES 

Black, slightly mottled with pale whitish. Head pale olive, black dotted ; sides of 
lower jaw black-banded ; muzzle broad, oblong, trigonal, rather dilated on the sides ; 
forehead high, broad, and flat, with a small tubercle at the front angle of the orbit. 
Nuchal shields strongly keeled, two in a cross line in two groups. Cervical shields six, in 
three pairs, all close together, the two anterior pairs of equal size, large, strongly keeled, 
and bent in on the outer sides, the hinder pairs much smaller. The vertebral series of 
doisal shield broad, square, scarcely keeled, with one, and in the front of the back two 
rows of oval, elongated, keeled shields on the side of them, and a few isolated, scattered, 
compressed, high, tubercular-like, small, ovate shields on the sides of the body. Shields 
of the upper arm oblong, trigonal, keeled, in six oblique cross series. The lines of the 
upper jaw sinuous, three-parted, the front with five, the second with seven, and the 
hinder vdth five teeth. 

The largest specimen I have seen is in the Free Museum at Liverpool, which is 
nearly 5 feet long, but I have no doubt it grows larger. The muzzle of this specimen 
from the tip of the nose to the orbit is 3^ inches, its width in front of the orbit 
2| inches, and at the notch of the canine teeth 1^ inch. The eyelid is obliquely 
divided from the front of the orbit to the back of the eye. 

The Black African Crocodiles appear to be a common species on the west coast of 
Africa ; for they are often brought to the Port of Liverpool by tlie palm-oil ships, and 
frequently in a li\ing state ; indeed I am informed there were some lately alive in the 
Society's Gardens in the Regent's Park. 

Mr. Andrew Murray, at my recommendation, has described it in the ' Proceedmgs ' of 
the Society as a new species of Crocodile under the name of C. frontntus ; for at that 
instant it did not occur to me that it might be the Black Crocodile of Adanson, noticed 
as an Alligator. It is to be observed that, though they have specimens of this 
Crocodile in the Paris Museum in such abundance as to part with the skeleton of it as a 
duplicate, it is not included as AUiijator palpehrosus, or under any name, in M. 
Auguste Dumeril's List of the Reptiles of West Africa, printed in the last volume of the 
Archives du Museum of Paris. 

This Crocodile has very much the external appearance of the Caiman with bony 
eyelids, Crocodilus palpehrosus, Cuvier ; and I think it very likely that Cuvicr mistook a 
specimen of it in the Paris Museum, which Adanson had marked with his own hand 
" Krokodile noir du Nii/C7%" for a specimen of that species. (See Cuvier, Oss. Foss. iii. 
p. 41.) Audit is still confounded with that species by the Frencli naturalists; for there 
is a skeleton in the British Museum, lately sent from M. Braconier, of the French 
Museum, under the name of Caiman a pavpieres osse\ises. 

Adanson, in his ' Voyage to Senegal,' at p. 10, mentions the occurrence of Crocodiles, 
and at p. 73 a second kind of Crocodile, which is as large as the other, and distin- 
guished by the black colour and by the jaws being much more elongated. It is moi-e 
carnivorous, and said to be fond of human flesh. 



OF EBCENT CEOCODILIANS. 155 

Cuvier, in his Essay on the species of existing Crocodiles, first published in the 10th 
■\'olume of the ' Annales du Museum,' and reprinted in his ' Ossemens Fossiles ' under 
the head of Le Caiman a paupieres osseuses {Crocodilus palpelrosus, nob.), after dividing 
tliis species into two varieties, expressed a doubt if they were not inhabitants of 
different continents. He observes, " One of my individuals, which has been for many 
years in the Museum, has on it the half-effaced name of KroJcodile noir du Niger in the 
hand-writing of Adanson," — and proceeds thus: — 

" This naturalist, in his ' Voyage,' speaks of two Crocodiles in the Senegal. M. de 
Beauvois adds that he saw at Guinea a Crocodile and a Caiman. It is thei-efore clear 
that there is a species with the form of a Caiman that inhabits Africa. 

" There remains still an embarrassment. Adanson says his Black Crocodile has the 
muzzle longer than the Oreen, which is certainly the same as the Crocodile of the Nile ; 
but we have a specimen ticketed by his own hand which has a much shorter muzzle 
than that from Egypt. 

" Has Adanson made a mistake in ^\Titing this phrase '\ or has he erroneously 
ticketed the specimen'? How are we to disentangle these errors]" &c., vol. v. p. 41. 

Dumeril and Bibron, in their ' Erpetologie Generale ' (vol. iii. p. 75) adopt and 
repeat all that Cuvier has said, and still doubt if these two varieties may not be found. 
the one in America, and the other in Africa. 

If Cuvier and his successors had examined the two specimens on which they founded 
the account of his second variety of C. palpelrosrts, they would have found that they were 
not only distinct species, but also species belonging to two genera or subgenera. The 
one which had served as the model for Seba, and which Seba, with the iisual inat- 
tention to true liabitats at that period, said came from Ceylon, was a true Alii- 
f/cctor, and a native of America; and the other, ticketed by Adanson as from the 
Niger, was really a Crocodile from Africa : so that the sarcastic observation which he made 
on travellers, and which may in some cases be true, in this instance was uncalled for, 
the traveller being in fact more accurate than the cabinet naturalist ; and Adanson only 
made a slip of the pen in saying the beak was longer instead of shorter than the common 
Green Crocodile ; and any one who compares the Black Crocodile of Africa with an 
American Caiman will not think that M. Beauvois was very much out when he called 
it a " Caiman." 

Cuvier, in his Essay, when describing Crocodilus biscufatiis, established on the Gavial 
du Senegal of Adanson, again refers to the Crocodile noir of that author. He states 
that among the drawings of Adanson there is the figure of a Crocodilus vulgaris, named 
Crocodile noir, and a Caiman a paujneres osseuses, inscribed the Crocodile vert. This 
must evidently have been an inadvertence, like the statement of the length of the nose ; 
but, as Cuvier observed, this is pardonable, as Adanson most probably named these draw- 
ings after he had forgotten them, and had been studying other things, long after his 
voyage, which occupied some of the first years of his youth. (See Cuvier, Oss. Foss. iii. 53. ) 



156 DE. J. E. GEAT'S STjS^OPSIS OF THE SPECIES 

A Caiman, in some of its characters, but which is nevertheless a true Crocodile, with 
the canines fitting into a notch, and not into a pit, in the upper jaw, is, there cannot 
be any doubt, the Crocodile that Adanson referred to ; for it agrees with his descrip- 
tion in colour and in its ferocious habits. And further that it is the Crocodile that the 
French naturalists refer to, is proved by the fact, already recorded, that we have received 
from one of the persons employed by M. Dumcril at the Paris Museum a skeleton 
of a young specimen of the Black Crocodile of West Africa as the skeleton of the 
American Alligator palpebrosus of Cuvier. 

** Face very lonr/, slender; nasal not reaching to the nostril. Gavialian Crocodiles. 

7. Mecistops. 
Face subcylindrical, scarcely dilated in the middle ; orbits simple. Nuchal shields 
numerous, small, in two cross series. Cervical disk narrow, containing two or three 
pairs of plates. Dorsal plates small, all keeled, in six longitudinal series, lateral 
one nan-owest. Intermaxillary produced behind, and embracing the front end of tlie 
nasal. 

Mecistops, Gray, Aun. & Mag. Nat. Hist. Srd series, x. 273; Cat. Tortoises & Crocodiles B. M. 58. 
Huxley, Proc. Linn. See. iv. 15, 1859. 

This genus has some resemblance to the Gavials ; but the structure of the skull and 
the position of the teeth are those of a true Crocodile. 

Professor Owen observes, " There is, however, a very close resemblance in the elon- 
gate, slender proportion of the skull and the elongated festooned border of the jaws 
between this species and the Crocodilus schlegelii from Borneo." — Loc. cit. p. 158. The 
Crocodihts schlegelii is a Gavial. 

Dr. Falconer observes, " The nasal bones (in Mecistojis) are extremely narrow and 
attenuated, but, as in the true Crocodiles, they descend between the maxiUaries so as to 
project into a notch between the intermaxillaries. The same holds good in C. schlegelii, 
where, as with Gavials, the nasal terminates a short way in front of the orbits, and does 
not enter into the formation of the anterior portion of the beak " (p. 363). " This cha- 
racter is a good diagnostic mark between the Crocodile proper and the Gavials, 
separating C. schlegelii from the latter genus,* under which MuUer ranged it" (p. 363). 

Dr. Balfour Baikie states, " In all essentials the skuU of the Mecistops shows it is to 
be properly classed as a member of the family Crocodilida; rather than the Gavialidte. 
The teeth are irregular, the sides of the jaw are not parallel ; there is a distinct swelling 
opposite the ninth remaining upper molar ; and the lower canines are received into 
notches in the upper jaw." — P. Z. S. 1857, p. 58. 



OF RECENT CEOCODILIANS. 



157 



Mecistops cataphractcs. (African False Garial.) (Plate XXXII. figs. 1, 2, 3.) 

Crocodilus bisndatus, Cuvier, Oss. Foss. iii. 53, 65, t. 5 (very young). 

Crocodihis bisulcattis, Bory, Diet. Class. N. H. v. lOB, misprint. 

Crocodilus cataphradus, Cuvier, Oss. Foss. v. t. 5. f. 1, 2 (erocodile k nuque cuLrassee) ; [copied 

A. Dum. Arch, du Mus. x. t. 14. f. 2]. Dum. & Bib. E. G. iii. 126 (young). Bennett, Proc. 

Zool. Soc. 1834, p. 110. Owen, Cat. Osteol. Spec. Mus. Coll. Surg. p. 155. n. 710 (Cuvier's type) . 
The Crocodile, Bowdich, Madeira, 232. 
Crocodilus leptorhynchus, Bennett, Proc. Zool. Soc. 1835, p. 129. A. Dum. Arch, du Mus. x. 252 & i. 

171, t. 14. f. 1. 
Mecistops cataphradus, Gray, Cat. B. M. 58. 
Mecistops benndtii. Gray, Cat. B. M. 57. 
Gavial of Senegal, Gray, Rep. Brit. Assoc. 1862, Sect. 107. 
Mecistops, Balfour Baikie, Proc. Zool. Soc. 1857, p. 58. 

Hah. West and Central Africa ; ? Fernando Po {Bennett), Gaboon, Lagos. Central 
Afiica, River Binue {Baikie). 

The species has been described from small young specimens. It grows to a laro-e 
size. There is an imperfect specimen which is scarcely adult, in the British Museum, 
that was sent from Fernando Po by Capt. E. F. Burton, which must have been L3 or 14 
feet long. Unfortunately it wants the head ; the body is 5 feet and the tail %\ feet long. 



Fig. 15. 




Fis. 17 




Fis. 16. 



Fig. IS. 





'Wir 



w* 



Figs. 15-18. Head and cei-vical shield of Mecistops cataphmctus. 



The specimen, originally sent by Mr. Bennett, was said to have come from Fernando 
Po; but Dr. Balfour Baikie observes that Fernando Po is a small volcanic island, 
totally without the muddy rivers delighted in by Crocodiles, and possessing nothing but 
streams, which duiing the rainy season are tumultuous mountain-torrents, with rocky 
beds.— Proc. Zool. Soc. 1857, p. 58. 

VOL VI. PART IV. Z 



158 DR. J. E. GRAY'S SYNOPSIS OF THE SPECIES 

Most probably Mi'. Bennett's specimen came from the coast, and was only received 
through agents at Fernando Po. 

C'uvier, in his Essay, described, under the name of Crocodilus hiscutatus, and figured 
the nuchal shields at t. 2. f. 6, a species of Crocodile founded on a specimen in the French 
Museum, which is labelled in Adanson's hand " Gavial du Senegal" and also on a 
very mutilated stuffed specimen which Cuvier found in the Museum of the Academy 
of Sciences at Paris (see Oss. Foss. v. 53, 65, t. 2. f. 6). He observes: — "the colour 
of these specimens is scarcely darker than that of the common Crocodile ; therefore it 
cannot be the Black Crocodile of Adanson." And he further specially remarks that 
" the jaws are a little longer and narrower than those of C. vulgaris, but not so long and 
slender as those of C. acutus." 

It resembles the latter in the dorsal shield of the vertebral line being only slightly 
keeled; but its peculiar character is that the middle of its nape is armed with two 
large pyramidal shields, and with two smaller ones in front of them. 

This Crocodile has been a paradox until this time. MM. Uumeril and Bibron 
regarded this mutilated specimen as only a specimen of the American Crocodile 
(C. americanus) with an anomalous development of the cervical and nuchal shields, 
observing that the specimens of this species are liable to variation in this respect ; but 
yet they do not describe any as exactly resembling C'uvier's description or figure. 

It does not appear that the specimen labelled by Adanson came under the 
examination of these naturalists; at least I cannot find any reference to it in their 
work. Cuvier unfortunately does not state its size ; but I have a strong opinion that 
it must have been a very young specimen of Mecisstops cataphractus before its elongated 
jaws were developed, and that the name of Gavial du Senegal was very applicable to 
it ; the back is grooved, by the flatness of the vertebral series of shields, as described 
by Cuvier, and as is characteristic of the American Crocodile (C. acutus) with which 
MM. Dumcril and Bibron compared it. But this is a question that can only be solved 
by the examination of the original specimens. 

Cuvier, in his Essay (vol. v. p. 58), observes, "When in England in 1818', I saw at the 

' I recollect this visit ■n'itli pleasure ; for I was deputed b}- Dr. Leach to show this celebrated naturalist and 
wavering politician some of tlie natm-al-liistory treasures, and also some of the social and political peculiarities of 
the metropolis, such as the Tower, the Bell and Lancaster and other schools, &c. Among the rest, I took him 
to the Westminster election, at Covent Garden. Being known to Sir Francis Burdett, I took M. Cuvier on 
to the hustings, and introduced him to some of the Westminster notabilities, whom he knew by reputation, 
and was anxious to see in person. Ho was so interested in these liygone saturnalia that we lingered tob 
long; for when Capt. Murray Maxwell attempted to speak, we were glad to " duck our heads " to avoid the 
cabbage-stumps, rotten eggs, and dead cats and dogs with which the Captain was assailed ; and when the mob 
attempted to take the hustings by storm, and were only driven off by the men-of-war's men who were retained 
by Capt. Murray's committee, wo found it difficult to retreat. Cuvier visited England again in 1830, during the 
short revolution which placed Louis riiilippe on the throne. WliUe here, the Zoologists in\-itcd him to a dinner 
at the All)ion Tavern : ho was greatly pleased with what he called the almost royal magnificence of the 



OF EECEXT CKOCODILIAXS. 159 

Museum of the College of Surgeons a dried specimen of a Crocodile." This he 
describes and figures under the name of •■'■Crocodile a nuque cuirassee"' {Crocodihts 
cataphractus, nob.). 

In 1834 Mr. Edward Turner Bennett (Proc. Zool. Soc. ii. p. 10) gave a notice of a 
specimen of Crocodihts cataphractus of Cuvier being alive in the gardens of this Society. 
At the meeting of the Society on the 22nd September, 1835 (Proc. Zool. Soc. iii. p. 129), 
after the animal had died, on more close examination, he described this animal as a new 
species, under the name of Crocodilus leptorhynchus ; and Mr. Martin added some notes 
on its internal anatomy. 

It is to be observed that Mr, Bennett and I were misled on this occasion by the 
erroneous breadth given to the animal in the figure published by Cuvier ; for he speaks 
of the length of the head "being to its breadth as 3 to 1," instead of as 2\ to 1. 

In my Catalogue of the Tortoises, Crocodiles, and Amphibians in the Collection of 
the British Museum, published in 1844, I formed a genus under the name MecistojJS 
for this animal, and for the first time described a full-grown specimen of it which we 
had received from the Gambia as M. bennetti; for Mr. Kendal considered it distinct 
from Cuvier's animal, but observed that they might be varieties. 

This might all have been avoided if we could have seen the original specimen ; but 
when I inquired for it, it could not be found. 

The specimen described and figured by Cuvier is fortunately now to be seen in the 
Museum of the College of Surgeons, referred to under No. 710 in the Catalogue of 
Osteological Specimens of that collection. It is a young dried specimen of the Crocodile 
which is now so frequently brought from the west coast of Africa, and it affords no 
ground for the supposition of M. Dumeril, expressed in his paper " On the Eeptiles of 
Western Africa " (Ai-ch. du Mus. v. 252), that these may be distinct species ; and it 
shows that the figure of Cuvier, though characteristic, is not very carefully drawn, and 
that any diffei'ence that may appear results from the want of accuracy in the figure, and 
is not to be found in the animal itself, — supporting the opinion that I expressed in my 
paper in the 'Annals and Magazine of Natural History,' 3rd series, x. p. 274. 

M. Auguste Dumeril, in his paper " On the Reptiles of Western Africa " (Archiv. du 
Mus. X. 271), gives a good figure of a half-grown specimen of this species under the 
name of Crocodilus leptorhynchus, 1. 14, and places by the side of it a tracing of Cuvier's 
figure of Crocodilus cataphractus, to show that they cannot be alike; but the 



eutertainment. Dm-iug the dinner the news arrived that the Orleans party had succeeded : he and his ste])- 
daughter, Miss Duvaucel (who was in the gallery wdth some ladies), immediately displayed the national 
colours. Cuvier's political predilections were not strong ; for he had held office under Napoleon and under the 
Boiu'bons, and he made no secret that he came provided so as to acknowledge the success of either party : 
he had a white and a tricolour cockade in his hat ready to show as the occasion required. ^Micn I visited 
him in after times, he more than once referred to the events of his visits. 

z2 



160 DB. J. E. GRAY'S SYNOPSIS OP THE SPECIES 

comparison of the specimens on which these species were founded shows how much 
better it is to refer to nature than to depend on figures and descriptions, which are 
liable to the imperfection attending human observation and record. 

Dr. Falconer, in the 'Annals and Magazine of Natural History ' for 184G (xviii. 362, 
t. 6), described and figured a skull of this species under Cuvier's name, which was in 
the Belfast Museum, and said to have been sent from Sierra Leone. 

Dr. Balfour Baikie described the skull of a specimen from the River Binue (see 
Proc. Zool. Soc. 1857, p. 58). 

Family III. ALLIGATORIDiE. 

The upper and eleventh lower teeth longer, like canines, the canines of the lower jaw 
fitting into holes or perforations on the edge of the upper jaw. 

AlIigatoridcB, Gray, Cat. Tortoises &c. B. M. 56, 18J4. Huxley, Journ. Proe. Linn. Soc. iv. 3. 
Alligator, Cuvier. Gray, Ann. Phil. x. 195. 

Teeth strong, unequal ; the hinder ones differ in shape from the anterior. The front 
pair of mandibular teeth, and the fourth pair- (canines) are received into pits on the 
edges of the prromaxilla and maxilla;. The mandibular teeth behind these pass inside 
and not between the maxillary teeth. The premaxillo-maxillary suture on the palate 
is straight or convex forwards. The symphysis of the lower jaw is short. 

Spix, in his work on Brazilian Lizards, gives very good figures of the Alligators, with 
the colours well marked. The Memoir on South-American Alligators by Natterer, 
contains very accurate and detailed figures of the head and the neck-shield of the 
different species. He has figured some varieties or species very nearly allied to those 
here noticed, wliich have not come under my observation. 

Spix divided the Alligators into two genera: — Jacaretinrja, with acute nose (1. J. 
inoscMfer,i.\ = CaimaniKdpd)rosus, p. 161 ; 2. J. punctulatus, t. 2=Jacare imnctulata, 
p. 159); and Caiman, or Jacare, with blunt nose (including 1. C. niger, t. i=Jacare 
nigra, p. 167; 2. C.Jissipes^Jacare latirostris, p. 107). 

His figures are very good representations of the species — indeed, the best knowli. 

MM. Dumeril and Bibron admit the three species described and figured by Spix, 
thus : — 

1. A. sclerops, p. 74 ; Caiman noir, Spix, Bras. t. 4. Head elongate, fiuttened, a ridge 
in front of each eye, the upper eyelid Jinely striated. Nape with two roivs of small 
oval compressed scales. Back with two central longitudinal ridges, the three last cross 
bands of six keeled scales. Black, yellow-banded. I have no specimen agreeing with 
the account of the nuchal scales and the eyelid of A. sclerops : according to Spix the 
dorsal scales are elongate. 

2. A. cijnoceplialus, p. 86, Caiman Jissipes, Spix, Bras. t. 3. Head short, broad, 
thick, a ridge in front of each eye, the upper eyelid rugose. Nape witli two roivs of 



OF RECENT CROCODILIANS. 161 

large square keeled shields. Back scale keeled, the three last cross lands of four scales 
Sides with some strong keeled scales. Back green, black-dotted. 

3. A. punctulatus, p. 91, Spix, Bras. t. 2. Head elongate, nose flattened, with a 
rounded point in front, without any j)reocular ridges, upper eyelid rugose. Nape with 
two rows of shields. Back flat, scarcely keeled. Sides with some larger scales. Yellow, 
black-dotted. 

John Natterer, in his "Beitrag zu den Siid-Amerikanischen Alligatoren," edited by 
Fitzinger, describes eight species of the genus Champsa : five have partly bony eyelids, 
and three have them entu-ely bony. The five former belong to the genus under con- 
sideration. 

The preorbital ridge distinct, beak broad with three lateral foveolsB, eyelid striated, 
beak broad and blunt. C. nigra, t. 21. 

The nuchal scutella many, in three series. C.jissijies, t. 22. 

The nuchal scutella many, in two series. C. sclerops, t. 23. 

The preorbital ridge evanescent, beak without lateral foveolse, eyelids rugose. The 
frontal ridge flexuous, bent in front. C. vallifrons, t. 24. 

The frontal ridge arched, bent back. C. punctulata, t. 25. 

M. Natterer gives the following proportional measurements of the heads : — 

Length of Width of Width of Beak 
Length of Width of Crown Crown above the 

Head. Head. before. before. eighth tooth, 

in. 1. in. 1. in. 1. in. 1. in. 1. 

Champsa nigra 10 8 3 6 4 !) 5 1 

fissipes 10 3 6 5 2 7 3 5 4 

sclerops 6 6 5 8 2 8 3 3 3 3 

vaUifrons 7 10 4 6 2 2 9 2 3 

punctulata 10 5 5 4 2 .5 3 2 2 5 

The figures of the heads of the last two species differ from that of C. sclerops chiefly in 
the nose being narrower (C. punctulata being the narrowest and very slender), narrower 
than in any specimens that have come under my observation ; the lower jaws in the 
figure also differ in shape, that of C. vallifrons being the most slender. Dr. Strauch, 
who had M. Natterer's specimens to examine, regards the two latter as the same species, 
but distinct from sclerops. 

Synopsis of Genera. 
I. The ventral scutella like the dorsal ones, bony and articulated together, forming a 
shield. The eyelids tvifh an infernal bony plate. The cervical scutella in pairs, 
forming an elongated shield. Nasal bone short. Tropical America. 

1. Jacaee. The orbits united by a bony cross ridge. Eyelids partly striated or 

rugose. 

2. Caiman. The orbits not united by a cross ridge. Eyelids bony, entirely smooth. 



162 DK. J. E. GEAY'S SYNOPSIS OF THE SPECIES " 

II. The ventral scutella thin, the dorsal scutella bony, not articulated together. The 
eyelids fleshy, smooth. The cervical scutella in pairs, separate. Nasal lone 
elongate, separating the nostrils. North America. 

3. Alligatok. The face broad, depressed. 

Section I. The ventral scutella like the dorsal ones, bony and articulated together, 
forming a shield. The eyelids ivith an internal bony plate. The cervical scutella 
in pairs, forming an elongated shield. Nasal bone short. Tropical America. 

1. Jacare. 
Head moderately high, shelving on the sides. Orbits united by a distinct bony 
cross ridge. Eyelids striated or rugose, strengthened by a small internal bone. The 
cervical scutella four or five pairs, forming a shield ; the dorsal and ventral scutella 
both consolidated together, forming a dorsal and ventral shield ; the gular and ventral 
scutella smooth. 

./«ca;-e, Gray, Cat. Tort.Croc.&c. B.M.64,18M'; Ann. & Mag. N. H. 3rd series, x. 327, 1862. Huxley, 

Proc. Linn. Soc. 1859, 4. 
.Jacuretinga, Spix, Lacert. 

The pits in the maxilla are the ca-\'ities left by the preorbital ridges as they advance. 
The intermaxillary bone short, truncated behind, with an elongate-oval or lanceolate 
ca^■ity between this and the front of the palate. 

The figures of Natterer are excellent to general appearance, but they do not agree 
with the measurements of our specimen ; that is to say, the nose of Champsa flssipes, 
from the ridge, is about the same length as the forehead, but in his figure it is 
represented as larger, and it is so in all the nthor figures: perhaps this is to allow for 
the perspective. 

A. Head elongate ; interorbital ridges strong. Dorsal scutella elongate, keeled, keels of 
vertebral series highest ; lumbar scutella in six longitudinal series. Nuchal scutella 
small, compressed. Eyelids striated, with a rather large internal bone. Back 
black varied with yellow. Melanosuchus, Gray, Ann. £■ Mag. N. H. x. 328. 

1. Jacake nigka. (Black Jacare.) 

CrocodUus sckrops, Schn. Ampli. 162. Blainv. Osteogr. Crocod. t. 3. f. 2, t. 4. f. 13. 

Crocodilus yakare, Daud. 

Alligator scler ops, Cuvier, Oss. Foss. v. 35, t. 1. f. 6 & 7, t. 2. f. 3. Briihl, Skclct. Krokod. t. 12. 

f. 3, 5, 6, 7, t. 19. f. 21. 
Alligator sckrops, var., Gray, Syn. Kept. 
Caiman niger, Spix, Bras. t. 4 (good) . 
Champsa nigra, Natterer, Beitr. t. 21 (good) . 



OF EECXENT CROCODILIAN S. 163 

Alligator niger, Owen, Cat. Osteol. Spec. Mus. Coll. Surg. p. 704. n. 166 (adult). 
Jacare nigra, Gray, Cat. Tort. & Croc. 65 ; Ann. & Mag. N. H. x. 328, 1862. 

Hah. Para, 13 feet long {Graham); Guiana {Owen). 

I think it better to adopt Spix's name, as sclerops has been used for all the species. 

B. Head short; orbits with diverging ribs in front to edge of Jatv. Dorsal scutella 
broad, slightly keeled, equal; the lumbar scutella in four longitudinal series. 
Nuchal scutella distinct, in two cross series. Eyelids rugose, tvith a small internal 
bone. Back olive, handed with brown. Cynosuchus, Gray, Ann. & Mag. N. H. x. 328. 

In many of the specimens the first scale of the nuchal shield has two keels, in others 
it has only one ; but in several specimens the scale has two keels on one side and only 
one on the other. 

a. Head short, broad, depressed, with very distinct preorbital ridges to the edge of tltf 
jaw. Cervical disk short, broad, formed of four bands of scutella. Sides of jaws 
pale, with a series of dark spots. 

2. Jacabe latirostris. (Dog-headed Jacare.) 

Dorsal shields in eight longitudinal series, four on each side. Ventral shields in 
twelve series. 

Crocodilus latirostris and C. yacare, Daud. Rept. ii. 407, 417. 

Caiman fissipes, Spix, Bras. t. 3 (good). 

Champsafissipes, "Wagner, Icon. t. 17. Natterer, Beitr. t. 22 (good). 

Crocodilus sclerops, Wied. Abbild. t. Blainv. Osteogr. Crocod. t. 3. f. 2, t. 4. f. 13. Schinz, Rept. 

t. 112. 
Jacare fissipes, Gray, Cat. Tortoises B. M. 64. 
Alligator sclerops, Pr. Max. Abbild. t. 
Alligator cynocephalus, Dum. & Bib. Erp. Gen. ii. 86. 
Jacare latirostris, Gray, Ann. & Mag. N. H. x. 328. 

Hah. Brazils ; Pemambuco (/. P. G. Smith) ; Surinam. 

The nose of the young specimen is as long as the width at tlie eiglith tooth. Tlie 
nose fi-om the ridges nearly as long as the back of the head ; width of the muzzle at 
the notch one-half the length of the head. 

Var. 1 (three young, in spirit). Head short; side of face pale, with a dark spot 
under each ear, and another larger under each eye. The lower jaw pale, five round 
spots on each side, the middle one, under the eyes, the largest. Back black, with inter- 
rupted or iiTegular pale brown cross bars. 

Hah. Pernambuco {J. P. G. Smith). 

The smaller specimen is peculiar for the very small size of the ventral shield in front 



164 DK. J. E. GBAT'S SYNOPSIS OF THE SPECIES 

of the vent. The spots on the side of the face and lower jaw are to be seen in the 
older specimens when they are between 3 and 4 feet long. 

Var. 2. Head rather larger and narrower. The nose from the ridge rather longer than 
the back of the head ; width of the notch two-fifths the length of the head. Cheek 
and side of the lower jaw with five large black spots. Ventral shields in twelve series. 
Dorsal shields four. 

Hab. South America ; Lake of Santa Cruz de la Sierra. 

3. Jacake multiscutata. (Brazilian Jacare.) 
With sixteen series of ventral shields ; hinder ventral shields very narrow. Dorsal 
shields in ten longitudinal series, five on each side. 
Hab. Brazil. 
A skin in the British Museum (46. 7. 10. 41). 

b. Head elongate, longer than the width at the eighth tooth, with none or only indistinct 
evanescent ridges from the front of the orbit. Cervical disk oblong, elongate, of 
five series of scutella. 

* Face depressed, broad ; sides of the jaws with a series of large coloured spots. 

4. Jacare longiscutata. (Long-shielded Jacare.) (Plate XXXIV.) 

Dorsal scutella elongate, longer than broad, uniformly keeled, in ten longitudinal 
series on the middle of the body; ventral scutella elongate, in fourteen or sixteen 
longitudinal series. Sides of the jaws pale, with five or six band-like spots ; the inner 
pairs of the first and second series of cervical scutella large and equal-sized. 
Jacare longiscutata, Graj', Ann. & l\Iag. N. H. x. 328, 18G2. 

Hab. South America. Brit. Mus. 

This is very like the following ; but the head is rather broader, and the dorsal and 
ventral shields are much larger, and more numerous. 

It is known from the young of Jacare nigra by its olive colour, the spots on the sides 
of the jaws, and the presence of the distinct nuchal scutella. 

5. Jacare ocellata. (Eyed Jacare.) (Plate XXXIII.) 

Dorsal scutella broad, uniformly keeled, in eight longitudinal series in the middle of 
the body ; ventral scutella in twelve longitudinal series, the hinder ones smaUcr, longer, 
and more numerous ; the central pair of cervical scutella in the first series smaller than 
those that follow. 
Jacare ocellata, Gray, Ann. & Mag. N. H. x. 329, 1862. 

Hah. Lake of Santa Cruz de la Sierra. British Museum. 



OF KECENT CROC'ODILIAXS. 165 

** Face affemtafed, raflier luijh on flie sides; sides of tJie jaios one-coloured. 

G. Jacare pu.vctulata. (Dotted-jawed Jacare.) 
Back yellow, banded with brown; the sides of the head yellow; upper and lower 
jaws yellow, one-coloured, or minutely speckled ; sides of the neck smooth, with flat 
scales. Nose rather high and square. 

Jacare sclerops, Gray, Cat. Tortoises B. M. 64. 

CrocodUus sclerops, Schn. Amph. 162. Cuvicr, Ann. Mus., & Oss. Foss. v. t. 1. f. 6 & 7, t. 2. f. 3. 

Tiedem. Amph. 60, t. 5. Gnerin, Icon. t. 2. f. 2 & ]0. Gray, Syn. Rept. 62. Dum. & Bib. 

Erp. Gen. iii. 79. 
CrocodUus americanus, Laur. from Seha, t. 101. f. 10. 
CrocodUus caiman, Daud. Eept. iii. 394. 
Caiman (Jacaretinya) punctulutus, Spix, Bras. t. 2 (goorl). 

Champsa sclerops, ^Yagner, Syst. t. 7. f. 1, 2, and f. 42. Natterer, Beitr. t. 22 (heads good). 
Alligator punctulatits, Dnm. & Bibr. Erp. Gen. ii. 91. 
Jacare punctuluta, Gray, Ann. & Mag. Nat. Hist. x. 329, 1862. 

Hah. Brazil [Sj)ix) ; Surinam ; Argentine Eepublic (//. Clirisfij). 

Natterer figures two other species, under the name of Champsa vaUifrons (t. 24), 
{Jacare vaUifrons, Gray, Cat. B. M. 05), and Ch. punctulata (t. 25) {Jacare punctnhifa. 
Gray, Cat. B. INI. 65), which seem to differ from the former in the head being narrower 
and more tapering. I have seen no specimens agreeing with these figures; but they look 
very like varieties of the above. At the same time, some of our specimens appear to 
have a more attenuated snout than others ; but when you apply the callipers to the 
nose and to other parts of the head, the absolute proportions of the parts are veiy 
nearly the same. 

A stuffed specimen from the Ai'gentine Eepublic measures 6 feet 9 inches long, the 
head from the occiput is IO5, and the nose from the ridge 6J inches. In another, from 
the Zoological Society's Gardens, 5 feet 10 inches long, the head from the occiput is 
10 inches, the nose ii-om the ridge 6^ inches long. A series of young specimens in 
spirits are pale brown, the back and tail with narrow brown cross bands, those on 
the back sometimes broken into square spots; the cheek and outside of the lower jaw 
pale yellow, without spots. The sides of the nuchal disk dark-coloured. 

7. Jacaee hirticollis. (Rough-necked Jacare.) 

The scales on the sides of the neck rough, spinulose, pale yellow ; back and tail 
brown, cross barred ; cheek and sides of the lower jaw yellow, not spotted. 

Hub. Demerara. B. M. 

I may observe that, characteristic as are the figures of Dr. Natterer's paper, none of 
them exactly agrees in measurements with the specimens in the British Museum. 

In some specimens of the Jacare the first and, sometimes, even the second cervical 

VOL. YI. PART IV. 2 A 



166 DE. J. B. GBAT'S SYNOPSIS OF THE SPECIES 

scutella have two keels, in others only one ; but this is no specific distinction ; it is not 
rare to find species with two keels on one side of the neck, and only one on the other. 

2. Caiman. 

Head high, flattened on the sides, angulated above. Orbits without any ridges. The 
eyelids smooth, strengthened with a large, single, internal bony plate. The dorsal and 
ventral scutella bony, articulated together, forming a dorsal and ventral shield ; the 
gular and lateral ventral plates keeled, the abdominal ones smooth ; the cervical 
scutella four or five paks, with sometimes one or a pair interposed between the second 
and third pairs. 

Skull with the superior temporal fossse obliterated, the circumjacent bones uniting, 
the eyelid with a single large bony plate covering the whole upper surface. Vomer 
not apparent on the palate. 

Caiman, Gray, Cat. Tortoises &c. Brit. j\Ius. 66, 1844. ; Ann. & :\Iag. Nat. Hist. 3rd series, x. 330. 
Huxley, Proc. Liiin. Soc. iv. 3. 

This genus has been divided into two species — one having the cervical shields two, and 
the other four in a cross scries ; in all the latter there arc two in a cross series, with 
one or two interpolated between the shields. 

I have seen no specimen which agrees in the the nuchal shields with cither of the 
figures in Cuvier, Oss. Foss., though our two species agree in other respects with his 
figures ; and how such species with distinct organic characters could be regarded as 
varieties, I am unable to learn. 

I cannot conceive what induced M. Cuvier in his ' Essay ' to consider the two South- 
American Alligators with bony eyelids varieties; for he justly observes, "The Crocodile 
of St. Domingo is not more distinct from the Crocodile of the Nile than these two 
varieties are from each other." In the Latin synopsis of the species, which is appended 
to the paper, they arc regarded as distinct, and the second one is called C. trigo- 
natus. Yet MM. Dumeril & Bibron, in their \a ork, ])ersist in following Cuvier's 
first idea of their being only varieties, and in regarding Adanson's specimens as 
belonging to the second vai'icty, and also in doubting if the " two varieties," are both 
from America. 

The specimen in tlie British Museum proves most distinctly that there are two very 
distinct 'Alligators with bony cj-elids found in Tropical America ; wliich agrees well 
with the character that M. Cuvier and MM. Dumeril & Bibron give to the two varieties 
of that species ; and these species are, as Cuvier observes, as distinct from one another 
as C. americanus from C. vulgaris. The heads of both these species are figured by Dr. 
John Natterer in his "Essay on American Alligators" in the Vienna 'Transactions.' 
This autlior also figured a third species, which he calls A. gihbicei)S, which, if it is 
separable from A. trigonatus, must be distinguishable from it by very slight characters. 

The Black Crocodile [Ilalcroiiia palj>ebrosa) of West Africa has so much resemblance 



OF RECENT CROCODILIANS. 167 

to this animal that Cuvicv considered Adanson's West-African specimen a variety of 
this species. 

Dumeril & Bibron evidently considered the African and the American animals 
the same species; and we a short time ago received from M. Braconier, of the Jardin 
des Plantes, a skeleton of the African species under the name of AUi(/ator paJpe- 
hrosus, var. 

A. Head shelving on tlie sides. Nuchal sciifella in a single cross series, cervical scutella 

Jive pairs; dorsal scutella highly keeled, irregular, in six series; the lumbar 
scutella in tioo longitudinal series; the gular and two outer lateral series of 
ventral scutella keeled. The flat upper disk at the base of tail broad and strongly 
crested. Paleosuchus, Gray, Ann. & Mag. N. H. x. 330. 

1. Caiman teigonatus. (Rough-backed Alligator.) 
Crocodilus triyonutus, Scliu. Ampli. IGl. 6. Tiedemann, Amph. 66, t. C7. 
Crocoditus ■palpehrosus, var. 2, Cuvier, Oss. Foss. v. 40, t. 2. f. 1. 

Caiman trigonatus, Gray, Cat. Tortoises &c. B. M. 66; Aim. & Mag. N. H. x. 330, 1862. 
AlUyator indpchrosus, Briihl, Skel. Krok, t. 19. f. 3. 
Chainpsu trlyonata, Natterer, Beitr. t. 26 (good). 

Ilab. Tropical America. 

The largest specimen in the Biitish Museum is rather above 4 feet long. The young 
specimens have the lateral ventral shields keeled. 

B. Head flat, and erect on the sides. Nuchal scutella many, in two cross series; 

cervical scutella three 2)airs ; dorsal scutella slightly keeled; the lumbar 
scutella in four longitudinal series; the gular, the ventral, and the lateral 
abdominal scutella keeled. The flat upper disk at the base of the tail elongate. 
Aromosuchus, Grray, Ann. & Mag. N. H. x. 830. 

2. Caiman palpebrosus. (Banded Alligator.) 

Brown ; tail black-banded. 

Crocodilus palpebrosus, -vax., Cuvier, Oss. Foss. v. t. 1. f. 6-17 and t. 2. f. 2. 

Champsa palpebrosa, Natterer, Beitr. t. 27 (good). 

Caiman (Jacaretinya) moschifer, Spix, Bras. t. 1 (skull). 

Caiman palpebro»us. Gray, Cat. Tortoises &c. B. M. 67 ; Ann. & Mag. Nat. Hist. x. 330, 1862. 

Crocodilus palpebrosus, Tiedem. Nat. Amph. t. 6. 

Alligator palpebrosus, Merrem, Syst. 35. Gray's Syn. Rept. 63. 

Hcd). Tropical America. 

Natterer figures the head of a species under the name of C. gibbiceps ; but 1 do not 
see how it differs fi-om the above, except that the head is a little higher — perhaps a 
sexual distinction. Dr. Strauch regards C. gibbiceps as the same as C. palpebrosus. 

2a2 



168 DE. J. E. (iKAT'S SYNOPSIS OF THE SPECIES 

Section II. The ventral scutella thin, the dorsal scutella honij, not articulated together. 
The eyelids fleshy, smooth. The cervical scutella in pairs, separate. Nasal hone 
elongate, separating the nostrils. North America, 

3. Alligator. 
Head depressed, broad, witliout any ridges in front of the orbit. Snout very broad, 
flattened and rounded at the end ; the ninth maxillary tooth the largest. The eyelids 
smooth, fleshy. The dorgal scutella not articulated together, in six longitudinal series ; 
the ^entral scutella thin ; the gular and abdominal shields smooth ; nuchal scutella one 
pair,' small ; cervical scutella three pairs, hinder smallest. Nostril separated by a bony 
septum. The feet webbed. Dorsal plates in six longitudinal series, the two vertebral 
closer together. The sides with a short series close to the others, sometimes reduced to 
only one or two shields. 

Alligator, Gray, Cat. Tort. B. M. 6G ; Ann. Mag. N. H. x. 330, 1862. Huxley, Proc. Linu. Soc. iv. 3. 
Champsa, Wagler, Syst. d. Amph. 140. 

Alligator jiississippensis. (Alligator.) 

Alligator, Catesby, Carol, t. 63. 

Crocodilus mississippensis, Daud. Rept. ii. 413. 

Crocodilus luciiis, Cuvier, Ann. Mus. x., and Oss. Toss. v. 1. 1 . f. 8 ; t. 2. f. 4. Tiedera. Amph. 58, t. 4. 

Merrem, Zool. 34. Owen, Cat. Osteol. Spec, in Coll. Surg. p. 165. n. 760, 761. Blainv. Osteog. 

Crocod. t. 2. f. 1, t. 5. f. 1. Bruhl, Skelet. Krokod. t. 8. f. 5, 6, t. 9. f. 3, t. 10. f. 3, 4, 1. 11. f. 2, 3, 

t. 20. f. 
Alligator mississippensis, Gray, Cat. Tortoises B. M. 66 ; Ann. & Mag. Nat. Hist. x. 331, 1862. 
Crocodilns cuvieri, Leach, Zool. ISIisc. ii. 117, t. 102. 

Alligator luciiis, Merrem, Tent. .34. Dum. & Bibr. Erp. Gen. iii. 75, t. 25, 26. 
Alligator cuvieri, Bory de St. Vincent, D. C. H. N. v. 104. 

Hah. North America, New Orleans, Texas. 
Yar. 1. The nose very broad and short. The largest specimen of this variety in the 

British Museum is nearly 4 feet long. 
\-A\-. 2. The nose narrower and longer. The largest specimen in the British Museum 
is of the same size as the former, which is nearly 4 feet long. Are they the two 
sexes ■? 
The young specimens in spirits have the back black, with narrow white cross bands. 
The head pale brown, black-varied. Ventral shields in eight or ten longitudinal rather 
irregular series. 

There is a very young specimen of this species in spirits, from New Orleans, in the 
British Museum. It is black, with white cross bands. The beak is short, rather 
slender, with a ridge of skin in front of each eye, giving the appearance of a frontal 
rid<>e. 



OF EEUENT CROCODILIAAS. 169 

EXPLANATION OF THE PLATES. 

PLATE XXXI. 

Figs. 1, 2, 3. Skull of Bombifrons indicus. Adult. 
Figs. 4, 5, 6. Skull oi Halcrosia nhjra. Half-grown*. 

PLATE XXXII. 

Figs. 1, 2, 3. Skull oi Mecistops cataphractus. Adult. Length 21 inches. 
Figs. 4, 5, 6. Skull of Molinia intermedia. Adult. Length 30 inchesf . 

PLATE XXXIII. 

Tdcare ocellata. Young : stuffed. Natural size. 

PLATE XXXIV. 

Jacare longiscutata. Young : stuffed. Natural size. 

* CuUed on the Plate " Halerosia frontdta.'' t Called on the Plate " Crocodihis intermedim." 



SnM%ol:^ocm6M'Jf 








K*<i;;v.' 






ft 







1,2,3. Bombifrojis indicus. 
4,5,6- Hadicrosia frontata. 



cfhimA7wc.Arm:6:fm. 





GEFcrd, 



1, 2, 3, LECISTOPS CATAPHRACTUS. 
4', b, b, CROCODILUS INTERMEDIUG. 



■WWest imp 










*A 

pq 
o 
o 






I 

a: 



-.1 



5: 




O 



o 



3 

-5 



M 



[ ni ] 



Vlll. ^ofe to MeriKJir on flip Induni ('efacfa roUcrfed hi/ Sii; Walter Elliot. 
By Professor Ow£X, F.B.S., F.Z.S.. ,ix: 

Eeatl May Srth, 1867. 

In relation to my paper on Indian Cetacea, read before this Society on the 2Gth of 
June 1S6-J, and published in the Society's 'Transactions'', I have received the following 
letter from Sir Walter Elliot, K.S.I., F.Z.S., to whom I was indebted for the specimens 
upon which my observations were based. 

Travellers' Club, 15th April. Is(j7. 

" Dear Peof. Ouex, — Soon after my arrival in town a few weeks ago, my attention 
was called to some of the details in your paper on Indian Cetacea, in the Zoological 
Society's ' Transactions.' In replying to some inquiries of Mr. Flower, at the College 
of Surgeons, regarding the skull of Fhijseter simus, I noticed that you had described 
two individuals, a male and a female, whereas I had never met with more than a single 
female specimen of this animal. I was puzzled to account for this ; but as Mr. Sclater, 
who was with me at the time, stated that the original drawings from which the Plates 
had been taken were at the Zoological Society's office, I took an early opportunity of 
referring to them. I also .sent to Scotland for a note-book in which I had entered 
remarks on specimens as they were obtained. On comparing these with your paper I 
found that the inaccuracies I had observed had been caused entirely by my own careless- 
ness in furnishing you with the scanty and imperfect materials on which your paper 
was founded, and by my omission to eliminate a faulty drawing. 

" You may remember that I fir-st brought the crania to you in 1863, to know whether 
you thought them of sufficient interest to be described. On my return to Scotland, I 
sent you drawings with some remarks of my own, but overlooked the faulty figure 
entirely, which thus remained in the packet with the true ones. In April 1865 you 
wrote to me for some further information with reference to the notes written on the 
drawings, and added tliat you could only find two skulls, although my notes referred 
to others. In reply I sent you copies of all the memoranda I could find, and said that 
the crania must be with you, as I had left them all at the Museum. I came to London 
some weeks later, and on calling to see you I found the crania were still missing ; but 
they were subsequently discovered, and your paper was prepared. 

" To account for the origin of the erroneous figure, I must premise that the office I 
held in Madras from 1849 to 1854 was a very laborious one. demanding my whole 

' Vol. vi. I). 17. 



172 PEOFESSOE OWEN ON INDIAN CETACEA. 

time, and leaving little leisure for other pursuits. Having always been fond of natural 
history, I kept a draughtsman continually engaged in depicting objects of interest. 
My house was on the sea-shore, and the fishermen from several miles along the coast 
used almost daily to bring me something or other which they considered to be rare 
or curious. But as I went to my office at 10 o'clock a.m., and did not return till 6 
or 7 P.M., my artist had orders to proceed with his sketches as soon as the speci- 
mens were brought to him. On my return home in the evening, my first business 
was to inspect his work. By dint of constant supervision, I brought him to the exercise 
of scrupulous accuracy. If I found the least mistake, I had another drawmg made the 
following day. He was principally engaged on naked mollusks, Crustacea, and the 
mutations of lepidoptera, which he drew with the aid of the microscope. To the 
exactitude of these, Messrs. Alder and Hancock's paper on the Indian Nudibranchs, in a 
former volume of the ' Transactions,' bears testimony. 

" The Wongu or Physeter was brought to my house on the 28th of February, 1853. 
On examining the sketch the same evening, I was not satisfied with it, and therefore 
directed a more accurate drawing to be made, which was done, under my inspection, 
early the following morning. I was much interested in the specimen, which I believed 
to exhibit an entirely new form ; and I made the following note in pencil on the back of 
the revised drawing, which is still legible : — ' If the description of the Porpoise-family 
is con-ect, this must be a very different genus. The mouth is smaU, very much under 
the rounded snout, not reaching so far back as the eye, which is far above it, in a line 
with the rounded snout. The blow-hole is in front of the eyes, and (in this individual') 
to the left of the middle line of the back, opening diagonally, with the points curving 
slightly backwards. Colour above shining black, smooth ; beneath pale, and in this one 
discoloured with blood. Fore part [i. e. in front of the dorsal) depressed ; behind the 
D. much compressed ; the part nearest the tail rising into a sharp ridge.' On the fiice 
of the drawing I wrote in ink ' new sp. Tel. name, tvomju: adult ? : Waltair, March I, 
1853;' and under the mouth I noted the dentition 'i-^.' On the first or cancelled 
sketch, the only writing was a note in the handwriting of the painter, ' Found at 
Waltair on the 28th Feby. '53,' and, in Telugu characters, the name ' wongu.' 

"Having completed the drawing, I made the following entry in my note-book : — 
'March 1, 1853. — A large Porpoise was brought by the Vizagapatam fishermen, of 
which the following is the description : — 

ft. in. 
Totid leiii,'tli 7 -2 

ft. in. fl. in. 

rriim muzzle to spiracle 7 

From spiracle to dorsal 2 'J 

From commcucement of dorsal to end of caudal . . 3 10 

7 2 

' I then thought this circumstance was accidental. 



PROFESSOR OWE\ ON INDIAN CETACEA. " 173 

t"t. iu. ft. ill. 

Length of dorsal from iascrtion in front to tip 1 

Breadth of ditto 11 

Liiteml measurements. 

Length from snout to insertion of pectoral 1 5 

Length from insertion of pectoral to vent 3 7 

J..ength from vent to centre of caudal 2 3 

7 3 

Inferior measurements. 

Length from snout to vent 5 

Length from vent to centre of caudal 2 3 

7 3 

From snout to eye 11 

Length of gape • 4 

Girth where largest (in front of dorsal) 4 4 

Length of pectoral 1 2 

Breadth of caudal 1 10 

Length of vent 7 

Length of small apertures at either side of ditto 2 

" ' This very remarkable animal does not agree with any known genus or species. 
The fishermen call it ivongu. The snout is rounded and blunt, the mouth, small and 
placed far below it, the teeth ^.^=20. The eyes considerably above the mouth, and 
neaily over the termination of or a little behind the gape ; the spiracle before the eyes, 
situated to the left of the dorsal or central line, obliquely placed as regards its length, 
slightly curbed and the points turning backwards. 

"'The colour is shining black above, growing gradually paler towards the belly, 
which is coloured in the drawing from the blood which had flowed over and stained it. 
The skin is quite smooth. The body in front of the dorsal large and much depressed. 
Behind the dorsal it becomes smaller and compressed more and more towards the caudal, 
the latter half (between the dorsal and caudal) being compressed into a sharp ridge, 
which runs into the base of the caudal fin. 

" ' This was an adult female, from which was taken a single perfect foetus with the 
same peculiarities as the dam, viz. the diagonal spiracle on the left of the dorsal line, 
the points curving slightly backwards. The skin of this was stuffed, but was unfor- 
tunately carried away by a Jackal, when being dried in the garden.' 

"On my return to England eight or nine years afterwards, I showed my drawings to 
several persons interested in such matters. Those of the mammalia were for some time 
in the hands of the late Dr. Coldstream, who exhibited them at a meeting of the 
Natural History Society of Edinburgh. They were subsequently lent to several other.'< 
in Scotland, among them to a medical gentleman, surgeon to a whaling-vessel, who had 
paid much attention to cetacea, and who had them in his posses.sion for some time. 

" In the course of these migrations some liberties appear to have been taken with 

VOL. VI. PART IV. 2 B 



174 ' PROFESSOR OWEN ON INDIAN CETACBA. 

the drawings themselves. On the cancelled figure of the Physefer, I now perceive a 
faint pencil-note in an unknown hand, "wongao," which is wrongly spelt, and quite 
unauthorized. This has been followed by the addition of a male organ to the figure, 
also quite gratuitous. No wonder, then, that you were led into error ; and I cannot 
sufficiently blame myself for not having made a more careful scrutiny of the drawings 
before despatching them. 

" It also occurs to me that, owing to the fragmentary form in which my notes were 
communicated to you at different and distant dates, you may have overlooked the fuller 
and more careful description forwarded in my last letter ; for the text corresponds more 
nearly with the pencil-note on the drawing ; and the proportions assigned by you to the 
several parts of the specimen, which differ slightly from my measurements, appear to 
have been taken from the drawing above, which, though made by measurement, was 
not exactly according to scale. 

" I have gone into greater detail than perhaps was necessary in explanation of these 
circumstances, to enable you to make such corrections as you may think necessary to 
your very valuable and interesting paper, with reference to matters of fact in which I 
unfortunately misled you. 

" Yours very truly, 

"WALTER ELLIOT." 

" To Professor Owen, F.B.Sr 



i 



[ 1-5 ] 



IX. Contriiutions toivards a more comjjlete knotvledge of the Skeleton of the Primates. 
By St. Geokge Mivart, F.L.S., Lecturer mi Comparative Anatomy at St. Mary's 
Hospital. Part I. The Appendicular Skeleton of Simia. 

Read December 13th, 1866. 

[Plates XXXV. to XLIII.] 

XHE skeleton of the Orang-Outang, besides earlier notices, has been more or less 
carefully described by Owen', De Blainville", and W. Vrolik^. These descriptions, 
however, were anterior to the discovery of the Gorilla, which has necessitated fresh and 
more detailed examinations of and comparisons between the animals most nearly 
resembling Man. 

Such detailed and elaborate investigations have been instituted, and similar descrip- 
tions published, by Professor Owen* with regard to the skull and spinal column of the 
Gorilla, the Chimpanzee, and the Orang ; and Professor Huxley^ has carried yet further 
investigations as to the condition presented by the skulls of those animals, and the 
modifications undergone by them during growth. 

The limb-bones also of the Gorilla and Chimpanzee have been thoroughly investigated 
by Professor Owen"; but the appendicular skeleton of the Orang has not yet been 
described with similar care and minuteness. 

Yet this highly interesting form, which in some respects resembles Man more than 
any other animal does, fully deserves to be made the subject of the most careful study, 
especially as it is more than probable that at no very distant date it will share the 
fate of the Dodo and Binornis, while we may hope that tropical geology will one da} 
cause a careful description and complete delineation of the bones of Simia to be much 
prized by some future palaeontologist. 

The opportunity of furnishing such a description and such delineations is presented 
by the rich osteological collection of the British Museum, containing as it does eleven 
skeletons of the Orang-Outang, four of these being fully adult. 

I propose, then, to describe each bone of the Orang in detail, and to compare its 
characters with those of the Chimpanzee, the Gorilla, and Man. 

Scapula. (Plate XXXV.) 
The bladebone of the Orang is a triangular plate of bone, in some respects, 

' Trans. Zool. Soc. vol. i. 1835. = Osteographie, " Primates : Pithecus," p. 27, 1839. 

^ Recherclies d'Anat. C'omp. sur le Chimpansc, 1841. * Trans. Zool. Soc. vols. iii. it iv. 

° Paper read before the Zoological Society on the 8th November, 1864. ° Trans. Zool. Soc. vol. v. 



176 ME. ST. G. MIVART ON THE SKELETON OF THE PEIMATES. 

and perhaps on the whole', resembling Man's more than does that of any other 
animal. 

If the bone is placed with the glenoid surface vertical, and compared with a similarly 
placed human scapula, the main difference will be seen to arise from the fact that, in 
the Orang, the inferior- vertebral angle is so much less produced downwards, while at 
the same time it extends more backwards, the angle formed by the axillary margin 
with the glenoid surface being only from 110° to 120°, instead of being from 135° to 
140°, as in Man ; while the prevailing direction of the vertebral margin, instead of being, 
as in him, nearly parallel with the glenoid surface, forms with it a marked angle open 
downwards. In both these respects Simia agrees more or less closely with Troglodytes ; 
but in the direction of the spine of the scapula, the former genus differs from Man in a 
way opposite to that in which Troglodytes differs from him ; for the angle (open upwards) 
formed by the spine with the glenoid surface, is from 65° to 70°, and therefore less than 
in Man, in whom it is about 82°; while in Troglodytes it is from 86° to 100°. Thus 
there is less obliquity in the position of the spine on the blade"' than in Troglodytes, 
and the proportion borne by the supraspinous fossa to the infraspinous one'* is much 
less, the latter sometimes^ predominating more than even in Man. 

The spine commences at the lower end of the uppermost fifth of the vertebral margin 
of the scapula, by a marked flat triangular space, which is sometimes larger both abso- 
lutely and relatively than the same part in Man, thus differing notably from Troglodytes, 
where the triangular sm-face is very indistinctly marked or absent (PI. XXXV. fig. 1 s). 

The spine, apart from the acromion, forms a more elongated triangular plate of bone 
than in Troglodytes, and slightly more so than in Man. Its upper surface is in general 
markedly concavo-convex'', and its under surface concave, to a degree never existing in 
Troglodytes, and which would not be found in Man but for the flattened and over- 
hanging free border of the spine. This projecting border is, in Simia, very rough, the 
rougliness continuing backwards almost to the triangular space before mentioned, and 
thus differing from the same part in Troglodytes (where the roughness is both less in 
degree and less extended) and more resembling that of Man. Simia, however, differs 
from Homo in that this rough free margin is much narrower, and that its lower margin 
much less overhangs the infraspinous fossa'. 

' Professor Huxley says of the Orang, " the scapula, on the whole, bears a greater resemblance to that of Man 
than it does in either the Chimpanzee or Gorilla " (Medical Times, ISO-t, vol. i. p. 565). W. VroKk also says 
it is " broader and more analogous to the scapula of Man " than in the Chimpanzee (Todd's Cycloptedia. 
vol. iv. p. 203). 

- In describing the skeleton of such an animal as the Orang apart from quadrupedal forms, I think it better 
to describe it as if in the erect attitude, and to speak of that as " inferior " wliich in ordinary mammals would 
be " posterior." ' Duvemoy, Archives du Museum, 1855, t. viii. p. 24. ' Duvernoy, loc. cit. 

' This is especially the case iu the variety described as Pithenis morio by Prof. Owen, No. 1179 b in the 
( )stcologicaI Collection of the British Museum. " Not so in the type of the variety Morio. 

' Owen, Trans. Zool. Soc. vol. i. p. 364. 



ME. ST. G. MIVAET OK THE SKELETON OF THE PKIMATES. 177 

There may or may not be a conspicuous foramen, for a nutritious vessel, towards the 
middle of the upper part of the infraspinous fossa ; and one or two such foramina may 
exist in the anterior half of the supraspinous fossa, near the base of the spine. 

This base, or attached border of the spine, sometimes extends forwards rather nearer 
to the margin of the glenoid surface than it does in Man, and always approaches it 
more nearly than in the Gorilla, though not more so than, sometimes not so much as, 
in the Chimpanzee. The antero-external border of the spine is concave, as in Man 
and Troglodytes, but, as in the latter, is somewhat shorter (apart from the acromion) 
than in the human scapula. The acromion is flattened in the direction opposite to 
that of the spine, but is longer and narrower' than in Man or Troglodytes ; its surface 
also is more roughened, and the facet for the clavicle is closer to the extremity of the 
acromion than in the last-mentioned genera. The degree of curvature of the process, 
and its prolongation towards a point over the middle of the glenoid surface, vary some- 
what from individual to individual (PI. XXXV. figs. 4 & 5). 

The supraspinous fossa is generally about equally deep at its anterior and posterior 
ends, the base of the spine (otherwise than in Man or Troglodytes) being nearly parallel 
to the upper margin of the scapula. Sometimes, however, it is decidedly deeper at 
its glenoidal end; and rarely the vertebral end -sery slightly exceeds the rest of the 
supraspinous fossa in depth. 

The infraspinous fossa is concavo-convex, as in Man and Troglodytes, the convexity, 
however, being sometimes more marked than in the latter genus, and always more 
extensive than in the Chimpanzee. Siniia, however, agrees with Troglodytes in that 
the ridge of the axillary margin does not expand, as in Man, into a wide flattened 
surface for the teres major, but, on the contrary, only into a very narrow one. The 
infraspinous fossa is always, in the Orang, narrower, vertically, than is the supra- 
spinous one at the glenoidal end of the spine, reversing the conditions w-hich exist 
in Man. This excess of the supraspinous fossa is greater than in the Gorilla, but 
rarely, if ever, so great as in the Chimpanzee. 

The subscapular fossa is not so concave as in Man, on account of the less inflexion 
inwards of the part of the blade which forms the supraspinous fossa. It is also less 
concave than in the Chimpanzee, and than in some specimens of the Gorilla. As in 
Troglodytes, the oblique ridges traversing this fossa are less marked than in Man. 

The superior border of the scapula is the shortest one, but is longer absolutely, 
and still more so relatively, than in Man and Troglodytes. 

It is slightly concave and nearly horizontal, instead of, as in Man and Troglodytes, 
sloping sharply down to the coracoid process ^ (PI. XXXV. fig. 2). There can scarcely 
be said to be a trace of the suprascapular notch I The vertebral margin is no longer, 

' Owen. Trans. Zool. Soc. vol. i. p. 36-i. 

■ This slope is much more gradual than is generally the case in Man, in the scapula of an Andaman Islander. 
No. 1 NB, in the British Museum. ' This notch is almost indistinguishable in the same Andaman Islander. 
VOL. VI. PART IV. . 2 C 



178 ME. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 

as in Man and the Gorilla, the most extended one, nor is it equal to the axillary one, 
as sometimes in the Chimpanzee ; but it is absolutely shorter. The part of it which 
is superior to the origin of the spine is much shorter, as compared with the part 
below', than is the case in Man, and veiy much more so than in Troglodytes, from 
three-fourths to five-sixths of the vertebral margin being below the origin of the spine. 
If not straight, this margin is very slightly convex, and scarcely ever^ presents a trace 
of sigmoid curvature, the part at or near to the origin of the spine being sometimes^ 
more prominent than the rest, instead of less so as in Troglodytes. 

The axillary margin is unlike that of the higher forms in that it is decidedly 
convex (PI. XXXV. fig. 1) in almost all cases, though in the variety Morio it is 
more nearly straight (PI. XXXV. fig. 3). The ridge near the glenoid surface, for the 
attacliment of the triceps, is prominent, and, as in Troglodytes, is continued a longer 
distance, and bounds externally a much wider groove than is the case in Man. 

The superior vertebral angle, though very distinctly marked, is much less produced 
than in Man and Troglodytes; hence the very slight concavity of the upper margin 
(PI. XXXV. fig. 2). 

The inferior vertebral angle is rather more obtuse ' than in the higher forms, i. e. 
than in Man and Troglodytes. 

The glenoid surface is pyriform (PL XXXV. figs. 4 & 5 </), with the broad end down- 
wards, as in Troglodytes ; it is more elongated in proportion to its breadth than in Man, 
but it is more concave vertically than in the Chimpanzee or Gorilla. 

The coracoid process is shorter and thicker than in Man, and is broadest behind^ 
being very much expanded and flattened at the posterior part of its upper surface. Its 
inclination towards the glenoid surface is much as in the Chimpanzee, and greater 
than in Man and the Gorilla, but it agrees with that of these Apes, and differs from 
Man's in being directed more downwards and less forwards (PL XXXV. fig. 2). 

This process is subject to considerable individual variation as to its length" (PL XXXV. 
figs. 4 & 5), and as to the presence or absence of a smooth groove in its inner surface^. 

The coracoid is entirely separate from the rest of the scapula when the first true 
molar of each jaw has come into use'. 

Clavicle. (Plate XXXVII. figs. 1-4.) 

The clavicle of the Orang is much elongated, and both absolutely longer, and longer 
as compared with the spine, than in Man or Troglodytes. Indeed in the Orang the 

' Archives du Mus. t. viii. p. 24. ^ There is a slight sigmoid curvature in the variety Morio. 

^ E.ij. ill the specimen No. .').'5. 12. 2U in the British Museiun. 

' Scarcely more so in the variety Morio than sometimes in the Gorilla. 

' Also the case in the variety Morio. 

' In the specimen 3 A in the British Museum it is very short, in 3 C, however, it is much longer. 

" Present in both scapula; of No. 3 C in the British Jfiiseum ; also in the Gorilla No. 1011 I. 

' Such is the case in the specimen No. 3 H in the British Museum. 



MR. ST. G. MIVAET OX THE SKELETON OF THE PEIMATES. 179 

clavicle decidedly exceeds one-fourth of the length of the spine (as measured fi'oni the 
atlas to the coccygeal end of the sacrum), while in Man and Troglodytes it always, as 
far as I have observed, falls short of that proportion. The clavicle of the Orang also 
more nearly equals the length of the scapula than in the higher forms'. 

Its curvature is very slight, much less than in Man and the C^himpanzee, and less 
also than that which often exists in the Gorilla. The sternal curve, convex forwards, 
is very mucli more exten.sive than is the backwardly convex curvature of the outer 
or acromial part". The acromial end bends decidedly somewhat do^^iiwards. The 
curvature docs not appear to be gi'eater in young than in old individuals^, nor in 
small adult specimens as compared witli larger ones (PI. XXXVII. figs. 3, 4). 

The two extremities of the bone are more or less flattened, the sternal articular 
surface being generally long and narrow, compared with the form it presents in jNIan and 
Troglodytes. The long axis of this surface is sometimes nearly parallel with the greatest 
diameter of the acromial end of the bone. 

The clavicle of Simia may be described as presenting three surfaces and three 
margins. Of these, the first or superior and the second or more or less anterior one 
extend the whole length of the bone ; but the third or inferior surface does not reach 
to the sternal extremity of the clavicle, being replaced by an extension of the second 
or more or less anterior surface, which here assumes a nearly inferior position. 

The first or superior surface is smooth and pretty well marked off' (along its posterior 
margin) from the thud or inferior surface by a ridge continued inwards from the 
tubercle for the conoid ligament, but which does not attain the sternal end of the bone, 
where the first surface is separated behind from the second one by the strong ridge for 
the rhomboid ligament. It is separated, towards the acromial end of the clavicle, 
from the second or more or less anterior surface by a very strong and rough ridge 
(PI. XXXVII. figs. 1 & 2 <Z) for the deltoid'^ ; but towards the sternal end of the 
bone a slight one (for the attachment of the iwctoraUs major^) serves as the line 
of demarcation between the first and second surfaces in front. 

The second, a more or less anterior surface, is wide and concave towards the acromial 
end of the bone, where it is limited above by the very prominent and rough ridge for 
the deltoid before mentioned, and below by the anterior boundary of the third or 
inferior surface ; towards the sternal end of the bone this surface becomes rather infe- 

' De Blainvillc has found it to exceed the scapula in length (Osteographic, " Primates: Pilhecvs," p. 30). 

- Archives du Mus. t. riii. p. 2.5. 

' As shown in the immature specimen Xo. 3 H, in the osteological collection of the British Museum. 

' Judging from the representation given by Professor G. Sandifort in his treatise on the anatomy of the 
Orang Outaug, in ' Verhandelingen over de Natuurlijko Geschiedenis der Nederlandsche overzecsche bizittingon,' 
Leyden, 1840, p. 48, and tab. 3. fig. 2 C ; also from that of Cuvier in the ' Eeoueil dc Planches de Myologie,' 
pi. 15. fig. 2^. 

' ilr. W. S. Church describes part of the pectoralis major as arising from the clavicle in the Orang (Xat. 
Hist. Review, vol. i. p. 513). Cuvier also so represents it (Kecueil de Planches de Myologie, pi. 15. tig. '2j). 

2c2 



180 ME. ST. a. MIVAET OX THE SKELETON OF THE PRIMATES. 

rior in position, and presents a more or less marked and depressed triangular surface 
(PI. XXXVII. fig. 3, between p and r), towards the sternal end of which there is 
generally a nutrient foramen. This triangular surface is bounded inferiorly and poste- 
riorly by a more or less marked ridge and roughened tract (PI. XXXVII. fig. 3 r), 
doubtless answering to the rough surface serving for the attachment of the costo-clavicular 
ligament in Man. Superiorly and anteriorly the surface is bounded by the much more 
faintly marked ridge already mentioned, which, no doubt, gives origin to the pectoralis 
major (PL XXXVII. fig. Ip). 

The third, or inferior surface, does not, as has been said, reach the sternal end of 
the bone, but terminates at a point about an inch and a half from that extremity. 
The first and second surfaces (which are sejDarated from each other behind and below 
by the third surface for the outer four-fifths of the bone) come into juxtaposition, 
behind as well as in front, at the point where this third or inferior surface has its 
sternal termination. This last-mentioned surface is more or less conca\'e till near the 
acromial end of the bone, where, in adults, it is very rough. There is a very large 
tubercle at the posterior margin of this surface (PL XXXVII. figs. 2 and of), serving 
for the attachment of the conoid portion of the coraco-clavicular ligament, immediately 
in front of which tubercle there is sometimes a very conspicuous nutrient foramen. 
Tlie line for the trapezoid portion of tlie same ligament is generally very prominent 
(PL XXXVII. fig. 3 f) ; and behind and external to it there is sometimes' a smaU, yet 
marked fossa; but the concavity beneath the acromial end of the bone is never so 
marked as it often is in Troglodi/tes^. 

The acromial end of the bone is very little expanded in Simia ; and in this the Orang 
presents a marked contrast to the GoriUa ; and it also difiers from Troglodytes in the 
presence of the second or mainly anterior surface, with its strongly marked ridge above, 
in the great roughness of the inferior surface of the acromial end of the bone, and in its 
generally elongated sternal extremity. 

If the Orang be compared with Man, it will be seen that his clavicle would resemble 
the Orang's, if it were much straightened, the sternal end compressed, and the front 
edge, towards the acromial end, widened out into a concave surface surmounted by a 
prominent ridge, the expansion of the acromial end restricted, but the tubercle for the 
conoid ligament considerably enlarged. 

Humerus. (Plate XXXVI.) 

This bone is of great size in the Orang, but nevertheless is not, as in Man and Troglo- 
dytes, the longest bone of the arm, being always exceeded in length by the ulna, and 
sometimes by the radius also. 

As in the Gorilla, it exceeds three-fifths the length of the spine measured from the 

' E. g. in Nos. 3 A and 3 C in the British Museum. 

' See skeleton of a Gorilla, No. 1011 I, in the British Museum. 



ME. ST. a. MIVART OX THE SKELETON OF THE PRIMATES. 181 

atlas to the lower end of the sacrum — a proportion decidedly exceeding that existing in 
the Chimpanzee, and greatly so that found in Man. 

It is nearly twice the length of the scapula, which is less than in Man, though more 
than in Troglodytes. 

The humerus of the Orang is not so straight as that of Man, the Gorilla, or Chim- 
panzee, but is more bent concave forwards, and sometimes' very much so (PL XXXVI. 
fig. 3). It is also more inclined ulnad at its lower end, the inner margin of the 
bone being decidedly concave (PI. XXXVI. fig. 2). 

As in Man and Troglodytes, the shaft may be described as consisting of three surfaces ; 
but these are not well defined, the lowest fifth of the anterior surface not being so 
sharply prominent as in Homo. Thus the shaft is not so decidedly triangular at its lower 
part as in Man, neither is it so compressed laterally towards its middle as is generally the 
case in him, the Orang in both these respects agreeing more with Troglodytes. 

The ridges proceeding upwards from the condyles are more marked than in Man or 
Troglodytes, especially the external one (or supinator ridge), which extends rather more 
than one-third up the shaft, and is sometimes limited above by a marked musculo-spiral 
groove. The ridge from the internal condyle extends about halfway up the shaft. 

The posterior surface of the humerus is convex above, much as in Man, not flattened 
as in the Gorilla. Below, it is flat, as in the superior forms ; it is not, however, turned 
so much outward as in them, but looks backwards, indeed almost equally with the 
upper part of the posterior surface (PI. XXXVI. fig. 2). 

The bicipital groove is sometimes more marked than in Man, and more sharply limited 
on each side at its upper part, though less so there than in Troglodytes. Lower down 
it is often more marked than in that genus, or than is generally the case in Man. 

The surface probably serving for the insertion of the coraco-brachialis is extraordinarily 
rugose, more so than it ever is, as far as I have observed, in Man, and sometimes 
more so than in the Gorilla, much more so also than in the Chimpanzee ; the rougli- 
ness sometimes extends so far downwards as to join the ridge extending upwards 
from the internal condyle (PI. XXXVI. fig. 4 c). 

Near the surface for the coraco-brachialis, and a little below or a little above the 
middle of the bone, is the medullary foramen opening distad, as in Man. 

The surface for the deltoid is not so much raised as is generally the case in Man ; 
and below it is the more or less distinct oblique depression marking the course of 
the musculo-spiral nerve and artery (PI. XXXVI. figs. 1 & 25). 

The head of the humerus is very large^ and rounded, its greatest diameter decidedly 

' As in both the humeri of No. 3 B iii the British Museum. 

- Professor Owen says, "it is larger in Simia satyrus than in man, its extent equalling a complete hemi- 
sphere " (Trans. Zool. Soe. vol. i. p. 364). De Blainville remarks, " sa tete artioulaire est surtout singulierc 
par son enormite, son diametre etant hien supe'rieur a celui dc la tete du femur " (Osteographie, " Primates ; 
Pithecus;' p. 30). 



182 MK. ST. G. MIVAET ON THE SKELETON OF TKE PEIMATES. 

exceeding the breadth of the two tuberosities, in which respect Simia differs from both 
Man and Troglodytes (PI. XXXVI. figs. 5 & 6). 

The posterior projection of the head is greater than in Man and the Gorilla, the 
bone immediately below the posterior part of the margin of the articular surface being 
more concave (PI. XXXVI. fig. 4). The anatomical neck is more marked than in the 
higher forms, especially than in Man, and the head rises more above the summit of the 
radial (greater) tuberosity. 

The angle formed by the groove separating the last-named tuberosity from the head, 
with a line cormecting the two condyles, is in general much more acute than in Man 
and Troglodytes. There is nevertheless considerable individual variation, the angle 
being sometimes as near a right angle as in the Gorilla (PI. XXXVI. fig. 5), while m 
other instances (PI. XXXVI. fig. 6) it is only about 34°; yet, in all the specimens of 
Simia which I have examined, the head looks more directly backwards and less 
inwards than in Man and Troglodytes. 

The radial tuberosity in the Orang is less prominent than in the Gorilla, and but 
little more so than in Man, though the surface for the teres minor is more sharply 
defined than in the higher forms (PI. XXXVI. fig. 3). The surface for the infra- 
spinatus looks more outwards and less upwards than in Man (PI. XXXVI. fig. 3). 

The ulnar, a smaller tuberosity, has its upper part next the bicipital groove less 
prominent than in Man or Troglodytes. Sometimes its lower part is more prominent 
than its upper portion. This tuberosity is more nearly ajiproached by the margin of 
the articular surface of the head than in the higher forms ; so that there is a small deep 
pit (PI. XXXVl. fig. 4) between them, instead of a rather wide and slightly concave 
surface. 

When the humerus is vertical, and its anterior surface opposite the observer, the 
ulnar tuberosity generally hides part of the lower margin of the neck of the bone, 
in which respect Simia resembles Troglodytes and differs from Man'. As in higher 
forms, no part of the head of the humerus is so hidden in the Orang. 

At the lower end of the bone the ulnar, or inner, condyle is not so prominent as in 
Man or Troglodytes, nor does it extend so much downwards as in Man and the Chim- 
panzee, but appears as if it had been truncated obliquely from below upwards and 
ulnad, though this appearance is not so marked as in the Gorilla, on account of the 
less prominence of the condyle in the Orang. As in Troglodytes, there is not that 
concavity on its posterior surface which is more or less marked in Man. 

The external, or radial, condyle is much as in Man and the Chimpanzee, and its most 
prominent point is situated lower down than in the Gorilla. 

The inferior articular surface of the humerus is almost quite as in Man, except that 
its innermost part genei-ally descends less below the rest of the surface than in him 
or the Chimpanzee, though more so than sometimes is the case in the Gorilla. 
' Owen, Trans. Zool. Soo. vol. v. p. 4, pi. 3. figs. 1, 5, 8. 



MB. ST. G. MIVART ON THE SKELETON OP THE PEIMATES. 183 

Whether, however, it descends more or less than in the Gorilla, it always differs from 
Man and agrees with Troglodytes in that the inner margin of the anterior surface of 
the trochlea (below the ulnar condyle) is vertical, and not inclined ulnad at its lower 
end as in Ilomo^. 

As in Man and the Chimpanzee, the surface above the capitellum, in front, is less 
concave than in the Gorilla. 

The surface above the trochlea is almost always perforated". 

The olecranal fossa is bounded on its radial side by a more marked and extended 
ridge than exists in Man and sometimes in the Gorilla (PI. XXXVI. fig. 2). This 
ridge is the continuation upwards and backwards of that part of the articular surface 
which projects between the radius and the ulna. 

Radius. (Plate XXXVII. figs. 5-8.) 

The radius is very elongated and sometimes slightly exceeds the humerus in length, 
in which respect, as has been already said in describing the last-named bone, the Orang 
differs from Man and Troglodytes. 

Its length as compared with that of the spine, measured as before, is much greater 
than in Troglodytes, being slightly upwards of three-fifths of that of the latter, instead 
of but little more or less than one-half Of course the Orang differs much more still 
from Man in this respect. 

The radius is always very nearly as much as, if not a little more than, twice the 
length of the scapula — a proportion not attained in the higher forms. 

The shaft of the bone is considerably curved^, mth the concavity ulnad ; but though 
much more so than in Man, the curvature is somewhat less than that which appears 
generally to exist in the Gorilla. 

The radius of the Orang is so rounded a bone that it can no longer be said to have 
the three surfaces and three margins existing in that of Man. 

The anterior face, however, is pretty well defined and expands distally ; the surface 
for the flexor longus pollicis, however, is very slightly marked, much less so than in 
Troglodytes, while it presents nothing like the concavity whence that muscle takes its 
origin in Man. 

The foramen for the nutrient vessels is situated towards the lower end of the upper 
third of the bone, and rather on the radial side of the anterior surface, instead of on its 
ulnar side as in Man and Troglodytes. It is, however, directed proximad, as in the 
higher forms just mentioned (PI. XXXVII. fig. 5). 

The lower end of the anterior surface is more concave transversely than in Man. 

' 8ec Trans. 7mo\. Soc. vol. v. pi. '■\. figs. 1, 5, 8. 

- It is imperforate in both humeri of No. 31 and in those of No. 3H 50. 8. 15. 1 in the British Museum. 
' W. VroUk remarks " II mo parait que cette courburc est un produit de Tage " (Rechci-ches d'Anat. C'omp. 
sur le Chimpanse, p. 1.3). 



184 ME. ST. G. MIVAET OX TILE SKELETON OP THE PEIMATES. 

though the concavity which is in him produced by the projection forwards of the distal 
margin of the bone is wanting, that margin in the Orang, as also in Troglodytes, not 
being similarly prominent. 

The posterior surface presents but a very slight flatness for the origin of the extensors 
of the pollex ; and sometimes, indeed, there is no flattening perceptible. The external 
surface, which generally in Siinia passes insensibly into the posterior one, presents a 
rough tract and a slight excavation for the siqnnator teres, extending downwards nearly 
to the middle of the bone. 

Of the three margins which exist in Man, the posterior one is, in the Orang, never 
more than faintly marked, and that only towards the middle of the bone. 

The external margin of the radius of Man may be said to have disappeared altogether 
in the Orang; but the internal margin, for the interosseous ligament, is distinctly 
marked, though it is never nearly so sharply projecting as in him and the Chimpanzee, 
and scarcely so much so as in the Gorilla. 

The bicipital tuberosity is much less prominent than in Man ; but its surface is more 
excavated, and, as in Troglodytes, is much more ulnad in position (PI. XXXVII. fig. 6 b). 

The bone is not so contracted at its neck as in Man and the Chimpanzee, and the 
rim or margin' of the head is not so sharply marked inferiorly as in them, the Orang 
in these points resembling the Gorilla. The proximal articular surface of the head is 
less concave than in the human radius. 

A little above the styloid process there is a very prominent and rough surface 
(PL XXXVII. flg. 5 a) for the insertion of the supinator longus. The styloid process 
itself is not so pointed as in Man and Troglodytes (PI. XXXVII. fig. bf). 

The grooves for the extensor tendons are quite similar to those of Man, except that 
they are sometimes more marked than in him ; and this is even the case with the 
groove for the extensor secundi internodii j^olliris and that for the extensor ossis 
metacarpi pollicis (PI. XXXVII. figs. 6 & 7). 

The articular surface for the reception of the ulna looks more backwards than in 
Man, especially when the ulnar angle of the anterior side of the distal end of the bone 
is much produced forward and ulnad, as is sometimes the case (PI. XXXVII. fig. 6(/). 

The carpal surface at the inferior end of the bone has the internal quadrate surface 
for the semilunare larger, in comparison with the triangular one for tlie scaphoides, 
than is the case in Man (PI. XXXVII. fig. 8). 

Ulna. (Plate XXXVIII.) 

This bone, which, unlike the radius, seems in the Orang to be constantly longer than 
the humerus, bears much the same proportion to the ulna of Man and Troglodytes that 
the radius of the Orang bears to the radius of those forms. 

When its anterior (flexor) surface is opposite the observer (PI. XXXVIII. fig. 1), 

' Owen, Trans. Zool. Soc. vol. v. p. 7. 



MR. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 185 

the shaft may be seen to have a sigmoid curvature, which is convex ulnad below, above 
concave. This curvature is more marked than is generally the case in Man, or than 
sometimes in the Gorilla ; it is less so, however, than in the Chimpanzee. When the 
bone is viewed laterally (PL XXXVIII. figs. 2 & 4), the shaft is seen to present a 
curve, convex backwards, which is slightly more marked than in Man, but not quite so 
much so as in Troglodytes. 

The body, or shaft, of the ulna is more rounded than in Man or Troglodytes, and can 
hardly be said to present the three surfaces and margins usually described as existing in 
the human ulna, the parts which correspond to the anterior and posterior margins of 
Man being so ill defined. The ulna tapers distally, but, on account of the length of 
the bone, more gradually than in the higher forms. 

The anterior surface of the shaft has a more or less flattened, and even sometimes deci- 
dedly concave (PI. XXXVIII. fig. 1) surface for 'Co.q flexor profundus digitorum; and 
the nutrient foramen, much more conspicuous than in Man or Troglodytes, is more or 
less remote from the radial margin of the bone, and rather below the uppermost third of 
its total length (PI. XXXVIII. fig. 1 b). As in the higher forms, its direction is proximad. 
The internal surface of the shaft is smooth, but more convex than in Man and Troglodytes, 
except at its summit, where the conca\ity is more extensive than in them, reaching as it 
does somewhat more nearly to the superior limit of the olecranon (PI. XXXVIII. fig. 2). 
The posterior, or radial, surface of the shaft is less strongly divided into two parts 
than in Man and the Gorilla, though the lower and much larger one (serving to give 
origin to the extensors of the poll ex and index) is generally as flat as in Man, and more 
so than in Troglodytes ; very rarely it is strongly concave. 

An anterior margin can sometimes hardly be distinguished, and never extends, as in 
Man, from the coronoid process to the lower extremity of the ulna. Sometimes, how- 
ever, it can be traced from that process down to somewhat below the level of the 
medullary foramen. Similarly the posterior margin of the human ulna (which extends 
fi'om the olecranon to the styloid process, and gives attachment to an aponeurosis com- 
mon to the flexor prof undiis digitorum, the flexor carpi ulnaris, and the extensor carpi 
ulnaris) is in the Orang represented by a prominence which ceases to be distinguishable 
at about the middle of the ulna. 

The external or radial margin begins above at the posterior margin of the lesser 
sigmoid cavity, and extends rather more than two-thirds down the bone. It is not so 
sharp as in man and the Chimpanzee, but it is more so than in the Gorilla. The sharp- 
ness, however, generally only extends along about the middle third of the bone, w^iich 
at that part is considerably roughened for a greater or less extent close to the radial 
border (PI. XXXVIII. fig. 1 a). Very rarely, however, the radial margin is enormously 
produced\ In the Orang, unlike the higher forms, the upper part of this margin does 

' As ia the specimen in the Collection of the British Museum, which bears the No. 32, from the MS. catalogue 
of the Zoological Society's CoUeotion. 

VOL. VI. PART IV. 2 D 



186 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 

not bound anteriorly the surface for the anconeus, but is separated from that surface by 
a flattened tract of bone interposed (PI. XXXVIII. fig. 4) between it and the ridge 
which does so Hmit the anconeal surface. 

The olecranon process is small and scarcely broader relatively than in Man, and, as in 
him, it does not project so much ulnad as it does in Troglodytes. It is not in any way so 
prominent as in that genus, being even less so than in Man (PI. XXXVIII. figs. 2 & 6). 

The greater sigmoid cavity (PI. XXXVIII. fig. 1 g) is formed nearly as in Man, except 
that it is broader in proportion to its length than in him, or indeed than in Troglodytes. 
The lesser sigmoid cavity is less extended from above downwards, and more from behind 
forwards, than in the Gorilla, thus resembling more the form it presents in Man and the 
Chimpanzee. 

The coronoid process is broader both absolutely and in proportion to its projection 
forward than in the two last-named forms, and it is' also relatively broader than it is 
sometimes in the Gorilla. 

The tubercle for the flexor sublimis digitoritm is, as in the Gorilla', well developed. 

The fossa for the hrachialis anticus is very marked and deeper than in Man or the 
Chimpanzee, or than is sometimes the case in the Gorilla (PI. XXXVIII. fig. 1 c). 

The fossa for the anconceus is much smaller relatively than in Man, and is less defined 
anteriorly by the ridge running .downwards and backwards from the hinder end of the 
lesser sigmoid cavity, that ridge being (as also in the Chimpanzee, but not in the Gorilla) 
much less marked than in Man. On the other hand the posterior margin of the anconeal 
fossa is much more sharply defined than in the higher forms (PI. XXXVIII. fig. 4/). 

As in the Chimpanzee, but not in the Gorilla, the surface for the supinator brevis is 
much less concave than in Man, and, indeed, is but slightly marked. 

In Man and Troglodytes this surface for the supinator brevis is contiguous for almost 
its whole extent with that for the anconmiis, the surface for the extensor ossis metacarpi 
poinds only slightly intervening between them inferiorly. In the Orang, however, a 
wide flattened tract of bone (serving most probably to give origin in part to the extensors 
of the poUex) extends up almost to the lesser sigmoid cavity (PI. XXXVIII. fig. 4). 
This tract is bounded in front by the upper end of the external or radial margin of the 
ulna ; posteriorly it is limited by the ridge running downwards and backwards from the 
last-named surface (PI. XXXVIII. fig. 4 e) and bounding anteriorly the surface for the 
anconaiis. 

At the lower end of the ulna on the inner side of its anterior face is a ridge serving 
to give attachment to the pronator quadratus- (PI. XXXVIII. figs. 1 & 2x). It is 
much more marked than in the higher forms. 

As in Troglodytes, the distal articular surface of the shaft of the ulna is relatively 

' See Owen, Trans. Zool. Soo. vol. v. p. 8. 

= Noticed by Professor Owen, Trans. Zool. Soe. vol. i. p. 304. He refers to Webster and Treadwcll's 'Boston 
Journal of Pliilosoph)',' vol. ii. p. 570, and the ' Philosophical Magazine,' vol. Ixviii. p. 186, 1826. 



MR. ST. Q. MIVAET ON THE SKELETON OF THE PRIMATES. 187 

more transversely extended than in Man and is more reniform ' ; the concavity also 
between it and the styloid process is deeper (PI. XXXVIII. figs. 5 & 6). This last- 
mentioned process appears to vary much as to size, from individual to individual 
(PL XXXVIII. figs. 2 & G s) ; but it is never so long, compared with the total length 
of the ulna, as in Man and the Chimpanzee^ 

The groove for the tendon of the extensor carpi iilnaris is generally very little marked, 
and less so than is the case in Man or the Gorilla, so far as I have been able to observe. 

Manus. (Plate XLII.) 
This segment of the skeleton attains, in the Orang, a greater absolute length than 
it does in Man or Tro(jlodytes. Its proportion to the spine (measured as before) is also 
greater ; but those borne by it to the rest of the pectoral limb and to the i-adius are less 
in the Orang than I have found them to be in the Chimpanzee, though gi-eater than in 
Man or in the Gorilla. In its slenderness the manus of the Orang more resembles that 
of the Chimpanzee than that of the Gorilla or of Homo. 

Carpus. 

This segment differs very importantly from that of the higher forms, in that, as is 
well known, there is a separate and distinct ninth carpal bone', the os intermedium. 

The proximal row of carpal bones forms a double arch, as in Man and Troglodytes. 

The vertical arch (with its convexity proximad) is rather more acute than in Man ; 
but the OS pisiforme being small, its outline is not interrupted by that bone, as it is in 
Troglodytes, and so far it resembles more the homologous arch of the human liand 
than does the vertical carpal arch of the last-named genus (PI. XLII. fig. 1). 

As in the higher forms, the carpus in the Orang articulates directly with the radius 
only. 

Scaplioides. (Plate XLII. figs. 2, 3, 4.) 
This bone is very much narrower antero-posteriorly (from dorsum to palm), and 
relatively much more transversely extended, than in Man, and there is no transverse 
dorsal groove ; so that the scaphoid of the Orang has very much the appearance that 
that of Man would have, if the part anterior to (or on the distal side of) his dorsal groove 
were cut away. Indeed the whole scaphoid of the Orang appears to answer to only 
the upper or proximal part of the human scaphoid' and of that of Troglodytes. It 

' Owen, Trans. Zool. Soc. vol. v. p. 7. 

- In a mounted specimen of the Gorilla in the British Museum, this process is very short indeed. 

^ Pointed out by W. Vrolik, Reeherches d'Ant. Comp. sur le Chimpanse, p. 13 ; and Todd's Cycloptedia, 
vol. iv. p. 203. 

■• See De BlainviUe, Osteographie, " Primates : Piihecus," p. 16 ; Professor G. M. Humphry, Limbs of Verte- 
brates, 1860, p. 4 J Professor Huxley, Hunteiian Lectures, Medical Times, 1864, vol. i. p. 565 ; and l)r. 
Lucae, Abhaudlungeu von der Senckenbergischen naturforschenden Gesellschaft, 1865, vol. v. p. 311. The 

2 D 2 



188 ME. ST. G. MIVART OX THE SKELETON OP THE PRIMATES. 

does not articulate with the trapezoides and os magnum, being separated from them by 
the OS intermedium. 

The proximal or superior side of the scaphoides articulates with the radius by a large 
rounded sm-face, which is decidedly less convex than the corresponding one of Man, or 
than that of the Gorilla, and is much like that of the Chimpanzee (PL XLII. fig. 2 a). 
It is less quadrate than in the Gorilla, and less transversely elongated than in Man. 

The radial tuberosity (PI. XLII. figs. 2-4, b) appears much produced, because of the 
narro^vness of the bone. It is not, however, really vei7 much more so than in Man, and 
is not so much so as in Troglodytes. There is a deeper concavity between the tuberosity 
and the radial articular surface than in Man. 

The distal, or inferior, side of the bone presents a strong concavity divided by a 
transverse prominence into two articiJar surfaces. The smaller and more proximal of 
these (PL XLII. fig. 3f) joins the semilunare, and, as in Troglodytes, is larger than the 
coiTespondmg surface of Man. The one nearer the palmar (or more distal) surface is 
more concave, and articulates mth the ulnar end of the os intermedium. Palmad and 
radiad of this is a small irregular surface, with several vascular foramina ; and radiad, 
again, of this last surface is another smaller articular one (PL XLII. figs. 3 & 4, e) for 
the radial side of the thii'd or proximal face of the intermedium. External again to 
this (i. e. on its radial side), and on the inferior aspect of the base of the tuberosity, is 
a small surface (PL XLII. fig. 4y) which joins the trapezium. 

On the upper part of the bone, towards its ulnar side and between the surfaces for 
the radius and semilunare, is a small irregular tract of bone with ^•ascular foramina. 

The interspace on the dorsal surface of the carpus, between the scaphoides and 
intermedium, answers to the dorsal groove of the scaphoid of Man and Troglodytes. 

Intermedium. (Plate XLII. figs. 5, 6.) 

The intermedium is a slightly crescentic bone, but not very dissimilar in shape to the 
cuneiforme, which about equals it in size ; its extension, however, is mainly in the 
transverse direction. 

It may be described as ha\ing three surfaces and three borders. 

The first of these surfaces (PL XLII. fig. 5), and the one which looks backwards, 
downwards, and more or less radiad, consists for the most part of a transversely extended 
and slightly convex articular sui-face for the trapezoides. The ulnar end, however, pre- 
sents a rough tract for the attachment of ligaments. 

The second surface, which looks palmad, downwards, and ulnad (PL XLII. fig. 6), 
is concave, and articulates with the rounded head of the os magnum. 

The third surface, the one which looks upwards and rather ulnad, is convex, and 

skclotou No. 5083 A, in the Museum of the Royal College of Surgeons, almost demonstrates this homology, as 
in that Chimpanzee the scaphoid has a development almost exactly corresponding to that of the scaphoid, plus 
the intermedium, of the Orang. See Philosophical Transactions, vol. clvii. (18G7) plate xiv. fig. 1. 



ME. ST. G. MIVAET ON THE SKELETON OP THE PRIMATES. 189 

unites with the distal conca^dty of the scaphoides by a transverse articular surface, more 
or less interrupted towards its middle by a rough non-articular part. 

The margin which separates the first and third of these surfaces has its middle portion 
produced into a more or less pointed process, projecting, proximad, over the groove which 
divides the bone from the scaphoides (PI. XLII. fig. 5 e). 

The radial end of the intermedium is obtusely pointed (PI. XLII. figs. 5 & 6, d), but 
its ulnar end is truncated (PI. XLII. figs. 5 & 6, c) and presents a small articular surface, 
more or less concave, which joins the semilunare. The angle at the jimction of the distal 
end of this small surface with the ulnar end of the border separating the first and second 
of the before described surfaces, projects somewhat over the radial border of the dorsum 
of the OS magnum. 

Semilunare. (Plate XLII. figs. 7-9.) 

This is very large in the Orang, and much larger, as compared with the scaphoides, 
than in either TNIan or Troghdijtcs. It is of about the same size as the semilunare of 
the Gorilla, and has very nearly the same shape. As compared with that of Man it is 
especially elongated from above downwards, and its proximal surface is more strongly 
convex from behind forwards, i. e. from dorsum to palm. 

The surface for the cuneiforme (PI. XLII. fig. 8 c) differs from the same surface in Man 
and Troglodytes., in that it is decidedly concave. The dorsum of the bone is narrower 
transversely than in Man or the Gorilla ; and the concave surface for the magnum passes 
insensibly into that for the cuneiforme, though between the two is a very small part which 
joins the unciforme. 

Cuneiforme. (Plate XLII. figs. 10 & 11.) 

This bone is much elongated as compared with its homologue in Man and Troglodytes. 
It is of about the same size as the intermedium, and, as has been before said, it is of 
somewhat similar shape. It is, however, extended rather from above downwards than 
transversely, and its large articular surface is concavo-convex, instead of concave only, 
as in the intermedium. 

It differs from the cuneiforme of both Man and Troglodytes in the convexity of the 
surface for the semilunare, and in the distance between the facet for the pisiforme and 
the proximal end of the bone. 

The rough portion for ligaments of the anterior surface is continued as a wide and 
roughened groo\e (PI. XLII. fig. 11 c), separating the articular surfaces for the unci- 
forme and pisiforme. That for the last-named bone is smaller, absolutely as well as 
relatively, than in any of the higher forms. The smooth surface for the fibro-cartilage 
of the wrist joint (PI. XLII. fig. 10 e) is more elongated than in Man or Troglodytes; 
but, as in the latter genus', the tubercle for the internal lateral ligament of the wrist is 

' Trans. Zool. Sou. vul. v. p. 10. 



190 MR. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 

small and less marked than in Man. The surface which joins the unciforme (PI. XLII. 
fio-. 11 «) is strongly concavo-convex, and thus differs much from that in Man, and more 
resembles the corresponding surface in Troglodytes. 

Pisiforme. (Plate XLII. figs. 12-14.) 
The pisiforme of the Orang is much smaller, both absolutely and relatively, than 
that of the Gorilla, and rather so than that of the Chimpanzee. It is always shorter 
than in the last-named form, but yet, sometimes at least, differs from the pisiforme of 
Man in being rather longer than broad. Its palmar surface (PI. XLII. fig. 13) is slightly 
concave, and the bone projects downwards and ulnad near the unciforme process. 

Trapezium. (Plate XLII. figs. 15-20.) 

A striking difference exists between this bone in the Orang and the homologous one 
of the GoriUa, inasmuch as the two large tuberous processes which exist in the latter 
form' are here wanting. It differs in the same way, though to a less degree, from the 
Chimpanzee's ; and even as compared with Man's, the tubercle and groove of its palmar 
aspect are somewhat less marked (PI. XLII. fig. 17). 

The saddle-shaped surface for the metacarpal of the pollex is always (as also in 
Troylodytes) much inferior in relative size to that of Man ; indeed, even in absolute 
size, it is much inferior to his. There seems, however, to be considerable variation as 
to the development of this part, as sometimes (PI. XLII. figs. 15 & 17, a) there is a very 
distinct, though small, saddle (the surface being strongly concavo-convex), while in 
other instances (PI. XLII. figs. 16 & 18, «) both concavity and convexity are very slight. 
This variation is not confined to the Orang, but exists also in Troglodytes-. 

The surface for articulation with the metacarpal of the index (PI. XLII. fig. 17 J) is 
generally very close to that for the metacarpal of the pollex — a circumstance in which 
the Orang differs from Troglodytes, and resembles Man. The surface for the index 
looks more palmad than in Man or Troglodytes, but, as in them, it is continuous with 
the articular surfaces for the trapezoides and scaphoides. The distal pair of these three 
surfaces form a more marked angle with each other than in Man and Troglodytes; 
w\\\\e the proximal pair (for trapezoides and scaphoides) generally meet together at a 
rather more open angle than in those genera. A sesamoid ' bone is interposed between 
the trapezium and the scaphoides on the radial side of those bones. 

' Owen, Trans. Zool. Soc. vol. v. p. 10. 

■ The saddle is unusually little marked in the skeleton of a Gorilla, No. 5779 K, in the Museum of the Royal 
College of Surgeons ; and in the detached and articulated manus of a Chimpanzee, No. 744, iu the same collec- 
tion, it is absolutely wanting. Professor Hiudcy has noticed the absence of a saddle-shaped surface in this 
species. Sec ' Medical Times,' 1864, toI. i. p. 428. 

' Figured by Prof. Vrolik in Todd's Cyclopaedia of Anat. and Phys. vol. iv. p. 204, fig. 124 i. Mr. W. H. 
Flower also informs me he observed its existence in the wrist of an adult male Orang iu the Loydcn Museum. 



ME. ST. G. MIVAET ON THE SKELETON OF THE PEIMATES. 191 

Trapezoides. (Plate XLII. figs. 21-23.) 

As in Troglodytes, the relative extent of this bone from the dorsum of the manus to 
the palm is very much less in Simia than in Man. The two articular surfaces for the 
metacarpal of the index are of very unequal size — that on the ulnar side being very 
greatly m excess (PI. XLII. fig. 22 a). The articular facet for the magnum is exceed- 
ingly small, and confined to the dorsal part of the ulnar side of the bone. That for 
the intermedium has its greatest diameter transversely extended, instead of from behind 
forwards [i. e. from dorsum to palm), as has the corresponding surface for the scaphoides 
in Man (PI. XLII. fig. 2.3 d). The surface for the trapezium is concave. The proximal 
radial angle is a little produced, but not so much as is the case in Wzxi. 

Magnum. (Plate XLII. figs. 24-28.) 
The distal part of this bone is much more transversely extended, as compared with 
the proximal part, than is the case in Man and Troglodytes ; also there is a more marked 
lateral constriction below the head. As in the Gorilla', the antero-posterior (from 
dorsum to palm) extent of the distal surface is much greater than in Man ; and, again, 
as in the Gorilla" and also in the Chimpanzee, the radial side of the distal articular 
surface is strongly notched (PI. XLII. fig. 28 a). On the ulnar side of the same surface 
is a similar notch, which is much more marked than either in Man or Troglodytes 
(PI. XLII. fig. 28 h). Indeed this distal articular surface has more the shape of 
the letter T than it has in the higher forms ; but it resembles that of Troglodytes in 
being more concave towards its palmar margm than is the case in Man. The head of 
the bone, as also in Troglodytes, has a more radiad aspect than in Man ; it articulates 
above with the intermedium and semiluuare. As is the case in the higher forms, there 
is a small articular surface for the metacarpal of the index, towards the palmar border 
of the distal end of the radial face of the bone (PI. XLII. fig. 26 c). The surface for the 
unciforme is, as in 'Troglodytes, more concave from above downwards than is the case 
in the human magnum. 

Unciforme. (Plate XLII. figs. 29-33.) 

This bone resembles its homologue in Troglodytes, and differs from that of Man in 
the large size and more downward production of the palmar process, also in the more 
acute angle formed by the siu'face for the magnum with that for the cuneiforme, and 
in the greater relative extent of the distal surface from before backwards (from the 
dorsum to the palm). 

It differs from that of the Gorilla, as well as from that of Man, in the greater length 
fi-om above downwards and the less relative transverse extent of the articular surface 
for the cuneiforme (PI. XLII. fig. 32 c). 

' Trans. Zuol. Soc. vol. v. p. 10. ' Loc. at. 



192 MR. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 

Metacai-pus. 

The length of this segment of the limb (estimated by the third metacarpal), compared 
with that of the spine, is greater than iu any higher species, namely about 18'2 to 100. 

The relative length of the same metacarpal, as compared with that of the entire manus, 
is very much the same as in Troglodytes, namely about 39 to 100, and greater than 
in Man, in whom I have found it to be about 34-6 to 100. 

The four outer Metacarpals. 

As in the higher forms, these metacarpals in the Orang are thicker at each end than 
in the shaft ; the distal extremities are wider than the proximal ends (though not so 
much so as in the Gorilla), and the shafts slightly broader distally. 

The heads have their antero-posterior diameter (from dorsum to palm) about equal 
to their transverse dimensions. 

The shafts ai'e much elongated ; and these metacarpals in the Orang are like the 
Chimpanzee's, and are more slender than those of the Gorilla or of Man. The shafts are 
also more rounded than in the higher forms, the dorsal flattening being less marked, 
while there are only faint traces (PL XLII. figs. 38 & 42) of those palmar tuberosities 
and ridges at the divergence of the interossei which are so marked in Troglodytes, espe- 
cially m the Gorilla'. The jirocesses on each side of the proximal ends of the palmar 
surfaces of the heads are mucli less marked than in any higher forms, especially than 
in Troglodytes. The fossae on the sides of the heads are also less marked than in that 
genus. These metacarpals increase, not only in length but also in projection distad, 
from the fifth to the second successively. 

First Metacarpal. (Plate XLII. figs. 35 & 36.) 

This metacarpal presents a saddle-shaped surface for the trapezium, very like that of 
Man. The proximal prominence on the palmar side is less enlarged than in Troglodytes' ; 
and the whole bone is more bent, with the concavity palmad, than in the higher forms. 
The shaft is sometimes slightly as it were twisted on its long axis. 

Second Metacarpal. (Plate XLII. figs. 37-40.) 
The shaft of this metacarpal is also somewhat twisted, and it is concave radiad. Its 
proximal end, like that of the same bone in the Gorilla, docs not, as in Man and the 
Chimpanzee, bifurcate for the reception of the trapezoides ; and its proximal surface is 
therefore less concave transversely. As in Troglodytes, the tubercle for the flexor carpi 
radialis (PI. XLII. fig. 38-40, a) is stronger than in Man, but that for the extensor carpi 
radialis longior is not more marked than in him. Between these tubercles there is a 
deep groove (in which vascular foramina open), which is contmued between the lower 
facet for the third metacarpal and the articular surface for the trapezoides. The ulnar 
' Owen, Trans. Zool. Soc. toI. v. p. 11. - Owen, he. cit. 



MR. ST. G. MIVART OK THE STCBI/ETOK OF THE PRIMATES. 193 

lateral facets are, as in Troylodytes, divided by a deep groove ; and, as in all the liighcr 
forms, the ulnar angle of the proxinal end of the palmar part is strongly inclined ulnad. 
Sometimes (PI. XLIIi fig. 34 ii.) the proximal articular surface has a crescentic form, 
witli the concavity of the crescent towards the dorsum. 

Third Metacarpal. (Plate XLII. figs. 41-44.) 
As in Troglodytes, so also in Shnia, the proximal radial angle of the dorsum is less 
produced than in Man (PI. XLII. figs. 41-43 c). Owing to this, the dorsal part of the 
proximal articular surface is less concave than in him, but the palmar part of that 
surface is more extensively convex. The Orang agrees with Man and the Chimpanzee, 
and, as far as I have observed, differs from the Gorilla, in having two facets on each 
side of this metacarpal, for articulation with the contiguous one (PI. XLII. figs. 43 & 44). 
The proximal articular surface (PI. XLII. fig. 34 iii.) somewhat approaches the form of 
the letter T. 

Fourth Metacarpal. (Plate XLII. figs. 45-48.) 
The proximal end of this metacarpal is narrower transversely than in Troglodytes or 
Homo. The dorsal part of the proximal surface is more concave than in Man, though 
scarcely so much so as in Troglodytes. The palmar part of the same surface is mucli 
more convex, and is longer, from dorsum to palm, than in Troglodytes, and still more 
so than in Man. There are two articular surfaces for the third metacarpal, and the 
single one for the fifth metacarpal is larger tlian in any above (PI. XLII. figs. 47 & 48). 

Fifth Metacarpal. (Plate XLII. figs. 49-51.) 

This metacarpal differs from the corresponding one in Man and Troglodytes in that 
its proximal surface is longer (from dorsum to palm), and is more convex, and at the 
same time less concave than in them (PI. XLII. figs. 34 v. & 51). 

Digits. ;_ 

The proximal phalanges of the four outer digits of the manus are much curved, with 
the concavity palmad (more so than in Man or Troglodytes), though not so much so as 
are the homotypal segments of the pes. ■ 

They are also very broad, and have projecting lateral ridges (PI. XLII. fig. 1), 
which are more developed than in Man, though not so ihuch so as in Troglodytes^ . 

The proximal phalanx of the pollex is more slender than in higher forms, and thus 
differs notably fi-om its homologue in the Gorilla. The several second and third 
phalanges are formed nearly as are their homologues in the Chimpanzee ; the second 
phalanges, however, are somewhat less conical. The distal ones, like those of Troglo- 
dytes, are more attenuated than those in the human manus, The proportion borne by 

' Owen, Trans. Zool. Soc. vol. i. p. .365. 

VOL. VI. — PAET IV. 2 E 



194 MR. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 

the first phalanx of the third digit to the length of the entire manus is greater than in 
Man or the Troglodytes. 

The pollex, with its metacarpal, as compared with the spine, is longer in the Orang 
than in Man or Troglodytes ; compared with the length of the manus it is, as in the last- 
named genus, much shorter than in Man. The proximal phalanx of the pollex is more 
slender than in Troglodytes or Homo, notably so as compared Avith that of the Gorilla. 

The index, with its metacarjml, as compared with the spine, is longer than in the 
higher forms, as also in the third digit. Without their metacarpals these digits, when 
compared with the length of the manus, are scarcely longer proportionally than in the 
Chimpanzee, and but little more so than in Man or the Gorilla. 

The difference between the length of the index and that of the pollex is greater 
than in the higher genera'. 

The fifth digit is the shortest, not counting the pollex ; the second may or may not be 
somewhat longer than the fourth ; and the third is the longest (PI. XLII. fig. 1). 

The order of projection is similar to that of length. 

As in Troglodytes, the proportion, in the Orang, borne by the longest digit (without 
its metacarpal) to the longest metacarpal is less than in man, though it is somewhat 
greater than in the Gorilla. 

The pollex does not reach to the distal end of the metacarpal of the index, but falls 
short hy about one-eighth of the length of that metacarpal ; it is therefore decidedly 
shorter, thus compared, than in the Chimpanzee, and still more so than in the Gorilla : 
and thus in this respect the Orang differs very widely from Man". 

Os innominatum. (Plate XXXIX.) 

This complex bone consists, as in the higher forms, of the ilium, ischium, and pubis 
anchylosed together. The ilium is wide, but less so in proportion to its height than in 
tlie Gorilla, and very much less so than in Man, being in fact much as in the 
Chimpanzee, though perliaps on the whole somewhat broader''. 

The external surface (PI. XXXIX. fig. 1) is convex anteriorly, concave posteriorly; 
but generally the concavity is very much more marked and extensive than is the 
convexity, in which the Orang agrees with Troglodytes — as also in the depth of the 
concavity, which is much greater than in Man. The curved lines found on the human 
ilium are not to be distinguished in Simia any more tlian m Troglodytes ; and the bone 
is somewhat less developed posteriorly than in that genus. 

' Lucac, ?oc. cit. p. 308. 

' As often before remarked or represented, Owen, Trans. Zool. Soc. vol. i. p. 365 ; De Blainvillo, • Osteo- 
graphie," Primates, Pithecus, p. 30; Huxley, 'Medical Times,' 1864, vol. i. p. 565; and Huxley ifc Hawkins. 
' Atlas of Comparative Osteology,' plate x. fig. 3. Also Duvernoy, ' Archives du Mus.' vol. viii. p. 27 ; and 
Lucae, he. cit. p. 305. 

' As mentioned by Professor Owen, Trans. Zool. Soc. vol. i. p. 363, and by Professor Huxley, ' Medical Times," 
1864, vol. i. p. 565. 



MR. ST. G. MIVABT ON THE SKELETON OF THE PEIMATES. 195 

The intemal surface (PI. XXXIX. fig. 2) looks foi-wards, but not inwards, thus 
agreeing with the Chimpanzee, and differing from Man and the Gorilla. The part of 
the internal surface which is above the ilio-pectineal line is flat, or with only a very 
slight concavity' ; this is sometimes supplemented, however (PI. XXXIX. fig. 4), by an 
inflection of the anterior superior angle of the ilium. The " auricular " surface is more 
elongated in proportion to its breadth than in Man, in which respect it resembles 
Troglodytes (PI. XXXIX. fig. 2 in). That part of the internal surface which is beneath 
the ilio-pectineal line is more convex than in Man and the Gorilla, the ilio-pectineal 
line itself not being so prominent as in these forms. This part of the inner surface 
(PL XXXIX. fig. 3) is also more elongated than in Man. 

The crest of the ilium sometimes describes a decided sigmoid curve (PI. XXXIX. 
fig. 5), though this is always much less marked than in Man, and is occasionally absent, 
namely, when the anterior end of the summit of the ilium is not at all inflected. In Man 
the crest is enlarged somewhat behind its anterior end, and more or less immediately over 
the acetabulum. In the Orang (as in Troglodytes) no such widening takes place ; but 
on the other hand, there is sometimes at the anterior end of the crest (PI. XXIXIX. 
fig. 4 & 5 a) a very marked enlargement, which may answer to the one above-mentioned 
of Man ; and if so, we may imagine that part of the ilium which in Man is anterior to 
it, to be altogether absent in the Orang. As in the higher forms, so also in Simia, 
the crest is always enlarged at its posterior end ; but the crest, as a whole, is (as in 
Troglodytes) narrower in proportion to its length than in Man. The vertical curvature 
(as in the Chimpanzee) is much less than in Man or the Gorilla ; but the crest is 
produced upwards somewhat suddenly at about the anterior end of its posterior tliird 
(PI. XXXIX. fig. 1). 

The anterior margin of the ilium is always concave, and often more so than in 
Troglodytes, though, as in the latter genus, the wide distance between the anterior 
spinous processes causes it to dificr much from the form of the anterior margin of the 
ilium of Man (PI. XXXIX. fig. 2). 

As in Troglodytes, the anterior superior spinous process in Simla is not so marked 
and distinct a process as in Man ; but the anterior inferior spinous process (PI. XXXIX. 
fig. 2 b) is sometimes almost, if not quite, as prominent as in him. Sometimes, however, 
it is not more marked than in Troglodytes. 

Within this process, and above the acetabulum, the ilium presents a smooth surface 
for the ^.so«s and iliacus muscles; and no ilio-pectineal prominence marks the junction 
of the ilium with the pubis. 

The upper part of the posterior margin of the ilium is, on the whole, nearly straight 

to the lower margin of the auricular surface ; and though its outline is irregular, there 

is scarcely any trace of the concavity which exists in Man between the posterior 

termination of the crest of the ilium (or posterior superior spinous process) and the 

' M. Duvernoy says " un peu creux,'" ' Archives du Museum,' toI. i-iii. p. 28. 

2e2 



196 ME. ST. G. JMIVAKT ON THE SKELETON OF THE PEIMATES. 

upper end of the auricular surfece. This upper part of the posterior margin is, as also 
in Troglodytes, myxcYi longer than in Man. 

The lower part of the posterior margin (below the auricular siu-face) is very decidedly 
concave, but, as in Troglodytes, the concavity is nothing like so strongly marked as in 
the human ilium (PI. XXXIX. figs. 1 & 2 g). Indeed it is rather less marked than 
appears to be generally the ca.se in Troglodytes. 

The ilium forms a considerable portion of the acetabulum, but not quite so much of 
it as does the ischium. It is altogether superior to the depressed surface (for fat and 
vessels), which surface is entirely formed by the last-named bone. 

The pubis has a horizontal ramus with three surfaces and three prominent lines, as 
in Man. 

The superior or horizontal surface is broader than in Troglodytes, though it is never- 
theless considerably narrower Ihan in Man. This surface presents a narrow groove 
running from without inwards, and concave from behind forwards, the concavity being 
much mcrcased by a very large process. This process (PI. XXXIX. figs. 2, 3, & 4^;), 
which exists in almost all adults, is entirely formed by the pubis, and, being situated 
at the internal termination of the ilio-pectineal line, is probably (as Professor Owen' 
names it) the spine of the pubis. It is nevertheless so remote from the symphysis 
and so near the acetabulum that it has rather the appearance of an iliopectineal 
eminence. 

The internal sm-fixce of the horizontal ramus (PI. XXXIX. fig. 3) is smooth, and 
(not counting the just-mentioned spine) is narrower vertically, above the obturator 
foramen, than in Man. 

The external or inferior surface is, as in Man, deeply grooved (PI. XXXIX. fig. 4). 
Indeed it is often much more so than is sometimes the case in him, the Orang in 
this differing markedly from the Gorilla and Chimpanzee ; in both of which (as far as 
I have observed) the groove is never more than slightly marked, while in the Chi7n- 
panzee it is often altogether absent. 

The external extremity of the bone forms but a very small portion indeed of the 
acetabulum. 

The body of the pubis is, as also in Troglodytes, thinner from witliin outwards, and 
more vertically extended than in Man ; the margin bounding the obturator foramen is 
also thinner than in him. 

There is no spine of the pubis other than tiie one idready mentioned, and no process 
whatever near the symphysis, which, as also in Troglodytes, is much longer than in Man 
(PI. XXXIX. fig. 2 s). 

The descending ramus of the pubis resembles that of the Gorilla and Chimpanzee in 
being much wider than that of Man. 

' Trans. Zool. Soc. yol. i. p. 363. W. Vrolik calls it " Epine pubieuue ou eminence ilio-puctinee," ' KecherchcB 
d"Anat. Comp. siu le Chimpanse,' p. 10. 



ME. ST. G. IVnVAIlT ON THK SKELETON OF THE PRUfATES. 197 

The ischium consists, as in Man, of a body and ramus ; and part of the external 
surface of the former constitutes the greater portion of the acetabuknn, including (as 
before said) the whole of its depressed tract. Below the socket for the femur the 
other part of the antero-extemal face (PI. XXXIX. fig. 4) presents (as in Troglodytes) 
a wide surface of bone— concave from above do\^^lwards, and strongly convex from 
behind forwards — in the place of the narrow groove which in Man separates the 
acetabulum from the ischiatic tuberosity. The postero-extenial surface of this part of 
the ischium (PI. XXXEX. fig. 1) is similarly elongated as compai-ed with Man's structure, 
but in all the forms it is smooth and bounded inferiorly (as is also the outer surface) by 
the margin of the tuberosity of the ischium. In the Orang this surface is not prolonged 
backwards, as in Man, by so prominent an ischiatic spine, though this process is 
considerably more developed (PI. XXXIX. figs. 1-4 h) than in the Gorilla, and 
sometimes than in the Chimpanzee also. 

The tuberosity of the ischium is formed very much as in Man, and is less flattened 
beneath, and has its margin somewhat less everted than in TrogJodytes. At the same 
time the Orang resembles the last-named genus, and differs from Man in that the 
rugose surface is prolonged more in the direction of the symphysis pubis (PI. XXXIX. 
fig. 6), and less in that of the spine of the ischium than it is in him. It is, however, 
decidedly more prolonged up backwards towards the last-named process, than in Troglo- 
dytes (PI. XXXIX. fig. 3 I). 

The ramus of the ischium in the Orang agrees with that of the Gorilla and that of 
the Chimpanzee in being very much more vertically extended than in Man. Its 
external sui'face is also more concave, and its inferior border more everted, while the 
margin bounding the obturator foramen is thinner than in him. 

The last-mentioned foramen is generally somewhat subtriangular, with one angle 
turned towards the outer end of the horizontal ramus of the pubis. 

The acetabulum is longer vertically, in comparison with its breadth, than in Man. Its 
depressed surface and the cotyloid notch are very much smaller than in the higher forms 
(PI. XXXIX. fig. 4/). 

The acetabulum is deepest superiorly, and more predominantly deep there than in 
Man, and somewhat more so than even in Troglodytes. 

The anterior part of the pelvis does not descend so much as in the last-mentioned 
genus, but is more like that of Man in this respect. 

The false pelvis is longer and more shallow than in the Gorilla, and still more 
so than in Man. As in Troglodytes the inlet of the true pelvis is "less' con- 
stricted anteriorly, less cordate, and more fully elliptical in shape " than in the 
human form. The ellipse, however, is, sometimes at least, less elongated than in 
Troglodytes. 

As in the Gorilla and Chimpanzee, but a small part of the acetabulum is visible 

' Owen, Trans. Zool. Soc. vol. v. p. 14. 



198 ME. ST. G. MIVAET OX THE SKELETON OF THE PRIMATES. 

when the outer surface of the ilium (PL XXXIX. fig. 1) is opposite the observer, 
instead of almost the whole of its cavity being so, as is the case in Man'. 

Femur. (Plate XL. figs. 1-7.) 

This bone is exceedingly short in the Orang, both absolutely and relatively, as com- 
pared with Man and the Gorilla. 

In Troglodytes it is more than half the length of the spine (measured as before), 
and in Man it is more than three-fifths of it ; in the Orang, however, I find it less 
than half. 

As compared with the length of the os innominatum the femur is somewhat longer in 
the Orang than in Troglodytes — as it is decidedly the longer of the two ; still the pro- 
portion very much more resembles that existing in the last-named genus than Man's, 
as his femur is about double the length of his os innominatum. 

A comparison of the femur with the humerus shows a greater difference from Man 
than that presented in Troglodytes, though even in the Chimpanzee the femur is slightly 
the shorter of the two, instead of very much the longer, as in Man. 

When the femur of the Orang is made to rest with both condyles on a horizontal 
surface, and placed as nearly as may be in a vertical position, the bone does not incline 
outwards (peronead) superiorly so much as does that of Man when similarly placed ; it 
does so, however, in a slightly greater degree than is the case in Troglodytes-. 

The body or shaft of the bone differs much from that of the femur of Man, and 
greatly resembles that of Troglodytes. This is the case as regards the absence of a 
strongly projecting linea aspera, the less transverse convexity of the anterior surface, 
the much greater antero-posterior compression of the bone, and its less degree of cur- 
vature convex forwards, the shaft being even straighter than in the Gorilla (PI. XL. 
figs. 3 & 4). It also differs from Man's, and agi-ees with that of Troglodytes, in tiie 
large proportion borne by the transverse diameter to the length — though in this 
respect it resembles the Chimpanzee, it being more slender than in the Gorilla. The 
lateral expansion downwards of the shaft, though more gradual than in Man, is much 
less so than in the Gorilla ; and the external margin of the same is more concave than 
in Troglodytes, and approximates, therefore, in its outline to Man's. 

In the Orang, as in the Gorilla and Chimpanzee, the external and internal surfaces 
of the shaft are much narrower from before backwards than in Man. This arises from 
the non-projection of the linea aspera, which seems to be, as it were, flattened out in 
both Simia and Troglodytes, though least so in the Gorilla. 

The anterior intertrochanteric line (PI. XL. fig. 1 e) is continued into the spiral line 
(PI. XL. figs. 2 & Sy), and can be followed downwards to the entocondyloid prominence 

' Owen, Trans. Zool. Soe. vol. v. p. 14. 

- There is a certain amount of individual variation in this respect : in the specimen No. 3 c in the British 
Museum, the inclination is considerable, approaching that of Man. 



MR. ST. Q. MIVAET OX THE SKELETON OF THE PRIMATES. 199 

(PI. XL. fig. 2 h). About halfway down it approximates more or less towards the 
middle line of the posterior surface of the femur. 

On the opposite side of the bone a more or less marked line, or rugose tract, extends 
from the base of the great trochanter downwards towards the external condyle'. The 
posterior surface of the femur, between these two lines, is more or less rough and 
irregular, and contrasts with the smoothness of the rest of the shaft. 

As in Troglodytes, so also in Simla, there is no strongly marked ridge descending 
quite to the ectocondyloid prominence (as is the case in Man), neither does any ridge 
run downwards from the lesser trochanter. 

Below the great trochanter, and more or less in the course of the long line descending 
towards the external condyle, there is a marked and rough depressed surface for the 
gluteus mcaimiis (PI. XL. fig. 4 g). The lower end of this depression does not reach to 
the middle of the bone's vertical extent. 

As in Troglodytes, the popliteal space is flatter than is the case in Man. 

The entocondyloid prominence (PI. XL. figs. 1 & 2 Z) is somewhat more developed 
than in Troglodytes, but not quite so much so as in Man. The ectocondyloid one 
is about the same as in the last-named genus, and therefore is more marked than 
in the human femur (PI. XL. figs. 1 and 2 k). 

I have not seen a conspicuous medullary foramen on the posterior surface of the 
femur in any one specimen of Simia. 

As in the higher forms, the neck is considerably more vertically than antero- 
posteriorly extended. It forms with the shaft an angle (open inwards and downwards) 
of about 155°, which is considerably gi-eater than that in Man, or than that in the 
Chimpanzee, and much more so than the corresponding angle in the Gorilla-. There 
is nevertheless some slight individual variation in this angle. 

The great or peroneal trochanter never attains so high a level, compared with the head 
of the bone, as in Man, and still less than as in Troglodytes. In shape it is very like 
the corresponding part in the higher fonns, except that it differs from that of the 
Gorilla, and resembles that of the Chimpanzee, and still more that of Man, in its 
peronead projection (PI. XL. figs. 1, 2, 4, 6 b) beyond the line of the external margin 
of the shafts. 

The lesser, or tibial, trochanter is shaped much as it is in the Chimpanzee, being less 
elongated than is sometimes the case in the Gorilla, and less conical than in Man. 
"Wlien looked at from above (PI. XL. figs. 1, 2, .3, 4, 6 c) it appears, as also in Man 
and the Chimpanzee, closer to the head of the bone than is the case in the Gorilla '. 

' Giving origin in part, probably, to the femoral portion of the Biceps, as Mr. W. S. Cliurch notes its origin as 
" extending from 2j inches below the great trochanter to within the same distance of the external condyle'' 
(Nat. Hist. Review, 1862, vol. ii. p. 86). 

- See Owen, Trans. Zool. Soc. vol. v. p. 1.5, plate 7. 

' Owen, Trans. Zool. Soc. vol. v. p. 1.5. ' Owen, he. cit. plate 7. 



200 MR. ST. G. MIVART OX THE SKELETON OF THE PRIMATES. 

The trochanteric fossa is deeper than in the Gorilla, but, as also in Man, it is less 
deep than in the Chimpanzee. The intertrochanteric line behind is rather more 
sharply and strongly prominent than in any of the higher forms (PI. XL. fig. 2 d). 

The head of the femur is very large, especially as compared with that of the 
Chimpanzee, though absolutely exceeded by that of Man, and also by that of the 
Gorilla when of large size. It is sharply defined by a prominent border all round, 
except sometimes for a short space near the intertrochanteric fossa (PI. XL. figs. 1, 
2, .8, 4, G, 7 «). 

The sharp projection of its anterior margin is more like what exists in Troglodytes 
than in the general structure of Man. 

It is commonly asserted that the ligamentum teres is absent in the Orang, as also 
the pit for its reception on the head of the femur'. I find no trace of the latter in 
either femur of any specimen, with one exception'-' ; but in that exceptional specimen 
each femur (PI. XL. fig. 7 i) exhibits a small but distinct depression on its head in 
the place occupied in other forms by the pit "for the round ligament. This absence 
has not, as far as I am aware, been noticed in Man or the Chimpanzee ; but in the 
Gorilla I have sometimes been unable to detect any trace of such a fossa on the head 
of the femur''. It may tlierefore be the case that this ligament is occasionally absent 
in the Gorilla, and occasionally present in the Orang. 

The rotular surface (PI. XL. fig. \o) does not, as in Man, -project higher on the 
peroneal than on the tibial side, but more resembles in this respect that of the Gorilla 
than even that of the Chimpanzee ; it extends, however, further up the shaft, and 
has its superior margin more acutely convex than in Troglodytes. It is even less 
concave transversely than in the Gorilla and Chimpanzee, and therefore still more 
widely differs in this respect from the rotular surfiice of Man than do the corresponding 
parts in them. 

The external condyloid articular surface is somewhat narrower than is the internal 
one, but the difference is less than in Troglodytes* (PI. XL. fig. 5 m «& n). 

The breadth of the intercondyloid fossa (which, as in the Gorilla, is generally 
shallower than in Man) is about equal at its anterior and posterior ends. 

As in Troglodytes, the wliole distal surface of the bone is broader in proportion to 
its antero-posterior extent than in Man^, and the external condyle projects backwards 
less than the internal one does — the external one being, as in that genus, the shorter 
one from before backwards, instead of rather the longer of the tAvo as in Man" 
(PI. XL. fig. 5 m & n). 

' Owen, Traus. Zool. Soc. vol. i. p. 3Gi5, and Do Bhiiuville, ' Osteographio,' Primates, Pitheciis, p. 31. 
- No. 3 i in the osteological collection of the British Museum. 

' E. g. in all four femora of the specimens Nos. 5179 a and 51 79 b in the Museum of the Koyal College of 
f^iu-geons. * Owen, Trans. Zool. Soc. vol. v. p. M, plate 7. fig. 3. 

' Owen, he. cit. p. IG, plate 7. fig. 3. ' Owen, he. cut. p. 18. 



MR. ST. G. MIVAET OX THE SKELETON OP THE PEIMATES. 201 

Patella. (Plate XL. figs. 8 & 9.) 

The patella of the Orang has its surface marked with vertical grooves, as in Man, 
but it agrees with that of Troglodytes and differs from the human patella in being 
more rounded and without the produced inferior apex, in having no median vertical 
projection on its posterior surface, and in the subcquality of thickness of its outer and 
inner edges, as well as of its superior and inferior ones. 

It differs from that of all the higher forms in its greater breadth, in the less convexity 
of its outer surface, in the almost complete flatness of its inner surface', and in its 
smaller size, as compared with the adjoining ends of the femur and tibia. 

Tihia. (Plate XLI. figs. 1-5, 8, 9.) 

The length of the tibia, as compared with that of the spine, is much as in Trorjlodytes, 
and nearly one-fifth less than in Man. The proportion borne by it to the femur I have 
found larger than in the Chimpanzee or Gorilla. As compared vnth the radius, the tibia 
of the Orang is much shorter than that of Troglodytes ; but yet the difference is much 
less than between the latter genus and Man, in whom the tibia is about half as long 
again as is the radius. 

Besides the relative length of the bone, the Orang differs from Man and agrees with 
Troglodytes in the great relative width and less lateral compression of the tibia, in the 
convexity, vertically, of its anterior surface, the vertical concavity of its outer or 
peroneal surface (PI. XLI. fig. 1), the shortness and bluntness of the crest, and the 
more rounded form of the shaft, which renders it somewhat difficult to describe 
according to the three surfaces and three margins which exist in Man. 

As also in Troglodytes-, the transverse diameter of the superior surface is greater, 
compared with the antero-posterior diameter of the same, than is the case in Man. 

In the greaterprojection, tibiad, of the internal tuberosity and in the stronger vertical 
conca^ity of the inner surface of the bone leading down from it to the shaft, tlie Orang 
resembles the Chimj^anzee, and differs from the Gorilla, and still more from Man. 

The tubercle, as also in Troglodytes, is less prominent than in Man ; but there is some 
individual variation in this respect. 

The external tuberosity is at least as large as, if not larger than, the internal one ; 
and its thickness between the articular surface for the femur and that for the fibula is 
(PI. XLI. fig. 2), as also in Troglodytes, relatively, and often absolutely, greater than in 
Man. The latter articular surface, again, as in Troglodytes, is also larger relatively 
than in Man (PI. XLI. fig. 5 k), but it is flat, instead of, as in the Gorilla, strongly 
convex. 

The groove for the tendon of the popliteus is very slightly marked ; but the depres- 
sion for the semimembranosus (behind the internal tuberosity) is very much so 

' W. Vrolik, ' Recherches d'Anat. Comp. sur Ic Chimpanse,' p. 15. 
- Owen, Trans. Zool. Soc. vol. v. p. 1 9. 
VOL. VI. — PAKT IV. 2-F 



202 MR. ST. G. MIV.AJRT OX THE SKELETON OF THE PRI31ATES. 

(PI. XLI. fig. 4 ?■), and, as in Troglodytes, is more rounded and less antero-posteriorly 
elongated than in Man. 

Of the two articular surfaces for the condyles of the femur, the internal one has its 
inner margin (as in the Gorilla and Chimpanzee) more convex and prominent than in 
Man (PI. XLI. fig. 8) ; and the concavity of its surface is mainly produced, as Professor 
Owen remarks of the Gorilla', by the elevation of that part of it which joins the spine. 

The external articular surface (for the external condyle of the femur), though more 
convex antero-posteriorly than in INIan, is less so than in the Gorilla ; and sometimes, 
indeed, it is decidedly, though very slightly, concave antero-posteriorly. 

The spine is also more human in its form than it is in either Troglodyte, being 
almost as bifid as in Man ; while the groove wliich descends backwards from its apex, 
and divides the posterior ends of the articular surfaces for the condyles, is considerably 
deeper, and more marked than in any of the higher forms (PI. XLI. figs. 2 & 8). 

The lower end of the tibia is inclined so that the anterior margin of its distal end is 
much more oblique (downwards and tibiad) than in Troglodytes, and very much more 
so than in the human tibia. This obliquity arises from the large size of the lower 
articular surface for the fibula, and from the inclination inwards of the articular surface 
of the tibial malleolus. 

The inner, or free, surface of the last-named process projects more strongly tibiad 
than in Troqiodytes — the inner surface of the shaft immediately above it being more 
concave vertically than in that genus, though scarcely more so than in Man. 

'The anterior margin of the distal end of the tibia is more prominent than in the 
higher forms, the surface of the shaft just above it being more concave, vertically, than 
in them. 

The distal end of the posterior surface of the tibia has a deeper groove for tlie tibialis 
posticus and flexor lonyus Jiallucis than I have obser\ed in any of the higher forms 
(PI. XLI. fig. 2;)). 

As in Troqiodytes, so also in Simia, the tibial malleolus is more truncated at its apex 
than is the case in Man. 

The distal articular surfiice of the shaft of the tibia (PI. XLI. fig. '■)) is, as in the 
Chimpanzee-, far more convex transversely than in the Gorilla or in Man. On either 
side of the strong median convexity there is (also as in the Chimpanzee) a slight 
transverse concavity; but the whole surface presents only a mere trace of an antero- 
posterior concavity, while, as in Troglodytes, its anterior margin descends as much (when 
the shaft is vertical) as does its posterior margin, instead of, as in Man, the latter margin 
descending further than the anterior one. 

The articular surface on the outer (peroneal) side of the malleolus foi-ms a more 
open angle with the distal surface of the shaft than even in Troglodytes, and its vertical 
extent is also mudi less (PI. XLI. fig. 1). 

' Trans. Zool. Soc. vol. v, p. 19. - Owen, he. clt. p. 20. 



ME. ST. G. MIVAET ON THE SKELETQN OF THE PKIMATES. 203 

The articular surface for the lower part of the fibula is much larger than in Man and 
'Troglodytes, being about the same size as the malleolar one for the inner side ot the 
astragalus (PI. XLI. figs. 5 & 9 s). 

The shaft of the tibia is, as in the higher forms, triangular above its middle ; but 
below that it is, as in TrofiJodytes, much more rounded than in ]\Ian. 

The internal surface is convex, except at its upper part and just within the crest and 
below the tubercle, in which last situation there is, as also in Troglodytes^ a rough 
and depressed surface (PI. XLI. fig. 1) for the insertion of the sartorius, gracilis, and 
semitendinosus muscles. 

The external, or peroneal, surface of the tibia is, as in Troglodytes, more strongly 
concave above than in Man, while from rather above the middle of the bone, it merges 
insensibly into the anterior surface. This external surface of the human tibia is much 
broader from behind forwards than is the part whicli corresponds to it in Simla, if the 
faint ridge (PI. XLI. fig. 5 /) running downwards from the front of the upper surface 
for the fibula is that to which the interosseous ligament is attached. 

The posterior surface of the tibia presents an oblique popliteal ridge, which, however, 
as also in Troglodytes, is much less strongly marked than in Man. 

The medullary foramen (PI. XLI. fig. 2 n) is much as in Man and the Chimpanzee, 
and more above the middle of the bone than I have observed it to be in the (jorilla. 

The anterior border, or crest of the tibia, is less sharp, much shorter, and inclines 
more markedly tibiad below than in Man, agreeing in these points with Troglodytes, 
except that in Slinia the crest is rather more sharp towards its upper end. 

The inner border, as also in Troglodytes, is so little marked as to be liardly distin- 
guishable, except for a short distance above the posterior border of the malleolus. 

There is a well-marked external or peroneal border, which, however, does not 
correspond with the external border of the tibia of Man, inasmuch as it descends from 
behind, and not from in front of, the upper facet for the fibula ; it becomes lost about 
halfway down the tibia. 

Another and less marked external ridge (PI. XLI. fig. 5 /) appears to correspond witli 
the external margin in Man. It springs from a point anterior to the upper articular 
surface for the fibula, and descends to the apex of the lower articular surface for that 
bone. This ridge is rather more strongly marked in Troglodytes. 

Fibula. (Plate XLI. figs. 1, 3, G, 7, 10, & 11.) 
As in Troglodytes^, this bone is much more distant from the shaft of the tibia than in 
Man ; it is also shorter, stouter, and straightcr than in him, and has the ridges and 
depressions on its surface less marked, — in all which points the fibula of the Orang 
agrees with that of the Gorilla and that of the Chimpanzee. 

The proximal articular surface (PI. XLI. figs. 7 & 10) is more rounded tlian in 
' Owen, Trans. Zool. Soc. vol. v. p. 19. - Owc-n. Joe. cit. vol. i. p. 30(i. 

2 f2 



204 ME. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 

Man, and, though slightly concave, has not the marked depression existing in 
Troglodi/tes. 

As in the Gorilla and Chimpanzee, the outer side of the head has a very marked 
prominence (PI. XLI. figs. 1, 3, & 6 ^) for the long external lateral ligament and the 
tendon of the biceps. There is no styloid process, that eminence being more completely 
absent than in Troglodytes. 

As in the last-named genus, so also in Simla, one single anterior ridge appears to 
answer to both the external and internal anterior lines of Man' (PI. XLI. fig. In). 
This ridge descends along the anterior surface of the bone, and bifurcates very near its 
inferior end. 

The margin answering to the posterior external one of Man (PI. XLI. fig. 3 v) is 
quite indistinct, except at the lower part of the bone. It extends obliquely, from the 
inner side of the posterior surface of the malleolus, in an upward and outward direction, 
over the postei'ior surface of the shaft of the fibula, for a greater or less distance towards 
its head. 

There is in the Orang, as in Man and Troglodytes, a posterior internal ridge (PI. XLI. 
figs. 3 & 7 w) which runs obliquely downwards and forwards from the inner aspect of 
the head of the fibula, and joins the anterior margin before mentioned. 

The medullary foramen (opening distad as in Man and Troglodytes) is placed more or 
less near to the middle of the bone (from above downwards) below and rather behind 
the posterior internal ridge (PI. XLI. figs. 3 & 7^). 

The contraction of the anterior surface of the fibula in Simia and Troglodytes, through 
the coalescence as it were of the external and internal anterior margins of that bone in 
Man, is more than compensated for by the wide surface for muscular attachment 
ofiered by the interosseous ligament. 

As in Troglodytes, the peroneal malleolus is shorter and blunter than in Man, and does 
not descend below the tibial one^ (PL XLI. fig. 1). 

Pes. (Plates XLI. fig. 12, & XLIII.) 

The absolute length of this segment in the Orang exceeds that of the higher forms ; 
as also its length as compared with the spine, with the rest of the pelvic limb, and 
with the tibia. 

The proportion borne by the length of the pes to that of the manus is, as also in 
Troglodytes, much less than in Man. 

Tarsus. 
Unlike the carpus, the tarsus consists of the same number of bones as in the higher 

' The fibula of Man is very lucidly described by ilr. A. T. Norton, in his coiivenient and carefully prepared 
' Osteology for Students,' recently published. - Owen, Trans. Zool. Soc. vol. v. p. 20. 



ME. ST. G. MIVAET OX THE SKELETON OF THE PRIMATES. 205 

forms. In absolute length this segment falls short of that of the Chimpanzee, and it is 
very much shorter than the homologous part in Man and the Gorilla. 

Compared with the length of the spine, that of the tarsus is veiy slightly less than in 
the Chimpanzee, but falls more short of that in Man and the Gorilla. The proportion 
borne by it to the whole length of the pes is more characteristic, as in the Orang it 
appears to be only as about 26-6 to 100, while in Troglodytes it is .36 or 40 to 100, and 
in Man is as about 46 to 100. 

Calcaneum. (Plate XLIII. figs. 2-7.) 

The OS calcis of the Orang is very unlike that of Man, or that of the Gorilla, and 
more resembles that of the Chimpanzee. The projection of the heel backwards 
beyond the hinder margin of the posterior articular surface for the astragalus, some- 
times about equals, sometimes falls short of the antero-posterior extent of that surface. 
The Orang in this differs widely from Man, and still more from the Gorilla, in which 
last the length of the os calcis behind the posterior margm above-mentioned exceeds 
that of all the bone anterior to it. In the Chimpanzee the length of the os calcis 
behind the posterior articular surface for the astragalus does not quite equal the 
antero-posterior extent of that surface, which, again, exceeds that of the bone in 
front of it. 

In the Orang the length of the os calcis anterior to the same surface (PI. XLIII. 
fig. 6) sometimes equals, sometimes falls short of that of the bone behind it. The 
upper surface of the last-mentioned posterior portion is more or less concave antero- 
posteriorly (PI. XLIII. figs. 2 & .3), more so than generally in the Gorilla, though 
not so much so as in the Chimpanzee. 

The posterior surface of the tuberosity is more narrow' transversely than in Man 
or Troglodytes, and is prolonged both upwards and downwards, the latter making the 
plantar surface much concave antero-posteriorly (PI. XLIII. tigs. 2, 3, & 5). Although, 
when the surfaces for the astragalus are horizontal, the tuberosity inclines strongly tibiad 
at its plantar end, — yet this inclination is (sometimes at least) not so great as in the 
Gorilla'. The outer or peroneal face of the calcaneum has a somewhat more human 
aspect than has that of the Gorilla, inasmuch as it is vertically less convex and more 
extended, relatively, than in the latter; sometimes even there is a slight vertical 
concavity. 

The posterior articular surface for the astragalus is not so convex as in Troglodytes, 
and it is more posteriorly placed, with respect to the anterior articular surface, than is 
the case in that genus or in Man. As in the Chimpanzee, there is no trace of a second 
posterior plantar tubercle ; that for the external lateral ligament is more posterior in 
position than it is in any of the higher forms, though in this the Orang more resembles 

' Professor Huxley remarks, " The calcaneal process is narrow from side to side," ' Medical Times,' 18f)4, 
vol. i. p. 50.5. - Dr. Lueae remarks this, he. cit. p. iS04. 



206 ME. ST. G. MIVART ON THE SKELETON OF THE PEIMATES. 

the Chimpanzee than it does Man or the Gorilla. Sometimes, in the Orang, there is a 
marked antero-posteriorly directed groove above this peroneal tubercle (PI. XLIII. 
fig. 2 e), but it is never bounded inferiorly by such a strongly projecting ridge as 
exists in the Gorilla. 

The tubercle for the calcaneo-cuboid ligament is distinct (PL XLIII. fig. 7/), but 
not prominent as it is generally in Man. 

The articular surface for the cuboid is nearly vertical ; but the depression at the lowest 
part of the tibial side is extraordinarily deep, forming a funnel-shaped cavity (PI. XLIII. 
fig. 4 c) for the reception of the very long conical and pivot-like prominence on the 
posterior surface of the cuboid '. 

Astragalus. (Plate XLIII. figs. 8-13.) 

This bone, in Simia, has a very different appearance from that of any higher form, 
owing to the great length and strong tibiad inclination of its neck (PI. XLIII. fig. 8 a). 

The superior articular surface (for the shaft of the tibia) is sometimes rather more 
concave transversely than in Man or Troglodytes, and it extends backward somewhat 
less. The head of the bone (for articulation with the naviculare) is also more compressed 
from above downwards than in them (PL XLIII. fig. 12 h). 

The difference in size between the articular surfaces for the two malleoli is greater 
than in Troglodytes or Man. That for the peroneal malleolus forms rather a slightly 
acute than a right angle, with the upper surface of the astragalus. That for the tibial 
malleolus forms, in the Orang, a very obtuse angle with the same, but yet not so obtuse 
a one as it does m the GoriUa. It encroaches more on the neck of the bone than in the 
liiglier forms. That part of the tibial face of the astragalus which is posterior to the 
surface for the malleolus, is much smaller than in Man or Troglodytes, but, as m the 
Gorilla, is separated from the latter by a deep groove (PL XLIII. fig. 10 d), behind which 
the surface is very smooth and rounded. On the other hand, that part of the peroneal 
face which is behind the surface for the outer malleolus (PL XLIII. fig. 11) is larger 
than in Man or Troglodytes, is more depressed, and has one or more small openings. 
The groove for the flexor tendon is wider than in Man or Troglodytes, and while more 
sharply limited on its peroneal side than in them, is less so on its tibial side (PL XLIII. 
figs. 9 & 13^). 

The posterior articular surface for the calcaneum has, as in the Gorilla, its anterior 
and inner margin more convex (PL XLIII. fig. 9 i), and its posterior and outer one more 
concave than in Man, its crescentic form bemg very marked. It is separated from the 

' This is doubtless connected with the great mobility, in the Orang, of the joint between the anterior and 
posterior tarsals, which has been noticed by previous observers. Professor Owen speaks of this, Trans. Zool. 
Soc. Tol. i. p. 367 ; and Professor Huxley remarks, " The mobility between the distal and proximal divisions 
of the tarsus is exceedingly great, and is the chief cause of the habitual turning inwards of the sole of the foot," 
• Medical Times,' 1864, vol. i. p. 50.5. 



MR. ST. G. MIVAET ON THE SKELETOX OF THE PRIMATES. 207 

anterior articular sui-face by a groove for the astragalo-calcaneal ligament. This groove, 
though somewhat deeper than in Man (PI. XLIII. fig. 9 k), is not nearly so much so as 
in the Gorilla. 

The anterior articular surflice for the os calcis (PI. XLIII. fig. 9 h) is much elongated, 
and is concavo-convex, but it is not very distinctly marked off from the rest of the 
articular surface of the head of the astragalus. 

Namculare. (Plate XLIII. figs. 14-16.) 

This bone in the Orang has a rather more marked proximal concavity tlian has its 
homologue in the Gorilla, and a much more marked one than that of Man. 

The vertical diameter of the same surface also predominates over the transverse one to 
a greater degree than in the human naviculare. Simia agrees with Homo, and differs from 
Troglodytes^ in having the plantar end of the hone less antero-posteriorly expanded ; and 
the tuberosity (PI. XLIII. fig. 15 e) is less produced than in Man, and much less so than 
in Troyloflijfes. 

The distal articular surface, as a whole, is narrower, in proportion to its vertical 
extent, in the Orang than it is in the Gorilla, and stiU more so than in Man. It agrees 
with that of Twrjlodi/tes, and differs from that of Man in the marked concavity of the 
sm-face for the ectocuneiforme (PI. XLIII figs. 14-16 5), and in the convexity of those 
for the mesocuneiforme and entocuneiforme. Yet the angle formed by the first of these 
with the surface for the mesocuneiforme is smaller than in the GoriUa, and stiU more 
so than in Man. In the Orang there is generally a larger facet (PI. XLIII. fig. 16 c) 
for the cuboid than in Tvof/lodi/fcs. The rough tract on the peroneal surface of the 
bone (between the facet for the cuboid and the peroneal margins of the proximal 
and distal articular surfaces) is much more extensive in the Orang tlian in Man or 
Troglodytes (PI. XLIII. fig. 16 d). 

Entocuneiforme. (Plate XLIII. figs. 22-26.) 
The entocuneiforme of Simia differs much from its homologue in Man or Troglodytes, 
its tibial surface being strongly concave, as also the margin connecting the surfaces for 
the first and second metatarsals. The articular surface for the metatarsal of the hallux 
has a greater vertical concavity than in any higher species, being sometimes, indeed, truly 
saddle-shaped (PL XLIII. fig. 23 & 24 a). Sometimes, however (PI. XLIII. fig 26 a), this 
vertical concavity is absent. This surface looks rather more tibiad than it does forwards ', 
and it cannot be said to be notched on its inner side — as is the case in Man and Tro- 
glodytes. The greater part of the anterior aspect of the bone is occupied by a rough 
non-articular tract which extends upwards and widely separates the surfaces for the 
first and second metatarsals' (PI. XLIII. figs. 2.3-26 c). 

' JI. Vrolik remarks that it is so placed " que Ic gros orteil doit s'ecarter des autres." ' Recherchcs d'Anat. 
Comp.' p. 1.5. = Jfoticcd by Professor Hu-vlej', see ' Medical Times,' 1864, vol. i. p. 565. 



208 ME. ST. G. MIVAET ON THE SKELETON OF THE PEIMATES. 

The bone, as a whole, tapers rather more upwards than in the higher forms, and 
would appear to do so more plainly but for the tibiad production of the anterior 
superior tibial angle, or summit of the surface for the hallux. 

The postero-peroneal face of the entocuneiforme is, in the Orang, almost entirely 
occupied by surfaces which articulate with the naviculare, the mesocuneiforme, and the 
second metatarsal. These articular facets are well defined by sharp margins (PI. XLIII. 
figs. 22-26 b, d, e), but are nevertheless continuous, and form an oblique band of 
articular surface extending from the posterior inferior angle of the bone to its anterior 
superior one. 

Mesocuneiforme. (Plate XLIII. figs. 27-31.) 

This bone in the Orang is less vertically and more antero-posteriorly extended than 
are its homologues in Man and the Gorilla. Its dorsum (PI. XLIII. fig. 27) also is 
larger antero-posteriorly (as compared with its transverse dimensions) than in Troglodytes, 
in which it more resembles that of man. As in the Gorilla and Chimpanzee, its posterior 
surface (PI. XLIII. fig. 31) is more concave than in the human mesocuneiforme. Its 
anterior surface (as also in the Gorilla) has its upper tibial part more bevelled off than 
in Man (PI. XLIII. fig. 30), and the rest of that surface is more concave than in him 
or in the Gorilla. The tibial surface of the bone (PL XLIII. fig. 28) presents one 
large articular surface (for the entocuneiforme) which is mainly extended from behind 
forwards, instead of the two distinct facets which exist in Man. 

The peroneal surface has, at its posterior plantar angle, a ratlier convex articular- facet 
(PI. XLIII. fig. 29 d) which articulates with the ectocuneiforme ; and there is also a more 
or less marked articular surface extending antero-posteriorly along the top of this 
peroneal face. 

Ectocmieiforme. (Plate XLIII. figs. 32-3G.) 
In the Orang the ectocuneiforme has its proximal articular surface (PI. XLIII. 
figs. 32 & 36 b) much more oblique and much more convex than in either Man or 
Troglodytes. As in the latter genus, the posterior extension of the bone, below the 
hinder articular surface, is greater than in Man ; and the posterior inferior angle is 
produced into a rounded head (PI. XLIII. fig. Me). The distal articular surface is 
somewhat T-shaped (PI XLIII. fig. 35) and more concave than in the higher forms. 
The tibial surface presents a strong convexity above, and near its posterior plantar angle 
is a small concave facet (PI. XLIII. fig. 33 c) for articulation with the mesocuneiforme. 
The peroneal surface either presents two distinct facets for the cuboid, or these may 
coalesce and form one continuous articular surface for that bone (PI. XLIII. fig. 34 d). 

Cuboides. (Plate XLIII. figs. 17-21.) 
This bone is shorter antero-posteriorly, as compared with its transverse extent, than is 



MK. ST. G. MIVAllT ON THE SKELETON OF THE PRIMATES. 209 

the case in Man ; in this tlie Orang resembles the Troglodytes, as also in the greater 
distinctness of the distal surfaces for the fourth and fifth metatarsals respectively 
(PI. XLIII. fig. 19 «, b). That for the fourth metatarsal is more concave in both the 
vertical and transverse directions than it is in the Gorilla, and therefore much more 
so than in Man. As in the Chimpanzee, this surface occupies a greater share of the 
distal surface than is the case iu the Gorilla and in Man. The posterior surface of the 
cuboid (PI. XLIII. fig. 21) is very much more concavo-convex than in the other and 
superior forms ; and its tibial plantar angle is produced into the conical and pivot-like 
process before alluded to (PI. XLIII. figs. 17 & 18 c). As in Troglodytes, the under 
surface of the cuboides, behind the prominent ridge, is less cxt(msi\e than in Man ; and 
(as also in Troylodijtes) a small deep fossa exists just inside the angle formed by the 
junction of the above-mentioned ridge with the posterior margin of the plantar surface 
of the bone. On the tibial side of the bone there is a large articular surface for the 
ectocuneiforme, which surface is continuous behind with that for the na\iculare 
(PI. XLIII. fig. 20 e, b). As in Troglodytes, so in Simia, this surface descends nearer 
to the plantar margin than it does in Man. 

Metatarsus. 

The metatarsus attains a greater absolute length in Shnia than in Man or Troglodytes. 
The length of this segment of the limb (estimated by that of the second metatarsal), 
compared with that of the spine, is considerably greater than in the higher forms — being 
as about 17-2 to 100, instead of from 10 to 12-5 to 100 as in Man and Troglodytes. 

The relative length of the same metatarsal as compared with that of the entire pes is 
very much the same as in the others, namely about 32-1 to 100, while in Man, the 
Gorilla, and Chimpanzee I have found the proportion to be as 29-7, 28-5, and 30-G to 
100, respectively. 

The four outer Metatarsals. (Plate XLIII. figs. 37 & 40-54.) 

As in the higher forms, these metatarsals in the Orang are thicker at each end than in 
the shaft ; but the distal extremities are almost as broad as the proximal ends, thus differ- 
-ing from the four outer metatarsals of Troglodytes, and still more from those of Man. 

The shafts continue of nearly the same width throughout, and thus differ from those 
of Troglodytes, which taper distally, and still more from tliose of Man. On the other 
hand they do not broaden even slightly from behind forwards, as do their homotypes in 
the manus from above downwards. 

The transverse diameter of each head does not nearly equal its vertical dimension ; 
but still the difference is not so great as it is in the higher forms. 

The metatarsals iu Simia are rather more curved, with the concavity downwards, than 
are the metacarpals ; they also diverge distad somewhat less. Their shafts are scarce!)- 
more laterally compressed than are those of the metacarpals; yet they are slightly more 

VOL. VI. PART IV. 2 G 



210 ]ME. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 

slender than the latter, though the difference is much less than in Troglodytes. They 
are not more flattened beneath than are their homotypes of the manus. Their distal 
articular surfaces are destitute of the dorsal transverse groove which limits each of them 
posteriorly in Man and Troglodytes. On their plantar aspect the same surfaces have 
not those lateral projections which exist in the other forms. 

As in Troglodytes, so also in Simia, the dorsum of each distal articular surface slopes 
downwards towards its anterior end more than it does in Man. The extremities of the 
same surfaces of the fourth and fifth metatarsals of the Orang have also a very slight 
inclination peronead. A line connecting the proximal articular surfaces of the metatarsals 
of the Orang is almost at right angles with one following the antero-posterior direction 
of their shafts, these greatly differing fi-ora the condition presented by Man ; and alto- 
gether the metatarsus and metacarpus are less differentiated in Simia than they are in 
Man and Troglodytes. 

First Metatarsal. (Plate XLIII. figs. 38 & 39.) 

As in Troglodytes, the proximal end of this bone in the Orang presents a much more 
decided concavity than does that of its serial homologue, and is very different in form 
from the corresponding part in Man. 

In shape this metatarsal is very like that of its homologue in the Gorilla, and the 
direction of the distal groove (by which it articulates with the convex surface of the 
entocuneiforme) is oblique as in Troglodytes, and extends from above downwards and 
tibiad when the dorsum of the bone is placed horizontally. The shaft is also slightly 
twisted on its long axis. This bone in the Orang is sometimes a little longer, but 
generally a little shorter, than is the metacarpal of the pollcx, being thus unlike the 
same bone in Man and Troglodytes, where it is much longer than is its homotype. 

As in the other forms, so in the Orang, the proximal end of this metatarsal is larger 
than that of the corresponding metacarpal, but the difference is much less than in Man 
and Troglodytes ; thus this metatarsal, like the four outer ones, has a greater resemblance 
to its homotype in the Orang than it has in the last-mentioned forms. As in Troglodytes, 
the process for the attachment of the tendon of the peronceus longus is largely developed 
(PI. XLIII. fig. 39 a). 

Second Metatarsal. (Plate XLIII. figs. 40-43j. 
The proximal surface of this metatarsal in the Orang is much shorter vertically and 
tapers less downwards than that of Man (PI. XLIII. fig. 37 ir.). It is also more 
concave transversely than in him, and is somewhat convex ^ertically on its peroneal 
side. Thus the posterior margin of the dorsum of this metatarsal presents a notch, 
and in this as in the preceding points it agrees, more or less nearly, with its homologue 
in Troglodytes. 



ME. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 2Il 

This proximal surface resembles that of the homotypal metacarpal one more in tlie 
Orang than in the other and superior forms; but it may be readily distinguished by 
the less inclination distad of its upper part, and by the large articular facet for the 
entocuneiform (PI. XLIII. fig. 37 ii. a). 

On the peroneal side of the bone there are two articular facets for the third meta- 
tarsal (PI. XLIII. fig. 43 b, V), whicli are sometimes connected at theii- proximal ends 
by a very small vertical articular surface for the ectocuneiforme. 

As in Trocjlodytes, the tibial articular smface (for the entocuneiforme) in the Orang 
is relatively larger than it is in Man (PI. XLIII. fig. 42). 

There is a more or less marked tubercle at the proximal end of the plantar surface of 
the bone, between the articular suiface for the entocuneiforme and that for the meso- 
cuneiforme. 

Third Metatarsal. (Plate XLIII. figs. 44-47). 

The proximal surface of the third metatarsal of the Orang is less concave than that of 
man, and is, indeed, partially convex, as also in the Gorilla. It is more notched on its 
tibial side (PI. XLIII. fig. 37 in.) than in the just-mentioned forms, by a deep groove, 
which divides the two articular facets for the second metatarsal. In the Gorilla the 
lower facet appears to be wanting. 

On the peroneal side of the bone (PI. XLIII. fig. 47) there are two distinct facets, 
instead of one as in Man and the Gorilla. These are divided by a groove, which, 
however, scarcely invades the peroneal margin of the proximal surface. This surface 
may be distinguished from the homotypal one in the manus by its becoming much 
narrower downwards and by the absence of a marked concavity on its upper part. 

Fourth Metatarsal. (Plate XLIII. figs. 48-51). 

In the Orang, as in the Gorilla, the proximal surface of this metatarsal is destitute 
of the concavity which exists in Man, and is besides much more convex. 

Its inferior margin is straighter than in Man or the Gorilla, and the tibial side of the 
bone (PI. XLIII. fig. 50) has two articular facets for the third metatarsal, instead 
of one as in them. These facets are separated by a wide and deep groove. The 
peroneal side of the bone (PI. XLIII. fig. 51) has one long surface (for the fifth 
metatarsal) which is more vertically extended than in Man or the Gorilla. Compared 
with the homotypal surface of the manus, the proximal one of this metatarsal is more 
convex, the convexity extending on the peroneal side up to the dorsum, instead ot 
being interrupted by a concavity as in the manus. 

Fifth Metatarsal. (Plate XLIII. figs. 52-54). 

The fifth metatarsal of Simia has a proximal surface which is both more convex 
vertically and concave tranversely than that of the Gorilla, and still more so than tha.t 

2 g2 



212 ME. ST. G. MIVART ON THE SKELETON OF THE PRIMATES. 

of Man. The external proximal process, or tuberosity (PI. XLIII. figs. 52 & 5.3 a) 
is smaller in the Orang than in the higher forms; and in this the fifth metatarsal 
differs less from the fifth metacarpal than in them. 

As in the Gorilla, the tibial articular surface is less antero-posteriorly extended thnn 
in Man. 

The proximal surface of this metatarsal is very much more concave and less conve?; 
than is the corresponding surface of the fifth metacar-pal. 

Digits. 

The proximal phalanges of the four outer digits are very much curved', with the con- 
cavity downwards (PL XLI. fig. 12), much more so than are their homotypes of the 
manus. They are also narrower transversely, less flattened below and rather shorter, 
than are the latter. In these last three points, however, they differ less from their 
homotypes than do the same proximal phalanges oi Troglodytes from their serial homo- 
logues, and of course very mucli less than do those of Man. The proximal phalanx of 
the hallux is much shorter than is the homotypal segment in the Orang", a circumstance 
in which it differs from all the higher forms. The second phalanges of the four outer 
digits are shorter and naiTowcr than are their homotypes ; but again the difference is 
less than in Troghdjjtes, and greatly less than in Man. 

The second phalanx of the hallux is often absent', but wlien present is much shorter 
than is its homotype of the poUex'*, in which respect the Orang again differs from the 
superior forms. The third or distal phalanges scarcely differ in length from those of 
the manus, and at tlie most they are but a trifle shorter, thus agreeing with Troglodytes 
and differing much from Man. 

The hallux with its metatarsal, when compared in length with the spine, I ha\ c found 
to be only as about 13-6 to 100, instead of from about 17 or 18 to 100 as in Man and 
Troglodytes ; compared with the length of the entire pes, it barely exceeds a quarter, 
instead of approaching one-half as in them. The index with its metatarsal, as compared 
with the length of the spine, is very much longer than in the higher forms, as also is the 
third digit. Without their metatarsals, these digits, when compared with the length of 
the entire pes, are not so very much longer than in the Chimpanzee (.39 and ■12 to 100, 
instead of 32 and 34), but they of course greatly exceed those of Man. 

The proportion borne by the whole hallux to the whole poUex is strikingly different 

ProfcBSor Huxley, ' Medical Times," 18C4, vol. i. p. .''iGo. 

' See Lucae, he. cit. plate iii. figs. 5 & 9. ' Sec lucae. he. clt. plate iii. figs. 5 i 9. 

' Camper found this to be the case in seven out of eight Orangs (ffiuvres, torn. i. p. 54). Two phalanges, 
however, are recorded in two cases by Professor Owen, Trans. Zool. Soc. vol. i. p. 367. W. Vrolik, he. cit. 
pp. 15 & 16, says, " One or two phalanges are present," and appears inclined to think that the distal one may 
disappear with age. De BlainviUe, he. cil. p. 32, records five cases, in each of which the hallux had two 
phalanges. 



ME. ST. Q. MI V ART ON THE SKELETON OF THE PRIMATES. 213 

from that existing in the superior forms, the former only being about four-tifths tlic 
length of the latter, instead of much exceeding it, as in Man and Troglodytes. 

The proportion borne by the whole index of the pes to that of the manus is greater than 
that existing in the higher forms, the former being almost quite as long as the latter. 

The difference between the length of the index and that of the hallux is vastly 
greater than that in Man or Troglodytes. 

The fifth digit is the shortest, not counting the hallux ; the second is somewhat 
longer, then the fourth, and the third is the longest (PI. XLIII. fig. 1). The order of 
projection is similar to that of length. 

The proportion bome by the longest digit (without its metatarsal) to the longest meta- 
tarsal, is greater than in Troglodytes, and of com-se very much greater than in Man. 

The hallux only reaches as far forwards as from three-fourths to five-sixths of tlie 
length of the metatarsal of the index ; and therefore the Orang differs from Man and 
Troglodytes in that its hallux does not reach so far forwards in relation to the contiguous 
digit as does its pollex, while in them the hallux, thus estimated, extends further 
forwards than does the pollex'. 

The appendicular skeleton of Simia, while in some respects it more nearly resembles that 
of Man than does the corresponding structure in Troglodytes, yet on the whole departs 
further from the human skeleton than docs that of the Chimpanzee, or that of the Gorilla. 
This divergence is most marked in the extremities ; and the small differentiation of the 
bony structure of the terminal limb-segments of the Orang is especially remarkable. 

In addition to the various resemblances and differences offered by the limb-bones of 
Simia to the same parts in Homo and Troglodytes, I find that the Orang, when com- 
pared with all the species of the order Primates^ presents the following more or less 
noteworthy conditions : — 

The proportion borne by the pectoral limb to the spine is greater than in any other 
genus except Tarsius and Ilylobates. 

The proportion of the length of the radius to that of the spine is greater than in any 
other except Hylobates. 

The length of the index, with its metacarpal, compared with the spine, is greater 
than in any except Tarsius and ITylohates. 

The length of the metacarpal of the pollex is greater, in proportion to that of the 
spine, than in any other except Hylobates and Tarsius. 

The spine of the ischium is more largely developed than in any other Primate 
except Man. 

' See Huxley and Hawkins, ' Atlas of Comparative Osteology,' plate x. figs. 1 , 2 & 3, and plate xii. figs. 
1 a, 2a, 3 a, & 4 « ; also Dr. Luoae's paper before referred to, plate iii. figs. 2, 5 & U, plate ii. fig. 5, plate i. 
fig. 2, and plate iv. figs. 1, 5 & 6. 

' See a paper " On the Skeleton of the Primates," communicated to the Royal Society on November 22, 
1S66, read on January 10, 1807, and to be piiblishcd in the 'Philosophical Transactions,' vol. dvii. (1S(;7) 
p. 209, plates xi.-xiv. 



214 



MR. ST. G. MIVAET ON THE SKELETON OE THE PRIMATES. 



The length of the pes is greater, in proportion to that of the spine, than in any 
except Ateles, Cheiromys, and Tarsius. 

The proportion borne by the longest digit of the pes to the spine is greater than in 
any other Primate except Tarsius. 

The length of the longest digit, without its metatarsal, compared with that of the 
tarsus, is greater than in any of the order except the Nycticebinw and perhaps Indris. 

The Orang differs from every other Primate without exception in: — the great 
absolute length of the pectoral limb minus the manus, of the manus itself, of its third 
digit both with and without its metacarpal, and of the metacarpal of the poUex ; the 
great difference between the length of the poUex and that of the index ; the large 
diameter of the acetabulum compared with the length of the spine ; the small pro- 
portion borne by the femur to the humerus ; the very obtuse angle formed by the 
neck of the femur with its shaft ; the all but constant absence of the pit for the 
ligamentum teres on the head of the femur ; the shortness of the tibia compared with 
the humerus ; the length of the pes compared with that of the rest of the pelvic limb ; 
the length of the pes compared with that of the tibia ; the absolute length of the three 
middle metatarsals ; the absolute length of the longest digit with its metatarsal ; the 
veiy small proportion borne by the length of the hallux to that of the longest digit of 
the pes ; the occasional absence of the second digit of the hallux ; the great length of 
the index, with its metatarsal, compared with the length of the spine ; the small length 
of the hallux (both with and without its metatarsal) compared with that of the whole 
pes; the great length of the index, without its metatarsal, compared with that of the 
whole pes ; the nearly equal length of the indices of the pes and manus, both with and 
without the metatarsal and metacarpal ; the shortness of the tarsus compared with the 
length of the pes. Thus the Orang is one of the most peculiar and aberrant forms to 
be found in the order Primates. 



DIMENSIONS AND PROPORTIONS. 
' Scapula. 





Length from top 
ijf glenoid surface 
to inferior ver- 
tebral angle. 


Length 

of axillary 

margin. 


Length 

of vertebral 

margin. 


Length 

of superior 

margin. 


Greatest 
length of 
glenoid 
surface. 


Greatest 

breadth of 

glenoid 

surface. 


No. 3 A" 

„ 3B 

„ 3C 

Average of the 1 
three J 


7-15 
7-00 
7-50 

7-31 


G-75 
6-45 
G-90 

G-70 


5-75 

5-20 
6-00 

5-G5 


3-G5 
3-70 
3-40 

3-5S 


1-G5 
1-55 
1-75 

1-G5 


1-07 
1-00 
1-17 

1-08 


Variety Morio . . 


5-49 


4-82 


3-98 


2.98 


1-22 


0-81 



' The skeletons which have been selected for measurement are those of three adult males in the Osteological 
Collection of the British Museum, and numbered respectively 3 A, 3 B, and 3 C. 



MR. ST. G. MIVART ON THE SKELETON OF THE PEIIVIATES. 





Posterior 

vertebral 

angle. 


Angled formed 

by glenoid 

surface with 

axillary margin. 


Angle formed by 


Angle formed by 

spine of scapula 

with axillary 

margin. 


Angle formed by 

spine of scapula 

with vertebral 

margin. 




with spine of 
sc^apiila. 


No. 3 A 

„ 3B 

„ 3 C 

Average of the' 
three J 


35 
35 

35 


no 

113 
115 

112 


o 
05 

70 

70 

68 


o 

42 
41 

40 

41 


105° 
103 

108 

105 


Variety Morio . . 


35 


120 


70 


■il j 106 



Average axillary margin : 100 : : average vertebral : 84-3. 
Average axillary margin : 100 : ; average superior margin : 53-3. 



215 



Clavicle. 





Length 
measured in a 
straight line. 


Length 

measured 

along curves. 


Breadth 

about 

middle. 


Greatest 

diameter of 

acromial end. 


Greatest 
diameter of 
sternal end. 


No. 3 A 

„ 3 B 

„ 3C 

Average of the! 
three J 


7-50 
G-55 
7-20 

7-08 


7-80 
6-75 
7-40 

7-31 


-61 
•60 
-60 

-60 


1-10 

1-02 
1-05 

1-05 


1-27 
1-30 
1-42 

133 


Variety Morio . 


5-28 


5-40 -44 


■77 


■78 



Average length of scapula from summit of glenoid surface to posterior vertebral angle : 100 : .- the average 
length of clayicle measured in a straight line : 96- 7. 



Humerus. 





Length measured 

in a straight line 

from summit of 

head to lower 

end of inner mar- 
gin of trochlea. 


Transverse 

diameter of 

middle of 

shaft. 


Extreme 
breadth of 
tubero- 
sities. 


Extreme 

breadth of 

head. 


Extreme 

breadth 

between 

condyles. 


No. 3 A 

„ 3 B 

„ 3 C 

Average of the 1 
three J 


14^00 
14^10 
14-40 

14-16 


-93 
-92 
-90 

-91 


159 

1-68 
1-07 

1-64 


1-72 
1-72 

1-82 

1-75 


2-50 
2-59 
2-69 

2-59 


Variety Morio . . 


11^35 


-67 


1-26 


1-30 


2-05 



Average length of scapula : 100 : : average length of humerus : 193-5. 
Average length of clavicle : 100 ; : average length of humenis : 200-1. 



216 



MK. ST. G. MIVAET ON THE SKELETON OF THE PEIMATES. 



Badius. 





Estreme length 

to end of Btyloid, 

^ 5, in a straight 

line. 


Diameter of 
head. 


Diameter of 
distal end. 


Diameter of 

middle of 

shaft. 


3 A 


14-1.5 
13-90 
14-70 

14-25 


91 
93 
92 

92 


1-44 
1-55 
1-60 

1-53 


-64 
-70 
-63 

-65 


3B 


3 C 


Average of the 1 
three j 


Variety Morio . . 


11-75 


72 


1-09 


■47 



Average length of scapula : 100 : : that of radius : 194-6. 
Average length of humerus : 100 : : that of radius : 100-5. 

Ulna. 





Extreme length to 
end of styloid pro- 
cess, measured in a 
straight line. 


Greatest 
breadth of 
olecranon. 


Length of sty- 
loid process. 


3 A 


14-70 
14-20 
15-20 

14-70 


-91 
1-00 
1-10 

1-00 


-34 
•29 

•28 

-30 


3B 


3 C 


Average of the 1 
three J 


Variety Morio . . 


11-98 


-66 


-24 



Average length of humerus ; 100 : : that of ulna, measured in a straight line : 103-9. 



Os innominatitm. 





Extreme 
length. 


From superior 
anterior to 
superior pos- 
terior spinous 
process along 
crest of ilium. 


From superior 

anterior to 
superior pos- 
terior spinous 
process in a 
straight line. 


Distance be- 
tween anterior 
spinous pro- 
cesses. 


Distance be- 
tween pos- 
terior spinous 
processes. 


Distance from 
posterior in- 
ferior spinous 
process to 
spine of 
ischium. 


Distance from 
spine of is- 
chium to is- 

chiatic tubero- 
sity. 


3 A 


9-90 

9-69 

10-25 

9-94 


6-25 
6-45 
6-42 

6-37 


5-10 
4-82 
4-60 

4-84 


3-50 
3-40 
3-25 

3-38 


2-65 
2-75 
2-40 

2-60 


2-62 

2-58 
2-50 

2-56 


1-90 
1-83 
2-10 

1-94 


3 B 


3C 


Average of the 1 
three ( 


Variety 3Iorio . . 


7-84 


4-55 


3-51 


2-75 


2-.50 


1-90 


1-75 



MR. ST. G. MIVAKT ON THE SKELETON OF THE PRIMATES. 



217 





Distance from 
anterior infe- 
rior spinous 
process to 
symplijsis. 


Length of 
eympbysis 
" pubis. 


Greatest ver- 
tical diameter 
of acetabulum. 


G-reatest hori- 
zontal dia- 
meter of aceta- 
bulum. 


Ilio-pubic 
angle. 


Ilio-iachial 

angle. 


Angle of ante-! 
rior margin of 

ilium with 

upper margin 

of pubis. 


3 A 


4-10 
3-90 
4^50 

4-16 


1-97 
145 
1-80 

1-74 


1-70 
1-68 
1-75 

1-71 


1-GO 
1-52 
1^68 

1^60 


130° 
132° 
140° 

134° 


175° 
163° 
167° 

168° 


145° 

137° 
150° 

144° 


3B 


3C 


Average of the 1 
three J 


Variety Morio . . 


3-25 


M7 


1-38 


1^30 


128° 


172° 


150° 



Femur. 





Extreme 
length. 


Greatest 

diameter of 

head. 


Transverse 

diameter of 

middle of 

shaft. 


Antero-poste- 

rior diameter 

of middle of 

shaft. 


Breadth be- 
tween the 

supracondy- 
loid emi- 
nences. 


Breadth be- 
tween the outer 
and inner mar- 
gins of the 
condyles. 


3 A 


10-40 
10-70 
11-20 

10-76 


1-40 
1-40 
1^45 

1-41 


1-05 
•95 
•95 

•98 


•76 
•72 

•78 

•75 


2-33 
2-27 
2-33 

2-31 


2-23 
2-11 
2-29 

2-21 


3B 


3C 


Average of the 1 
three j 


Variety Morio . . 


8-90 


1-12 


■71 


•57 


1-79 


1-70 



Average length of femur : 100 : : that of os innominatum : 92-3. 
Average length of humerus : 100 : : that of femur : 76-0. 



Tibia. 



Fibula. 





Eitreme 
length. 


Breadth of 
proximal 
surface. 


Antero-poste- 

rior diameter 

of middle of 

shaft. 


Greatest 
transverse 
diameter of 
distal end. 


Extreme 
length. 


3 A 


9-20 
9-05 
9-80 

9-35 


2-30 
2-22 
2-39 

2-30 


•83 
•78 
•78 

•79 


1-60 
1^70 
1^72 

hG7 


8-75 
9*12 


SB 


3C 


Average of the l 
three J 


Variety Morio . . 


7-24 


1-77 


•59 


1-25 


7-23 



Average length of femur ; 100 : : 
Average length of radius : 100 : 



that of tibia : 86-8. 
that of tibia : 65-6. 



VOL. VL PART IV. 



2 H 



218 



MK. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 



Maims. 





Length of 1st 
metacarpal. 


Lengtli of 
second. 


Length of 
third. 


Length of 
fourth. 


Length of 
filth. 


Length in Length follow- 
straight line of ing curve of 
longest proxi- ' longest proxi- 
mal phalanx. | inal phalanx. 


3B 

3C 


1-90 
2-00 


415 
4-22 


3-96 

4-18 


3-84 
3-97 


3-54 
3-60 


2-85 
3-OU 


315 
3-55 


Average . . 


1-95 


4-18 


4-07 


3-90 


3-57 


2-92 


3-35 



Pes. 





Length of 1st 
metatarsal. 


Length of 
second. 


Length of 
third. 


Length of 
fourth. 


1 Length in 
Length of istraight line oi 
liltli. 1 longest proxi- 
] nial plialanx. 


Length iuUow- 
in^ curve of 
longest proxi- 
mal phalanx. 


3B 

3C 


1-85 
1-92 


3-90 3-80 
3-83 3-78 


3-63 
3-63 


3-45 275 
3-32 2-75 


314 
315 


Average . . 


1-88 


3-86 i 3-79 


3-63 


3-38 


2-75 


3-14 



DESCRIPTION OF THE PLATES. 

PLATE XXXV. Scapula of Shma. 

I'ig. 1. Outer surface. 

Fig. 2. Internal surface. 

Fig. 3. Outer surface of the variety Mono. 

Fig, 4. Glenoid surfiice of the specimen No. 3 A in the British Museum. 

Fig. 5. Glenoid surface of the specimen No. 3 C in the .same collection. 

In all the above figures the letters indicate respectively : — a, the acromian process ; 
c, the coracoid process ; g, the glenoid siu-face ; s, the triangular surface at the origin 
of the spine; t, the tubercle for the trapezoid ligament; vl, the superior vertebral 
angle ; v 2, the inferior vertebral angle. 



PLATE XXXVI. Humerus. 
Fig. 1. Anterior surface of the right humerus of the specimen No. 3 C in the British 
Museum. 
Posterior surface of the same. 

Outer surface of the humerus of the specimen No. 3 B. 
Inner surface of the humerus of No. 3 A. 
Superior surface of the humerus of No. 3 B. 
The same of that of No. 3 C. 
Fig. 7. Distal surface of the humerus of No. 8 B. 



Fig. 2 
Fig. 3 
Fig. 4 
Fig. 5 
Fig. G 



ME. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 219 

In these last six figures, a represents the articular head ; h, the bicipital groove ; 
c, rough surface, probably for the coraco-hrachialis ; rZ, the lesser, or ulnar, tuberosity ; 
e, the greater, or radial, tuberosity ; h\ the external condyle ; m, the perforation in the 
coronoid fossa ; o, the internal condyle ; p, the capitellum ; q, the trochlea ; s, musculo- 
spiral groove. 

PLATE XXXVII. Clavicle and Badivs. 
Fig. 1. Anterior surface of the right clavicle of the specimen No. 3 C in the British 

Museum. 
Fig. 2. Superior surface of the same. 
Fig. 3. Inferior surface of the same. 
Fig. 4. Anterior surface of left clavicle of the variety Morio. 

In these figures, a represents the acromial end of the bone; c, tlie tubercle for tlic 
conoid ligament ; d, the prominence for the deltoid ; p, ridge probably for pedoralis 
major ; r, rough surface for costo-claviciilar ligament ; s, sternal end of the bone ; 
t, ridge for trapezoid portion of coraco-clavicular ligament. 
Fig. 5. Anterior surface of the right radius of specimen No. 3 B. 
Fig. 6. Posterior surface of the same. 
Fig. 7. Posterior sm-face of distal end of the same, showing the grooves for the extensor 

tendons completely. 
Fig. 8. Distal surface of the same. 

a, process for the supinator longus ; h, tuberosity ; f, the styloid process ; y, surface 
for ulna ; h, the surface for the scaphoides ; k, groove for extensor ossis mefacarpi 
poinds ; /, that for ex. carpi radialis longior ; m, that for the ex. carpi radialis hrevior ; 
0, that for the extensor secundi internodii ])ollids ; ])^ that for the extensor communis 
digitorum ; u, the surface for the semilunare. 

PLATE XXXVIII. Ulna. 

Fig. 1. Anterior aspect of right ulna of the specimen No. 3C. 

Fig. 2. Ulnar (or inner) aspect of the same. 

Fig. 3. Posterior aspect of the same. 

Fig. 4. Radial (or outer) aspect of the same. 

Fis. 5. Posterior surface of the olecranon of the same. 

Fig. 6. Side view of the distal end of No. 3 A. 

Fig. 7. Distal surface of ulna of No. 3 A. 

a, radial or outer margin; b, nutrient foramen; c, fossa for hrachialis aiiticus; d, 
lesser sigmoid cavity ; e, ridge running downwards and backwards from that cavity ; 
/', surface for anconceus ; g, greater sigmoid cavity ; h, the distal articular surface ; 
s, styloid process ; x, ridge for attachment oi pronator quadratus. 

2 n 2 



220 ME. ST. G. MIVAET OK THE SKELETON OF THE PRIMATES. 

PLATE XXXIX. Os innominatum. 

Fig. 1. Outer aspect of right os innominatum of No. 3 C. 

Fig. 2. Inner aspect of the same. 

Fig. 3. Posterior aspect of the same. 

Fig. 4. Anterior aspect of the same. 

Fig. 5. Superior aspect of the same. 

Fig. 6. Inferior aspect of the same. 

a, anterior superior spinous process ; h, anterior inferior spinous process ; c, posterior 
superior spinous process ; d, posterior inferior spinous process ; e, ilio-pectineal line ; 
/, cotyloid notch of acetabulum ; g, greater sciatic notch ; h, spine of ischium ; ?, lesser 
sciatic notch ; I, tuberosity of ischium ; m, auricular sui-face ; o, obturator foramen ; 
p, spine of pubis ; r, subpubic groove ; s, symphysis pubis. 

PLATE XL. Femur and Patella. 

Fig. 1. Anterior aspect of right femur of No. 3 B. 

Fig. 2. Posterior aspect of right femur of No. 3 A. 

Fig. 3. Inner (or tibial) aspect of the same. 

Fig. 4. Outer (or peroneal) aspect of the same. 

Fig. 5. Distal end of right femur of No. 3 B. 

Fig. 6. Proximal end of the same. 

Fig. 7. Articular head of femur of No. 3 I, in the British Museum, showing the 

presence of a pit as for the Ugamentum teres. 
Fig. 8. Anterior surface of patella of No. 3 C. 
Fig. 9. Posterior surface of the same. 

n, articular head; h, greater (or peroneal) trochanter; c, lesser (or tibial) trochanter; 
(I, posterior intertrochanteric line ; e, anterior intertrochanteric line ; f, its continuation 
as the " spiral line ; " r/, depression of the gluteus maxinms ; h, line leading down 
towards the entocondyloid prominence ; i, pit for Ugamentum teres ; k, ectocondyloid 
prominence ; I, entocondyloid prominence ; m, external condyle ; «, internal condyle ; 
0, rotular surface; p, popliteal space; s, pit iox jaopliteus. 

PLATE XLI Tibia and Fibula. 

Fig. 1. Anterior aspect of right tibia and fibula of No. 3 C. 

Fig. 2. Posterior aspect of right tibia of No. 32, Z, S. 

Fig. 3. Posterior aspect of right fibula of No. 3 I. 

Fig. 4. Internal (or tibial) aspect of right tibia of No. 3 A. 

Fig. 5. External (or peroneal) aspect of the same. 

Fig. 6. External aspect of right fibula No. 3 C. 

Fig. 7. Internal (or tibial) aspect of the same. 



ME. ST. G. MIVAKT ON THE SKELETON OF THE PRIMATES. 221 

Fig. 8. Proximal surface of right tibia No. 3 C. 

Fig. 9. Distal surface of the same. 

Fig. 10. Proximal end of right fibula of No. 3 C. 

Fig. 11. Distal end of the same. 

Fig. 12. Lateral view of proximal phalanx of index of pes. 

a, crest of tibia ; b, tubercle ; c, external tuberosity ; rf, internal tuberosity ; e, arti- 
cular surface for external condyle of femur ; f\ articular surface for internal condyle 
of femur; ^, spine; /*, groove behind spine; ?, pit for semimembranosus; k, upper 
surface of tibia for fibula ; /, ridge leading downwards from the front of that surface ; 
»t, ridge leading downwards from behind that surface ; n, medullary foramen ; o, internal 
malleolus ; p, groove for tendons of tibialis posfi cits and Jlexor loncjus diyitorum ; q, arti- 
cular surface for superior face of astragalus ; r, articular surface of malleolus ; s, lower 
articular surface of tibia for fibula ; t, process of head of fibula for biceps ; ?/, anterior 
margin of fibula ; y, posterior external margin of fibula ; w, posterior internal margin 
of fibula ; x, peroneal malleolus ; y, medullary foramen ; z, articular surface of fibula 
for tarsus. 

PLATE XLII. Manus. 

Fig. 1. Palmar surface of the manus No. 5076 in the Museum of the Royal College of 

Surgeons, the bones of whicli are united by their ligaments only. 
Fig. 2. Left scaphoides of skeleton No. 8 B in the British Museum: its proximal 

surface. 
Fig. 3. Distal surface of the same. 
Fig. 4. Dorsal (or extensor) surface of the same. 

a, surface for articulation with the radius ; b, radial tuberosity ; c, surface for arti- 
culation with the semilunare; d, surface for the intermedium; e, that for the radial 
end of the intermedium ; f, that for the ti-apezium. 
Fig. 5. Left intermedium of No. 3 B : first surface. 
Fig. 6. Second surface of the same. 

a, surface for articulation with the trapezoides ; b, that for the magnum ; c, ulnar 
end of the bone; d, its radial end; e, the process which projects over the junction of 
the intermedium with the scaphoides. 
Fig. 7. Left semilunare of No. 3B: its radial side. 
Fig. 8. Ulnar side of the same. 
Fig. 9. Palmar side of the same. 

a, surface for articulation with the radius ; J, that for the scaphoides ; c, that lor 
the cuneiforme ; fZ, that for the magnum. 
Fig. 10. Left cuneiforme of No. 3B: its ulnar surface. 
Fiff. 11. Radial surface of the same. 

a. Surface for articulation with the unciforrae ; b, that for the pisiforrae ; c, groove 



222 ME. ST. G. MIVAET OX THE SKELETON OF THE PRIMATES. 

separating these two surfaces; d, surface for the semilunare; e, that for the fibro- 

cartilage of the wrist-jomt. 

Fig. 12. Left pisiforme of No. 3B: its dorsal (or extensor) surface. 

Fig. 13. Palmar aspect of the same. 

Fig. 14. Its articular surface for the cuneiforme. 

Fig. 15. Left trapezium of No. 3 B: its outer (or extensor) surface. 

Fig. IG. The same aspect of the right trapezium of the skeleton No. 3 A in the British 

Museum, 
Fig. 17. Left trapezium of No. 3B: its palmar surface. 
Fig. 1 8. The same aspect of the right trapezium of No. 3 A. 
Fig. 19. Left trapezium of No. 3 B: its proximal end. 
Fig. 20. Distal end of left trapezium of No. 3 B. 

r/, saddle-shaped surface for articulation with first metacarpal ; 5, surface for second 
metacarpal ; e, surface for trapezoides ; d, that for scaphoides. 
Fig. 21. Left trapezoides; its dorsal (or extensor) surface. 
Fig. 22. Distal surface of the same. 
Fig. 23. Proximal surface of the same. 

(7, the larger (or ulnar) surface for articulation with the second metacarpal ; h, smaller 
(or radial) surface for articulation with the same metacarpal ; c, facet for the magnum ; 
r/, surface for the intermedium. 

Fig. 24. Left magnum of No. 3 B : its dorsal surface. 
Fig. 25. Palmar surface of the same. 
Fig. 26. Radial aspect of the same. 
Fig. 27. Ulnar aspect of the same. 
Fig. 28. Distal surface of the same. 

a. notch on radial side of distal surface ; h, notch on ulnar side of the same surface ; 
c. palmar articular surface for second metacarpal; d, dorsal articular surface for the 
same metacarpal ; e, surface for the trapezoides ; /'/', surfiice for the unciforme. 
Fig. 29. Left unciforme of No. 3 B : its dorsal surface. 

Palmar surface of the same. 

Radial aspect of the same. 

Ulnar aspect of the same. 

Distal surface of the same. 

palmar process ; b V, articular surface for the magnum ; c, that for the 
cuneiforme. 
Fig. 34. Proximal surfaces of the four ulnar metacarpals of the left manus of the 

skeleton No. 3 B. 
Fig. 35. Dorsum of first metacarpal of the same manus. 
Fig. 30. Radial side of the same (first) metacarpal. 
Fig. 37. Dorsal aspect of second metacarpal of the same manus. 



Fig. 


30. 


Fig. 


31. 


Fig. 


32. 


Fig. 


33. 


«, 


its 



]VrR. ST. a. MIVART ox THE SKELETON OF THE PRIMATES. . 223 

Fig. 38. Palmar aspect of the same metacarpal. 
Fig. 39. Eadial side of proximal end of same. 
Fig. 40. Ulnar side of proximal end of the same. 

a, tubercle for \k\e flexor carpi radialis; b, groove between the two surfaces for the 
third metacarpal. 

Fig. 41. Dorsum of third metacarpal of the same manus. 
Fig. 42. Palmar aspect of the same metacarpal. 
Fig. 43. Kadial aspect of proximal end of the same. 
Fig. 44. Ulnar aspect of proximal end of the same. 

c, radio-proximal angle of dorsum. 
Fig. 45. Dorsum of fourth metacarpal of same manus. 
Fig. 46. Palmar aspect of the same metacarpal. 
Fig. 47. Radial aspect of proximal end of the same. 
Fig. 48. Ulnar aspect of proximal end of the same. 
Fig. 49. Dorsum of fifth metacarpal of the same manus. 
Fig. 50. Palmar aspect of the same metacarpal. 
Fig. 51. Eadial aspect of proximal end of the same. 



PLATE XLIII. Pes. 

Fig. 1. Plantar aspect of the jjes No. 5079 in the Museum of the Royal College of 
Surgeons, the bones of which are united by the ligaments only. 

Fig. 2. Right OS calcis of skeleton No. 3 B in the British Museum : its peroneal 
aspect. 

Fig. 3. Tibial aspect of the same. 

Fig. 4. Anterior (distal) aspect of the same. 

Fig. 5. Posterior-aspect of the same. 

Fig. 6. Dorsum of the same. 

Fig. 7. Plantar surface of the same. 

a, anterior articular surface for astragalus ; b, posterior articular surface for astra- 
galus ; c, funnel-shaped cavity of articular surface for cuboides ; d, tubercle for external 

lateral ligament ; e, antero-posterior groove above the last. 

Fig. 8. Dorsum of right astragalus of No. 3 B. 

Fig. 9. Plantar surface of the same. 

Fig. 10. Tibial aspect of the same. 

Fig. 11. Peroneal aspect of the same. 

Fig. 12. Anterior (distal) aspect of the same. 

Fig. 13. Posterior aspect of the same. 

a, neck ; b, articular surface for naviculare ; c, surface for tibial malleolus ; tl, gro(jve 

behind the last ; e, rounded surface posterior to the groove ; /', surface for peroneal 



224 MR. ST. G. MIVAET ON THE SKELETON OF THE PRIMATES. 

malleolus ; g, groove for flexor tendon ; A, anterior articular surface for os calcis ; i, pos- 
terior articular surface for os calcis ; k, groove for astragalo-calcaneal ligament. 
Fig. 14. Right naviculare of No. SB: its anterior (distal) aspect. 
Fig. 15. Tibial aspect of the same. 
Fig. 16. Peroneal aspect of the same. 

«, surface for entocuneiforme and mesocuneiforme ; b, surface for ectocuneiforme ; 
c, articular facet for cuboides ; d, rough surface of peroneal side of bone ; e, tuberosity. 
Fig. 17. Right cuboides of No. 3B: its dorsum. 
Fig. 18. Its plantar surface. 

Fig. 19. Anterior (distal) surface of the same bone. 
Fig. 20. Tibial aspect of the same. 
Fig. 21. Proximal aspect of the same. 

a, surface for fourth metatarsal ; h, surface for fifth metatarsal ; c, pivot-like process 
of tibio-plantar angle ; rf, ridge of plantar surface ; e, articular surface for ectocuneiforme ; 
/', articular surface for naviculare ; g, groove for tendon oi percmceus longus. 
Fig, 22. Left entocuneiforme of No. 3 B : its anterior (distal) aspect. 
Fig. 23. Peroneal aspect of the same bone. 
Fig. 24. Tibial aspect of the same. 
Fig. 25. The same bone seen from above. 
Fig. 26. Tibial side of right entocuneiforme of the skeleton No. 3 C. 

f(, articular surface for first metatarsal ; b, articular surface for the second metatarsal ; 
c, surface between the articular surfaces for the first two metatarsals ; d, surface for 
mesocuneiforme ; e, surface for nanculare. 
Fig. 27. Right mesocuneiforme of No. 3 B : its dorsum. 
Fig. 28. Tibial aspect of the same bone. 
P'ig. 29. Peroneal aspect of the same. 
Fig. 30. Anterior (distal) surface of the same. 
Fig. 31. Posterior sui'face of the same. 

n, articular surface for metatarsal ; I, articular surface for naviculare ; c, articular 
su)face for entocuneiforme ; d, articular surface for ectocuneiforme. 
Fig. 32. Right ectocuneiforme of No. 3 B : its dorsum. 
Fig. 33. Tibial aspect of the same. 
Fig. 34. Peroneal aspect of the same. 
Fig. 35. Anterior (distal) surface of the same. 
Fig. 36. Posterior aspect of the same. 

a, articular surface for metatarsal; b, articular surface for naviculare; c, facet for 
mesocuneiforme ; d, articular surface for cuboides. 

Fig. 37. Proximal surfaces of the four peroneal metatarsals of the left pes of the 
skeleton No. 3 B. 

a, facet for entocuneiforme. 



ME. ST. G. MIVAJiT ON THE SKELETOX OF THE PKU[ATES. 

Fig. 38. Dorsum of first metatarsal of No. 3 B. 
Fig. 39. Lateral view of the same metatarsal. 

«, process for attachment of tendon oi peronmus lonf/iis. 
Fig. 40. Dorsal aspect of second left metatarsal of the same skeleton. 
Fig. 41. Plantar aspect of the same metatarsal. 
Fig. 42. Tibial side of proximal end of the same. 
Fig. 43. Peroneal side of pi'oximal end of the same. 

a, articular facet for entocuneiforme ; b, b', articular facets for third metafarsal. 
Fig. 44. Dorsal aspect of third left metatarsal of the same skeleton. 
Fig. 45. Plantar aspect of the same. 
Fig. 46. Tibial aspect of proximal end of the same. 
Fig. 47. Peroneal aspect of proximal end of the same. 
Fig. 48. Dorsal aspect of fourth left metatarsal of the same skeleton. 
Fig. 49. Plantar aspect of the same. 
Fig. 50. Tibial aspect of proximal end of the same. 
Fig. 51. Peroneal aspect of proximal end of the same. 
Fig. 52. Dorsal aspect of fifth left metatarsal of the same skeleton. 
Fig. 53. Plantar aspect of the same. 
Fig. 54. Tibial as])ect of proximal end of the same. 

a, tuberosity. 



VOL. VI. PART IV. 2 I 



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X. Description of the Memains of three extinct Species of Elephant, collected by Capt. 
Spratt, C.B., R.N., in. the Ossiferous Cavern of Zehhug, in the Island of Malta. 
By George Busk, F.B.S. ; partly from the Xotes of the late H. Falconer, M.I)., 
F.R.S. 

Bead June 27th, 1865. 

[Plates XLIV. to LIII.] 

§ I. Introductory. 

±N the following observations I have confined myself principally to the anatomical 
characters of certain proboscidian remains which were collected some years since (1859) 
by Captain Spratt, C.B., R.N., at that time in command of H.M. Surveying Ship 
' Medina,' in an ossiferous cavern some distance inland in the Island of Malta, under 
circumstances which will be fully detailed by that gentleman elsewhere *. 

As any extended geological account of the locality would be here out of place, it will 
suiRce for the present occasion to state, from information supplied by Captain Spratt, 
that the cavern in question is situated on the north side of a rocky valley separating the 
town or casal of Zebbug from another town about a mile distant, called Siggieni. 
The cavern, when first opened, was filled to the roof with yellow and grey sandy clay, and 
it had no stalagmitic floor. Amidst this deposit, which had evidently been washed in 
by water, were numerous fragments of bones and teeth of at least two species of 
Elephant, manifestly widely distinct in size, some of the bones indicating an animal of 
very dwarf dimensions, as compared with all other known forms, recent or extinct. 
Besides these elephant-remains, those of other apparently extinct animals were also met 
with, all mingled with subangular but not waterworn fragments of the limestone rock. 
The cavern ran nearly horizontally from the face of the ravine or inland cliff, having a 
short branch terminating in a small chamber. It varied in height from 4^ to 5J. and 
in width from 1| to 2\ feet. 

Amongst the other bones associated with those of the Elephant were many of 
aquatic Birds, and especially of a large extinct species of Swan [Cygniis falconen. Park.), 
a few jaws and other bones of perhaps more than one species of Dormouse, one of which 

* Since this paper was read, Captain Spratt has communicated a paper on the " Geological relations of the 
Zehbug Cavern," which will be found in the 23rd volume of the Quarterly Journal of the Geological Society (1867). 
Two Reports, also, on other ossiferous Carcnis in Malta, in which numerous remains of Elephants and other 
animals have been collected by Dr. Leith Adams, have appeared in the Reports of the British Association for 186.5 
and 1866 ; and I should also state that Dr. Leith Adams, who has been for some years indefatigable in his re- 
searches in the caves and iissures of Malta, has made an immense collection, more especially of Elephantine 
remains, an account of which, when they have been more fully worked out, will add very considerably to our 
knowledge of the various species, and more especially, as it seems to me, of the largest one, and of E. melitensis. 
— June, 1867. 

VOL. VI. PART V. 2 K 



228 ME. BUSK OX THE REMAINS OP 

was of gigantic size {Myoxus rnelitensis, Falc), and, together with these, numerous 
remains of a land Tortoise, not as yet fully determined. 

It should be mentioned also that, in several other localities in the island, caverns of 
a similar character have been discovered, containing chiefly bones of one or more species 
of Hippopofamus unmixed with those of the Elephant, and regarded by Captain Spratt, 
for certain geological reasons, as having been introduced into tlie caverns and fissures at 
a somewhat earlier period than those of the proboscidian pachyderm. 

The occurrence of such abundant remains of large herbivorous mammals in so 
limited a spot as the present Island of Malta, taken in conjunction with other consi- 
derations of a geographical or, rather, hydrographical nature noticed by Captain Spratt, 
indicates beyond all doubt a former connexion of a very close nature with the African 
continent, and also points to the former existence in the site of the Maltese Islands 
of extensive currents of fresh water. It consequently becomes an extremely in- 
teresting problem to determine, with as much accuracy as the materials will allow, 
the zoological relations of these quaternary pachyderms to those at present existing, 
and also to those which have either been contemporary with them in other parts, or 
may have preceded them in order of time in the Mediterranean region. 

The latter question, however, is one of such great magnitude, and so much more 
purely of palseontological interest, that, even were I able to do it justice, which is wholly 
out of my power, I should not here attempt to enter upon it — leaving its solution 
to more competent hands, to whom tlic present paper may, perhaps, be of some 
assistance. 

With respect to the other extinct animals associated with the Maltese Elephants in 
the Zebbug cavern, I am happy to say that the determination of the birds, and more 
especially the description of Cyynus falcmeri, has been undertaken by Mr. W. K. 
Parker, F.R.S., whilst there is reason to hope, from a letter from Dr. Leitli Adams 
which I find amongst Dr. Falconer's notes, that the Gigantic Dormouse will find an 
able historian in him, who has met with the remains of that genus ui extreme abun- 
dance in other localities in the island, in the exploration of which he is at present 
engaged, with the promise of the most fruitful results *. 

Captain Spratt's valuable collections were originally committed by him to Dr. 
Falconer, whose irreparable loss we i-ecently had so deeply to deplore; and a short 
notice respecting them was given, by him and Captain .Spratt, to the Geological 
Section of the British Association, at the Cambridge Meeting in 1SG2. Iir this commu- 
nication (a mere notice only of which appears in the published volume of Reports), Dr. 
Falconer announced the discovery, among these remains, of those of a diminutive or 
•' pigmy " species of Elephant, for which lie proposed the name of Elephas rnelitensis. 

* The account of the Gigantic Fossil Dormouse, which was anticipated at the time this paper was read, has 
since been furnished by Dr. Leith Adams, and is given below, p. 307. That of C yijmts fulcone ri is already pub- 
lished, aiitea, p. 110. 



THEEE EXTINCT SPECIES OF ELEPHANT. 229 

Naturally much struck with sucli an extraordinary deviation from the otherwse univer- 
sally received conception of tlie proboscidians as animals of colossal size, Dr. Falconer 
entered upon the study of its remains with his usual zeal ; and he appears to have con- 
templated the presentation to the Royal Society of a paper on the subject, including 
observations on the Mediterranean Cave-fauna generally, — a design the non-execution of 
which by such a man cannot be too deeply regretted. But, so far as I have been able 
to trace in his notes, he has left behind him no record of observations upon any of 
the bones except the teeth, doubtless having reserved the remainder for subsequent 
study. With respect to the teeth, however, he has left some very interesting and 
important remarks, the substance of which, and, so far as is possible, the words, will be 
recorded in the following pages. 

Since the decease of Dr. Falconer, the collection of remains from the Zebbug Cave 
has been placed in my hands by Captain Spratt, with a request that I would undertake 
the completion of the task left unaccomplished by that distinguished palaeontologist. 
This I have endeavoured to perform, though hand pari passu, and fully sensible of the 
loss that science has sustained from the change of hands into which the description has 
fallen, and especially since the only assistance I can derive from the notes of my la- 
mented friend is, as I stated, on the subject of the teeth ; but, in addition to these 
notes, I would remark that Dr. Falconer had had careful drawings made by Mr. 
Dinkel of some of the principal bones, under his own inspection, some of which will 
form part of the illustrations appended to this paper. 

Under the circumstances, tlierefore, it is scarcely necessary to observe that I am alone 
responsible for nearly the whole of the descriptive part, so far as it relates to the bones 
of the skeleton, and that any errors or misconceptions contained in it must be laid to my 
charge alone. 

It is proper also to mention that Dr. Leith Adams has, for the last year or more, been 
engaged in the exploration of fossiliferous caverns ir various parts of Malta, and has been 
very successful in procuring abundant elephantine remains, both large and small, the 
examination of which will, doubtless, in his hands, supply mauy deficiencies in the 
account I now venture to lay before the Society. 

Captain Spratt's collection of proboscidian bones and teeth is very considerable ; but 
a great part of it is made up of broken and often much-rolled fragments ; still among 
the remainder are several well-marked and characteristic specimens of many of the more 
important bones, and a very fine collection of milk- and permanent teeth in excellent 
preservation. 

When I began to arrange the collection for examination, I found, to my extreme 
surprise, that it apparently comprised the i-emains of not less than three distinct species 
of Elephant — two of diminutive, and one of tolerably large dimensions. With the 
presence of the latter form, I was aware that both Dr. Falconer and Captain Spratt 
were acquainted ; but neither of them were, I believe, at all cognizant of the existence 

2 K 2 



230 MB. BUSK OX THE REMAINS OF 

of more than one dwarf species. Further attentive examination and comparison of tlie 
bones has only served to confirm this impression ; and I hope in the following pages to 
be able to show that the Zebbug proboscidian remains, strange as it may seem, embrace 
those of not less than three species, two of which must be regarded as pigmy or dwarf 
forms (though one probably exceeded the other in size), whilst the larger one equalled 
in stature the smaller forms of the existing African or Asiatic species. It will 
doubtless be regarded as a remarkable circumstance that the distinction between the 
two smaller forms should have escaped the penetrating and long-experienced eye of Dr. 
Falconer ; but I think this may be readily explained by the consideration that, so far as 
his notes show, he had not as yet entered upon the critical study of the bones of the 
skeleton, but had confined himself to that of the teeth alone, parts to which, as is well 
known, he attached such paramount importance in the study of the Proboscidia. And 
I have little doubt that, had he li\ed to resume his investigation of the Maltese fossils, 
wliich for the last two years or more had been completely interrupted by the attention 
he had devoted to the fossil remains from (jibraltar, he would, on turning to the bones 
of the skeleton, have become aware of the existence of more than one " pigmy" Elephant. 

But under the cu'cumstances, and having convinced myself of the existence of two 
such forms, I have felt some doubt as to the names that should be given to them. Both 
cannot of course be IJ. melitensis of Falconer ; and I propose therefore to limit that 
name to the larger of the two small forms, and to designate the other by the name of 
one to whom paleontology, especially as regards fossil proboscidia, is so deeply indebted, 
and to term it E. falcoiieri. 

As regards the large form associated with E. melitensis and E. fulconeri, there are not 
in the present collection, as it appears to me, sufficient materials for the drawing of an 
accurate comparison between it and several other extinct species ; and I shall therefore 
not venture at present to suggest any name for it, preferring to leave this in suspense 
until better-marked remains of its teeth and other parts may justify its being either 
referred to some already described species, or distinguished definitively from all with 
which we are at present acquainted. Reasons will perhaps be apparent, in what 
follows, for the suggestion that it may be identical with E. antiquns; but the evidence as 
yet in our possession is far too scanty to allow of this being affirmed with any degree of 
certainty *. 

The collection, I would remark, is made up partly of the bones of adult, and partly 
of those of young or even, perhaps, foetal animals; and these immature bones, like the 
mature ones, are plainly divisible into three sets, each of which it is fair to assume 
belongs to one or other of the adult forms indicated by the mature bones. No diffi- 
culty, of course, exists in referring the young bones of the large form to their proper 
place ; but with respect to the other two, owing to their much nearer correspondence in 
size, the question of allotment is not so easily settled, and I am quite willing to believe 

* Vide note, p. 227. 



THEEB EXTINCT SPECIES OP BLEPHAXT. 231 

that any determinations I have made in this matter are open to future rectification. 
I have, however, in the following paper thought that it would be most convenient to 
consider what I regard as the remains of each species separately, in its mature and im- 
mature states, and I shall therefore commence with the large species. 

§ II. Elephas, sp. 1. (Plates XLIV., XLV.). 

There are between twenty and thirty fragments of bones belonging to an Elephant of 
considerable size, which in fact may be judged to have attained neaily, if not quite, eight 
feet in height. But of these fragments there are only three or four which it will be 
necessary or useful to describe in any detail. The remainder consist of various-sized 
irregular fragments of long and flat bones, including the cranium and pelvis, which are 
too imperfect to allow them to be turned to any useful purpose. 

The more readily identifiable portions of the adult skeleton consist only of (1) the 
symphysial portion of the lower jaw, (2) a large portion of the head of the left (1) 
humerus, and (3) a nearly complete spine of probably the 17th or 18th dorsal 
vertebra. Beside these may be briefly noticed : — a large fragment of a femur, appa- 
rently of even greater proportional dimensions than the other bones ; a considerable 
fragment of the spinous process of another dorsal vertebra ; and a considerable frag- 
ment of a tusk near the base. 

1. Of these, by far the most important fragment is the portion of the lower jaw 
(PI. XLIV. fig. 1). It is evidently that of a mature, if not aged, animal ; and it consists of 
the entire symphysis, and a portion of the ramus on either side, about 4 inches in length, 
measured along the lower border. The upper border on each side is broken off on a level 
with the large mental foramen and canal, which is thus represented on either side by a shal- 
low groove, more than 0"-5 in width. The rostrum, or prolonged beak of the symphysis, if 
it existed, is broken ofl", the fracture extending chiefly to the left side. The fractured sur- 
face looks as if it were in part of ancient, and in part of recent date ; and there are several 
other marks on the under surface of the bone, which show that it has been recently sub- 
jected to rough usage with a sharpish or pointed instrument, probably a. pickaxe or geolo- 
gical hammer. Owing to the circumstance that the fracture at the apex of the symphysis 
is chiefly on the left side, the right border of the symphysial gutter remains almost 
entire, as does also a considerable portion of the left border above and behind the 
broken part ; an accurate measure, therefore, of the width of the gutter can be taken, and 
its exact form perceived, whilst at the same time the angle at which its borders descend 
can be determined. The width of the gutter at the highest point at which it can be 
measured, and nearly on a level with the mental foramen, is about 2" ; and its borders 
descend in front almost vertically downwards, as in Elephas pvimirjenhis and in old 
E. indicus. As before said, it is extremely doubtful, from the appearance of the borders 
of the gutter, whether there was any rostral prolongation whatever. The depth of the 
symphysis, measured in a -vertical direction from the bottom of the symphysial gutter, 



232 MR. BUSK ON THE REMAINS OF 

is 2' -15. In two lower jaws of a mature African Elephant, at the College of Surgeons, 
tlie width of the symphysial gutter at the same point as in the Maltese fossil is 2""6, 
and in a third (female) 2" "3. In the Indian Elephant the mean ■width of the gutter at 
tlie same point is about 1"'9, varying from 1"'8 to 2'''0. As regards the width of this 
part alone, theiefore, the Maltese specimen would seem to correspond more closely with 
the Asiatic than with the African species ; but in the former the vertical depth of the 
symphysis is about 3"-5, and in the African 2"'7 ; so that, as compared with either of these 
species, the symphysial gutter is wider in the Maltese specimen in proportion to the depth 
of the symphysis. But when compai'ed with a very perfect mandible oi E. primi genius 
in the College of Surgeons, the width of the gutter is precisely the same, viz. 2"-0, 
and the depth of the symphysis also pretty nearly equal. Consequently we may conclude, 
so far as such a cliaracter will allow us, that the Maltese large Elephant had, as regards 
the mandible, more of the characters of E. primigenins than of either of the existing 
species. The mental foramen, which, as has been said, must have been about 0"*5 
in diameter, is placed about 1""0 behind the edge of the sj-mphysial gutter. The 
distance between the two openings in a transverse line is about 2"*7, whilst in the 
Indian Elephant the corresponding distance is about 4"-0, and in the African 6"'5 ; 
whilst in the Mammoth already referred to it is 6'''8. It may here be remarked that 
the mental foramina are placed much nearer the border of the gutter in the Indian 
than in the African species, or, I believe, than in the Mammoth ; consequently in their 
comparatively distant position from the edge in the Maltese fossil, the latter approaches 
the African more than the Indian Elephant. 

From the very obtuse angle at which the rami meet at the symphysis (nearly 90°) it 
may be concluded that the jaw was broad and short. 

There are no vestiges of any alveolar cavities. 

2. The next well-marked fragment is a large portion of the articular head of the 
humerus. The remains of the antero-posterior arc of the articular surface indicate that it 
formed the segment of a circle having a radius of 2"-8, and, in the transverse direction, 
of 2"'7. The antero-posterior diameter of the head may therefore, in accordance with 
what obtains in the humerus of other Elephants, be regarded as nearly C". 

In the table of measurements given by Cuvier * the diameter of the head of the 
humerus in an adult female Indian Elephant about 9 feet high (2"'-76) is stated to be 
7"-28 (O^'ISS), and the length of the humerus 32"-87 (0"-835) ; whilst in a young but 
well-grown African Elephant in the British Museum the corresponding measures are 
8" and 36" ; in a still younger (but not very young) specimen of the Sumatran Elephant 
they are 5"'2 and 28"'2 ; and in a very fine and perfect humerus of E. primigenius, in 
the British Museum, they are 7"-3 and 32". According to these measurements the 
proportionate diameter of the head to the length of the entire humerus would stand in 
the respective cases as follows : — 

* Osscm. fossiles, 4"' ed. 1834, 8vo, torn. i. p. 504. 



THEEE EXTINCT SPECIES OF ELEPHANT. 233 

E. africanus (young) 1 to 4-00 

E. indicus {{em?\e) 1 „ 4'53 

-E. /«f/?V»s (Sumatra) 1 „ o-42 

E. imniiijenius 1 „ 4-50 

From this we may probably consider that the usual length of the entire humerus in a 
fully grown mature Elephant is rather more than four times (4-3) the antero-posterior 
diameter of the head. In the young animal, in which the head has probably not attained 
its full size, the length would seem to be greater in proportion. From these data as 
compared with those afforded in Cuvier's table, it may be concluded that in all pro- 
bability the Maltese Elephant to which the above-mentioned fragment belonged was 
about 8 feet high. 

3. A third well-characterized bone, belonging apparently to an animal of the same pro- 
portions, is the nearly entire spinous process of the 17th or 18th dorsal vertebra (fig. 2). 
The fragment is 5''-61ong; and the neural spine itself measures along the anterior border 
5"-2, and along the posterior 4"-7. Its smallest antero-posterior diameter is l"-3, and at 
the base 2"-3. Its least tr. d. is 0"'5 ; and the ap. d. of the expanded outer end l"-65. It 
is grooved behind for about half its length, whilst the anterior edge is acute nearly 
throughout. The right articular facet is present and entire ; it measures 1"'2 * in length, 
by 0"-0 in breadth, and it is of an oblong form. The corresponding spinous process in 
the African Elephant (No. 708 h, B.M.) with which comparison has before been made, 
measures 5"'5 along the anterior, and 3"-8 along the posterior border, its least ap. d. being 
l"-2, and at the base 3"-7, the least tr. d. l"-0, and ap. d. 8"-8, at distal end l"-5. 

Of the teeth of this species, I have been able to detect only innumerable fi-agments 
of the tusks, which from their size must be referred to an animal of considerable 
bulk. The majority of these are too imperfect for description ; but amongst them is 
a fragment, nearly five inches long, of the solid part of a tusk, 2"-8 in diameter. The 
fragment has been split off nearly down the middle ; and the interior is thus shown to be 
quite solid ; the portion, therefore, of the tusk to which it belonged was some distance 
above the base, which must have been greater in circumference. It is also to be ob- 
served that the outer surface of the fragment is strongly sulcate, showing that the outer- 
most layer or lajers have been removed ; we may conclude, therefore, that the fragment 
does not represent the real diameter of the tuslv, which may consequently be regarded as 
ha\dng been of considerable size, and quite commensurate with an animal 8 feet high 
or more. 

Among the immature bones, are two well-marked fragments which, from their size 
appear to belong to the young of the same species as that whose remains have just 
been described. The portions in question are — (1) a left exoccipital, and (2) a con- 
siderable portion of the sliaft of the left femur. 

The exoccipital bone (figs. 3 and 4, PI. XLIV.) corresponds pretty nearly in size, and 

* This facet appears much too small in the figure, owing probably to its having been drawn foreshortened. 



234 



MR. BUSK ON THE KEMAIKS OF 



entirely in the appearance of the surface of the bone, with that of a young African 
Elephant in the British Museum, in which specimen all the bones of the cranium and 
face are perfectly separate ; and in the lower jaw the 3rd molar is in full wear, no 
vestiges remaining of the 2nd molar. The dimensions of the two bones are as under :— 

Table L— Measurements* of Exoccipital Bone, in Maltese and African Elephants. 



Height or 



length of 
exoccipital. j 



Breadth of 
exoccipital 



Young African Ele- ] j ^_- 
phant, No. 70S, B.M. J 
Young Maltese fossil. ! 4-7 



3-5 
3!) 



Width be- 
tween bor- 
ders of con- 
dyloid fossa 
at foramen 
magnum. 



Length 
of basiocci- 
pital syn- 
chondrosis. 



1-6 
1-6 



1-9 

1-8 



Width of basi- T'"'^'™.^^^ 

occipital syn-'" "§■<>" 

chondrosis. of mastoid 



•95 
1-15 



1-3 

1-9 



Length 
of condy- 
loid arti- 
cular sur- 
face. 



Greatest 

width of 

condyloid 

articular 

surface. 



2-2 
2-35 



1-35 
1-55 



It will thus be seen that in general dimensions and proportions the two bones 
are remarkably alike ; but they present certain differences, which would appear, as I 
think, clearly to indicate that they belong to different species. 

In the first place, notwithstanding the apparent similarity of age, it will be observed 
that the fossil is very much thicker in the part occupied by the paramastoid cells, and 
that the proportions of the condyloid articular surface are not the same. But the most 
strikino- distinction consists in the circumstance that, in the African Elephant, the 
cerebellar fossa is very concave or deep, and that the sulcus for the lateral sinus is also 
very deep, and separated from the opening of the paramastoid cells by a sort of vertical 
wall ; whilst in the Maltese fossil the cerebellar fossa is only slightly hollowed, and 
there is scarcely any trace of a sulcus for the lateral sinus. And in another very 
young (or perhaps fcetal) cranium of the African Elephant (No. 708^', B.M.), in an 
exoccipital having a greatest diameter of 3"-l, and least of 2"-9, the sulcus for the 
lateral sinus is quite as well marked as in the above, — whence it may be concluded 
that this character is not dependent upon age, and may probably be relied upon as 
indicating a distinction between the Maltese form and E. africamts. I have not 
had an opportunity of comparing the exoccipital of the Indian Elephant of the same 
aoe. In the young African Elephant the opening into the paramastoid cells is tri- 
angular, and a transverse septum may be obscr\ed within, dividing the main cavity 
into two primary loculaments, of which the posterior is shallow, and the anterior very 
deep, communicating at the bottom with two deeper cells. In the Maltese fossil the 
same primary division into two chambers is observable; but the slender trabecular 
septum between them is absent, and replaced simply by an angular ridge. It should be re- 
marked in addition, that in the African species the concavity or sulcus above the 
condyle is much deeper than in the Maltese, and that the curve of that part of the 
foramen magnum which is formed by the exoccipital is different in the two cases. 
* Throughout this paper the measures are given in inches and tenths. 



THREE EXTINCT SPECIES OF ELEPHANT. 235 

The portion of the shaft of a femur (fig. 5) like the exoccipital, and probably belonging 
to the same animal, presents all the external characters of a very young bone. It is broken 
at either end at some distance from the epiphysial termination. The entire fragment 
measures 8"-4 in length, and its least tr. d. is 2"-15, and least circumference 5"'9. In 
a young E. indicus in the British Museum, in which all the epiphyses are separate, and 
the ossification of the articular ends themselves very incomplete, the length of the 
shaft is 21", the least tr. d. 2"-4, and the least circumference 6"-7. In general form 
the two bones resemble each other very closely, except that in the Maltese fossil the 
posterior surface rises more into an angle than it does in the other, in which it is uni- 
formly rounded and even. The nutrient foramen in both is on the inner side, about 
the junction, as it may be estimated, of the upper and middle third, or a good way 
above the middle of the bone, whilst in the African species it is much lower down ; and 
this I am inclined to believe will be found a constant and not unimportant character. At 
any rate, so far as it goes, it further tends to sliow a distinction between the Maltese 
and existing African species. 

Besides the above bones belonging to the largest of the three Maltese forms, there 
are numerous fragments of others, most of them apparently of an old animal or animals, 
and including portions of the cranium, jjelvis, and of some of the larger long bones. All 
manifestly indicate a species of comparatively large size ; but as they afford no special 
characters, I have not thought it necessary to enter into further details respecting them. 

§ III. ElEPHAS MELITENSIS. 

Bones or fragments of bones belonging to the larger of the two dwarf species of 
Elephant to which I ha\'e assigned the name of E. mehtensis constitute a very consi- 
derable part of the collection. They are exceedingly numerous ; but amongst them 
are ^■ery many much broken and scarcely recognizable portions of the cranium, with 
respect to some of which it is impossible to determine whether they belong to this or to 
the next species. But, as in their present condition these broken fragments offer no 
distinctive characters, their determination is not a matter of any great importance. 
The remaining fragments amply suffice at any rate to indicate the comparative bulk 
and many of the distinctive characters of E. melitensis. 

Separating the mature from the immature bones, and excluding the teeth, tlicre are 
about 18 fragments which it will be necessary to describe: — 

These are: — (1 & 2) portions of the right ascending ramus of the mandible; (3) the 
right half of the atlas ; (4, 5, & 6) the 7th cervical, 7th dorsal, and 3rd lumbar vcr- 
tebrse ; (7) the neural spine of one of the anterior doi-sal vertebrae ; (8, 8") the proximal 
end, including the head of the second rib on the right side, and a portion of the body 
of one of the larger ribs; (9) a portion of the left scapula; (10) the nearly entire head 
and part of the shaft of the right humerus; (11) a small fragment of the articular 

VOL. VI. PART v. 2 L 



236 MB. BUSK ON THE BEMAINS OF 

heart of the humerus (side unccrtam) ; (12) the upper end, minus the olecranon of the 
right uhia; (13) a detached olecranon of the left ulna; (14) a fragment of the shaft of 
the left ulna; (15) a portion of the right os innominatum, including part of the aceta- 
bulum; (16) a portion of the articular head of the femur; (17) a great part of the 
shaft of the right femur ; (18) the lower end of the left tibia. 

1. Bones of Cranium and Face. 

The ascending ramus of the mandible in the Elephant appears to afford \cry dis- 
tinctive characters, at any rate between the African and Indian species ; and the ac- 
quisition, therefore, of a considerable portion of this part of the skeleton of E. melitensis 
is of great value. The two fragments both belong to the same side. They are of exactly 
the same dimensions, and, so far as they can be compared, of the same age, and 
that probably of a fully mature animal in which the third molar had advanced into 
the horizontal ramus. The larger of the two fragments (PL XLVII. tig. 13) is about 
4"-6 long, and about 2"'2 in ap. d. at tlie lower part, or at about three inches below the 
upper border of the dental foramen. The condyle or head appears to have been broken 
off through the smallest part of the neck ; and this is at a level of about an hich above the 
middle, or thereabouts, of the dental foramen. The bone is broken on the external border 
at this part, so that the entire width or tr. d. of the neck cannot be determined; but it may 
be estimated at about 1"3. The ap. d. of the ramus on a level with the lowest part of the 
dental foramen is 1""7. The posterior or, rather, interior angle is rather acute, and it 
descends evenly till the angularity merges in the inner surface of the bone about l"-5 
below the dental foramen ; immediately exterior to this angular border the surface 
presents a shallow sulcus, bounded on the outside by a second ridge, beyond which the 
posterior and outer surface is flattened or slightly concave. The anterior, and internal 
angle is somewhat acute, especially at the lower part. The anterior surface is too much 
broken to demand any special description ; but it may be remarked that the ])cculiar 
pock-like pitting which is exhibited on that surface within the base of the coronoid 
process in all Elephants, but which is nuich more pronounced in the African than in the 
Indian species, is very well shown in the jaw of E. melitensis. As these markings 
represent a muscular insertion, their distinctness in the present case is an additional 
indication, if the proof were required, that the bone was that of a mature animal. The 
inner surface is concave, and presents at the upper part the inferior dental foramen. As 
nearly as can be estimated, this opening is about 1" in ap. d. It is very oblique ; and 
the inferior margin is interrupted by a deep and wide fissure. The anterior border is 
very thick, the posterior very acute. 

Compared with the corresponding part of the mandible in the African and Indian 
Elephants, that of E. melitensis exhibits striking peculiarities beyond its size, though 
on the whole its resemblance may be said to lean much more towards the African than 



THREE EXTIXCT SPECIES OF ELEPHANT. 237 

towards the Indian type. In order to make this plain, it will be as well to point out 
one or two of the differences which exist in this part of the skeleton between the two 
existing species. In the first place, as above remarked, the inner and posterior angle of 
the neck in the African Elephant descends evenly till it is lost, below the level of the 
dental foramen, in the general surface of the ramus; at most it exhibits, in older 
animals, a slight roughness about the level of the dental foramen. But in tiie Indian 
Elephant, of all ages, this border or angle, at about the level of that foramen pro- 
jects into a distinct sort of crochet, which, as it were, protects the dental foramen from 
behind. 

This striking difference of form, seen on viewing the ascending ramus of the mandible, 
between the Indian and African species, is shown in the accompanying woodcuts *, 





The tr. d. of the neck at the smallest part, as compared with that of the condyle, 
is rather less in the African than in the Indian species. In two specimens in wliich 
the comparison was made, the tr. d. of the head in the Indian Elephant was 4", 
and that of the neck l"-9 ; whilst in the African the head was 3"-7 in tr. d., and the 
neck l"-5. In general form also, a considerable difference may be remarked. Slewed 
laterally, the ascending ramus in the Afi-ican Elephant is more rounded than in the 
Indian, in which it is comparatively straight in the vertical direction. The coronoid 
process rises much higher, in fact nearly to the level of the condyle, in the Indian 
Elephant ; and its anterior border is nearly vertical, which in the African overhangs 
very much, and is at the same time much thicker and rougher, whilst it descends very 
rapidly from the condyle to a level considerably below it. A striking difference is also 
seen in the configuration of the dental foramen. In the Indian Elephant this orifice 
looks, as it were, directly upwards, owing to the distinct elevation of the inner border, 
which forms, in fact, a sort of spine or projection opposite to the posterior crochet 
above described, the border of the opening between these two points being interrupted 
by a deep angular notch. In the African Elephant the dental foramen, which is pro- 

* A. E. indicus. B. E. dfrkanits. 

2 l2 



238 ME- BUSK ON THE REMAINS OF 

portionately also of much larger size, is so much bevelled off below as to look, as it were, 

directly inwards instead of upwards ; and the anterior and lower borders are thin and 

continuous. Several other distinctions might be pointed out ; but the above are sufficient 

for the purpose of comparison with the jaw of E. meUtensis. This will be found to 

exhibit the comparatively slender neck, and the obliquely bevelled dental foramen of the 

African, together with the absence of any posterior crochet, the presence of which is so 

strikingly characteristic of the Indian species. But it differs from the African m the 

presence of the sulcus on the posterior border, in the much thickened anterior margin 

of tlie dental foramen, and in the deep emargination of its lower border. With respect 

to comparative dimensions, it may be stated that the tr. d. of the ramus on a level 

with the middle of the foramen, in the African Elephant, is 4"- 3, and in E. melifensis 

l"-8 ; so that the general dimensions of the bone may be taken at about half those of 

the African species. 

2. Bones of the Trunk. 

1. The portion of the atlas (PI. XLVII. tig. 12) is unfortunately very imperfect ; and 
what remains is much injured, presenting some appearance of its having been gnawed. 
It consists of the right half, including the entire superior and the greater part of the 
inferior articular surfaces. The transverse process is broken off, leaving only a deep and 
\\ide sulcus to represent the foramen for the vertebral artery, but which is continued 
into a perfect posterior condyloid foramen. The ap. d. of the fragment, which is 
probably pretty nearly that of the entire bone, is about 3"*5. The ap. d. of the 
superior articular facet is 2"-5, and its tr. d. about 1" 8. The inferior facet is not 
sufficiently complete to admit of accurate measurement. The greatest height of 
the bone, measured just behind the roof of the transverse process, is 2"'4 ; the 
diameter of the posterior condyloid canal about 0"-4*. 

In a young Indian Elephant (No. 2G78, C. S.) the ap. d. of the condyloid facet is 
4", and its tr. d. 3", the former being nearly, and the latter exactly, twice the 
corresponding measures in E. meUtensis. In a rather younger specimen of the Indian 
Ele]3hant, termed E. sumatrensis, in the British Museum, the same measurements 
arc 3"-3 and 2"-4 respectively ; and in an African Elephant of mature age, 4"-7 and 3"-2. 
lu the latter instance a considerable difference is apparent in the form of the facet. In 
E. indicus (var. sumatrensis) the surface is kidney-shaped, having a deep sinus on the 
inner border, wliilst in E. africmuts that border is entire; but I am not aware that this 
difference exists in all cases between the Indian and African species. 

2. Q'he seventh cervical vertebra (PI. XLVI. fig. 9) is a beautifully perfect specimen of 
that important and highly characteristic clement of the vertebral column. Its principal 
dimensions, contrasted with those of the same bone in an African Elephant, and in the 
B. M. specimen of .£. sumatrensis, are as follows: — 

* other firagments of the atlas of this specios are shown in I'l. LI. tig. 35, and, as I believe, but am not sure, 
in fig. 33. 



THREE EXTIKCT SPECIES OF ELEPHANT. 
Table II. — Dimensions of 7th Cemcal Vertebra. 



2.39 





Height. 


! Transver.'se j Transverse 
Lengtb of diameter at diameter at 
Spine. 1 articular i transverse 
processes. ] processes. 


Transverse 
diameter 
at costal 
facets. 


Transverse 

diameter 

of bodj 

anteriorly. 


Vertical 
diameter 
of body. 


Transverse Vertical 

diameter of diameter of 

vertebral vertebral 

canal. canal. 


E. mditensis .... 


6-8 


2-5 


3-55 


57 


3-5 


2-9 


2-85 


2-0 


1-7 


E. (ifriia lilts. . . . 


8-20 


4-9 


715 


9-8 


61 


6-35 


5-4 


3-85 


2-1 


E. SHiiuttrciisis . . 


7o 


2-5 


5-6 




5-4 


4-65 


3-5 


2-8 


1-9 


E. bi'licus .... 


140 


7-0 


6-5 


10-25 


6-8 


5-5 


4-25 


3-75 


3-0 



The body, which is about l"'-3 in its greatest thicl<ness, is almost circular, slightly 
con^■ex in front, and well hollowed behind, especially towards the lower part. The 
neural canal is triangular and, viewed anteriorly, very nearly equilateral ; the laminae 
or neurapophyses flat and very thin, about 0"-8 wide, and not more than about 0"'15 
thick. The neural spine is also thin and slender, of the same width at the base as the 
laminae, and it is curved very slightly forwards. The transverse processes are broad 
and strong, and the outer surface of the process of bone connecting them with the 
articular processes is flat, or very slightly concave. The posterior sulcus for the exit of 
the spinal nerves is 0"'4 across, at the base of the posterior articular process. The 
costal articular facets are subtriaugular, the sides of the triangle being about 1 inch in 
length. The entire bone presents all the appearance of mature, if not of advanced age. 

Compared with the corresponding vertebra of the young African Elephant, many 
points of great dissimilarity, besides those shown in the Table, at once present themselves. 

In the African species the neural spine is flattened and sulcate behind, whilst in U. 
melitensis it is acute and without any sulcus. In both, the neural spine inclines a little 
forwards. Inspection of the measurements will show another remarkable distinction, in 
the comparatively much greater lowness of the neural arch in the African as com- 
pared with the Maltese species. Had the respective diameters of this arch borne the 
same proportions to each other in E. melitensis that the)' do in E. africamts, the height, 
instead of l'"7, would have been only l"-09. There is reason, however, to believe that 
the lowness of the arch in the specimen of African Elephant employed for the purpose 
of comparison is in part owing to its younger age. 

When we compai-c the 7th cervical vertebra of E. nielitensis with that of the Sumatran 
Elephant (younger than the African example), the diflerences are still more striking, 
especially in tlie form and proportional dimensions of the neural arch, neurapophyses, 
and spine. The arch, instead of being triangular, is more of an oval form ; and the 
vertical diameter is little move than half its transverse width. The neurapophyses or 
laminae, instead of being thin and flat, or riband-shaped, are very thick and square, and 
the neural spine in proportion very slender, its base not being nearly equal in ap. d. 
to the width of the neurapophyses; and it is curved slightly backwards instead of 



240 



IHE. BFSK ON THE REMAINS OF 



forwards as it is in E. africanus and E. melitensis. The transverse processes also are 
very convex in front, instead of concave as they are in the latter species ; the outer 
surface of the process of bone connecting the transverse and articular processes is 
rounded in the Suniatran Elephant instead of concave. Many of these differences are 
doubtless attributable to difference of age ; but on the whole we may presume that 
the 7th cervical vertebra of E. melitensis has more of the African than Asiatic charactei". 
In order to render more distinct some of the diversities presented by tliis vertebra in 
the different instances cited, I have subjoined tlie accompanying woodcuts, which are 
drawn to a scale of one-third the tiatural size — 





E. tif rial tuts. 



E. (var.) sumatfemis. 




E. itldiftts. 




liteiisls. 



ill of which represent the posterior view of tlie neural arch. 
■"). A dorsal vertebra (Pi. XI>VI. fig. 10), either the Cth or 7th, but in all probability 



THREE EXTJXCT SPECIES OK ELEPHANT. 241 

tlie fonnei, if tlie ribs are articulated as in the Indian Elephant, seeing that the remains 
of an articular surface are visible on the anterior aspect of tlic left transverse process. 
The bone is remarkably perfect, wanting only the extremity of tlie neural spine and 
a small portion of the right transverse process. 

Its principal dimensions aie as follows: — Transverse width from the end of one 
transverse process to the other, as they are, 4"-7, but in the perfect state probably 5" or 
more. Height of body l"-8o. Thickness l"-5. Width of anterior surface of body 
2'-15, of posterior 2"-2 ; anterior costal facets l"-lx0"-9; of posterior l"05x0"-65; 
ap. d. of neurapophyses 1" 2 ; extreme distance between the outer borders of the 
two posterior articular facets 1"-C. The neural arch, especially when viewed from 
behind, is cordiform, about 1" high, and 1"'4 wide; the body is also cordiform in figure, 
very concave behind, and but slightly convex in front. The neural spine is inclined 
backwards almost to a horizontal position. It is sharply carinate above, with a deep 
irregular hollow on one side only ; beneath it is deeply and widely sulcate beyond the 
expanded base, whilst between the posterior articular facets it presents an elevated 
ridge. 

4. The second or third lumbar vertebra (fig. 11), evidently belonging to the same 
animal as the other two. It is unfortunately not quite so perfect as either of the 
others, but still suflSciently so to afford a very good idea of its characters. Its dimen- 
sions are: — Height of body l"-7; thickness or ap. d. l"-5 ; width of anterior surface 
2'''1, and of posterior 2"-4 ; ap. d. of neurapophyses 1 '"1 ; extreme distance between 
the outer borders of the posterior articular facets 1"'5. The neural arch is depressed, 
its height about 0"-8, and width in front l"-65. The body is suboval, very concave 
behind, and nearly flat in front. Both ti'ansverse processes are broken short off, as is 
also the greater part of the apparently small neural spine. 

In the second lumbar ^■ertebra of E. (var ) sumatreiisis, which approaches the nearest 
in size to the Maltese specimen, the diameter of the body behind is 3"'45, and its 
thickness 2"-8 ; whilst the distance between the outer borders of the posterior articular 
surfaces is l"-95, and the transverse diameter of the canal 2"-2, and its height l"-68. 

5. The only other fragment belonging to the spine, and appearing from its dimensions 
to correspond very closely with the three vertebrse just described, is one of the anterior 
dorsal spines (PI. XLV. fig. 7). It is broken oiF through the roots of the neurapophyses, 
so that a small segment of the medullary canal is left. Measured from this point to 
the extremity, which, though chipped on one side, yet shows very distinctly that it was 
tipped with cartilage, the length of the spine along the anterior border is about 5". 
It is very slender and subtriangular in shape, with an acute angle in front, and rather 
obtuse ones on the sides. Behind, it exhibits a shallow groove towards the outer end ; 
but below the middle the surface rises into a ridge which descends nearly, but not 
quite, to the border of the vertebral canal. 

6. A portion of the second rib of the right side (PI. XLV. fig. 8) measuring about 4"-0 



242 MR. BUSK ON THE KBMAINS OF 

in length. The epiphyses are perfectly united, and no trace whatever of the junction 
remains, so that the bone must be regarded as mature. The head measures 1" in its 
lono-est and 0"-85 in its shortest diameter. The distance between the inner border of 
the head and the outer surface of the tubercle is 2". 

The corresponding rib in the young Asiatic Elephant in the B. ^L, denominated 
K sumatrensis, has the greatest diameter of the head l"-5, and least 1"-1. And in 
that species the tuberosity is differently formed, having a considerable elongation at 
the bottom, whilst in E. melitensis (a much older animal), the neck is fully as thick 
as in the so-termed E. sumatrensis. It is also to be remarked that in E. melitensis the 
notch or depression between the head and tuberosity is deeper than in the Asiatic form, 
in which also there is no depression below the tuberosity, such as is seen in E. melitensis. 
The comparison between the second rib of E. melitensis and that which I refer to 
E. falconeri will be drawn when I come to the description of the latter. 

7. Another, and in some respects a most important and interesting, fragment belong- 
ing to the bones of the trunk is a small and much mutilated portion of the pelvis 
(PI. XLVIII. fig. 26). It consists of part of the right ischium, including a small 
segment of the acetabulum, and a length of about three inches of the body of the bone. 
The surface on the outer, anterior, and inner aspects is almost entire or uninjured ; but 
posteriorly there is merely an apparently fresh fracture. The fragment, however, broken 
as it is, is amply sufficient to afford some very important characters. 

From the small remaining segment of tlie acetabulum it is evident that the curve of 
the articular surface must have had a radius of about l"-6. The cup was consequently 
fitted to receive a femoral head of about 3^ inches in diameter. The anterior surfiice 
of the bone is slightly hollowed, excepting quite at the upper edge, where the border 
of the acetabulum projects considerably and forms one margin of a wide and shallow 
sulcus or excavation, wliich was continued upwards into the cotyloid notch. The outer 
surface exhibits part of a large rough tubercular elevation. The inner is smooth and 
nearly even, marked only by a slight eminence. Tlie outer angle is round and smooth ; 
and the inner, or that which forms the ischial border of the obturator foramen, thougli 
not so obtuse, is also rounded and quite even. 

It is to the latter character more especially that I should wish to direct attention, in 
drawing a comparison between this part in E. melitensis and the corresponding part, 
fortunately also preserved, in E. falconeri., as it shows, perhaps as strikingly as any other 
single part, a considerable distinction between the two forms. 

I am not aware that the circumstance has been prenausly noticed ; but it is neverthe- 
less the case that a considerable difference in the form of the obturator foramen exists 
between the African and Indian species. 

In both the foramen has somewhat of an oval shape ; but in E. africanus the wider 
part of the oval is towards the upper or inner end, whilst it is towards the lower end in 
E. iiidicus. In E. africanus, again, the margin is tolerably even all round, whilst in 



THEEE EXTINCT SPECIES OF ELEPHANT. 243 

E. indicus the upper or anterior part of the oval appears to be constricted, as it were, 
by an eminence, usually on both sides, ischial and pubic, but at any rate on the former. 
It is to be observed, moreover, that in the African species the ischial border is thick 
and rounded, whilst in the Indian it is thin, or might almost be termed acute. The 
diiference in the outline of the foramen in the two species is shown in the accompanying 
figures, taken fi-om specimens of E. africanus and (so termed) E. sumatrensis in the 
British Museum. 

7 





E. indicus. E. africanus. 

I have had no opportunity as yet of examining this part in E. ])rimigenms, or in any 
other fossil species; but, from the figures of the Elephant's pelvis given in pis. xiii. 
& xvi. of the ' Ossemens fossiles,' it would seem that the ischial eminence above 
noticed and the peculiar constriction of the upper part of the obturator foramen are 
as well marked in fossil bones which in all probability belonged to the Mammoth 
as they are in the figui-e given, pi. xiii. fig. 4, of the pelvis of the Indian Elephant. 

Should the distinction here pointed out be found to hold universally, it would follow 
that, so far as the apparent form of the ischial border of the obturator foramen is con- 
cerned, E. melitensis very closely resembles E. africanus. It will afterwards be seen 
that in E. falconeri the configuration of this part more nearly resembles that of 
E. indicus and E. ])rimigenitis. 

3. Bones of the anterior extremity. 

The well-recognizable fragments of bones belonging to the anterior extremity of 
E. melitensis, comprise; — 

1. A fragment of the left scapula. 

2. The entire upper end of the right humerus. 

3. A portion of the upper end of the right ulna. 

4. The left olecranon. 

5. A fragment of the shaft of the left ulna, of probably a younger animal. 

VOL. VI. — r.VRT V. 2 M 



244 JIE. BUSK ON THE REMAINS OF 

1. The portion of scapula (PI. XLVIII. fig. 23) consists of the greater part of the neck 
and the adjacent bone, about three inches in its gieatest length. It fortunately retains 
the greater part of the glenoid fossa, of which perhaps the lower two thirds remain 
entire. The remaining portion is about 2 inches long in a vertical direction, whilst the 
greatest width of the fossa is about l"-2. The articular surface is perfectly smooth, 
and its curve in the vertical or longest du-ection has a radius of about 2", and in the 
transverse of about l"-75. The entire fossa may be estimated at about 2"-3 long by 
l"-7 broad. Its sides are pretty nearly parallel, and the lower margin is accurately 
semicircular ; the upper margin *(as already stated) is wanting ; but it may be concluded 
that when entire the fossa was of a broad oblong form, and had none of the constriction 
on the sides which is usually seen in the Asiatic Elephant and, I believe, also in 
E. primigenius. In this respect therefore it would seem more to resemble the glenoid 
fossa of the African than that of the Indian species. The following are the dimensions 
of this part in different specimens of Elephant taken for the purpose of comparison : — 

E. indicus (young) 3'5 X 2'6 

E. indicus (mature) 

E. indicus ( var. sumatrensis, young) 4-7 x 3 -0 

E. jmnii genius (Cuvier) 8'5 X 4-4 

E. africanus (mature) 6-7 X 4'3 

E. meJitensis (mature) 2'4 X 1'7 

The part of the bone immediately supporting the articular fossa is thick, massy, 
and rugose, especially on the dorsal aspect. The entire bone is compact and heavy, 
and it has manifestly belonged to a perfectly mature animal. 

2. The portion of humerus (PI. XLVIII. fig. 22) is in many respects one of the most 
instructive specimens in the entire collection. It is the entire head and upper part of 
the shaft of the right humerus of an animal which had arrived at full maturity ; for the 
proximal epiphysis is completely united to the shaft, although the line of junction is 
still apparent, except to a small extent on the inner side, where it is completely oblite- 
rated. And this is an important circumstance as indicative of the maturity of the 
individual, since the proximal epiphysis of the humerus would appear to be one of the 
latest to become united to the shaft. The fragment is remarkably perfect ; it appears 
to have been recently broken from the shaft ; and a small fragment has been chipped off 
the anterior part of the head, probably at the same time. It is also sliglitly and, to 
all appearance, recently chipped at the hinder border of the head ; and the tuberosity 
is slightly abraded. The bone presents no distinct trace of rolling or morsure. The 
articular surface of the head is somewhat remarkable for its comparative narrowness 
in the transverse direction ; so that, had it been completely detached from the rest of the 
bone, it might very readily have been regarded as fitted more for a ginglymoid than an 
enarthrodial joint. In all Elephants the head of the humerus is somewhat compressed. 



THEEE EXTINCT SPECIES OF ELEPHANT. 245 

or, as it were, elongated in the antero-posterior direction, and perhaps more so in the 
African than in the Asiatic species, as is, in fact, in some measure, shown in pi. vii. 
fig. 3 of the ' Ossemens Fossiles ; ' but in the present case this compression appears to 
be carried to the extreme. There does not appear to be anything distinctive in the 
form or size of the tuberosity ; but the bicipital groove, even as compared with that of 
the African Elephant, is remarkable for its great width and extreme shallowness. In 
fact, after making every possible allowance for the portion of bone which has been de- 
tached from the anterior border of the head, it would seem that there could scarcely 
have been any distinct bicipital groove, and certainly none at all comparable in depth 
with that in E. primigenius, iiidicus, and africanus, as may be seen in Cuvier's 
figures above cited. And from these, as well as from an observation in the text 
(tom. ii. p. 218), it would appear that this groove is .still narrower in the Mammoth 
than it is in E. indicus. It should also be mentioned that both borders of the groove 
arch over it in the mature E. hidicus ; whilst in E. africanus, and still more .so in 
E. nielifensis, there is no incurvation of the kind on either side. 

3. Ulna. — Of this bone the collection contains seven well-recognizable fragments, 
three of which from their size would appear to belong to E. melitensis, and four to the 
smaller species. One of the best-marked specimens of the former is represented in 
PI. XL VIII. figs. 24 and 24 a. It is a portion, about four inches long, of the upper part 
of the right ulna, which has been fractured transversely through the shaft, at about that 
distance below the articular surface. The olecranon is broken off obliquely downwards, 
on a level vnt\i the horizontal part of the articulation. The remaining part of the 
articular surface is nearly entire, being only slightly eroded at the anterior part of the 
outer cusp. The surface of the bone elsewhere, except at the upper part of the pos- 
terior angle, as above noticed, is quite uninjured. 

The bone is evidently that of a fully mature animal ; and from its colour and con- 
dition, both externally and within, it is not unreasonable to consider that it may have 
belonged to the same individual as that which owned the upper end of the humerus 
just described. The form of the articular surface is shown in the figure. The tr. d. 
of the inner condyloid facet, measured from the middle of the radial sulcus, is 
l"-65, and itsap. d. at right angles to the same line, 1"-G5 ; so that it is circular; whilst 
the radius of the curve of the concavity is about l"-3*. The transverse diameter 
of the upper end of the bone at the level of the lip of the articular surface is 2"-6. 
The transverse section at the lower end of the fragment exhibits a nearly equilateral 
triangle, the anterior side [a, fig. 5) of which rises into an angular eminence towards the 
inner side ; the outer side of the triangle is slightly convex, and the internal is nearly- 
straight or slightly convex, — the respective lengths of the three sides being : — internal 

* The tr. d. of the outer facet cannot be exactly determined, as it is partly destroyed ; but its extreme 
length, measured from the bottom of the radial sulcus, may be estimated at 1"-1, and its width, from the same 
point in a line at right angles to its length, is about 0"-9.5. 

2 M 2 




246 ■ MR. BUSK ON THE EEMAINS OF 

l"-25 ; external 1"-15 ; and anterior 0"-75, and 0"-5 on either side of the radial ridge. 
The anterior aspect of the remaining portion of the shaft presents at the upper part a 
very deep and spacious radial fossa, from which is prolonged obliquely downwards and 
inwards a shallow uneven sulcus, about 0"-6 wide, the outer border of which is formed 
below by the prominent ridge, the situation of which is indicated by the letter (r) in the 
accompanying figure, which is intended to show the outline of the transverse section of 
the shaft at a distance of about 3"-6 below the middle of the 
radial sulcus, or at a distance equal to about twice the trans- 
verse diameter of the internal condyloid facet. The external 
surface of the bone is smooth and concave above, slightly 
convex below; the internal is also very smooth, even, and 
nearly flat below, passing above into a rather deep sulcus be- 
tween the inner articular head and the base of the olecranon. 
The posterior angle is very acute, but above it is broken ob- 
liquely off. The internal angle is thick and rounded, and 
the external rounded and slightly carinate. 

4. A second well-marked fragment of the ulna is shown in fig. 25. It is the olecranon- 
process of the left ulna. Anteriorly it exhibits nothing but an irregularly fractured 
surface, and no vestige of the articular surface. It is also slightly broken on the inner 
face, and below it is fractured transversely 3" below the summit. The ap. d. of the 
upper end, measured at a point where the bone is entire, on the outer side of the 
median line, is about 2" ; but it doubtless projected considerably in front of this in the 
median line when entire. In size, colour, and general condition this fragment closely 
corresponds with the one just described, and it may probably be regarded as belonging 
to the opposite ulna of the same individual. 

5. A third portion of a left ulna, corresponding in dimensions with the above, is also 
contained in the collection. Though clearly referrible to the same species, it would 
seem to have belonged to a younger animal ; and as it is a good deal injured, apparently 
by recent fracture, it is needless to enter into any particular description of it. 

If we compare the characters of the ulna of E. melitensis as displayed in the specimens 
above described, with those of the same bone in E. africanus, very considerable differ- 
ences, besides mere size, will at once be perceived. 

(1) Avery striking dissimilarity exists in the form and proportions of the articular sur- 
face. In an ulna of the African Elephant 31" long, the transverse diameter of the 
upper articular end is 7"'6 ; the tr. d. of the inner facet measured from the middle 
of the radial sulcus 4"T, and its ap. d. 3"-l ; whilst the length of the outer facet is 
.3"-4, and its width 2"-l. 

These dimensions of the facets therefore, as compared with the transverse diameter 
of the articular head, taken at 1*000, in the respective cases, stand thus : — 



THREE EXTINCT SPECIES OF ELEPHANT. 247 

E. africanus. E. melitensis. 

Transverse diameter of inner facet 539 -034 

Antero-posterior diameter of inner facet -400 -634 

Length of outer facet -447 -423 

Width of „ „ -276 -360 

(2) The differences between the ulna of the African and Indian species, as regards the 
upper end are not very striking, but so far as they go they tend to show a nearer 
approximation, in E. melitensis, to the African form. These differences are — the 
radial sulcus is more rounded and shallower in E. africanvs, and the inner articular 
facet wider at what may be termed the neck, though in both the existing species that 
facet is much more elongated than it is in E. melitensis. The outer facet in all three 
is much alike, except that in E. indicus it has a small prominent tuberosity in front. In 
both species also there is a rather deep pit or fossa in front of the inner condyle, for the 
insertion probably of the brachialis anticus, which is scarcely indicated in E. melitensis. 

(3) Another character in which E. melitensis approaches E. africanus is in the deeper 
concavity at the upper part of the outer surface of the shaft, which part is nearly flat 
in E. indicus. 

As is well known, the lower articular surface of the ulna varies very materially in the 
existing species ; but as no means exist of comparing this part in E. melitensis, it is 
needless here to notice it further. 

4. Hinder Extremity. 

1. The principal fragment belonging to the hinder extremity of E. melitensis is a 
considerable portion of the shaft of the right femur, represented in PI. XLV. fig. 6. 

It measures 9"'2 in length ; and its least transverse diameter, which is at a distance of 
about 2 inches below the nutrient foramen, is 1"'9 ; whilst the antero-posterior at the 
same part is l"-5, and the circumference 5"'5 ; from which dimensions it may be com- 
puted, according to the data given in the Table of comparative measurements, that the 
total length of the femur was somewhere about 20 inches. The upper extremity, inclu- 
ding all trace of an epiphysial suture, has been broken off irregularly about two inches 
above the nutrient foramen, which is situated on the inner side of the bone, close behind 
the anterior and internal angle of the shaft. The lower end is also broken off in the 
same irregular manner, just where the shaft is beginning to expand ; and there is con- 
sequently no trace of the distal epiphysial surface. The compactness and thickness of the 
cortical substance, together with the well-marked muscular and vascular impressions, 
and the general aspect of the bone, all show that it is that of a mature animal, though 
it is not possible to determine whether the epiphyses were fully united. Its compara- 
tive dimensions, actual and computed, in relation to those of the humerus and portion 
of pelns &c., already described, leave no doubt that it must have belonged to an 
animal of the same size as that indicated by those bones. 



248 MB. BUSK ON THE EEMAINS OF 

V 

The data upon which this conclusion is based will be found in the Table of compa- 
rative measurements. 

The shaft presents all the general characters distinctive of the elephantine femur, but 
at the same time exhibits, in several respects, differences (besides its size) which distin- 
guish it from that of either the Indian or African species. 

In order to render this more evident it will be necessary to say a few words with 
respect to the distinctive differences of the femur in those species, concerning which but 
little seems to have been recorded. 

On this point all that Cuvier * remarks is—" that in the African Elephant the bone 
is slenderer [^fes grele] and has a shorter neck, in consequence of which the upper end 
is not so wide transversely as in the Indian species ; " whilst M. de Blainville f, on the 
other hand, states that the African femur is thicker {plus gros), less flattened, and 
rather more convex in front, with a shorter and more upriglit neck. He also remarks 
that it presents the rudiment of a trochanter minor, and that the external side is straighter, 
and the trochanter major not so high, and furnished with a less-expanded {evasee) fossa 
behind. He states that, at the lower end, the condyles are more equal, especially in 
length, closer together, and consequently separated behind by a narrower sinus. 
Neither of these statements conveys much information ; and that of M. de Blainville 
contains several particulars which are in direct variance with what I have myself been 
able to observe. In the first place, there is every reason to believe, and it will be seen 
from the figures in the Table of measurements, that the African femur is, as remarked 
by Cuvier, rather slenderer in proportion to its length than the Indian, though it does 
not seem that its transverse diameter across the head and trochanter is at all less. 
With respect to the rudimentary trochanter minor, noticed by Blainville, I believe it will 
be found usually considerably more developed, or, rather, more prominent, in the 
Indian than in the African femur, and that the digital fossa behind the trochanter is 
much deeper (and certainly more prolonged downwards) in the latter ; whilst, as regards 
the condyles, there can be no doubt that they are far more unequal in length, if by that 
expression be meant the antero-posterior diameter, in the African than in the Indian 

species. 

The general form and characters of the femur of Elephants are too well known to 
require remark ; but for convenience of description in what I have to say respecting the 
comparison of that of E. melitensis with those of the existing species, it is as well to 
observe that, notwithstanding its great compression in the upper part, and comparative 
rotundity below, the shaft is more or less quadrangular. It consequently presents four 
surfaces (anterior, posterior, internal, and external), separated by four corresponding 
angles. Taking these surfaces and angles in the same order, it will be found that in the 
Indian Elephant the anterior aspect is nearly straight in the vertical direction, whilst in 
the African it is slightly concave, The principal other difference in this aspect is that, 
* On. cit. i. p. 571. t Osteographie dc3 Mnnimiftres, iii. p. 42. 



THREE EXTINCT SPECIES OF ELEPHANT. 249 

ill the lower part, the shaft is more rounded or convex, as may be gathered from the 
circumstance that at the line of the lower epiphysial suture the antero-posterior diameter 
is to the transverse, in the Indian Elephant, as about GOo to 1000 ; whilst in the African 
it is about 661. Besides which the anterior outline of the section at that part is more 
angular in the Indian femur. The difference in question is roughly shown in the ac- 
companying figures, of which fig. 9 represents the transverse section at the lower 
epiphysial suture of ^. africanus, and fig. 10 that of E. iiidicus of about the same age. 




2. On the posterior aspect the diiference is considerably greater. In the African 
femur the bone throughout is flatter, and it is also much straighter in the vertical di- 
rection. In the Indian, commencing at about the termination of the upper third, the 
surface is much more rounded, and the shaft is convex in the vertical direction. The 
digital fossa is much deeper and prolonged further downwards in the African. 

3. The internal surface or aspect in the Indian Elephant is less rectangular, owing to 
the comparatively greater rounding off" of the anterior and internal angle, and the 
greater prominence inwards of the posterior and internal, more especially in the situation 
of the rudimentary trochanter minor, or adductor tuberosity, causing a prominence at 
that part, in the outline of the inner border, which is wanting in the African. The 
whole of the inner border is more rounded in the Indian, and also, in consequence of 
the greater projection inwards of the posterior and internal angle beyond the anterior 
and internal, more oblique in the upper part. 

4. The chief difference observable in the outer border of the shaft arises from the 
ciixumstance that the anterior and posterior surfaces, in the upper part, are less 
parallel to each other in the Indian than in the African femur, in consequence of which 
the outer surface is narrower in the former than in the latter, in M^hich, owing to the 
parallelism of those two surfaces of the bone, both borders are of about the same 
width. 

5. Of the four angles, the anterior and internal is much more rounded off" in the 
Indian than in the African, in which species, moreover, it is marked with a far 
deeper vascular groove. The anterior and outer angle also, in the upper part of the 
bone, is more pronounced in the African. The greater prominence of the posterior and 
internal angle in the Indian femur has already been noticed, to which may be added 



^50 ME. BUSK ON THE REMAINS OF 

that the inner condyloid ridge, which is a continuation of the angle in question, is more 
acute in that species. The posterior and external angle in the Indian Elephant runs 
in a nearly straight direction from the trochanter major to the outer condyle, and rises 
a little below the middle of the shaft into a considerable prominence or rudimentary 
third trochanter, below which it is continued into an acute external condyloid ridge. 
In the African, on the other hand, this angle forms a considerable curve inwards in the 
upper part of the bone, and presents scarcely any projection in the site of the third tro- 
chanter, presenting instead a rather broad rough surface, which is moreover placed 
lower down on the shaft ; and below this the bone is rounded and with scarcely any 
distinct condyloid ridge. 

It may also be remarked that in the Indian femur the surface is hoUowed behind 
the third trochanteric tuberosity, whilst in the African it is not at all so. 

6. With respect to the condyles, as has already been said, they are more unequal 
in length in the African than in the Indian species, as will be seen from the 
measurements given in the Table. 

7. Another distinction which will probably be found constant, arises from the 
situation of the nutrient foramen, which in the Indian Elephant is placed on the inner 
surface, sometimes near the anterior, and sometimes near the posterior angle, but always 
high up on the shaft or in the upper third ; whilst in the African it would seem to be 
situated below the middle, though on the same aspect. 

8. The patellar sulcus is wider in the African. 

With these observations we may proceed to consider the distinctive characteristics 
of the femur of E. meliteiisis. 

1. It differs from the African, and agrees with the Indian, in the convexity in the 
vertical direction of the posterior surface, which is greater than it is even in the latter 
species. 

2. It resembles the Indian, and differs from the African, in the slight degree of 
hollowness behind the rudimentary third trochanter. 

3. It agrees with the Indian, and differs from the African, in the prominence of the 
rudimentary third trochanter, and the development of the external condyloid ridge. 
But it differs from the Indian, and agrees with the African, in the curvature of the upper 
part of the posterior and external angle. 

4. It resembles the African in the depth of the vascular groove on the anterior and 
internal angle, and in the rotundity of the anterior surface of the shaft at the lower part. 

5. It resembles the Indian in the high position of the nutrient foramen. 

G. It differs very considerably from both, but more especially from the African, in the 
want of parallelism between the anterior and posterior surfaces in the upper or com- 
pressed part of the shaft, and the consequent great disparity in width of the internal 
and external surfaces. Other minute differences are perceptible when the different 
bones are placed side by side ; but the above will suffice, perhaps, to show that the 



THREE EXTINCT SPECIES OF ELEPHANT. 251 

femur of E. melitensis presents certain distinctive specific characters in some respects 
intermediate between those of the two existing species. With regard to the points of 
difference between it and that of E. falconeri, I will i-eserve what I have to say until 
I come to that bone. 

§ IV. Elephas falconeei. 

Of the remains referred to this second diminutive species, the following have been 
selected for the purpose of conveying some notion of the characteristics of the mature 
animal. 

1. Portions of an atlas. 

2. Portions of several vertebrae. 

3. A portion of the second rib. 

4. A portion of the scapula. 

5. The greater part of the right humerus. 

6. The lower extremity of the left humerus. 

7. 8. The upper part of the right and left ulna. 

9. The entire proximal phalanx of the 3rd manual digit. 

10. A portion of the pelvis, including the entire acetabulum. 

11. A small portion of the upper part of the shaft of the left femur. 

12. The entire shaft, without the epiphyses, of the left femur of a younger animal. 

13. Astragalus. 

14. The 4th left metatarsal bone. 

Besides these, the collection contains numerous fragments of bone clearly referrible 
to an animal of the same size ; but those above enumerated are sufficient for the present 
purpose. 

1. Atlas. 

Of this important bone, figures of two fragments are given in PI. LI. (figs. 32 & 33). 
which at first sight appear to belong to animals of the same size ; and until I came to 
examine them very closely, I thought that they both belonged to E.falconeri. But 
upon due examination it will be found that one only, represented in fig. 32, really aj)- 
pertains to that species, and that the other is a portion of a very young atlas, belonging, 
as I believe, to E. melitensis. 

The fragment shown in fig. 32 is the greater part of the left half of the atlas of ap- 
parently a perfectly mature animal, as shown by the strongly developed muscular anil 
other impressions, the general density and aspect of the bone, and the complete ossifi- 
cation of the terminal epiphyses of the transverse processes. From its various dimen- 
sions the height of the animal to which it belonged, supposing its proportions to be 
like those of Cuvier's E. indicus, may be computed at about 41", or, if we take the 
presumed height of the African Elephant in the British Museum as the standard, at 

VOL. VI. PART v. 2 N 



252 MB. BUSK ON THE KBMAINS OF 

about 40 inches. This would be a height rather greater than that which I have assigned 
to H. falconeri, computed from the dimensions of the femur and humerus ; but as in the 
latter, at any rate, the upper epiphysis was not united, the height deduced from those 
bones whose growth was not completed may be regarded as somewhat below the mature 
stature. And it will afterwards be seen, when I come to speak of the pelvis of JE. 
falconeri, or what I deem to be such, that that bone also indicates larger proportions 
than those which may be deduced from the humerus and femur, and agreeing very 
closely with those derived from the atlas. I consider therefore that the discrepancy, 
which after all is by no means more than might be looked for in any species of Elephant, 
may be explained upon the supposition that the Zebbug collection contains remains of 
a fully mature E. falconeri, and of one of younger age and somewhat lower stature, oi', 
it may be, of individuals of different sexes, — a supposition for which some support may 
be found in the cu-cumstance that we have in the collection a tusk which from its 
size may be deemed that of a male, whilst at the same time it contains numerous 
bones of extremely young or, perhaps, in some cases, of fetal animals, which must have 
been in immediate dependence upon the mother. 

The present fragment, as I have said, is remarkably perfect, and from it all the infor- 
mation that can be wished for with respect to the atlas is readily obtainable. And it 
is a fortunate circumstance that we are also in possession of a considerable portion of 
the opposite half of the atlas of the lai'ger form, to which I have restricted the name of 
E. melitensis, and which has been already described, although the contrast between the 
two has been reserved to the present place. 

The fragment, from the middle of the inferior tubercle of the ring (which fortunately 
exists) to the point of the transverse process, measures about 2"'5, so that the entire 
breadth of the vertebra was 5"-0. The distance between the outer inargins of the two 
anterior articular facets, or what may be termed the transverse diameter of the con- 
dyloid cup, may be estimated at about 3""3, which of course will be about the transverse 
width at the base of the occipital condyles of E. falconeri, should they ever be met 
with. 

The anterior articular facet presents a very shallow and very small sinus on its inner 
border, whilst the outer margin is quite entire and with very little sinuosity. The 
radius of its longest curve is 0"'85, and that of the surface for articulation with the 
odontoid process 0"-6, whence it may be concluded that that process was rather more 
than an inch in width at the base. It may be observed that the relative proportions of 
the long and short diameters of the anterior facet are exactly the same as in a young 
Indian atlas (No. 2678, K. C. S.), the latter being to the former as 75 to 100; whilst 
in the atlas of E. melitensis the proportion is as 67 to 100, which, curiously enough, is 
precisely the proportion the facet presents in the African Elephant in the British 
M\iseum, — and that a similar difference in the same direction, though apparently to a 
considerably greater amount, is seen in the measurements taken from M. de Blainville's 



THREE EXTINCT SPECIES OF ELEPHANT. 253 

figure of the same species. And this is a coincidence of perhaps some moment when it 
is regarded in relation with the difference already pointed out between the ischial border 
of the obturator foramen in the two small Maltese Elephants— a difference which also 
indicates an approach toward tlie African type in E. meUtensis, and a corresponding 
resemblance to the Indian in E. falconeri. 

2. Other Verfebne. 
Several fragments of the spines of dorsal vertebra; are contained m the collection, of 
which two are represented in PI. LI. figs. 34 and 36 (the largest and most instruc- 
tive of which is shown in fig. 34). It consists of the base of the neural spine, and a 
small portion of the arch. On the under suiface tlie two posterior articular facets are 
left quite entire, and on the dorsal aspect a great part of the left anterior articular facet 
also remains. The perfect facet measures about 0"-6 x 0"-4. As well as can be judged 
from such an imperfect fragment, the vertebra to which it belonged was probably the 8th 
or 9th, or 10th ; and the specimen consequently admits of easy comparison with the 7th 
or 8th dorsal vertebra of ^. meUtensis before described, and which is figured in PI. XLVI. 
The corresponding facets in that vertebra measure 0"-7 x 0"-5 ; whilst the transverse 
width across from the outer edges of the facets, at the widest part, is, in the one case, 
1"-1, and in the other l"-55, proportions corresponding with those of the other bones. 
The other fragments agree in all respects so closely with the one described that there 
can be little doubt of their belonging in all probability to the same individual. 

3. Bibs. 

The only portion of the ribs distinctly recognizable fi-om its dimensions is a fragment 
(neariy three inches in length) of the second right rib (fig. 37). The fragment is a good 
deal worn, and the surface is much eroded, as if by weathering. It is consequently not 
in nearly so advantageous a condition for comparison as the corresponding portion of 
the second left rib oiE. meUtensis, described in page 241, and figured in PI. XLV. fig. 8. 
It nevertheless affords several distinctive characters beyond its mere size, which is at 
once obvious. 

1. In the rib of E. meUtensis the upperside of the neck is excavated into a large 
and deep fossa; whilst in that of E. falconeri it is rounded and without any excavation 
whatever. 2. In E. meUtensis the anterior surface of the bone in the expanded portion 
is very concave, and ui E. falconeri nearly level *. 3. In E. meUtensis a very acute and 
prominent ridge or angle descends for a considerable distance from the anterior part of 

* In the Indian Elephant, both in the very young animal and in one nearly full-grown (Chiuiy), the anterior 
surface of the rib at the part indicated is, if any thing, rather convex, and quite unlike the condition presented 
in E. meUtensis. As in that species, however, the anterior and inner border of the bone in the upper or cui-ved 
part is acutely angular, whilst on the other hand the neck is compressed and evenly rounded on the upper 
aspect, and not thick and hollowed as it is in E. meUtensis. 

2n2 



254 ME. BUSK ON THE EEMAINS OF 

tlie head, which renders the inner or concave border of the rib, for the distance of about 
3 inches below the head, acutely angular ; whilst in E. falconeri this part is rounded. 
On the posterior aspect there is no marked difference, nor is there any on the outer or 
dorsal aspect. 

4. Scapula. 

The only representative of this bone, apparently belonging to the smaller of the two 
dwarf Elephants, is a small portion of the right (PI. XLVII. figs. 14°, 14'). It presents the 
entire glenoid cavity, with the neck, together with the commencement and about two inches 
behind it of the spine ; the remaining portions of the supra- and infraspinous fossa? are 
very small. The glenoid fossa is narrow, elongated, and pyriform in shape. It measures 
about 1""7 X 1", and the border slightly overhangs at the upper end. The radius of 
the longitudinal curve is l"'l, and of the transverse 1", — curves that would seem to 
correspond pretty closely with the computed size of the head of the humerus of U. 
falconeri. The remaining portion of the spine (which shows no sign of an epiphysial 
surface on its edge) rises to a height of about 1""25; and it commences about 1""5 behind 
the margin of the glenoid fossa, at first gradually and then abruptly, the anterior edge 
being smooth and sharp. The dorsal edge is much expanded ; and at the end of the 
fragment, or at a distance of fully 3"'5 behind the edge of the glenoid fossa, there is no 
indication whatever of a descending apophysis, which would therefore seem to have 
been situated further back than it is even in the Indian Elephant, in which it springs at 
a distance of not more than about twice the length of the glenoid fossa behind its poste- 
rior border, whilst in E. africanus it is placed not more than one length behind the 
glenoid fossa. In this respect therefore E. falconeri would seem to approach the Asiatic 
rather than the African type, if indeed it may not have differed from both in the entire 
absence of the descending apophysis. The subscapular surfiice is smooth and evenly 
convex, in a line parallel with, but rather below, the level of the spine. The glenoid 
fossa is narrower below than above ; and the bone at that part is narrow and wholly 
without any of the coracoid protuberance on tlie dorsal aspect, which is so strongly de- 
veloped in the scapula of E. melitensis and all other known species. The bone is 
obviously that of a young animal, as shown by the pitted surface of the articular fossa ; 
and to tliis circumstance the narrowness of the glenoid fossa below, and the slenderness 
of the neck at the lower border may perhaps be in part assigned *. 

* The fragment of scapula above described was regarded by Dr. Falconer as belonging to the dwarf Elephant, 
and it would be very difficult to assign it to any other known animal. But repeated consideration of it since 
the above was written makes me more and more uncertain on the matter. The differences between it and the 
scapula of any known species of Elephant arc so considerable as to be in appearance almost insurmountable. 
Amongst the most marked of these are: — 1, the absence of any descending apophysis within the usual 
distance from the glenoid fossa, as above noticed ; 2, the form of the glenoid fossa itself, though this 
varies perhaps a good deal in the Elephant : 3, the entire absence of the least trace of a coracoid tube- 



THEEE EXTINCT SPECIES OF ELEPHANT. 255 

5, 6. Humerus. 

Of this bone the collection affords several well-marked specimens, two of which are repre- 
sented in PI. XLIX. One of these(fig. 26) is amongst the most perfect and instructive of all 
the bones collected in the Zebbug Cavern. It is a nearly complete left humerus, wanting 
only the proximal epiphysis and great part of the internal condyle with the corresponding 
part of the shaft above it. The upper epiphysial surface, however, remains perfect and 
wholly iminjured; so that we may conclude that the capitular epiphysis was naturally de- 
tached. The fractured surface at the inner condyle appears to be recent ; and the bone 
has been broken obliquely across the shaft, and through the condyloid extremity 
probably at the same time ; but the fragments having been carefully united, the integrity 
of the bone is very satisfactorily restored, with the exceptions above indicated. 

Although the upper epiphysis was not united, there is no trace whatever remaining of 
the line of junction of the lower epiphysis, not even on the exposed fractured surface. 
Nor is there a trace left of the non-ossification of the epiphysial cartilage on the supinator 
ridge, which is late in becoming completed in the Elephant ; we may conclude, conse- 
quently, that the animal to which the bone belonged had nearly, if not quite, reached 
its full maturity and stature. And the maturity of its age, at any rate, may also be 
inferred from the deep and strong muscular impressions, and from the density and 
weight &c. of the bone generally. The specimen, as it is, measures 9 inches in length — 
that is to say, from the highest point of the upper epiphysial surface to the lowest 
point on the condyloid extremity. But from its various dimensions, which will be found 
in the Table, its length when entire may be estimated at about 12 inches, which would 
give a height of about 3 feet to the Elephant to which it belonged. According to the 
same data I estimate the antero-posterior diameter of the head at 2"-2, which is exactly 
proportionate also to the length of the head in E. melitensis, when that dimension is 
measured in relation to the antero-posterior diameter of the upper epiphysial surface. 
As the part which is wanting in the present specimen is precisely that of which we have 
so excellent an example in the upper extremity of the humerus of E. melitensis (fig. 22), 

rosity : and, 4th, the form of the acromial end of the spine, which, so far as I know, is always hooked over, as it 
were. This peculiarity in the Maltese scapula is shown in the subjoined fi^ire, in which the spine is repre- 
sented in a vertical position. 




256 Jnt. BUSK ON THE KBMAINS OK 

and we possess no other well-marked or recognizable portions of the humerus of that 
species, no means exist of instituting a direct comparison between the humerus of the 
two forms ; but the present affords very abundant means of comparison with those of 
other species. 

With regard to the differences between the humerus of the Indian and African Ele- 
phant but little information is to be found in osteological works. All that Cuvier 
remarks on the subject is, that in the African Elephant the bone is of slenderer propor- 
tions, that the deltoid crest descends lower, that the supinator or external condyloid 
ridge is less salient, and that the bicipital groove is wider. 

M. de Blainville, on the other hand, as in the case of the femur, states that 
the African humerus is stouter and shorter, and the condyloid ridge more salient. 
These two writers concur, however, in describing the deltoid crest as descending lower, 
and the bicipital groove as being wider. 

For the purpose of comparison of the fossil bones, I have contrasted the humerus of 
the African Elephant in the British Museum with those of the same length belonging 
to the Indian species to which I have had access — vni\\ the following results : — 

As regards differences in the proportions, expressed numerically, it would seem («) that 
the antero-posterior diameter of the head is about the same, {b) that the transverse di- 
ameter of the head in the Indian is as about 102 to 92 in the African, (c) that the antero- 
posterior diameter of the tuberosity, as well as that of the head and tuberosity together, 
are also nearly equal, [d) that the least transverse diameter of the shaft is about the same, 
but {e) that in the African the antero-posterior diameter at the same part is considerably 
less, or as 60 to 49, whilst (/) the cuxumference at the same part of the shaft is in the Indian 
as 16 to 13 in the African, {g) that the transverse width of the condyles, being 84 in the 
Indian, is 78 in the African, and (/() that the antero-posterior diameter of the inner condyle 
is 63 in the Indian against 50 in the African, and of the outer as 56 to 48, and (/) that the 
proportionate antero-posterior diameter of the inner to the outer condyle is, in the 
Indian as 100 to 88, and in the African as 100 to 85, whilst {j) the antero-posterior 
diameter of the middle of the trochlea between the condyles is the same in both. In 
either species the length of the supinator ridge is the same, and equal to about one- 
third of the entire length of the bone. It may be said therefore that the head and 
tuberosity together are rather more compressed in the African, and that the shaft is, as 
stated by Cuvier, more slender in proportion to its length, and that there is, as in the 
African femur, a greater difference in size or antero-posterior diameter in the condyles 
in the African than in the Indian species. 

With respect to the extent to which the deltoid crest descends, I cannot perceive any 
material difference ; nor do I find that there is any marked difference one way or an- 
other in the salience of the condyloid ridge. Other differences remain to be pointed 
out ; amongst these is the much lower position of the nutrient foramen, which (as has 
been said before with regard to the femur) is placed much lower in the African than in 



THKEE EXTINCT SPECIES OF ELEPHANT. 



257' 



the Indian, being usually considerably above the middle in the latter, and as low as 
the commencement of the lower third in the former. But, besides the difference in the 
proportionate size of the condyles to each other, they differ not inconsiderably in form 
in the two species. The contour of the outer condyle in the African humerus is more 
globose or rounded towards the outside ; and, owing to this and to the circumstance 
that, whilst the middle part of the trochlea is of about the same diameter as in the 
Indian, and the condyles themselves are rather smaller, the transverse contour-line of 
tlie articular surface at the lowest part is widely different in the two cases, as may be 
seen in the subjoined reduced outlines. 




12 & 14. — Elephas afrimitus. 





13 & lo.—EUi,h 



'las ind'icits. 



And it is also to be remarked that another important difference exists in the transverse 
contour-line of the condyles, behind or across what may be called the intercondyloid 
sulcus, which line in the African humerus forms an easy sigmoid curve, whilst in the 
Indian the intercondyloid depression is bounded on either side by an abrupt border, as 
may be better seen in the accompanying iigures. 

In the African humerus the surface of the shaft above the olecranon-fossa is more 
concave than it is in the Indian, and the internal condyloid ridge more acute. And in 
the Indian humerus there is a considerable depression about the middle of the supinator 
ridge behind, which does not exist in the African specimen examined. A further im- 
portant distinction also is seen on the posterior aspect of the bone in its upper half. In 



258 MR. BUSK ON THE REMAINS OF 

the Indian Elephant, of whatever age, an angular ridge is continued more or less dis- 
tinctly from the supinator or external condyloid ridge to the middle of the back of the 
'head (or, more properly speaking, of the shaft), up to the line of the epiphysial junction. 
In consequence of this the upper part of the shaft in the Indian humerus appears angular 
behind, or in some cases almost carinate, whilst at the same part the African humerus 
is rounded and even, the angle continued from the condyloid ridge usually not reaching 
beyond the middle of the shaft. 

Now, with respect to the humerus of E. falconeri, as we have no means of actually 
measuring the head, we are unable to compai-e its proportions with those of the existing 
species. And as the upper epiphysis is wanting, we have no direct means of measuring the 
proportionate diameters of the head and tuberosity ; but, to judge from the form of the 
epiphysial surface, it may be considered probable that the transverse diameter (in pro- 
portion to the antero-posterior) was less than in the Indian, or even African, and 
very much less than in E. primigenius. The subjoined figure gives the outline of this 
surface in E. falconeri, of the natural size. 



In the shaft the chief peculiarities consist: — (1) in the presence of a very deep elon- 
gated fossa on the outer aspect, immediately behind, and overlapped, as it were, by 
the upper part of the deltoid crest; (2) in the comparative shortness of the supinator 
ridge, which equals little more than one quarter of the entire length of the bone, 
instead of one- third, as it does in the Indian and African Elephants ; in this respect 
the boue shows a resemblance to the humerus of E. primigenius ; (3) in the great 
relative disparity in the size of the articular condyles, which is greater even than in 
the African humerus, as the antero-posterior diameter of the mner condyle stands to 
that of the outer in the ratio of 100 to 77. This of course gives the contour of the ar- 
ticular trochlea from below a distinctive character, as may be seen in the reduced figures 



THEEE EXTINCT SPECIES OE ELEPHANT. 259 

adjoined, of which 17 represents the inferior contour of the trochlea as viewed in front, 
and 18 as seen from below, so as to exhibit the posterior intercondyloid fossa. As in the 




great disparity of the condyles, so also in the concavity of the surface above the olecranon- 
fossa and, apparently, in the accompanying elevation of the internal condyloid ridge, 
does the humerus oi E. falconeri resemble that of E. africanus; but in it the concavity 
in question is even still greater. On the other hand, again, in the strongly marked 
angularity of the upper half of the shaft behind, the bone exactly resembles that of E. 
indicus. Scarcely enough of the bicipital groove remains to enable us to determine 
whether it was wide and shallow as in the African, or deep and narrow as in the Indian ; 
but, so far as can be jjidged from the way in which the outer border of the groove arches 
over it, it may be concluded, perhaps, that in that respect it resembled the Indian humerus 
rather than the African, in which neither border ever arches over the groove. And 
in the same particular does the bicipital groove in E.falconeri differ from that in 
E. melitensis, in which, as before said, the groove in its shallowness and width fully 
equals, if it does not exceed, that of the African humerus. In other respects also, so 
far as can be judged from the small portion we possess of the humerus of E. meli- 
tensis, it appears to present several other important points of difference: — The lateral 
compression of the upper epiphysial surface is much more marked in E. melitensis, 
the extreme tr. d. of the surface in the specimen standing in the ratio of not 
more than 63 to 100, whilst in E. falconeri it is 76 to 100 ; whence we may conclude 
that the upper epiphysis, including the head and tuberosity, was proportionally broader 
in the latter. It is to be observed, also, that the small remaining portion of the outer 
surface of the shaft below the epiphysial junction in E. melitensis shows no indication 
of the existence on that side of the shaft of the very peculiar deep and elongated fossa 
which is so striking a feature in the humerus of E falconeri. 

From all that has been stated, it appears to me that, besides its diminutive size, the 
remarkable humerus assigned to E. falconeri exhibits abundant evidence of specific 
distinction from either of the living species of Elephant, as well as from E. melitensis 
and E. primigenius. 

The second portion of the humerus (represented in fig. 27, PI. XLIX.) fortunately 
replaces what was wanting of the lower extremity of the specimen just described. It 
presents the entire condyloid articulation ; and it is from it that the measurements and 
figures just cited, relating to this part, have been taken. It is of slightly larger propor- 

VOL. TI. PART V. 2 O 



260 ME. BUSK ON THE EEMAINS OF 

tions than the shaft ; and it appears to have belonged to an older, or at any rate to a 
more robust animal than that whose humerus has been above described. 

7, 8. Ulna. 

Two well-marked portions of the ulna of E. falconer! are contained in the collection, 
one belonging apparently to a younger animal than the other, which from its colour 
and general aspect would appear to appertain to the same individual as the condyloid 
articular extremity represented in lig. 27 ; whilst the former specimen, fig. 28, in like 
manner agrees in colour and appearance, and probably also in age, with the shaft of the 
humerus represented in fig. 26. But, except in their colour and apparent disparity in 
age, the two specimens exactly resemble each other ; and it is very satisfactory to hnd 
that the peculiar characters shown in them are not limited to a single individual, and con- 
sequently that those characters cannot be regarded as accidental. The specimens are por- 
tions of the upper end of the right and of the left ulna. The longer and more perfect 
fragment is that shown in figs. 28 and 28^*. It is about 3""7 long, from the summit of the 
olecranon, on which is observable a considerable part of the epiphysial surface. The shaft 
is broken irregularly across about 2"'8 below the level of the articular surface, above 
which the olecranon rises about l"-2. The olecranon is about 1"T in transverse diameter 
at the base, and its greatest antero-posterior diameter is nearly the same. The trans- 
verse diameter of the head on the level of the articular surface is about 1"'2 or l"-3, 
and the ap. d. at the same level 2"-2. The internal articular facet is 0"-95 in its 
widest transverse diameter, and the same in the antero-posterior, measured from the 
anterior border to a line drawn across it at right angles from the bottom of the radial 
sulcus. The radius of the curve of the articular facet in the antero-posterior direction 
is 0"-9, and that of the prominent part of the articular surface on the olecranon 0"-625. 

The anterior surface of the bone is hollowed, as usual, immediately below the notch 
for the attachment of the head of the radius; but tiiis hollow is very ciicumscribed, 
and immediately below it the surface is flat from side to side, and a little lower down 
convex. T'he outer surface is concave and quite smooth ; the internal, exce[)t between 
the articular head and the olecranon, flat, or slightly convex. The internal angle is 
rounded and smooth, and without any fossa ; the external very acute, but it does not 
jn-oject at all in front. The adjoining figures repiesent the outlines of a transverse 
section of the shaft, in fig. li) at about 2"T, and hi fig. 20 at about 2"-7 below the 
level of the articular surface at the bottom of the jg „j. go 

radial notch ; and the comparison of tliese with 
the outline of a transverse section of the shaft 
of E. melitensis at a rather lower point (p. 246) 
will serve to show how cl&sely in form they all 
resemble each other. In other respects also the 
ulna o{ E. falvoneri, save in size, appears to agree 





THREE EXTINCT SPECIES OF ELEPHANT. 261 

very closely witli that oi E. nielitensis^ except in one very important particular, which 
alone, as it seems to me, would be amply sufficient to indicate a specific distinction 
between the two forms, even had we no other bones for comparison. Had we been 
in possession of only a single specimen of the ulna of E. falconeri, its remarkable 
character in the respect alluded to might well have been deemed perhaps an accidental 
or individual deviation ; but when we are furnished with two well-marked instances in 
bones belonging to animals of different ages, and also tind that the deviation from the 
ordinary elephantine type is connected with a special characteristic of the humerus 
referred to the same species, it is impossible not to regard the character in question as 
normal, and therefore distinctive. 

One of the great peculiarities amongst tlie many others of the elephantine ulna, as is 
well-known, is the mode in which its articulation with the radius, more especially at the 
upper end, is effected, the comparatively diminutive head of that bone being, as it 
were, embraced between two arms or lobes of the head of the ulna, whose articular 
surface, as remarked by Blainville, thence acquires a trefoil form, the two lateral 
folioles or facets corresponding with the respective condyles of the humerus; whilst 
the central one ascends on the front of the olecranon and fits on the middle part of the 
humeral trochlea. The two lateral facets will therefore naturally differ somewhat in 
their relative dimensions, according to the size of the corresponding condyle. "We 
consequently find that in the African ulna the outer facet is, as compared with the 
inner, of somewhat smaller size than in the Indian ; and it has already been pointed 
out that in the ulna of E. melitensis the disparity is still greater in the same direction. 
In E. falconeri it is carried to the extreme, and it may almost be said that the outer 
foliole of the trefoil is wholly aborted, as may be seen in the figure (fig. 28 a). It 
is true that a small splinter has been broken off the external angle in front, just below 
the articulation, and also that the extreme anterior angle of the facet itself is abraded ; 
but it does not appear that either the fracture or abrasion encroaches much, if at all, 
upon the actual articular surface itself. At any rate in E. falconeri the outer facet is 
reduced to a minimum ; and it is interesting to observe with relation to this diminution 
that the outer humeral condyle, also, as compared with the inner, is smaller in that 
species than in any other with which it was compared. This abortion of the outer 
facet, and the attenuation of the corresponding part of the bone upon which it would 
be supported, give the ulna of E. falconeri so peculiar a character as, even when 
compared with that of E. melitensis, at once to strike the attention and to distinguish 
it from the corresponding bone in any other known species or form of Elephant, either 
recent or fossil *. But it is nevertheless interesting to institute some compai'ison between 
it and that of the Indian and African species in other particulars. Unfortunately, 
owing to the want of any other part of the bone except the upper extremity, and 
especially to the absence of the lower articular surface, which seems to afford excellent 
* The subjoined figures will convey an idea of the difference in form pf the upper articular surface in 

2o2 



262 



ME, BUSK ON THE KEMAINS OP 



distinctive characters, little can be said on this subject. It may be pointed out, however, 
that the bone, on the anterior aspect, differs very widely from the African in the com- 



E. afrimnns, E. hicHnis, E. melitensis, and E. faleonei-i. The two latter are of the natural size, and the others 
enlarged to the same width from the figures in the ' Ossemens Foseiles.' 





E. indiriis. 



E. afrieanus. 



23 



1 <t C.::- -Vii 





E. meliteiisis. 



E. falconeri. 



THEEE EXTINCT SPECIES OF ELEPHANT. 263 

parative flatness or rather convexity of the surface below the radial fossa, and in tlie 
consequent want of elevation of the outer and inner angles which constitute the borders of 
a deep concavity in E. africanus. From the same species it also differs in the rotundity 
of the inner angle, in which it in the same degree resembles the Indian. It also differs 
from the African in. the absence of any flattening on the inner face of the inner head, 
and from both the Indian and the African in the want of any pit or depression in front 
of the inner head (for the insertion, as I suppose, of the hraclnaUs anticus). But in the 
apparent arching inwards of the olecranon it presents a decidedly African character. 

As regards the dimensions of the bone when entire, if we take the comparative length 
of the humerus and ulna in the Indian Elephant, as exemplified in the skeleton of 
Chuny in the Koyal College of Surgeons, the length of the ulna in E. falconeri would 
be about 10"-3 ; according to the skeleton of the Ceylon Elephant in the British Museum, 
9"-9 ; according to Cuvier's measurements of the Indian Elephant, 9"-8; and according 
to the African Elephant in the British Museum, 9"*2. We may conclude therefore 
that the probable length was about 10 uiches. Assuming this as the length, it would 
seem that the transverse width of the upper articular head, allowing the utmost for loss 
by abrasion &c. is at least 0"-5 less than it ought to be, had it stood in anything like the 
same proportion to the length of the bone that that dimension does in the four- instances 
cited, in which the diameter in question would seem to equal about one quarter of the 
entii'e length of the bone. 

9. Bones of the Fore Foot. 

The only other bone belonging to the anterior extremity that admits of satisfactory 
identification is the proximal phalanx of the third digit of the left side (PI. LI. fig. 
41). The bone, which appears to be that of a mature animal, as the epiphysis is per- 
fectly united, without any trace of the junction, is quite entire, and presents no trace 
of weathering or wear. It is exactly I" long, and the same in tr. d. at the upper end, 
which is 0"-9 in ap. d., whilst the lower end has a tr. d. of 0"-9, and an ap. d. of 0"-5o. 
In form and proportions it differs in no respect from the corresponding bone in E. indi- 
ctis (Chuny), which has a length of 3"-0. Assuming the proportionate lengths of the 
bones to be similar to those in E. indicus, as shown in the specimen above named, 
this proximal phalanx would give for the humerus of E. falconeri a length of between 
11" and 12", or about the same as that which I have deduced from the other data whicli 
have been already discussed. 

10. Pelvis. 

The pelvis oiE. falconeri is represented by a considerable portion of the left os innomi- 
natum, which is shown in PI. L. fig. 31. The fragment includes the entire acetabulum, 
with a small portion of the body of the ischium, and a still smaller portion of the body of 
the pubis, — very important parts of the bone, inasmuch as they form the boundaries of the 
upper or anterior part of the foramen ovale, the value of which as affording a distinctive 



2M ME. BUSK ON THE EEMAINS OF 

character between the pelvis of the Indian and African Elephants has been already fully 
referred to (p. 242). Above the acetabulum is a large portion of the ilium with its anterior 
curved margin, extending to a height of more than three inches above the upper border of 
the acetabulum. The general character of the bone is that of a fully mature animal ; 
and. from its brown colour, condition, and comparatively large size, it corresponds very 
exactly with the larger and older portions of the humerus and ulna already described 
and referred to an individual of larger size and more mature age than that to which the 
shaft of the humerus and the more perfect portion of the ulna probably belonged. 

The dimensions of the bone, so far as they are afforded in the specimen, are as 
under : — Width of acetabulum (inside) 2"-l, length 2"-3 ; radius of concavity 
about 1"-1. The cotyloid notch is about 0"-5 in width ; and the channel continued from 
it on the anterior surface of the body of the ischium ceases immediately beyond the 
cotyloid border. The width of the contracted part of the ilium, above the acetabulum 
is l"-8. The body of the ischium is unfortunately broken off obliquely on the outer 
side ; but about l"-25 of the obturator border remains. This border is acute, and presents 
about 0"-5 below the summit of the obturator foramen a slight elevation representing 
Avhat I have termed in a previous part of this paper the ischial obturator spine, which is 
so strongly developed usually in the Indian Elephant, as well as in the Mammoth. The 
leno-th remaining of the pubic border of the foramen is too short to exhibit any trace 
of the corresponding pubic obturator projection ; but sufficient is left of the ischial 
border to show the important difference in form between that part and the corresponding 
part in E. meJitensis, which in that respect, as before pointed out, more resembles the 
African than the Indian species. Another particular in wliich the present specimen 
approaches the Indian and differs from the African type is in the comparatively great 
width of the cotyloid notch, which, as is well known, is much narrower in the African 
acetabulum. The internal or pelvic surface of the ilium is smooth and equably concave, 
and the outer is also smooth and evenly convex. Posteriorly the triangular surface of 
bone forming the back of the acetabulum is much less concave from side to side than 
in either the Indian or African species; and the two borders consequently are indistinct 
and rounded. And, corresponding with this general flatness of the surface in this part, 
the excavation continued upwards from the obturator foramen is extremely sliallow. 

11, 12. Femur. 

The collection contains at least three well-recognizable portions of the femur of E. 
falconeri, belonging to individuals of widely differing ages. Two of these specimens are 
shown in PI. L. figs. 29, 29 a, & 30. The latter of these is the upper portion of the shaft 
of a left femur, to all appearance of at least tolerably mature age ; and it corresponds in 
all respects as regards colour and condition with the shaft of the humerus (fig. 2(j). The 
fragment measures 3"-4 in length ; and it is broken irregularly across at the upper end, 
just above the inferior termination of the posttrochanteric fossa. At the lower end it 



THREE EXTINCT SPECIES OF ELEPHANT. 



205 



has been sawn across, I believe by Dr. Falconer ; and this part is probably, to judge from 
analogy, not very far above the point of least circumference of the shaft. No indication 
of the nutrient foramen is to be perceived in the fragment ; and its situation, therefore, 
was in all probability lower down the shaft than is usually the case in the Indian 
femur. The various dimensions afforded by the specimen are : — tr. d. at upper end 
2"-4., ap. d. about 0"-95 ; tr. d. at lower end 1"'25, ap. d. U"-85 ; circumference 3"-4 ; 
whilst the outline of the transverse section at that point, which, as before said, cannot 
be very much, if at all, above the point of least circumference of the shaft, is shown 
in the accompanying figures, contrasted with that of the femur of £. melitensis, taken, 
as nearly as can be judged, at the same part of the shaft. The anterior surface of the 





E. melitensis. 

bone presents a slight elevation in the middle, with a very shallow depression internally, 
and a much deeper and larger one (a pretrochanteric fossa, as it may be termed) 
externally, the outer boundary of which is formed by a well-pronounced, rough, elon- 
gated tuberosity. On the posterior aspect, at the upper end and outer angle, is seen 
the strongly projecting base of the trochanter major, within which is the lower part 
of a deep digital or posttrochanteric fossa. The inner and outer surfaces are of very 
nearly the same width from before backwards ; and they both have the rectangular form, 
peculiar more especially to the African femur. 

This fragment, compared with the corresponding part of the femur of E. melitensis, 
exhibits such marked differences, in almost all respects, as to afford, perhaps as strongly 
as any other of the remains, as striking a proof as can be desired of the, at any rate, 
specific difference between the two dwarf Elephants. In the first place the transverse 
section of the shaft, shown in the two figures given above, is widely different at corre- 
sponding points. On the anterior aspect the surface is totally different in the two 
cases. In E. -melitensis it slopes obliquely backwards and outwards, from the anterior 
and internal angle, with an even, slightly convex curve ; whilst in E. falconeri (owing to 
the anterior and posterior surfaces in the upper part, and till very near the lower end 
of the fragment, being parallel to each other, and the outer and inner faces conse- 
quently of equal width) the anterior surface is not oblique. But a still greater pecu- 
liarity in this respect, consists in the presence of the remarkable pre- or, more properly, 



266 ME. BUSK ON THE REMAINS OF 

infratrochanteric fossa, the existence of which, so far as I am aware, is peculiar to E. 
falconeri. On the posterior aspect the base of the trochanter major, or rather the 
posterior and external angle of the shaft going to it, is very much more elevated, as is 
also the surface of the bone on the inner side of this face. The central part, consequently, 
just below the level of the lowest point of the digital fossa, is much more concave in 
E. falconeri. In fact, the most cursory inspection of these two portions of the femur is 
sufficient to demonstrate the extreme difference between them. It will have been ob- 
served that in some particulars the femur of E.falconen exhibits African tendencies, 
which is a curious circumstance when we remember the numerous instances, in other 
parts of the skeleton, in which the contrary tendency would seem to be manifested. 

With respect to the dimensions of the entire femur, as deduced from those of the 
fragment, and from the length of what I regard as the corresponding humerus, its length 
may probably be taken at about 13" or 14", and the diameter of the head in the fully 
mature animal at about I"'8, or 2". The latter dimension is less than would accord 
with the size of the acetabulum in the portion of pelvis here referred to the same 
species, which demands a head of about 2" in diameter at least ; and I am able to 
explain the apparent discrepancy only on the ground, either that the proportion of the size 
of the head to that of the shaft was rather greater than usual in E. falconeri, or, as is 
perhaps more probable, that the pelvis in question belonged to an older and larger in- 
dividual of the same species. But the fact that the comparative size of the head in 
proportion to the length of the shaft varies very greatly, not altogether in accordance 
with age, is apparent in the circumstance that, if we deduce the size of the head in 
E. falconeri from the proportions exhibited in the femur of Chuny, it would be about 
1"'9 ; whilst if we take the proportions in a somewhat younger, but very much smaller, 
Sumatran Elephant in the British Museum, it would be only I"'5, and, from those of 
a Ceylon Elephant also in the British Museum, l"-7, and, from those of the African, 
l"-6. And these examples have been selected as being, I believe, those of animals 
as nearly as possible (except the Sumatran) of the same age, or with the dentition in 
nearly the same stage — that is to say, with the three molars in wear. In ail of them the 
femoral and many of the other epiphyses are still not united. Upon full consideration, 
therefore, I think it not unsafe to conclude that the apparent discrepancy between 
the size of the acetabulum and the computed size of the head of the femur represented 
in fig. 30 is not greater than may be looked for within the limits of one and the same 
species. 

A second instance of the femur of E. falconeri is that represented in figs. 29 & 29 a. 
It consists of the entii-e shaft of the left femur with both epiphyses detached ; and it 
is also, from its taper form and all its other characters, manifestly that of a very young 
animal. On the anterior aspect the great concavity on the outer side, immediately 
below the base of the trochanter, at once marks the peculiarity of the bone and its re- 
semblance to the older femur last described. On the same aspect is seen the nutrient 



THEEE EXTINCT SPECIES OF ELEPHANT. 



267 



foramen in an unusual situation for an Elephant — that is to say, in the middle of the an- 
terior surface, about 3" -5 below the summit of the shaft. Since in the former instance 
there is no trace of this foramen in the more usual situation, which is on the inner 
aspect and (as in the case of the Indian Elephant) at, or but little below, the upper third 
of the entire length of the bone, it is interesting to find its situation indicated in the 
present specimen ; and this situation, if it be not a mere individual variation, will further 
indicate an important distinctive character in the femur of E. falconeri *. At the lower 
end the form of the anterior surface is subtriangular, and in that respect more like the 
corresponding surface in the young Indian than in the African femur, in which, as 
before remarked, the anterior aspect of the bone is more rounded. On the posterior 
aspect the chief peculiarity consists in the great projection backwards of the postero- 
external angle above, by which the surface is rendered concave. At the lower end 
may be noticed a rather deep groove on the internal condyloid ridge. 

In order to give as complete an idea as I can of the distinctive peculiarities of the 
femur of E. falconeri, and of the manifest relation the present specimen bears to that 
last described, I have added the subjoined outlines of the transverse section in a line 
immediately below the base of the trochanter major, and as nearly as possible at the 
corresponding level in all three instances. From the more imperfect condition, however. 




1. 
2. 

3. 

as. 
ps. 



ea. 

atf. 



Transverse section of femiir of E. melitensis 

(PI. XLV. fig. 6). 
Transverse section of femiir of E. falconeri 

(PI. L. fig. 29). 
Transverse section of femur of E. falconeri 

(PI. L. fig. 30). 
Anterior surface. 
Posterior surface. 
Internal surface. 
External surface. 
External angle. 
Pretrochanterio fossa. 



* It may be remarked that, in this instance, the nutrient foramen occupies the same situation that it does in 
the greater number of Ruminants. 



VOL. VI. PART V. 



2p 



268 MR. BUSK ON THE EEMAINS OF 

of the femur of E. nielitensis, the secticm in that case is taken at a rather lower level, for 
which some allowance must be made. 

In these figures will again be seen the very different form of the femur in the two 
dwarf Elephants, and at the same time the great peculiarity of the anterior surface in 
A', falconeri, especially in the presence of the pretrochanteric fossa (indicated by the 
letters a tf), and of tlie posterior, in the great elevation of the postero-exteraal 
angle e a. 

The accompanying cut represents the outline of tlie lower epiphysial surface. 




The various dimensions of the bone are given in Table V. ; and, taking tliese data in 
comparison with tiie corresponding measurements of the femur of a young Indian 
Elephant of probably about the same age, in the British Museum, it would seem that 
the proportions are pretty nearly the same in both cases. The actual length of the 
shaft, without the epiphysis, in the specimen is 9"-5 ; and by computation from the di- 
mensions of the shaft of the femur of the young Elephant above referred to, which 
measures 21" in length, it would be 9''-6, the utmost deviation in any direction not 
being more than 0"-4. This coincidence perliaps affords some gi'ound for believing that 
the general proportions of the length of the limbs of E. falconeri to its height, at any 
rate when young, were not widely different from those of the Indian Elephant. 

1.3, 14. Bones of the Hind Foot. 

(1) Astragalus. 

Tlie only specimen of the Astragalus, contained in the Zcbbug collection, is that 
whose upper surface is shown in PI. XLVII. fig. 1 4. 

The bone is that of an immature animal ; and a portion is broken off on the outer 
.side, so that the greater part of the peroneal facet is removed. The true characters, there- 
fore, of the mature bone are not fully displayed in the specimen. 

The immature condition of the astragalus is shown by its generally light and porous 
condition, the thinness of the cortical layer, and by the remains of an epiphysial surface, 
marking the site of the unossified internal tuberosity, which, in the Elei)liant's astragalus, 



THREE EXTINCT SPECIES OF ELEPHANT. 209 

it would seem, is devuloped from a distinct point of ossification, or remains much 
longer in the condition of cartilage than in most other animals. It is also further 
evidenced in the condition of the internal calcaneal facet, which is not yet formed into 
a single articular surface, but consists of two small ones, separated by a shallow depres- 
sion — exhibiting in fact exactly the same condition as that shown in M. de Blainville's 
figure of the under surface of the astragalus of E. africanus, and regarded, apparently, 
by that author as a distinctive character between the Indian and African astragalus. 
In truth, however, it only indicates an immature condition, since in tiie mature African 
astragalus no trace of such a division of the facets exists, any more than it does, I 
believe, in any other species. It is not improbable, nevertheless, that the completion of 
this articular surface may be effected later in the African than in the Indian Elephant ; 
for in a very young astragalus of the latter the surface in question is quite entire. 
And as this, from other circumstances, seems to be an astragalus of about the same 
age as the Maltese specimen, it may be supposed that the latter may have resembled 
the African in the comparatively late completion of the internal calcaneal facet. 

In its general form the astragalus strongly resembles that of a young Indian Elephant in 
which the internal tuberosity is still unossified. The chief peculiarity observable in it, 
irrespective of the proportionate dimensions of the different parts, is in the existence of 
a large and deep pit towards the anterior part of the sulcus for the interosseous or 
calcaneo-astragaloid ligament ; no such pit exists in any other astragalus that has come 
under my notice. But it may, of course, be merely an individual peculiarity. In the 
comparative length of the neck the bone resembles the Indian rather than the African 
astragalus, the upjjer edge of the scaphoid facet projecting in front of the middle of 
the anterior border of the tibial facet exactly one-half of the median antero-posterior 
diameter of that facet. 

With regard to the proportionate dimensions of the bone in its various diameters, and 
of the different facets as compared with the same measurements in the existing species, 
in most particulars no marked difference is observable, as will be seen from the measure- 
ments in Table IV., from which it will also be perceived that with respect to its somewhat 
greater proportionate breadth, again, the astragalus oi E. falconer i shows a resemblance to 
the Indian rather than the African type. But in the proportionate dimensions of the 
various facets rather more important differences are observable. For instance, whilst 
in the Indian astragali, measured by me, the mean vertical diameter or height of the 
scaphoid facet, as compared with its length across, was about as 626, and in E. africaniis 
as 602 to 1000, in E. falconeri this dimension is not more than 500. And, again, 
whilst in the Indian Elephant the antero-posterior diameter of the tibial facet stands in 
relation to its transverse diameter as about 758 to 1000, and in E. africanus as 900, in 
the mature E. falconeri it is 941, showing a rather remarkable difference, more especially 
from the Indian species. But of all these differences the most striking appears to be 
in the transverse or longest diameter of the scaphoid facet, wiiich, in the Indian and 

2 p 2 



270 



ME. BUSK ON THE EEMAINS OF 



African Elephants, as compared with the total width of the astragalus, stands in the 
ratio of about 758 to 1000 ; whilst in E.falconeri it is about 900 *. 

If we proceed to compare the size of the astragalus with the computed and actual 
dimensions of the humerus and femur referred to E. falconeri, the result would, at 
first sight, seem to indicate an animal of rather greater stature than was assigned to 
that species from other considerations. For instance, in the Sumatran Elephant in the 
British Museum, which may be taken to represent the type of a somewhat diminutive 
variety of E. indicus, the Midth of the astragalus is about 4", and the length of the 
humerus 28", and of the femur 33"-5, in accordance with which the humerus of E. 
falconeri should be about 14", and the femur between 16" and 17". Again, in E. 

* The subjoined figures, taken from specimens in Dr. Leith Adams's collection, which has come into my hands 
since this paper was read and the ahove account of the astragalus drawn up, represent what I regard as the 
astragali of E. melitensis and £. falconeri. They have belonged to fuUy mature animals, and are drawn of 
the natural size : — 




E. fnlconeri 



THKEE EXTINCT SPECIES OF ELEPHANT. 271 

africanus the breadth of the astragalus is 5"-2, and the length of the humerus and femur 
respectively 3G" and 42", according to which, in E. falconeri these bones should measure 
14"-5 and 16"-9, or nearly the same as above. But if we take the proportions presented 
in the skeleton of Chuny, the result is different. In that instance the astragalus 
measures about 6" across, the humerus has a length of 35", and the femur one of 42", 
proportions which would make the lengths of those bones in K falconeri respectively 
12"-2, and 14"-7; and these dimensions do not differ very widely from those already 
derived from other data. And even this difference, such as it is, may perhaps be in some 
measure explained, as regards the Sumatran Elephant, and still more as regards the 
African, by the circumstance that the Maltese astragalus is comparatively wider in pro- 
portion to its size than it is in either of those forms, and very nearly corresponds with 
that of Chuny. 

(2) Fourth Metatarsal. 
A second bone which I refer to the hind foot of K falconeri is the fourth left metatarsal 
(PL LI. figs. 40, 40 a, 40 b). The bone is quite entire, and only shghtly chipped 
(m some of the prominent edges. It is of a dark-brown colour and obviously that of a 
fully mature and, as I should judge from its proportionate size as compared, for instance, 
with the proximal phalanx (fig. 41) which has been already described (page 263), rather 
large animal. As it entirely corresponds in its somewhat peculiar colour, compara- 
tive size, and other characters of age &c. with the condyloid end of the humerus 
represented in fig. 27 (which, as has been already stated, is of robuster pj-oportions than 
most of the other bones referred to E. falconeri), and also with the portion of tlie 
pelvis, fig. 31 (likewise distinguished by its comparatively large dimensions), we might 
perhaps venture to surmise that the metatarsal may have belonged either to the same 
individual or to one of similar size. The corresponding metatarsal in R indicus (Chuny) 
is 4"-8 long, corresponding with a length of 42" for the femur, and of 35" for the 
humerus. From these numbers we obtain, for the humerus of E. falconeri to which the 
metatarsal belonged, a length of about 13", and for the femur one of 15"-7— figures 
probably not far from representing the extreme size of those bones in that species. 
But any computation of the lengths of the long bones from those of the metatarsals may 
be regarded as liable to error, when we consider the remarkable difference in the pro- 
portionate size of the scaphoid facet of the astragalus in E. falconeri and E. indicus, 
which can hardly be unaccompanied by corresponding differences in the other bones of 
the tarsus and metatarsus. It is possible therefore that the metatarsal bones in E. 
falconeri may have been proportionally longer than in E. indicus, and consequently 
from their sizes those of the humerus and femur computed somewhat above the reality. 

§ V. Very young or Immature Bones. 
Having adduced the evidence afforded by the mature or nearly mature bones in 



272 Mfi. BUSK ON THE REMAINS OF 

Capt. Spratt's collection, as to the presence in it of the remains of three distinct species 
of Elephant, I will proceed to inquire what confirmation of the plurality of species is 
Added from the study of the remains of the very young (or, perhaps, in some cases, of 
absolutely foetal) animals. 

The number of these immature bones is very remarkable ; and many (although neces- 
sarily of very fragile texture) are in excellent preservation, though the majority are more 
or less worn or otherwise injured. 

In noticing the largest species of the Maltese Elephants, I have already described a 
very young exoccipital bone (figs. 3 & -4), and a portion of the shaft of a femur, 
obviously that of a very young animal, and which may have belonged to the same indi- 
vidual. Besides these juvenile remains of the large Elepliant, there are several other 
fragments apparently referrible to the same species, though of much younger age. As 
the mere size of these specimens is sufficient to distinguish them from the remains of 
either of the dwarf species, it will be unnecessary perhaps to enter into further par- 
ticulars concerning them, beyond what have already been given. 

With respect to the immature bones of the two smaller Elephants, however, it will 
be requisite to go into some detail, in order to point out what I conceive to be dis- 
tinctive specific characters in them, although I do not pretend at present to be able to 
assign the different forms to the respective species with any approach to certainty. 

1. Bones of the Cranium and Face. 
Of the cranial bones the only ones sufficiently perfect to be of much use in the 
inquiry are three or four exoccipitals, two of whicli are represented in PL LII. figs. 42', 
42' «, and 44, 44 a. These two bones are of the same size, and, to all appearance, of 
exactly the same age. Each is also broken precisely to the same extent, having lost 
the posterior angle, including the whole of the thickened border which joins the supra- 
occipital *. In all other respects the bones are entire and but very little worn, especially 
that shown in fig. 42'. One belongs to the right, and the other to the left side ; but 
they are obviously not a pair ; and it is curious that the collection also affords a third 
exoccipital belonging to the left side, not so perfect as either of those which have been 
figured, but sufficiently so to prove that it has exactly the same characters as fig. 42'. 
We have thus, from these bones alone, evidence of the presence in the ossiferous cave 
of Zebbug of three very young Pigmy Elephants. In general dimensions, as has been 
said, the two exoccipitals very closely agree, the principal difference remarkable between 
them being that fig. 42' f is rather higher, as it may be termed, in proportion to its an- 
tero-posterior diameter than fig. 44. The exact antero-posterior width cannot be positively 
given, owing to the imperfect condition of the bones ; but as the posterior part of each 

* Tke fact is, that both are broken across the weakest part of the bone. 

t To save words, as I am unwilling to employ a specific name, I use the Nos."of the figures to designate the 
two bones. 



THUEE EXTINCT SPECIES OF ELEPHANT. 273 

is broken oiF immediately behind the condyle, their respective breadths across the con- 
stricted part admit of fair comparison. Compared in this way, fig. 42' measures in 
extreme height l"-95, and at the constricted part above the condyle 0"-9 ; whilst fig. 
44 in the corresponding directions measures l"-85, and 1"-1. It is true that the latter 
is a little more worn on the upper edge ; but, making every allowance for this, it is un- 
doubtedly the wider of the two in the antero-posterior direction. Again, in tig. 42' the 
condyloid articular facet measures 1 " X 0"-5, and in Xo. 44 1"-1 x 0"-5. And in No. 42' 
the surface of the exo-basioccipital synchondrosis is 0"-G X 0"-.3, and in No. 44 0"G x 0''-4. 
But, besides these differences in measurements, which in such small dimensions are not 
so inconsiderable as they may seem, the two bones present others, as it appears 
to me, of even greater importance. In the first place, on the inner aspect of No. 42' 
(42' a) the cerebellar fossa is much more concave, and the sulcus for the lateral sinus 
much more pronounced ; whilst in 44 a the cerebellar fossa is but slightly concave, 
and no discernible trace of the lateral sulcus can be perceived. In consequence of 
this difference in the internal aspect, the opening of the paramastoid cells {p c), in 
fig. 42' «, is separated, as it were, from the cerebellar fossa, or ratlier from the lateral 
sulcus, by a steep or abrupt wall, which is wholly wanting in 44 a. On the outer aspect 
the chief thing remarkable is the greater flatness of the surface. The anterior margin 
immediately above the exo-basioccipital synchondrosis, or at the part where it forms the 
posterior boundary of the jugular foramen {j f) (the jugular sulcus, as it is termed in 
human anatomy), is very acute in both specimens; but the bone itself, immediately beyond 
the border, is very much thicker in 44. And this difference is so great upon viewing 
the bones edgeways, though not readily described in words, as of itself to give a diff'erent 
character to the two bones when viewed in this aspect. I am moreover particularly 
desirous of directing attention to this part of the bone, inasmuch as it is here that a 
very important difference exists between the exoccipital of the Indian and that of the 
African Elephant at the same age as the Maltese specimens. And it fortunately happens 
that we have the materials for comparison in the British Museum and Royal College 
of Surgeons, the former aff"ording the cranial bones of an African, and the latter that 
of an Indian Elephant, of apparently the same age as the Maltese bones we are discuss- 
ing. Comparison of these shows that, whilst at first sight they more closely correspond 
than might have been anticipated, in many respects they diff'er to about the same extent 
as the two Maltese specimens. Amongst these differences the most striking is the form 
of the anterior border at the jugular sulcus : in the African this border is very thick 
and rounded, whilst in the Indian it is thin and acute. And, as might be supposed, there 
is to some extent a corresponding difference in the depth of the lateral sulcus, and the 
general concavity of the cerebellar fossa, both of which are considerably greater in the 
Indian than in the African, though perhaps not to the same extent as in the two 
Maltese exoccipitals. On the same aspect also there may be observed in No. 44 two 
ridges close behind the edge of the jugular sulcus, where in No. 42' the surface is 



274 



ME. BUSK ON THE EEMAINS OF 



perfectly even. In No. 42' the exo-basioccipital synchondrosis projects more in front 
than in No. 44, as it does in the African as compared with the Indian. 

On the whole it would seem that the distinction between the two dwarf Maltese 
forms is pretty nearly, though certainly not quite, as well marked as is that between the 
two existing species, and that in some respects the form represented in fig. 44 exhibits 
rather a tendency towards the African type. 

As it may perhaps occur to some that the two small exoccipitals represented in PI. LII. 
might possibly have belonged to extremely young foetuses of a lai-ger Elephant, the 
characters which go to disprove such a supposition may be briefly pointed out. The 
subjoined woodcut represents the outlines of the inner and outer aspects of the exocci- 




pital of a very young uterine foetus of E. africanus in the British Museum. The bone 
itself, as will be observed, is on the whole of pretty nearly the same dimensions as those 
of the Maltese pigmy Elephants, whilst the condyloid facet {cf) and the exo-basiocci- 
pital synchondrosis [b o s) are both considerably smaller. It will also be seen that, 
although the expanded portion of the bone is at least as wide as in the Pigmy exocci- 
pitals, it exhibits scarcely a trace, or merely a rudimentary commencement, of the para- 
mastoid cells (p m c) which are so largely developed in the others. The outlines 
also of the symphysis and adjoining part of the mandible belonging to the same foetus 
will suffice to show that, even at a very much earlier stage of development, those bones 
are very much larger than the corresj)onding part in either of the pigmy Elephants, 
in which there is reason to believe the second milk-molar had been well used. Besides 
this difference in the degree of development of the peripheral part of the African foetal 
exoccipital, the condition of the bone itself is very different. When received at the 
Museum the bones were preserved in spirit ; and when in the moist state they were quite 
soft and almost cartilaginous, and now when dry are exceedingly light and fi-agile ; 
whilst in the Maltese specimen the bone is firm and solid, and evidently much further 
advanced in ossification. 

For the purpose of comparison, as to size, with other bones of the pigmy Elephants, 
1 have thought it might also be useful to give the subjoined outlines of various other 
bones of the same African uterine foetus. 



THKEE EXTINCT SPECIES OF ELEPHANT. 



275 



Prsemaxilla 




Various bones of uturiue lujtus of E. iifrlcanus. 



VOL. VI. PART V. 



2q 



276 ME. BUSK ON THE REMAINS OF 

2. Uj/per Jaw. 

The collection affords two fragments of the premaxillary bone, and none apparently 
of any other part of the upper jaw. 

1. One of these is a fragment rather more than 3 inches long, and about 1| wide at 
the lower end, evidently (from the texture of the bone) of a young animal, and from its 
dimensions not improbably belonging to the same individual, or to one of the same age 
and size, as that whose exoccipital is represented in PL XLIV. fig. 4. It is a portion 
of the right premaxillary ; and on the inner side the socket of the permanent tusk is 
exposed, in the form of a sulcus about 2 inches long, and about 0"-5 in diameter. 

2. The second specimen of the same bone is represented in PI. LIT. fig. 46, with what 
appears to be the point of a very young permanent tusk in situ. It should be observed, 
however, that there is no reason, so far as I know, to believe that the tooth and bone 
were actually found in conjunction. The former seems to have been fitted and cemented 
into its place by Dr. Falconer ; it is therefore impossible to say whether it really belonged 
to the bone or not. The fragment which belongs to the left side is of dense and close 
texture ; and the bone, to all appearance, is not that of a young animal, although un- 
doubtedly one of very small size. I should be inclined to refer it to E.falconeri ; and 
in colour and condition it exactly accords with some of the other bones already referred 
to that species. 

The portion of the tusk attached to it is nearly 2 inches long, and about 0"-4 in 
diameter. Its surface is marked by well-pronounced parallel ridges ; and the exposed 
part is partially coated with a thin layer of very hard semi transparent substance of ferru- 
ginous colour, and apparently of the nature of enamel. At one part of the circumference 
this enamel crust, if such it be, terminates naturally in a very thin edge. The greater 
part has been scaled off, and at the apex it, as well as the ivory below it, has been worn 
away by attrition ; but whether during life or not is uncertain, though, from the obliquity 
and smoothness of the worn apex, the former is by far the more probable. I am not 
aware that the existence of enamel on the permanent incisor of the Elephant has ever 
before been noticed, although, as is well known, the deciduous tusk always has a toler- 
ably thick cap of that substance ; so that its occurrence upon what is undoubtedly the 
permanent incisor, in the present case, is worthy of note. 

3. Lower Jaw. 

1. The collection contains seven or eight fragments of the lower jaw. One of these, 
consisting of the entke symphysis, has been already described as belonging to a mature 
animal of considerable size. A second fragment, apparently belonging to the same 
species, though of younger age, is a portion (about 5 inches long) of the anterior part of 
the right ramus. The whole of the outer table is broken away, and the exposed surface 
is somewhat worn ; but on it may be perceived tlie very shallow remains of the alveolus 
of the 2nd milk-molar, and further back a smooth depression, probably an indication of 



THREE EXTINCT SPECIES OF ELEPHANT. 



277 





Radius. 



Femur. 
Various bones of uterine fcctus of E. africanns. 



9 n 9 



278 JIB. BUSK ON THE REMAINS OF 

the socket of the 3rd milk-molar. From the size of the bone, it undoubtedly belongs to 
the largest of the Maltese Elephants. 

2. A second fragment of the mandible is represented in Fl. LTI. fig. 4-3. It is also a 
portion of the anterior part of the right ramus, which is broken across very obliquely 
from before backwards and downwards, in such a way that the two fractured surfaces are 
parallel with each other, and portions of the upper and of the lower border of the 
ramus are left of about the same length. The upper or, rather, anterior border is very 
acute, and represents a portion of the diasteme immediately anterior to the alveolus of 
the 2nd milk-molar. Close to the edge, on the external aspect, are three openings, of 
which the anterior and largest is the mental foramen, and the others also vascular or 
nervous channels leading into the dental canal, the open orifice of which is seen at the 
hinder end of the fragment, and is about 0"-25 in diameter. 

The alveolus of the 2nd milk-molar is widely exposed by the fracture, and, as usual, 
consists of sockets for two fangs. The anterior fang must have been about 0"-6 in dia- 
meter at the base, and larger than the posterior. Both sockets curve backwards ; but the 
hinder is much more oblique in its direction than the anterior. In size and form the 
alveolus would seem to correspond very exactly with such a tooth as is represented in 
fig. 4, Plate LIII.,the penultimate milk-molar, as I should presume it to be, of JE. meli- 
tensis ; but I am by no means certain of this. The tliickness of the ramus, opposite to 
the point of the posterior fang of the penultimate milk-molar cannot be very satisfac- 
torily determined, but may be estimated at rather more than 1", whilst its height at 
about the same part was probably 2"-5 to 2"'75. 

•3. The three remaining portions of the mandible are of much smaller size, and all 
apparently of uniform character. The most perfect of these is shown in fig. 45, and 
consists of the entire symphysis with the diasteme ; on each side the entire alveolus of 
the 1st milk-molar remains, and on the right an indication of that of the 2nd milk-molar, 
suflficient to show that the anterior fang of that tooth must have been of about the same 
diameter (0"-6) as that in fig. 43 ; so that there may be reason to presume that the jaw 
shown in fig. 45 may represent a younger state of the one shown in fig. 43. 

The portions of the two rami were not, I believe, found in connexion ; but there can be 
little doubt of their belonging to the same individual ; at any rate they correspond 
very exactly. The distance from the anterior alveolus to the extremity of the beak, or 
what remains of it, is about 1"'7, and the height of the ramus in a vertical line immedi- 
ately in front of the socket is l"-25, and its thickness about 0''-7. The small mental 
foramen is quite upon the border of the diasteme ; and on the right side there is only a 
single secondary foramen behind it, whilst on the left there are two. 

4. The remaining two fragments of the mandible are of particular interest, as showing 
what appears to me to be a distinction, apparently of specific importance, in that bone, 
even at a very early period of life, between the two dwarf species. In those specimens 
referrible to one or other of these smaller forms, which have been already described, I 



THEEE EXTIKCT SPECIES OP ELEPHAKT. 279 

have been unable to perceive any gi-eater distinction than can be accounted for by dif- 
ference of age ; but in the two specimens I am now about to refer to, and which appear 
as nearly as possible in the same stage of development, there is a difference in propor- 
tionate thickness which cannot be so accounted for. One of the two specimens is 
figured in PI. LII. figs. 42 and 42 a ; the other has not been figured. At first sight these 
two fragments might be supposed to be the right and left sides of the same sym- 
physis, each being about 2 inches long, and broken off, as would seem to be usual, at 
the part where the ramus is necessarily weakened by the presence of the alveolus of 
the 3rd milk-molar. Closer inspection, however, of the fragments shows that they do 
not correspond as the opposite portions of the same jaw would. 

The left fragment is thicker than the right, measuring at the smallest part behind the 
symphysis 0"-5, and the other 0"-4 ; and the distance between the mental foramen and 
the accessory foramen behind it (single in either case) is, in the left, 0"-3.5, and, in the 
right, 0"'45, though this is perhaps not a very important particular. The symphysial 
facet in the left portion is 1" x 0"-5, and in the left l"x 0"-l. 

The general antero-posterior curve, including that of the diastemic edge, is more 
rounded in the left fragment. And in consequence of this difference, and from tlie 
greater incurvation also, as it were, of the diastemic border, the left fragment, when 
viewed from above, appears much more rounded on the outer face. 

The differences, in fact, between the two fragments are amply sufficient to show not 
only that they cannot have belonged to the same individual, but, in my opinion, to 
indicate an important and, perhaps, a specific distinction, when it is considered that 
the bones are both in the same stage of development. 

4. Bones of the Extremities. 
I have been unable to detect in the collection any bones belonging to the trunk of very 
young animals ; but numerous specimens of various bones belonging to the fore and 
hinder extremities occur, amongst some of which, as it seems to me, significant indications 
of two distinct forms may be perceived. 

(1) Anterior Extremity. 
Of very young humeri we are in possession of three specimens, sufficiently entu'e to 
afford some diagnostic characters ; two of these are figured in PI. LII. figs. 49 & 50. The 
former is the almost enthe shaft of the left humerus ; it is apparently slightly rolled at 
each end, and an angular fragment has been recently broken oft" obliquely at the upper 
end in front. No part of either epiphysial surface is left ; but it is nevertheless clear 
that the specimen represents very nearly the entire length of the interepiphysial shaft. 
With due allowance for the great difference in age, the general characters of this speci- 
men correspond so closely with those of the humerus figured in PI. XLIX. fig. 20, that 
little doubt can be entertained with respect to their belonging to the same species. One 



280 ME. BUSK ON THE REMAINS OF 

character, the value of which has been already adverted to in speaking of the larger 
humerus, appears to me of great importance in this comparison, viz. the angularity on 
the posterior aspect of tlie upper part of the shaft, and the well-marked depression 
on the inner side of the posterior angle, both of which are also well marked in the 
larger humerus, fig. 26. The nutrient foramen, in this little bone, is situated on the 
inner border, about 0"-75 above the inner condyle. As the corresponding part in the 
larger humerus is broken away, the site of the foramen cannot be positively determined ; 
but as it does not exist in any other part of the bone, it could not have been placed very 
far from the same spot as in the foetal bone. I have little hesitation, therefore, in refer- 
ring this humerus, fig. 49, to a very young E. falconeri. The young humerus shown in 
fig. 50 is less entire at the upper end ; but at the lower a considerable part of the epiphy- 
sial surface remains. What is left of the shaft is sufficient to show that in the upper 
part it is more rounded, as in the African Elephant, and that the supinator, or external 
condyloid ridge, is not continued upwards, as it were, into a posterior angle as it is usu- 
ally in the Indian Elephant. So far as can be judged from the imperfect condition of 
the humerus fig. 49 at that part, the inner condyloid ridge is much thicker in the 
antero-posterior direction, and the inner border of the bone consequently more rounded, 
in fig. 50. In front also the anterior surface of the deltoid crest is much more oblique 
than it is in the humerus fig. 26, in which a well-marked angle, prolonged downwards 
from the anterior or external border of the bicijjital groove, bounds internally a perfectly 
fiat surface in front of the deltoid crest. The same angularity, it should be stated, is 
visible, though of course less marked, in the young liumerus of E. falconeri (fig. 49). The 
nutrient foramen is in the same situation as in the former specimen. The humerus 
fig. 50 shows slight marks of gnawing, as by a small rodent, on the hinder surface. 

Another fragment of a much older but still young animal, is a fragment of the lower 
end of the right humerus, broken obliquely through the shaft about 4"-5 above the 
middle of the lower epiphysial surface, a portion of which remains. So far as it admits 
of comparison, its characters accord with those of the humerus of E. falconeri ; and it is 
not unreasonable to suppose that the bone may have formed part of the skeleton of the 
same animal as the femur fig. 29. In it the want of roundness in the internal condy- 
loid ridge behind is well marked. 

The only other young b(jne belonging to the anterior extremity of which there is 
any specimen is the radius, of which bone there are two examples. They are figured in 
PL XLVII. figs. 18 & 19, both belonging to the same side. One is of larger proportions 
than the other, and not improbably, though by no means certainly, of rather more 
advanced age. Both bones are broken across at the corresponding point, which is the 
slenderest part of the shaft, or about three transverse diameters of the lower epiphysial 
surface above that end. At this point the contours of the transverse sections diffier a good 
deal, and show that the shaft of the larger radius (fig. 18) is much more compressed 
than the other, and particularly that the outer or, rather, anterior edge is very 



THESE EXTINCT SPECIES OF ELEPHANT. 



281 



much more acute, leading to the supposition that at the upper end the bone might have 
presented the acute angle which is exhibited in the African and not in the Indian radius. 
The lower epiphysial surfaces again exhibit different contours, as may be seen in the 
Plate, where however, unfortunately, one of the bones is represented on the anterior, 
and the other on the posterior aspect. I have therefore subjoined the outlines of this 
surface taken in corresponding positions of the bones, so as to show at a glance the not 
inconsiderable difference they present. 

42 

Us) 



{19) 





19 




41. Transverse sections of shaft of radius. 



42. Outline of distal epiphysial surfaces. 



From what has been said, it cannot be denied that the same distinction exists between 
the very young dwarf radii as I have attempted to point out in the exoccipital bones 
and humerus. And considering the large size, at what would seem not very different 
ages, and the African tendency faintly exhibited in the radius fig. 18, I should be 
inclined to refer that to the young of E. melitensis. That neither of the small radii 
just described is a foetal bone of a larger form of Elephant, is abundantly sho\vn by their 
dense texture and aspect of greater age, as compared with the far larger radius of the 
uterine foetus of E. nfricanus, of which an outline woodcut (No. 37) is given in p. 277. 

(2) Hinder Exfremity. 

A portion of the shaft of a very young femur of the largest Maltese Elephant has 
been already described and figured ; and I have already noticed the almost entire inter- 
epiphysial shaft of E.falconeri. No specimens of that bone of younger age, corre- 
sponding with the very young humeri, radii, &c. above noticed, occur in the collection ; 
but of the tibia numerous specimens, of various ages, and some very young, exist. Of 
these, however, all it will be worth while here to notice more particularly are those 
represented in PL XLVII. figs. 15, 16, 17, 20, and 21. Of these I regai'd figs. 16 and 17 
as belonging to a different type from that represented in figs. 20 and 21. 

Figs. 16 and 21, each having both epiphysial surfaces almost entire, are pretty nearly 
of the same length, and to all appearance, so far as can be judged from the condition of 
the surface, of about the same age. They admit therefore of tolerably fair comparison. 
In the first place, as the figures will show, the bones differ a good deal in proportionate 
thickness ; stated in numbers the differences in the various dimensions are as under : — 





Length. 


Upper end. 


Lower end. 


Least 
circumference. 


Fig. 16.... 

rig.2i.... 


2-5 
2-4 


1-0 X -7 
•9x-6 


•9 x -7 
■7x-6 


1-6 
1-4 



•282 



ME. BUSK ON THE EEMAINS OP 



The anterior angle of the shaft is more acute in fig. 21, and it is continued down 
towards the inner malleolus in a more pronounced manner. On the posterior aspect 
fig. 21 is much more concave at the upper part ; and the outer posterior angle is con- 
tinued, tolerably distinctly, quite to the lower end, whilst in fig. 16 it is not continued 
below the middle of the shaft. The posterior surface of the bone, consequently, in 
fig. 16 is more evenly rounded on the outer side than it is in the other, as shown in fig. IQb. 
The difference in the contours of both the upper and lower epiphysial surfaces is shown 
in the subjoined outlines, and is, as it appears to me, much greater than can be attributed 




CJz) 





Zl.aj) 



(^ 



to mere individual variety or to difference of age. It may in addition be remarked that 
the nutrient foramen is placed much higher up in figs. 20 and 21 than in the other. 
In all the respects above referred to, except in size, the bone represented in fig. 17 agrees 
with fig. 16, and fig. 20 in like manner with fig. 21. 

If, upon such imperfect data, one might speculate as to the species to which these two 
forms of tibia should be assigned, it would seem most likely that the type sho^vn in 
figs. 16 and 17, from its greater robustness, belongs to -E. melifensis, and the other to 
E. falconeri ; but upon this I refrain from expressing any positive opinion. But, besides 
the five specimens thus disposed of, the collection contains three other equally young, 
if not younger, tibiae, of a totally diff'erent type. 

Two of these tibiae are shown in PI. LI. figs. 38, 38 a, and 39, 39 a ; and their 
peculiar character as distinguished from the others will be at once obvious. The one 
represented in fig. 39 appears to have had a sort of spongy exostosis springing from the 
inner side of the head, which gives it a very bizarre appearance in the back view 
fig. 39 a* ; the smaller specimen, therefore, shown in fig. 38, affords better materials for 

* The artist has inadvertently drawn the inner face of the bone in fig. 39 instead of the proper anterior 
view. 



THREE EXTINCT SPECIES OF ELEPHAXT. 283 

comparison. Both bones are so much worn at eitlier end, apparently by water-rolling, 
that no portion of either epiphysial surface remains ; but tlie portions removed at either 
extremity cannot be very great, so that the length of the epiphysial shaft in fig. 38 may be 
estimated perliaps at about 3"-2 ; on comparing wliich with its other dimensions it will be 
seen that the bone is proportionally much more robust than that provisionally referred 
to E. melitensis. In other respects also it differs so remarkably, not only from that bone, 
but from all other tibice of any age belonging to the Elephant that have come under my 
notice, that I think it impossible to refer these bones to that genus at all. Had tlie 
means existed, which unfortunately they do not in this country, it would have been 
interesting to compare these immature tibiae with those of the irqjpoj'Otamus, to which, 
at a guess, one might be inclined to assign them*. 

§ VI. Dextition. (Plate LIII.) 

The only part of the Zebbug Collection respecting which the late Dr. Falconer has 
left any written observations, beyond a few brief and scattered notes, is that ^vhich com- 
prises the teeth. 

It is well known that tliat distinguished palaeontologist had devoted very great atten- 
tion to the odontography of the Proboscidia, and that he assigned paramount import- 
ance to the study of the teeth in the discrimination of species. It is with the greatest 
satisfaction therefore that, with respect to the dentition of the Maltese fossil forms, I 
find myself in possession of his copious and careful notes, and am thus, on this subject, 
enabled to rely upon his great and undoubted authority. 

Although in some points I have been led to form an opinion apparently differing from 
his, yet, as I feel that all pala:ontologists must desire to have the actual opinions and 
verbal descriptions, as he left them, of my lamented friend, I propose to give all that I 
can find of what he has written concerning the Zebbug fossil teeth in his own words, 
and to reserve to the end, or to notes, the few remarks I may have to offer myself. I 
would also add that the figures in PL LIII. have all been lithographed from Mr. Duikel's 
drawings, which, as they were made under Dr. Falconer's immediate supervision, may 
be taken to convey what he deemed the more important characteristics of the various 
specimens in the Collection. 

" Among the most interesting of the Zebbug fossils is a series of molar teeth and 
fragments of tusks. The molars comprise specimens ranging from the first milk-molar 
of very young animals up to what appear to be adult teeth ; and they are at once charac- 
terized, besides other differential marks, by the singularly small size of the species which 
yielded them. A^'arned by the great blunders committed by Nesti, Fischer de "S^'ald- 
heim, and other palaeontologists, who have been misled by the cliaracters of milk-teeth 
to identify them as the remains of pigmy species of Elephant, I have been chary in 

* With reference to tliis, it should be remembered that a dimiimtive species of Hippopotanms still exists. 
VOL. VI. — PART V. 2 R 



284 MR. BUSK ON THE REMAINS OF 

admitting the convictions which the specimens forwarded by Captain Spratt forced upon 
me when I first examined them. 

" 1. Milk-incisors. — The series fortunately includes a very perfect milk-incisor, which 
confirms the line of specific affinities indicated by tiie molars. The specimen is repre- 
sented of the natural size by figs. 1 & 1 «, 1 5. It differs from the permanent tusk in 
having the crown and fang portions distinctly defined. The crown forms an obtuse, 
flattened, rounded, and irregularly indented body, invested with a thick shell of enamel, 
supported in a long cylindro-conical fiing, part of which is broken off near the end. 
From the diameter of the broken end and of the central canal, it is manifest that at 
least a third of the entire lengtli of the fang is wanting. The specimen was compared 
with the corresponding tooth of a foetal African Elephant, belonging to a skull trans- 
mitted to the British Museum by Dr. Livingstone, in which the milk-molars are quite 
unworn. The two agree very closely in the dilated blunt form of the crown, investing 
shell of enamel, and defined fang. The chief difference detected between them was in 
the form of the latter, which in the young African Elephant forms a rather short and 
compressed cone, terminating in a sharp and slender point, while in the Zebbug fossil 
the fang is stouter, more cylindrical, and much more elongated. The dimensions of the 
specimen are : — 

Entire length l"-4 

Length of crown 0"'6 

Width of crown 0"-4 

Thickness of crown 0"-3 

Diameter at the collar 0"'3 

Diameter at broken end 0"'25 

"These minutice are given, and in such detail, in order to mark the affinity which the 
Malta fossil bears throughout in its dentition to the African Elephant. A shell of enamel 
has not yet, so far as I am aware, been detected on the milk-incisors of any species of 
the subgenus Euelephas. It occurs on those of the African species ; and I have detected 
it forming a sheath upon the young permanent tusks of E. insignis, belonging to the 
group Stegodon*. 

* In the British Museum, besides the fcetal African Elephant referred to by Dr. Falconer, there are nume- 
rous bones of another, very much smaller and obv-iously a very young uterine foetus ; outline figures of some of 
the bones belonging to this specimen have been given in a preceding part of this paper. Among its remains is 
the milk-incisor in the germ state ; that is to say, the fang is still incomplete, and the enamel cap probably very 
thin. The milk-incisor of the older foetus alluded to by Dr. Falconer is completely formed, and presents a dif- 
ferent appearance, owing to the increased thickness of the caj), which appears to be formed of two layers, an 
external (probably of ostcine), and an interior (the enamel). The fang comes to a fine point ; and when the upper 
portion of the cap (which is quite loose) is removed, the ivory nucleus of the crown, of nearly the same size and 
form as the crown of the uterine tooth, is exposed. The entire length of the tooth is l"-8, and the greatest 
diameter of the crown about 0"-4.j, and its length about 0"-6. The tooth, therefore, would seem to correspond 



THREE EXTINCT SPECIES OF ELEPHANT. 285 

" 2. Permanent Incisors. — The collection contains numerous fragments of Elephants' 
tusks, for the most part amorphous pieces or splinters of the outer layers, many of them 
bearing distinct marks of having been gnawed, but indicating tusks of very considerable 
diameter and out of all proportion to the small Zebbug molars. These fragments, which 
appear to indicate another and larger species of fossil Elephant, will be noticed in the 
sequel. 

" There is only one determinable specimen which will admit of being referred to the 
smaller form, and that only conjecturally. It consists of a portion of the distal end of 
a slightly curved tusk, about 5 inches in length. The greater part of the outer layer, 
which is weathered of a greyish tint, has disappeared by dislamination. The but-end 
yields a round section slightly compressed at the sides The outer layer is smooth, and 
throughout a line of thickness shoAvs no appearance of engine-turning. Beneath it the 
ivory surface is very distinctly channelled longitudinally and regularly ; and thus the 
section inwards to the cone exhibits very distinct engine-turning, more pronounced even 
than is commonly seen in proboscidian tusks, the inequalities being nearly as marked as 
in a tailor's thimble. The specimen tapers to a conical point. The dimensions are: — 

Length 5"-0 

But-end l"-15xl"-l 

This tusk would correspond in size with the true molars of the Malta form *. 



very closely, both in size and proportion, with the Maltese fossil. This circumstance may perhaps render it 
doubtful whether the latter really belongs to either of the dwarf Elephants, and may not rather belong to the 
largest extinct form. 

* It is a rather curious circumstance that the specimen above described by Dr. Falconer, and which, from some 
words wliich I have omitted, he seems either to have had or to have intended to have figured, was not to be 
found in Captain Spratt's collection when it came into my hands ; nor is there any figure of it to be found. 
But, strangely enough, another permanent tusk of preciscl}- corresponding dimensions, and with a ticket upon it in 
Dr. Falconer's handwriting, " Ehiihas melitensis, tusk, Zebbug Cave," is in the collection ; it is the one figured 
in PI. LII. fig. 48. The specimen, as wiU be seen, is far more perfect than that described by Ur. Falconer ; 
but in dimensions and all other characters (excepting colour, which is mottled with brown and black instead of 
being " grey ") the two so fully accord that we might almost suppose that the missing specimen and the one ex- 
tant in tlie collection may have belonged to the same individual. It is true that the specinu'n figured in PL LII. 
has been broken across ; but the fracture, instead of five, is more than seven inches from the truncated extremity, 
and probably fuUy eight inches from the entire conical point, which is stated to have existed in the missing 
specimen. And it should be noted that the fractured surface at the apex is not a recent one ; so that the speci- 
men cannot be the one described by Dr. Falconer with the point subse<iuently broken oft'. In the ]ircseuee, 
therefore, of this more complete specimen of an obviously similar tusk, the loss of the one described by Dr. Falconer 
will be the less felt. In the existing specimen the outer layers, as will be seen in the figure, are detached 
towards the point, exposing a subjacent surface vcrj- strongly sulcate ; and at the fractured end the coarse 
engine-turning described by Dr. Falconer is plainly visible. The diameters of the tusk, at the distance of 5 inches 
from the estimated real point, are 1"-I x 1", or very nearly the same as those given by Dr. Falconer, whilst an 
inch or so lower down they exactly correspond, viz. 1"-15 x 1"-1 ; and this is the greatest diameter down to the 
alveolar end, the maximum circumference being 3"-5. The extreme length of the specimen measured along the 
outside curve is about 10"-.5, to which may be added, to complete the point in its natural state, about another 

2 r2 



286 ME. BUSK ON THE REMAINS OE 

" 3. Lower Milk-molars. — Fig. 2 and fig. 2 a, represent, of the natural size, the ante- 
penultimate milk-molar (m.-m. 2) of a very young animal, and, I believe, the smallest 
Elephantine molar, fossil or recent, that has hitherto been met with, figured, or described. 
The outline of the crown is broad oval, being narrow in front and wide behind. It is 
composed of three collines, with a posterior talon. It is clear that the tooth did not 
belong to a fcctal individual, as the crown exhibits the most distinct proof of having 
been in use, and worn against an opposed tooth. Further, the posterior talon bears 
a well-marked disk of pressure, from the contact of an advancing tooth behind it. The 
disks of wear of the crown surface are broad, but not much advanced in wear. One 
large elongated and conical fang only remains, connected with the anterior and middle 
portions ; but the base of the tooth shows a doubtful appearance of there having been 
a small fang below the posteidor talon. The dimensions are : — 

in. 

Extreme length 0-40 

"Width of crown in front 0'23 

Greatest width 0-32 

Greatest height of crown 0'40*. 

inch, making tlic entire loiigtU of the tnsk between 10"-5 and V2", whilst tlie depth of the pulp-cavity is not mure 
than 2"-7 in the present state of the sjjccimen ; and as this is rcDiarkahly i)erfeet even at the thin alveolar edge, it 
probably could not have exceeded 2"'9, or 3". As this is rather less, I believe, than the usual proportion of the 
depth of the pulp-cavity to the length of the tusk as ivorj- is brought to the market (i), it probably indicates that 
the tooth was of considerable age and consequently belonged to a mature animal. Dr. Ealconcr further states 
that the specimen described by him was the only instance of the permanent tusk in the collection ; but in this, be- 
sides that I have just noticed, I find two other fragments of what I conceive to be very young permanent incisors. 
One of these, or rather a portion of one of these, is seen attached to theprcmaxillar}- bone represented in fig. 4G, 
to which reference has been already made. The other is the basal portion of another young tusk, of exactly 
the same diameter, about 2 inches long ; the outer end is broken obliquely off a short way beyond that part of 
the tooth which, to judge from the colour, was lodged in the alveolus, whilst the other was exposed to some 
reagent which has given it a lirowii colour. The remaining depth of the pulp-cavity is C'S) ; and when the tooth 
was perfect it was probably 1" or rather less; so that the entire original length of the tusk may be estimated 
at about 3 inches. Its circumference is l"-3, and greatest diameter 0"-4. At first sight it does not appear 
altogether impossible that the two fragments of the small t\isk may be parts of one and the same; but, in the 
iirst place, the fracturi'd surfaces do not fit, nor is the colour of the interior the same in both fragments ; and if 
the two were placed together even without any intermediate missing portion, the tusk would be too long in pro- 
portion, as it seems to me, for the prcmaxillaiy bone, and would project from the alveolus much more, in pro- 
portion to its thickness, than the young tusk of an elephant does. Though no appearance of engine-turned 
marking can be discerned in either fragment, the external longitudinal striation, where dislamination of the 
outer layer has taken place, is as coarse as it is in the larger tusk. I have already mentioned that Dr. Falconer 
had cemented one of these small tusks into what remains of the alveolar cavity in the premaxillary bone; but 
on close examination I find that the basal portion fits much more closely ; and I have therefore substituted it in 
the specimen for the other. 

* Tlie dimensions above given are not quite half those of the corresponding tooth in E. africaniis, and as nearly 
as ])ossible half those of the second mUk-molar in E. indlcus, E.primii/enius, and E. antiqma ; so that, admitting 
it to be reaUy the second and not the true first milk-molar, it is obvious that the Maltese specimen must have 



THREE EXTINCT SPECIES OF ELEPHANT. 287 

"Do Blainville (' Osteographie,' Elephans, pi. ix. fig. 1) has given a figure of a lower 
jaw of a very young African Elephant, in whicli a pre-nntepenultimate or theoretical 
first milk-molar was developed on one side of the lower jaw ; and in the ' Fauna Antiqua 
Sivalensis ' another example of the same kind is also figured*. The milk-tooth in botli 
these cases was very rudimentary ; and it is possible that the Zebbug specimen might be 
conjectured to be an equivalent tooth. But it appears to me that this view is distinctly 
negatived by the fact that the Zebbug milk-molar was supported upon a large fang, and 
that its crown is well worn, proving that it had served an alimentary function, and that 
it was not a case of unusual or monstrous development of a theoretical tooth which is 
commonly suppressed. In the instances of the African Elephant above referred to, the 
pre-antepcnultimatc milk-molar was restricted to one side of the lower jaw, and was not 
developed in the upper jaw. It is difficult to say of the Malta tooth whether it 
belonged to the upper or lower jaw. 

" Fig. 3 of the same plate represents the portion of the crovwi borne upon the large 
anterior fang of a milk-molar. It is composed of three distinct disks of wear, which 
are very open, resembling in this respect the characters yielded by fig. 2 ; indeed they 
are as much expanded as in the existing African Elephant. The crown is narrow in 
front, and widens very rapidly backwards, the dimensions being : — 

in. 

Width in front (of anterior ridge) 0" 3 

Greatest width behind O'b 

Length of crown-fragment (of three front disks) 0'54 

" The anterior end of the fragment bears halfway up a distinct smooth pit, being the 
disk of pressure against an anterior tooth that had been in contact with it. The enamel 
plates surrounding the worn disks show no marks of crimping. It is not possible to say 
what was the precise number of ridges entering into the composition of the cro\\ii of 
this tooth ; but judging from a germ specimen, to be described in the sequel, it con- 
belonged to a dimiuutiTe species. It is a eurious circumstance, however, and one well worthy of noto with 
respect to this tooth, that its fangs must have differed widely from those of the second milk-molar in all other 
known instances, in which they are suhequal in size and strongly divergent. Dr. Falconer states that there is 
some indicatiou of the existence of a distinct small anterior fang — though I am myself by no means satisfied of 
this, but on the contraiw conceive that the existing fang, as shown in the figure, is in fact composed of two con- 
nate ones. In any case it is obvious that, even had an anterior fang existed, it must have been very much 
smaller than the posterior ; and it is equally clear, from the direction of the remaining fang, that they were not 
divergent. Another circumstance, however, goes strongly to show that the existing fang is really a double one. 
In the foetal mandible, represented in fig. 45, the alveoli of a small tooth immediately in front of the third milk- 
molar remain ; and of one of these I have taken a wax cast of the interior, which shows that the fangs of the 
tooth occupying it were also connate and non-divergent. From this circumstance, if confirmed by further 
instances, it would seem probable, either that the true second milk-molar, in at least one of the pigmy Elephants, 
had connate, non-divergent fangs, or (what is perhaps equally probable) that that tooth was normally suppressed 
and replaced by a functionally developed first milk-molar. 

* PL xiv. fig. 4, left .side, a. 



288 ME. BUSK ON THE EBMAINS Of 

sisted of five lidges with front and back talons. From the very narrow width in front, 
and the rapid increase backwards, it is mfeiTed that it was a lower milk-molar, and 
probably the penultimate (m.-m. .3). 

" Fig. 4 represents the crown and side aspects of a beautifully preserved specimen, com- 
prising nearly the entire length of the crown, of an inferior milk-molar, left side. The 
crown presents the disks, well worn, of iive ridges with a small posterior talon. The 
disks are wide and open in the antcro-posterior direction, and somewhat rhomboidal 
in outline, as in the African Elephant, and they bear a close general resemblance to 
those of figs. 2 and 3. A large fang supportmg the last three ridges and talon is pre- 
sent, nearly entire ; but the fi-ont fang is broken off, together with a small portion of the 
anterior talon. The fractured surface from which the fang had been broken off is 
distinctly marked below, and shows that the crown is all but complete in length. The 
anterior part of the crown appears to have been worn down close to the level of the 
fiing. The giinding-plane is slightly concave in the antero-postei-ior direction, proving 
it to have been an inferior molar ; and I infer that it is the equiv alent too th of the 
specimen last described, i. e. the left lower penultimate milk-molar (m.-m. 3), and that 
when entire it was composed of five ridges with front and back talons. The dimensions 

are : — 

in. 

Extreme length of fragment 1"30 

Width of crown at front ridge 0-57 

Greatest width of front ridge OwO 

" The enamel plates in this, as in the two other specimens above described, are very 
thin, with no tendency to crimping, the appearance which looks like this being simply 
the vertical gi-ooves in contact with the cement. It is important to add that there is a 
broad and well-defined smooth depression upon the posterior end, indicating the pressure 
of a contiguous tooth bearing against it from behind. 

" Fig. 5 represents, of the natural size, the top and side aspects of a finely preserved 
milk-molar of the same series, inferred to be the last of the lower jaw, right side 
(m.-m. 4). Its crown surface is concave from back to front, proving it to be lower; 
and the oblique direction of the disks of wear determines the side. With the exception 
of a little damage to the anterior end, which has removed a portion of the front talon, 
the crown is quite perfect ; and the whole of the fangs are also present, more or less 
fractured. The crown was composed of eight ridges, all of which have been affected by 
wear. The disks bear the closest resemblance in form to those of fig. 4 ; and it will be 
seen by a comparison of the figures, that they were in a nearly corresponding condi- 
tion of wear. In three of the intermediate disks (viz. 4, 5, and 6) there is a slight 
tendency to an angular rhomboidal expansion in the middle ; but, as in the teeth above 
described, the enamel plates are very thin, and the edges in contact with the ivory- 
depressions are straight and perfectly free from any tendency to plication or crimping. 



THREE EXTINCT SPECIES OP ELEPHANT. 289 

There is a small talon process appended to the last ridge, enveloped by cement. The 
crown is worn low in front, and differs from those of the preceding teeth in maintaining 
nearly a uniform width throughout, the others being nan-ow in front, and widening 
suddenly backwards. The front fang is thick and massive, supporting two or three 
ridges ; between it and the large back fang there are the remains of a series of smaller 
fangs, more or less confluent with the latter, presenting characters widely different (in 
the gi-cater amount of complexity) from those yielded by fig. 4, and indicating, in 
harmony with other points, that they were not teeth of equivalent age. A part only of 

the anterior talon remains. The dimensions are : — 

in. 

Length of crown 22 

Width hi front 0-7 

AVidth in the middle Ow 

Width at eighth ridge 07 

Height of crown at the last ridge 0-8 

•' So far as I am aware, no milk-molar of an Elephant, fossil or recent, has hitherto 
been observed with so complex a crown, conjomed with such small dimensions. 

" 4. Upper Milk-molars. — Of upper milk-molars the series is less complete than of 
lower. Of the antepenultimate (m.-m. 2) or, as commonly called, first milk-molar, there 
is no determinable specimen. But of the penultimate (m.-m. 3 ) a very beautifully pre- 
served germ is represented by fig. (i of the same Plate, top and side aspects. It consists 
of the entire shell, before the ivory nuclei had become ossified, and without fangs ; the 
layer of cement had not been completely formed, and is denuded from the sides. The 
crown is of an oblong form, a little wider behind than in front, and is composed of five 
principal ridges, with a chstinct talon m front and behind. The tips of the digitations 
of the first ridge and talon are alone affected by wear, and that only to a slight degree. 
Taking into account the difference of upper and lower teeth in form, and the difference 
in the stage of wear, it agrees closely in size and proportions with the mferior penulti- 
mate shown ui fig. 4 and already described. The ridges are seen to be separated by 
rather wide intervals, and they are high relatively to the other proportions. The 
digital terminations of the ridges speedily become confluent below the apex, this con- 
stituting the principal cause of the absence of crimping in the enamel plates, noticed in 
the description of the lower milk-molars The dimensions are : — 

in. 

Length of crown 1'4 

Width in front 0-6 

Greatest width behind 0'8 

Height (greatest) of ridge plates O^Oo 

The tooth is of the left side. There are m the collection a number of detached plates 
of an unconsolidated germ tooth of the same age. 



290 



MR. BUSK ON THE EBMAINS OF 



" Of the last upper milk-raolar ( m.-m. 4) there are numerous fragments, but unluckily 
no entire tooth, in the collection. The most perfect of the series is the specimen re- 
presented by figs. 8 and 8 «, which consists of the intermediate portion, comprising six 
ridges, but mutilated both in front and behmd. It is e^ddent that of the hind portion 
only one or t\\'o coUines are wanting, these beuig the last ridge and posterior talon ; 
and as regards the anterior end, the fractured section is seen to pass vertically through 
the middle of the large front fang, indicating that the front ridge and talon alone are 
there wanting. The tooth, when entire, must have been nearly in a germ-state, as the 
tips alone of the front remaining ridge are aifected by wear. The height falls off very 
rapidly from the front backwards; and the ridges are high in proportion to the width. 
This tooth is inferred to have been the upper milk-molar corresponding ui age with 
fig. 5 of the lower jaw. The dimensions are : — 

in. 

Length of fragment of crown 1"4 

Width in front 0-95 

AVidth behind 0-85 

Extreme height of crown-ridges ' 2-3 

There are no means of determining with certamty what was the precise number of 
ridges that entered into the composition of this tooth*; but assuming from the data 

* Besides the teeth uoticed by Dr. Falconer, the collection contains an entire upper molar which appears to 
correspond so closely in dimensions with the fragment above described, and represented in fig. 8, as to leave no 
doubt in my mind of its being a corresponding tooth in the series, whatever its place may be. So far, also, 
as can be judged from the little-worn condition of the macJuerides, and from the thickness of the plates, it would 
seem to belong to the same type as fig. 9, from which tooth, however, it differs most remarkably in the height of 
the crown, though nearly corresponding in aU other dimensions. As the specimen is one of great interest, and 
in nearly perfect condition, I have thought it might be useful to add figures showing its main characteristics. 





The tooth is clearly an upper molar of the right side ; but whether it is to be regarded as a mUk- or as a 
true molar, opinions may be diWded. 



THREE EXTINCT SPECIES OF ELEPHANT. 291 

given ubove that only a single ridge and talon have been broken off at cither end, the 
perfect tooth would have presented eight ridges, besides talons. 

" Figs. 7 and 7 a represent, of the natural size, another fragment of a germ of the 
same tooth, comprising four of the middle and posterior ridges. It was proportionally 
smaller than fig. S ; but the form and size are irreconcilable with fig. 6 ; it is therefore 
mferred to be a part of m.-m. 4 ; it presents no .special characters for a description. 

" 5. True Molars. — The evidence above adduced from vari(jus instances of milk-molars 
jointly goes to prove the former existence in Malta of a small form of Elephant : this 
inference is fully corroborated by the remains of true molars ; and first, as regards tho.se 
of the lower jaw : — 

" Lower True Molars. — Fig. 12 represents, of the natural size, a specimen comprising 
the greater part of a lower molar of the right side. The anterior part of the crown. 

The erowu part is entire, excepting a small portion of the anterior talon, which has been recently chipped off; 
it measures 2"-9 in length by 1"-1 in extreme width, which is at the second plate; the greatest height (at the 
7th plate) is 1"'S5. The length of the grinding-surface is about 2" ; and on it are exposed the machcerides of 
part of the front talon and of six plates, together with the extreme point of a single median cusp of the seventh. 
Only two of the plates, however, are worn into complete rings ; and the sixth presents no less than seven minute 
annuli crowded into a space of 0"'5. The tooth is composed of eight plates and an anterior and posterior talon, 
i. e. of ten elements. The hinder end is hollowed and flattened below the talon ; but there is not the slightest 
indication of jiressure by a succeeding tooth, cither in this tooth, or in that shown in fig. 9. The average thick- 
ness of the plates is about 0"'27; and, as far as can be seen in the few spots where the cementum has been removed 
(apparently by attrition), the surftice of the enamel is finely and irregularly fluted, and in some places, though 
very rarely, an extremely faint indication of transverse vninkling is exhibited, but by no means so clearly as in 
the teeth shown in figs. 7, 8, 9. 

With respect to the position of this tooth in the series it is not very easy to arrive at anj- satisfactoi-y deci- 
sion. As I have said, it seems to resemble so closely in all respects that represented in fig. 8, that the two 
may, I think, be safely regarded as corresponding teeth, diSering only in the degree of wear they have undergone. 
The latter tooth is regarded by Dr. Falconer as the fourth upper milk-molar, and as representing an upper tooth 
corresponding to the lower milk-molar, fig. 5. From its general characters, as regards its form and the thick- 
ness of the jjlates, as shown more clearly in the entire tooth than in the fragment figiired by Dr. Falconer, it will 
also be seen to bear a strong resemblance to fig. 9, which tooth Dr. Falconer appears to have been inclined to 
regard as the second true molar, though not certain that it might not be the first, which I think is ecpiiilly (if 
not more) likely. 

■\Ve have to consider therefore whether the tooth described in this note, is m-ni. 4, or m. 1. 

First, with respect to its being m-m. 4 of the same species as m. 1, or m. 2, fig. 9. 

If we compare the relative lengths of the m-m. 4 and m. 1 in E. imticus, they will be found on the average, 
as regards length, to measure about 5"-l and 6"'6 respectively, or to stand in relation to each other as -'T'J to 
1-000 ; in E. primiyemus, 3"-6 and 5"-2, or as -692 to 1-000 ; in E. antiquus 5"-3 and G"-7, or as -791 to 1-000 : 
and in E. merldiomdis 4"-6 and 6"-2, or as -741 to 1-000; whilst the relative lengths of the tooth we are dis- 
cussing and that shown in fig. 9 are 2"-9 and 3"-0, or nearly identical; and in fact, when it is remarked that 
fig. 9 has an additional plate ((/), the other is actually quite as long, If not the longer. Again, if we take as the 
term of comparison the thickness of the plates, it will be found to be nearly identical in the two teeth ; whilst 
if we take the relative thickness of the plates in m-m. 4 and m. 1 in the above-named species of Elephant, a very 
con3ideral)le difference will be found to exist. For instance in E. indicus, the last milk-molar plates are about 



VOL. VI.' 



-PART V. 2 S 



292 MR. BUSK ON THE EBMAINS OF 

supported upon tlie large front fang, has been ground down, and removed by advanced 
wear: the remaining portion is complete back to the hind talon. The fragment 
presents all the characters of the residuary part of a last true molar, more especially in 
the very significant circumstance that the last ridges become gradually less and less 
vertical, and diverge in a fan-shaped fashion until the hindmost become nearly hori- 
zontal. This portion of the tooth is completely enveloped in well-preserved cement, 
and there is not the slightest indication of a disk of pressure caused by a hinder tooth 
pushing it forwards. This is the typical form which the last true molar (m. o) com- 
monly assumes m the existing Indian Elephant and in fossil species of the same sub- 
genus Euelephas. It is irreconcilable with that of a last milk-molar, or of either of the 
intermediate true molars, as it would necessarily imply the absence of any other 



0"-44, and those of the first true molar 0"-.54 in thickness, — in E. antirpius 0"-52 and 0"-0, in E. iirimi(jenms 
0"-33 and 0"-43, and in a single instance of E. mcridionalis 0"-5G and 0"-68, — showing that, as a rule, the plates 
of the first molar are considerably thicker than they are in the last milk-molar. I have no direct measurement 
of the relative thickness of the plates in these teeth of E. afi-kanus, but should estimate the difference as much 
greater even than in any of the species above mentioned, or as about 0"-6.5 and 0"-85. In both these particulars, 
therefore, of the length of the tooth and the thickness of the plates, the tooth under discussion, were it the 
ra-m. 4 of the same species as No. 9, would have proportions widely different from those which that tooth 
possesses in aU other known species of Elephant. 

In the same way, if we assume with Dr. Falconer that the tooth represents an upper m-m. 4 corresponding 
with such a m-m. 4 as fig. 5, we are met with similar objections. For upon comparing the relative dimensions 
of the upper and lower m-m. 4 in other species of Elephant, the latter wiU, I believe, in all, or at any rate in 
most cases, be found to be the longer and generally the narrower, though the difference in the latter respect is 
never anything like as great as it is between the subject of this note and the tooth shown in fig. 5, the one being 
1"-1, the other only 0"-7, in \Nadth. The former, again, is 2''-9 long and the latter 2"-2, showing an ecjually 
great difference, but in a direction exactly opposite to what it should be did the teeth stand to each other in the 
relation of upper and lower. But as the tooth fig. .5 is clearly a m-m. 4, it follows that the other cannot be 
an m-m. 4. 

Of course, if the tooth fig. 9 be regarded as m. 2, the above arguments against the smaller one being m-m. 4, 
are veiy much strengthened. 

Tlie second question to be discussed is whether, supposing fig. 9 to represent an m-m. 2 , the tooth figured in 
this note represents the m. 1 of the same species, or of one of the same size. This point I think may be briefly 
decided by a reference to the comparative dimensions of m. 1 and m. 2 in other species of Elephant. 

In E. indims the average length of these teeth may be stated as about G"-8 and 8"-8, in E. africanus as G"-4 
and 7"-3, in E. antiquns as 6"-" and S"-3, in E. primujenins as about 5"-3 and 8"-2, and in E. merifHunalts as 
(i"-5 and 8"-S, — showing that in round numbers the second molar, as regards length, stands in relation to the 
first pretty nearly as 63 to 83. 

But the tooth I am describing and that shown in fig. 9 are of nearly the same length ; so that here again, 
unless we assume the existence of entirely different proportions between the length of the teeth in the dwarf 
Elephant and that of those of all other species, we are compelled to the conclusion that the teeth in question both 
occupy the same place in the scries — that is, arc both either the m. 1, or m. 2 ; but which, it is not very easy to 
say ; and on the supposition that they are so, the very great difference between them in the height of the crown 
is vcrv remarkable. 



THREE EXTINCT SPECIES OF ELEPHANT. 293 

successional tooth advancing behind it. The outline is bowed very considerably side- 
ways, being convex upon the inner side, and concave on the outer, to a greater degi-ee 
even than is indicated by the figures. The residuaiy worn surface of the crown 
exhibits seven abraded ridges, of which tlie four anterior disks show rather wide and 
open depressions, with a defined angular expansion in the middle. The plates of 
enamel are rather thick and uniformly straight, presenting not the slightest degree of 
crimping or plicature, with the exception of the sharply angular, little, mesial expansion. 
The three next hinder ridges are but slightly abraded. The anterior disks are very 
oblique in their direction, which runs from the inside outwards and backwards. The 
unworn hind portion is so completely enveloped by cement, and the plates so nearly 
horizontal, that it is hardly possible to reckon exactly the number of ridges composing 
this part of the crown. But approximately the fragment is estimated to have in all 
nine ridges, with a small posterior talon. There are no means of determining exactly 
how many ridges have disappeared in front, as all remains of the great front fang are 
wanting. The width of the cro\Aai gradually diminishes back to the talon, as normally 
occurs with the last true molar of the Elephants. The dimensions are : — 



in. 



Extreme length of fragment 4'2 

Width of crown at second ridge 1-3 

^\'idth of crown at seventh ridge 1-2 

Width of crown behind 0*85 

Space occupied by second, third, and fourth disks IT 

Length of residuary grinding-siu'face, including seven ridges 2'3 

Extreme height of crown near middle, where unworn 2T 

The fangs are broken off along the base near the roots ; the contour of this part of the 
tooth when reversed is very mucli and nearly uniformly curved, like a bow on the 
stretch. 

"The specimen next to be noticed confirms the inference drawn from fig. 12. It 
consists of the posterior half of a lower molar, right side, including six ridges and the 
greater part of the posterior talon. It is represented by fig. 13. Like fig. 12, what 
remains of the fragment is concave on the outside, and convex inwards in the longi- 
tudinal direction. The three anterior ridges are worn ; but the grinding-surface is -serj- 
distorted, descending nearly vertically in a spoon-shaped conca^dty from the outside 
inwards and downwards: this peculiarity had evidently been caused by the crown 
having been opposed to an abnormally developed or diseased upper molar. The cUs- 
tortion is attempted to be shown by the contrasted shades of fig. 13, the dark tint 
showing the higher side. But the amount is best expressed by measurements, — 

ill. 

The height of the crown at the outer side of the distorted portion being 2-9 

And of the imier 195 

2 s 2 



294 MR. BUSK ON THE REMAINS OF 

making a difference of nearly an inch. A series of discoloured bands upon the surface 
of the cement of the outer side near the top, disposed concentrically, have been caused 
by this distorted wear. The three hindmost ridges are intact, and, instead of being 
vertical, they are in a certain measure retrofracted, the convexity being directed 
forwards — a character which is commonly presented by the hindmost of the true molars 
in the Elephants. The posterior talon appears to have consisted of a single flattened 
digitation, the back plate of which has been removed by a fracture. This circumstance 
lias deprived us of any direct evidence as to the presence or absence of a disk of 
pressure behind ; but the characters presented by the specimen, regarded in the aggre- 
gate, are consistent only with determining it to have been a penultimate or last true 
molar, probably the latter. The dimensions are : — 

in. 

Extreme length of fragment 3'3 

Width at anterior plate 1".S5 

Width behind 1-2 

Extreme height of crovm 2'9 

These dimensions are comparatively larger than those yielded by fig. 12; but the 
difference is not greater than may be fairly attributed to distinct individuals, or 
difference of sex. The distortion in the grinding-plane, noticed above, is rarely, if 
ever, seen in molars of the milk or adolescent stage, but is occasionally met with, 
variously modified, in teeth belonging to the period of old age. 

" A still more perfect specimen is represented, top and side aspects, by figs. 11 and 11 «, 
which is inferred to be another example of a last lower molar of the left side, in a 
diffei'ent stage of wear from the preceding two. What remains of the crown consists of 
ten ridges and a posterior talon. All of these are more or less affected by wear. The 
seven anterior ridges had been ground down into transverse disks, of which the two 
first are confluent into a uniform surface, from which all trace of enamel has dis- 
appeared ; and they are confluent also by a narrow isthmus with the third disk. The 
disks correspond in form exactly with those of the teeth already described, being broad 
in the antero-posterior du'ection, vrith a slight tendency to sharply angular expansion in 
the middle, which is more or less developed upon the fourth, fifth, and sixth ridges. 
The enamel edges in contact with the ivory depressions of the disks are straight, and 
entirely free from any tendency to plication or crimping, this being clearly a distinctive 
character of the species. The ridges, besides their considerable breadth, are separated 
by rather wde intervals of cement: this is well shown on the side aspect of fig. 11 «. 
The posterior talon forms a prominent splent, consisting of about a couple of digita- 
tions. All the fimgs have been removed by fracture close to the base. The crown 
surface ui front has been ground down to the level of the fangs ; and there is no trace 
rcniuining of the large anterior fang, or of the portion of the crown supported by it, 
which must have borne at least two additional ridges, which would give a total of 



THREE EXTINCT SPECIES OF ELEPHAJ^T. 295 

twelve or tlmtecn to the entire length of crown. The crown surface bears a very con- 
siderable degree of resemblance to that of the existing African Elephant, and greater 
probably than tliat presented by any other species, except certain varieties of Elqihas 
antiqnus. TJie dimensions are : — 

in. 
Length of crown '. 4-4 

Width in front at fourth ridge 1-4 

Greatest width of fourth ridge 1-4.5 

Width at eighth ridge 1-2 

Width at last ridge 1-1 

Height of crown at seventh ridge 2-05 

'• There is a certain amount of retrofraction in the vertical direction of the last ridges, 
but less considerable than in fig. 13, and nothing approaching the fan-shaped divergence 
and horizontality of the hinder ridges of fig. ] 2. There is not the slightest indication 
of a disk of pressure anyivhere upon the liind talon, the surface of which is perfectly 
preserved with its coating of cement from the tang upwards. This circumstance is of 
important significance in determining the tooth to have been the last true molar, as, 
considering its advanced stage of detrition, it is difficult to conceive that it could have 
been followed by an older tooth driving it forwards, without leaving the usual mark of 
pressure. It is further clear that the three specimens last described must have belonged 
to distinct individuals. 

" UpiJer true molars. — The specimen represented in fig. 9 is a very finely preserved 
molar of the upper jaw, right side, complete in every respect, with the exception of the 
ends of the fangs, which are more or less broken. The crown is composed of nine prin- 
cipal ridges, together with a front and back talon. The anterior fang is distinctly present 
and supports the two front ridges and talon. The front talon, or what remains of it, 
appears to be composed of four minute digitations, the greater part of it having been 
ground away*. The three anterior disks are transverse; the next five are only sliglitly 
worn, showing the tips of the digitations abraded into annular detached islands, or in 
three divisions. The anterior disk is expanded in the middle, and narrows at either side, 
presentmg only two or three flexures in the enamel plate, without any crimping. The 
second and third are of nearly similar form with uncrimped enamel and narrowing at 
the sides. The fourth, fifth, and sixth are each in three divisions ; and the seventh and 
eighth only show the tips of the digitations worn across. The enamel plates are deci- 
dedly thick for the size of the tooth f, and the ridges are very high relatively to the 
length. The layer of cement at the anterior end has been removed, and with it all 
appearance of a disk of pressure. The hind talon forms a gibbous projection beyond 

* The enamel cannot be said to be uncrimped, as it is decidedly crimped on the hinder edge of the anterior, 
and on both the anterior and posterior edges of the two succeeding machfridcs. 
t They are not quite so thick as they are represented in the tigure. 



296 MK. BUSK ON THE EEMAINS OF 

the vertical plane of the posterior fang. There is no distinct disk of pressure upon the 
crown portion of the talon ; but there is an obscure depression at the basal part, near the 
fang, which may be of this nature*. The following are the dimensions : — 

in. 

Extreme length of crown . . 2-9 

Width in front 1-35 (2nd plate) 

A^'idth in the middle 1 "3 

Width behind I'l 

Length of surface occupied by the eight anterior disks in wear 2-2 

Extreme height of crown at unworn portion, 9th ridge . . . 2-8 

" From the above dimensions, the contraction of the crown posteriorly and the consi- 
derable height of the ridges relatively to its length are well shown. 

" Had this specimen been discovered isolated, little or no hesitation would have been 
entertained by the palaeontologist in referring it to the age of a milk-molar of some 
species of Elephant. But when regarded as part of a series in connexion with the un- 
doubted milk-molars figs. 2-6 inclusive, and more especially with fig. 8, the whole of 
which are of such unusually small proportions, and when further compared with the 
adult molars of the lower jaw (figs. 11, 12, and 13) and an upper molar belonging to 
the Public Libraiy at Malta, it is manifest that it maintains its place consistently as a 
true molar of the same series. I am at present unable to decide with confidence 
whether it had best be regarded as an antepenultimate or penultimate f. 

" Of the antepenultimate upper true molar (m. 1) no perfect specimen is to be foimd in 
the collection. One fragment, inferred to be a part of this tooth from its size, form, and 
proportions, comprises the two anterior ridges, together with the large fong that supports 
them. The coiTesponding molar of the lower jaw is equally wanting as an entire speci- 
men ; but there are fragments referrible to it also." 

With respect to the Maltese tooth in question Dr. Falconer remarks: — 
" One of the most characteristic of the specimens is an upper molar of the left side 
bearing the following label : 

" ' Dente che si conserva nella pubblica Biblioteca di Malta e trovato in Novembre 
1859 in Malta.' 

" The tooth is a well-worn upper molar of the left side, perfect so far as the crown goes, 
with the exception of the front portion supported upon the large anterior fang, which 
portion had been worn away by continued grinding action. This is distinctly pro\ed by 
the circumstance that the gi-inding-plane of the crown intersects the most anterior of the 
extant fangs. The rest of the fongs from this point backwards to the posterior talon are 

* There can, I think, be no doubt that the deep hollow below the hind tabm is due to the pressure of the 
succeeding capsule. 

t There can be little doubt as to the true position of this tooth as ni. 1. In form and general character it is 
the exact counterpart, except in size, of an m. 1 of E. anticiuus—v!tf:ni\&<:(\. as such by Dr. Falconer. 



THEEE EXTmCT SPECIES OF ELEPHANT. 297 

all present, but more or less fractured or abraded. The molar is vertically fractured 
across through the middle, invohdng the loss of the greater part of one coUine ; but as 
the fragments fit at the base, this circumstance does not interfere with the precise appre- 
ciation of tlie crown-characters. What remains of the crown is composed of ten ridges, 
of which nine are more or less worn, the rest being intact. The posterior talon consists 
of a single flattened gibbous digitation appended to the last ridge, which is composed 
of three or four digitations. The most anterior disk of wear is vertically divided through 
the middle, so that its posterior half only is present. The seven anterior disks form 
oblong transverse depressions bounded hy parallel bands of enamel, there not being the 
slightest tendency in any of them to digital subdivisions forming secondary undulations. 
These disks are of nearly uniform width across, parallel and without any indication of 
the retroflected comua at the sides, such as are commonly seen in Elejjhas anfirjuus. 
Their most striking character is the nearly entire absence of anything approaching 
crimping (or primary undulations) upon the edges of the enamel plates, as they appear 
in relief on the surflvce of the crown. There is a slight appearance of vertical grooving 
upon the cement aspect in these enamel plates, but considerably less than is exhibited 
in the molars of any species of Elephant, fossil or recent, with wliich I am acquainted. 
The enamel of the plates is rather thick, quite as thick in proportion as in the existing 
Indian Elephant or E. antiquus. There is the slightest possible tendency to mesial 
angular expansion in some of the anterior disks, but it is barely appreciable, while in 
some other of the specimens this character is somewhat more pronounced. The talon 
consists of but a single flattened digitation ; and there is this remarkable circumstance 
about it, that it nowhere bears the slightest indication of any disk of pressure upon it 
arising from the protrusion of another molar advancing from behind. The last or tenth 
ridge of the specimen I have reckoned as such, and not as a talon appendage, from 
the fact that it is continued vertically down into the large posterior fang and distinctly 
within its bearing. The crown, in proportion to the height of the plates, is narrow. 
The disks of wear, where much abraded in front, are in close contact, the enamel plates 
nearly touching each other ; but they are well separated in the hinder part of the tooth, 
and the whole of the crown is enveloped by a coat of cement, which, at the sides, is seen 
to be of considerable thickness. 

" I have reckoned that what remains of the crovm is composed of ten ridges ; and, 
taking into account that the most anterior portion, supported upon the large front fano', 
had disappeared by age, and that it was probably composed of at least two ridges, this 
would yield for the ridge-formula of the molar a total of twelve coUines, exclusive of 
talons. 

" What was the age of this molar in the dental series of the animal ■? At the first 
glance it might be supposed from its size to be a third or last milk-molar ; but this infer- 
ence is at once negatived by the fact already remarked on, that the posterior talon bears 
nowhere upon it, nor does the end of the tooth exhibit the slightest indication of, a 



298 ME, BUSK ON THE KEMAINS OF 

depression arising from the pressure of a tooth advancing behind it. As the same result 
is yielded in a still more decided fashion by inferior molars noticed above*, I see no 
alternative to the inference that it was an adult tooth of a dwarf species of Elephant. 
The following are the dimensions : — j^ 

Extreme length of crown, measured from back talon to anterior edge, exactly 4-0 

Width of ditto at second ridge 1'4 

Width of ditto at third ridge 1-5 

Width of ditto at sixth ridge 1-4 

Greatest width of crown 1'55 

Width at 9th ridge 1-2 ■ 

Width at last ridge 1-0 

Greatest height of crown, taken at reflection of 10th ridge 2'95 

Length occupied by five disks, from second to sixth inclusive 1'8 

Width at middle of third disk, taken between the enamel-edges . . . . 0'23 

" With reference to the alimentaiy characters, the disks of ivory, and the cement-hollows 
between the enamel ridges are but slightly excavated ; in fact the most anterior portion 
of the crown exhibits the flat and nearly uniformly smooth surface which is commonly, 
presented by Elephants reared in the domestic state and fed upon potatoes and other 
soft food. The inference to be drawn from this is, that the food of the Maltese species 
was more herbaceous than woody. 

6. " liidge-formida. — It now remains to consider how the data furnished abo\e by 
the molars bear upon the determination of the ridge-formula, which of all the characters 
is the most significant in pointing out the affinities of the species. 

" (1) Milk-molars. — The antepenultimate milk-molar m.-m. 2 (fig. 2) is seen to 
have been composed of three coUines, like the corresponding tooth of the African 
Elephant f , while in E. primigenius, E. indicus, and other species of the subgenus 
Euelephas, it presents four collines. 

" The penultimate milk-molar m.-m. 3 is clearly proved by the upper germ-specimen, 
fig. 6, and by the lower, fig. 4, to have had five collines besides front and back talons. 
In the African Elephant it is composed commonly of five ridges in the upper jaw and of 
six in the lower ; whilst in the species of EueJephas the number ranges from seven to 
eight, seven being the complement in E. antiquus, and eight that in the Indian 
Elephant and Mammoth. 

" Of the last milk-molar, m.-m. 4, the specimen shown in fig. 5 fortunately presents the 
crown of an inferior tooth in perfect integrity, composed of eight ridges in addition to 
a front and hind talon ; the African Elephant commonly yields the same number, while 

* Those .shown in figs. 11, 12, 13. 

t In the only specimen to which I have had access of a foetal African Elephant, in the British Museum, and 
which is the one referred to by Dr. Falconer (page 284) as having been brought by Dr. Livingstone, the m.-m. 2 
in both upper and lower jaws have distinctly four collines and two talons (six elements). 



THREE EXTINCT SPECIES OP ELEPHANT. 299 

in E. antiquHS the number is ten, and in E. jmmif/enhis and E. indicus it amounts to 
twelve. 

" (2) True molars. — Of the antepenultimate true molar (m. 1) tliere is no perfect speci- 
men in the collection*. But as in all the species of Elephant and Mastodon this tooth 
invariably repeats the composition of the last milk-molar, we have no difficulty in fixing 
the normal number of its ridges at eight, besides talons. In E. antiquus the number is 
ten, and in the Mammoth and Indian Elephant twelve. 

" Of the penultimate true molar (ra. 2) there is no entire specimen of a lower tooth ; 
but we have the upper beautifully preserved, as shown in fig. 9 {vide note p. 296). It 
exhibits a crown distinctly composed of nine ridges, besides a front and hind talon. 
In the African Elephant the same tooth is commonly made up of nine ridges. In 
E. antiquus the normal number is twehe, while in E. jirimigenius and E. indicus it 
amounts to sixteen. 

"Of the last true molar (m. .3) there are fortunately specimens belonging to both the 
upper and lower jaws ; and although the portion supported on the anterior fang is 
wanting in both, as that constantly corresponds in all the species of Elephant with what 
is borne upon the same fang of the penultimate, we have little difficulty in restoring the 
missing part of the teeth. 

" The upper molar exhibits the remains of ten ridges ; and adding two for the part 
corresponding with the anterior fang, we get a complement of twelve ridges for the 
crown of the last molar. In the African Elephant the same tooth in the upper jaw- 
ranges with from nine to ten ridges, and in the lowe) fiom nine to twelve. In E. anti- 
quus the number is sixteen ; and in E. i)rimi genius the number of plates reaches twenty- 
four f. 

" From the above data the ridge-formula of the molar series is deduced to liave been 

Milk-molars. True molars. 
3 + 5 + 8 . 8 + 9 + 12 
3 + 5 + 8 ' 8 + 9+12' 

This formula at once brings the small Zebbug species within tlie subgeneric group of 
the Elephants which I have called Loxodon, along with E. africanus. The affinity of 
the fossil to the existing species is further clearly indicated by the narrow crown and 
mesial expansion of the disks of wear, together with the point already alluded to of the 
milk-incisors being invested at the crown with a layer of enamel. But, though allied, the 
two forms are specifically very distinct. Besides the signal difference of size, the forms 
of the disks of wear, although belonging to a common pattern, present broad marks of 
distinction. In the African species the di.sks are angularly dilated into rhombs in the 
middle, and the angles terminate in a round loop caused by a single outlying digital 
element, which in the progress of abrasion becomes confluent with the disk of the rido-e 

* Unless, a8 I believe, we may regard fig. 9 as such. 
t The numbers of plates were not flJlcd in in Dr. Falconer's 5IS. 
VOL. VI. — PAET v. 2 T 



300 MR. BUSK ON THE REMAINS OF 

to which it is appended. In the Zebbug form there is never a trace of this outlying 
loop ; and the disks, although open, exhibit only a very slight tendency to angular 
expansion. The character is most pronounced in the two milk-teeth, figs. 4 and 5, 
whilst it is entirely wanting in the penultimate upper molar, fig. 10. In fact this tooth 
differs more from the ordinary type of the Afi-ican species than does the corresponding 
molar of E. antiquus. The amount of agreement and of difference in the molars of the 
two species is best appreciated by comparing the last lower molar (figs. 11-13) of the 
Zebbug form mth the corresponding molar of the African species." 



Having thus given Dr. Falconer's descriptions of the teetli, and his valuable observa- 
tions concerning them, it only remains to inquire whether any evidence is afforded by 
these parts of the existence of more than one species of Elephant of dwarf size. This 
is a jioint to which Dr. Falconer has nowhere adverted ; and it was therefore a matter of 
considerable interest to me, after I had been led from the study of the other bones to 
the conclusion that the Zebbug collection contained the remains of two small species, to 
ascertain whether similar e^adencewas afforded by the teeth. And it seems to me that 
they do indubitably present sufficient evidence to that effect. 

When the teeth are placed side by side, it is at once quite obvious that they may be 
divided into two groups, at any rate so far as the true molars are concerned. These 
groups differ very markedly, more especially in the thickness of the plates, or in the 
number comprised within a given length, as well as in the form of the mac/ueriden. 
What the difference may be, if any, in the numerical formula, I am not prepared to say, 
as the materials are too scanty to allow of the solution of the question, which must wait 
to be decided by the very abundant materials since collected by Dr. Leith Adams. I 
shall content myself here simply with pointing out the striking differences exhibited 
between the teeth of the two species as they are represented in the Zebbug collection, 
and shown in the figures in PI. LII. 

If we compare, for instance, the m. 1 represented in fig. 9 with either of those shown 
in figs. 11-13, but more especially witii the first, of which a side view is also given, such 
a difference will at once be perceived in the form of the machserides or disks of wear, 
and in the thickness of the plates, as to stamp them as totally distinct forms. In the 
tooth fig. 9, nine ridges are comprised in a length of about 2"'5, which gi\'es an average 
thickness of each plate of about 0"'27, whilst in fig. 11 seven plates occupy a length of 
o'-O, equivalent to an average thickness of about 0"-43. Again, if we look at the form 
of the machserides in the two cases, the comparative narrowness of the disks and the 
disposition to true crimping of the enamel-edge, with the complete absence of any median 
angular expansion, in fig. 9 cannot fail to be at once perceived*. Again, in fig. 11 it 
will be seen that the hinder two or three plates, which are just coming into wear, exhibit 

* The difference is perhaps more marked in the actual teeth, owing to the circumstance that the enamel-edges 
are represented rather too thick in fig. 9. 



J 



THEEE EXTINCT SPECIES OF ELEPHANT. 301 

not more than two annuli, which are of large size and with thick enamel, whilst in fig. 9 
the corresponding plates show five or six annuli, of small size, and a corresponding thin 
enamel. The above differences appear to me to be quite as important as those which 
exist between the true molars of the Indian and African species, and infinitely greater 
than those which distinguish the former species from E. j^rimi genius. 

In fig. 10 is shown the crown surface of a tooth from Maccagnone, which would seem 
in its size and characters to approach very nearly to the type of fig. 9 ; and I presume 
that Dr. Falconer may have introduced the figure with the view of showing some relation 
between the Sicilian and Maltese teeth. He has, however, left no observations on this 
point ; and as the tooth itself is not now with the others, I am unable to say more about 
it than is shown in the figure. 

With regard to the milk-teeth, we have not the same facility of judging of their 
relations from the thickness of the plates alone as we have in many cases in the true 
molars. In this respect little or no difference will be observed among the various 
milk-teeth of which figures have been given; and some might thence, under the 
circumstances, be led to conclude that these teeth must all belong to one only of the two 
species whose molars differ so widely in the thickness of the plates. But such a con. 
elusion is by no means warranted by what we know of the milk-teeth in different species 
of Elephant, in which, notwithstanding very great differences in the thickness of the 
plates in the true molars, little or none will be found in that of the plates of the 
respective milk-teeth, whilst in some cases the difference in this regard will even be in 
an opposite direction to that in which it might be expected to show itself. For instance, 
the mean thickness of the true molar plates in E. africanus varies from 0"-85 to 1" or 
more, and in E. indkm is about 0"-55, whilst the thickness of the plates in the m.-m. .'1 
of the two is pretty nearly the same, or about 0"-3^--33; in E. pnmi genius, although 
the thickness of the plates inm. 1 is not more in most cases than about 0"-43--45, those 
of the 3rd m.-m. average about 0"-34, or rather more than in E. africanus, though con- 
siderably less than in E. antiqims, in which the thickness may be taken at about 0"-41, 
As we cannot, therefore, rely solely upon the thickness of the plates in the milk-molars 
as a diagnostic character in species so widely distinguished as E. indicus and E. africanus, 
it is impossible from that character to say whether or not all the milk-molars in the 
Zebbug collection belong to one or more species. Had they all been so worn as to 
afford a good view of the form of the macha?rides, the decision would probably have been 
easy enough ; but it will be observed (leaving out of the question the m.-m. :^, fig. 2) that 
only three of the specimens were worn sufficiently for this purpose ; and as, from the 
form of the machaerides in these instances, and the general condition and colour &c. of 
the teeth, it is not improbable that all belonged to one and the same individual, we must 
have recourse to other characters to determine the question of the true relations of the 
unworn specimens. 1 am not sure how far such a character may be depended upon ; 
but, in the case more especially of milk-teetii uncovered with cementum, I think what 

2 T 2 



302 MB. BUSK ON THE BEMAINS OF 

may be termed the sculpturing of the surface may be regarded as of considerable value. 
Takino- this, however, as a test, it will be found that whilst the surface of the plates in 
the m.-m. 3, fig. 6, is very coarsely and irregularly fluted in the vertical direction, and 
presents no trace of transverse wrinkling, that of tlie exposed plates in tig. 7 is very 
finely fluted, and at the same time very minutely wrinkled transversely*; and the same 
condition may be seen (though much less plainly, owing to a thick covering of cementum) 
in the fragment represented in fig. 8, or in the corresponding tooth described in the note 
in page 290. In the worn milk-teeth, figs. 3, 4. and 5, the surface is entirely covered 
with a thick cement. I am inclined therefore to assign fig. 6 to the same species as 
figs. 3, 4, and 5, whilst figs. 7, 8, and the corresponding tooth would belong to the 
same species as fig. 9, and perhaps as fig. 10. 

How these teeth are related to the other bones of the two dwarf species, the jjresent 
collection affords no certain means of positively deciding ; but, from the more general 
tendency towards the African type which is exhibited in so many respects in £. meli- 
teiisis, 1 should be inclined to assign the teeth represented in figs. 11, 12, and 13, with 
the corresponding milk-molars, to that species. This, again, however, is one of the 
questions which remain to be decided wlien we are furnished with more ample materials. 

Among the teeth undoubtedly belonging to the Zebbug collection when it came into 
my hands, was one about whose source some obscurity exists. It was not marked as 
coming from Zebbug, nor is it entered in the rough list of the collection. Captain 
Spratt, however, though uncertain about its relation to Zebbug, is pretty confident that 
it was brought from Malta. I have therefore not included it in my account of the 
Zebbug collection, but think it should be noticed for future reference, as connected 
with the island. 

The tooth is the m.-m. 3 of the right side, and it is tolerably entire, though consi- 
derably worn in front, so that the two anterior plates are worn down to the common 
base, and the third very nearly so ; but the diminished base of the anterior fang, appa- 
rently much absorbed, still remains. There have been six plates, and probably two talons. 
The length is about 2', and the extreme width (5th plate) 1"'3. The tooth, therefore, 
in general dimensions, differs but little in its size and proportions from the corresponding 
tooth in E. indicus and E. primigenius; whilst it is shorter in proportion than the 
3rd m.-m. of E. anfiquus, and proportionally a good deal wider than the same tooth in E. 
africanus and E. meridionalis. It almost exactly resembles, though rather more worn, 
a 3rd m.-m. from Long Hole, in Gower, and which is labelled by Dr. Falconer E. antiquus, 
though it seems to me to exhibit much more the characters of E. primigenius, to which 
species, I should, with the greatest deference to so high an authority, be inclined to 
assign both. 

* The transverse wrinkling appears to me, from subsequent obsei-vation, to be a very uncertain character ; but 
the huge vertical markings afford, undoubtedly, characters of some importance. 



THREE EXTINCT SPECIES OF ELEPHANT. 303 



DESCRIPTION OF THE PLATES. 



PLATE XLIV. 

Fig. 1. Symphysis of mandible of the largest Maltese Fossil Elephant (Elephas 1) 

(p. 231). 
Fig. 2. Neural spine of the seventeenth or eighteenth dorsal vertebra of the same 

species (p. 233). 
Figs. 3, 4. The outer and inner aspects of a very young exoccipital bone of the same 

(p. 233). 

PLATE XLV. 

Fig. 5. Fragment of shaft of femur (young) of the largest species, and not improbably 
belonging to the same individual as that which afforded the exoccipital 
(p. 235). 

Fig. 6. Shaft of femur of mature E. melitensis (p. 247). 

Fig. 7. Fragment of neural spine of dorsal vertebra of ditto (p. 241). 

Fig. 8. Portion of second rib (right side) of ditto (p. 241). 

PLATE XLVI. 

Fig. 9. Seventh cervical vertebra of E. melitensis (p. 238). 
Figs. 10, lOff. Sixth or seventh dorsal vertebra (p. 240). 
Figs. 11, 11«. Second or third lumbar vertebra (p. 241). 

PLATE XLVII. 

Fig. 12. Portion of atlas oi E. melitensis (p. 238). 

Fig. 13. Portion of ascending ramus of mandible of ditto (p. 236). 

Fig. 14. Upper surface of astragalus oi E. falconeri (p. 268). 

Fig. 14 (his). Dorsal aspect of fragment of scapula referred doubtfully to E. falconeri 

(p. 254). 
Fig. 14 {bis)\ View of the glenoid fossa. 

Fig. 15. Posterior view of very young tibia, probably of ^. melitensis. 
Fig. 15ff. Proximal articular surface. 

Figs. 16, 16«. Anterior and posterior views of a foetal tibia (p. 281) 
Fig. 17. Anterior view of an older tibia (pp. 281, 282). 
Figs. 18, 19. Two fcetal radii (p. 280). 
Figs. 20, 21. Two fcetal tibia; (pp. 281, 282). 



:104 MB. BUSK ON THE REMAINS OF 

PLATE XLVIII. 

Fig. 22. Proximal end of hunif>rus of E. melitensi.s (p. 244). 

Fig. 23. Fragment of scapula of ditto (p. 243). 

Fig. 23a. The glenoid fossa. 

Fig. 24. Proximal end of ulna of ^. melifensis (p. 245). 

Fig. 24 ff. The proximal articular surface. 

Fig. 25. An olecranon process of E. melitensis (p. 246). 

Fig. 26. Fragment of ischium of ditto (p. 242). 

PLATE XLIX. 

Fig. 26 [his). Anterior and posterior views of humerus of E. falconeri (p. 255). 
Fig. 27. Distal articular surface of humerus of ditto (p. 259). 
Fig. 28. Proximal end of ulna of ditto fp. 2G0). 
Fig. 28a. Articular surface. 

PLATE L. 

F'igs. 29, 29a. Anterior and posterior aspects of femur of E. falconcn (p. 266). 
Fig. 30. Fragment of upper end of femur of ditto (p. 2G4). 
Fig. 31. Portion of pelvis of ditto (p. 263). 

PLATE LL 

Fig. 32. Posterior aspect of left half of atlas of E. falconeri (p. 251). 

Fig. 32a. Anterior aspect. 

Figs. 33, 3oa. Similar aspects of another atlas of much younger age and doubtfully 

referred to E. melitensis (p. 251). 
Figs. 34, 34a. Anterior and posterior views of a fragment of an eightli to tenth dorsal 

\ ei tebra of E. falconeri. 
Fig. :)5. Fragment of atlas of E. meUtensis (p. 238). 
Figs. 36, 36 ff. Lateral and posterior aspects of neural spine of a dor.sal >ertebra of 

E. falconeri. 
Fiff. 37. Proximal end of second rib of ditto. 
Figs. 3S, 38 a. Anterior and posterior views of a very young or fcrtal tibia (? ///^y/jo- 

potannis). 
Figs. 39, 39a. Lateral and posterior views of another tibia of nppaiently similar 

character. 
Fig-s. 40, 40a, 40/;. Fourth metatarsal of E. falconeri (p. 271). I 

Fig. 41. Proximal phalanx of third manual digit of ditto (p. 263). 



THREE EXTINCT SPECIES OF ELEPHANT. 305 

PLATE LII. 
Figs. 42, 42«. Fragment of right ramus of mandible of very young animal. 
Figs. 42', 42'a. External and internal aspects of a very young or fuetal oxociipital 

(p. 272). 
Fig. 43. Fragment of right side of mandible, rather older than that shown in Fig. 42 

(p. 278). 
Figs. 44, 44ffl. External and internal aspects of a very young or foetal e.xoceipital 

apparently different from that shown in Fig. 42' (p. 273). 
Fig. 45. The entire symphysis of a very young or foetal mandible, displaying the socket 

of the m..-m. 1 and, on the right side, part of that of m.-m. 2 (p. 278). 
Fig. 46. Portion of tlie premaxillary with germ of permanent incisor (p. 270). 
Figs. 47, 47«. Distal extremity of fibula] oi E. falconeril 
Fig. 48. Tusk of E. falconeri (p. 285). 
Fig. 49. Very young humerus oi E. falconeri (p. 279). 
Fig. 50. Another very young humerus (p. 279). 

PLATE LIIL 

Figs. 1, \a, \b. Deciduous incisor of ^. melifenfiial (p. 284). 
Figs. 2, 2a. Antepenultimate lower milk-molar, m.-m. 2 (p. 286). 



Figs. 3, 3«. Fragment of (probably) the penultimate lower milk-molar, m.-m. 3 (p. 287). 

Figs. 4, 4a. Left lower penultimate milk-molar, m.-m. 3 (p. 288). 

Figs. 5, 5«. Last lower milk-molar of the left side, m.-m. 4 (p. 288). 

Figs. 6, 6ff. Penultimate upper milk-molar in germ, m.-m. 3 (p. 289). 

Figs. 7, 7a. Fragment of germ of last upper milk-molar, m.-m. 41 

Figs. 8, 8a. Portion of last upper milk-molar, m.-m. 4 ? (m. 1, ini/ii) (p. 290) ! 

Fig. 9. First upper true molar (m. 1)? oi E. falconeri I (p. 295). 

Fig. 10. Crown surface of tooth from Maccagnone (p. 301). 

Fig. 11. Last(!) lower molar of the left side, m. 3 (p. 294). (E. incUtcHxis f) 

Fig. 12. Greater part of the last lower molar of the right side, m. 3 (p. 2'.ll). 

Fig. 13. Posterior half of last lower molar of the riglit side, m. 3 (p. 293). 



500 



ON THEEE EXTINCT SPECIES OF ELEPHANT. 



The following Tables give the principal measurements of some of the Bones and Teeth 
which have served as data for the comparison of the Maltese with other forms of Elephant. 

Table III. — Atlas. Various measurements. 







s 


kM 


ss 


<D 




g 


t3 


§ 


1 


14- 




■a 

1 


bD 


.d. outside 
terior articu 
processes. 


.d. outside p 
terior articu 
processee. 




i 

a, 


•E 

O 


o . 

. & 


O g 

■II 


•g 

■a 

c 

o o 


adius of Ion 
curve of coil 
loid facet. 




w 


w 


H 


H 


■< 


-A 


-4 


^ 


n 


w 


« 


E. tndicus (Chuny) 


16-0 






















„ (Sumatran) .... 


10-5 


6-4 


7-3 


5-2 


2-4 


1-y 


3-4 


fi-5 


3-3 X 2-4 


.3-2 X 2-4 


1-7 


(2(578, E-C.S.) .. 


12-5 


7-0 


7-8 


6-5 


2-7 


21 


3-8 


7-6 


4-0 X 3-0 


3-.5 X 3-0 


2-0 


,, (Olivier) 


13-5 






















(Blaiiiville) .... 


U-4 


7-7 


7-8 












4-4 X 2-6 






E. africmius, B.M 


13-0 


8-5 


9-5 


8-8 


3-6 


2-5 


4-i 


8-3 


4-5 X 3-0 


4-3 X 3-2 


2-4 


(BkimdUe) .. 


11-8 


6-6 


7-6 












4-1 X 2-4 






E. meliteusis 




3-5 














2-5 X 1-7 




1-5 


J^ ^alconeri 


5-0 


2-9 


3-3 


2-6 


1-2 




1-7 




1-6 X 1-1 


1-8 


■85 





Table IV. — Dimensions and Proportions of Astragalus. 



. 




■S 


> 


-J 


■© 


i 

1 
1 


i 

■s 
1 

c: 

S 

O 


Inner calcaneal facet. 


s 

•a 

OS 

o 

a§ 


1 

a 
s 


i 
1. 

°.l 

•1-2 

11 


Proportion of v.d. of scaphoid 
facet to its tr.d. 

Proportion of tr.d. of scaphoid 
facet to total tr.d. of bone. 


Eadii of curves of scaphoid 
facet. 


E. indicas {C\\nny).. 

„ ( Sumatran) . . 

„ (2707,H.C.S.; 

,, (young) B.M. 
E. afrieti mix, B.M. . . 

E. meli'ensis 

E. f(d:on?i'i 

E.' „ (?) ■■•• 


5-1 
3-4 
4-3 
3-0 
4-G 
2-7 
1-9 
1-8 


6-0 
3-9 
4-9 
35 
5-2 
31 
2-2 
21 


2-4 
2-8 
2-1 
31 
2-0 
1-4 
1-2 


4-0 X 

3-1 X 2-6 2-9 X 1-8 
3-7 X 3-3 1 3-4 X 21 
2-8 X 2-1 2-7 X 1-8 
4-0 X 3-7 4-2 X 2-5 
2-4 X 2-2 3-0 X 1-9 
1-7 X 1-6 2-1 X 10 
1-7 X 1-4 1-9 X -95 


2-5xi-5 
2-6x1-3 
2-0x1-1 
3-5x1-8 
1-9x1-1 
1-4 X -6 


:-850 ' . . 

2-9x1-6 2-2X1-8 -871 -615 
3-5 X 2-2: 2-7 X 1-4 -871 -573 
2-5x1-0 1-6 X -75 -856 -600 
3-0x2-51 3-0x2-7 -884 -596 
1-8 X 1-4 1 2-1 X 1-0 -870 •945 
1-4 X -8 1-5 X -65 -863 -636 

1-4X-70 -857 -571 


■733 

■891 
•750 
-900 
-912 
•941 
•874 


.. ^666 
■620 ^743 

■610 ^696 
■648 ^770 
•602 '^740 
■633 ,■967 
•500 ■954 
•500 J-9U4 


1^8 & 55 
1^8 & 5-5 
2-0 & 3-0 

2-5 & b's 
1^7 & 1^5 
1 & 1^5 
1-0 & 2-3 



Table V.— The Principal MeasuiL'iiienls 



of the Humerus and Femur, as affording data for computing the lleigiit and Pvpnition of Elephants. 



[TofaftpOijtaOQ.) 



















Euraorus. 


























Femur. 






1 


Height of AniniBl. 






i 


1 


i 
1 


5 


1 
1 


i 

i 

i 

i 


I 

1 
i 

4' 


1 

•i 

'C 


1 

■= 

1 

■5. 

H 


i 
1 


1 
8 

,1 

1 


f 
1 

5 


■3 

- 
i 


i 

1 

1 


•3 
j 


i 

1 


i 


] 

■1 

C-1 


I 


1 

1 

i 

S 

1 


; 


1 
1 


■3. 
•3 

s 

h 


i 

! 
i 


t 

,1 

i 


! 
1 

i 

Q. 


1 


1 


i 


1 
1 

> 


data. 




i 

It 

■< 


1 


i 


1 
1 


1 
1 


i 

1 


i 


X o«Bnii (Chnnj) .... 


350 


8-0 


101 


S-0 


120 


3-0 


00 


15-0 


ll-O 


8-2 


li'3 


5-5 


40 


120 




• ■ 


42 


4-7 


.■i-1 


12'1 


13-0 


00 


8-1 


8'0 


7-8 










• 
llfl 


• 

115 


103 


103 


02 


• 

96 


103 


„ „ 2-MB.8.&S. 


35-0 


7-9 


10-4 


9'S 


110 


3-9 


00 


17-0 


11-2 


8-5 


0-3 


6-7 




120 


4-1 


4'3 


43 


00 


5-1 


38 


14-3 


13-0 


01 


70 


80 


7'8 










• 
120 


lis 


• 
105 


• 
103 


• 
95 


• 
96 


« 
105 


„ . (CoricT) .... 


32'S 


7-3 






0-7 


40 








? 
0-3 














380 


51 


4-7 






















■■ 


108 


108 


« 
95 


• 
06 


• 
85 


* 
80 


108 


„ „ OefloD, B.1L . . 


310 


6-5 


7-0 




8-2 


2-6 


4-4 


U-5 


9-0 


7-3 








11-5 






88'S 


6-0 


3-76 


2-7 


10-0 


110 


7-9 


70 


7-2 


02 










• 
107 


• 
104 


« 
94 


• 
91 


85 


85 


85 


„ „ Sonaliw. B.U. 


27-8 


5-2 


60 


a-8 




2-3 


3-6 


10-0 


7-2 


5-4 








10-0 


20 


2-4 


33-7 


4-1 


3'S 


2-3 


8-0 


90 


5-8 


50 


5-7 


51 










94 


• 
91 


• 
81 


• 
81 


• 
74 


• 
7« 


• 
83 


„ „ (r«uif)2723C.S. 


181 


3-0 


40 


32 




1-4 


2-2 


5-D 


51 


4-7 


2-5 


24 


,. 




10 


!■» 









































50? 


. „ (j™.«)BJL.. 


20-8 


4-5 


50 






1-7 


2-8 


7-3 


a-2 


50 


31 


3-2 




7'6 










2-2 


1-8 


00 




5-4 








27-5 


40 


4-7 


1-9 


« 
5(J 


,f. 


• 
55 


* 
01 




• 
57 


• 
00 


£. afivmw, B.U 


35-5 


7-9 


0-2 


9-3 


11-0 


33 


4-0 


13-1 




7-8 


50 


48 


•' 


120 


40 


4-4 


42-5 


00 


4-2 


31 


11-0 


131 


8-3 


80 


8-4 


0-9 










• 
119 


118 


• 
103 


• 
103 


• 
93 


• 
97 


• 
107 


E. Uilta (laip) 


• 




























2-8 


2-7 


33 


























-- 














s()-*15 


E. aulilfiuU. fig. 22 


• 
10-17 


3-5 


:n 


:1T 


i r. 




















1-8 


1-6 


• 
20 


























.. 


• 
50 


53 


• 
49 


• 
48 


44 45 


• 
50-55 


- „ fig" ..-. 


15-10 
































18-20 




I'O 


1-6 


5-6 




















o' 




• 
40 




il.. 


50-55 


£./it™wW.fis».2(!. 27 


10-12 


• 
22 






























• 
13 


* 
1-8 


























• • 

31) 30 


• 
32 


• 
32 


• [ « 
29 1 30 


• 
30-30 


_ „ 65.30 


10-12 










1-0 


1-5 


41 




25 


1-8 


1'4 


1-2 


3-2 






1^13 


1-3 


00 


3-6 


2-3 












«T, 


1 ''" 


i '" 


u:r2 












g.fnmSsm^, B.U 


320 


7-3 




98 


10-3 


4-3 




13'6 


9-5 


81 


5'0 


5'2 


311 


120 






• 
38 






































100-110 



N.B. The norobun raaiked vith ui • oro computed, the others takea from direct rocMnrunioiit. They indicate inches. 



J 







^ 
















^ 
















~" 
















~- 
















— 
















-~ 






Slix^ 


p. 


I. 


< 


Sii^ 


p. 


T. 


j 


Sin. 


P. 


T. 


1 


Sin. 


P. 


T. 


1 

< 


Siu. 


P. 


T. 


1 


Siso. 


P. 


T. 


1 
■4 


BiM. 


P. 


T. 


< 


Si>o. 


p. 


T. 


1 


Slio. 


P. 


T. 


1 


Silo. 


P. 


T. 


< 


Siio. 


p. 


T. 




-> X -.15 


4 
4 


2 


•20 


-75x-4i 
'«X'4£ 
-«5 


4 
3 

4 


2 

a 


-20 


2-7x1-3 
2-7x1-5 
»4xl-3 


8 
S 
8 


1 

1 

1 


-37 

-37 


2-6 X 
2-8 xl-3 
265x1-3 


8 
D 

8 


2 

I 
1 


■31 
-31 
-32 


6-2x1-6 
4-5x1-5 
4-7x10 


12 
12 
11 


2 
1 
2 


-40 
.-38 
■40 


4^4 Xb3 

6'3xl7 

63x1-0 
6-2x1-0 
5-1 X 17 


13 

12 

12 

12 
13 
13 


1 
1 
2 
1 
1 


-35 

■40 

■41 
-40 
-10 


0-3 X 2^1 


12 


2 


-48 


0-8x2-0 


12 


2 
2 

1 
2 


■52 
•00 

-46 
-48 
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XI. On a Species of Dormouse (Myoxus) occurring in the Fossil state in Malta. 
By A. Leitu Adams, M.B., F. G.S. 

Kead May 9th, 1867. 

[Plate LIV.] 

IN the 'Journal of the Royal Dublin Society' for November 1861, I figured the dental 
aspect of this Dormouse, which I have proposed to call Myoxus melitensis. But as the 
figures there given do not fully indicate the characters of the animal, I have deemed it 
requisite to furnish the following further illustrations, taken from the numerous speci- 
mens that have since come under my notice. The contour of the cranium, the relatively 
small size of the anterior and posterior molars compared with the intermediate ones 
(which are about as long as they are broad on the crowns, the bold machcerides pre- 
senting well-defined, undulating ridges), and the absence of the small grinder in the 
upper jaw separate it from the Sciurina, and assimilate it to the subfamily Myoxina, 
whilst its large proportions represent a species distinct from any other known Myoxus, 
recent or fossil. 

Among the abundant remains discovered by me in the caves, fissures, and alluvial 
deposits of Malta were several lower jaws comparatively more slender, and presenting a 
more marked concavity on the lower border, whilst they did not seem to diflPer in any 
other respect from the others (see op. cit. plate 2. fig. 11). To the form to which these 
belong I have given the name of Myoxus cartei ; it may be doubtful, however, whether 
the above characters are really sufficient to create a distinct species from the other, which 
I have named Myoxus meliteTisis. 

This Eodent seems to have existed in enormous numbers, inasmuch as its remains are 
met with in abundance throughout the cavern- and fissure-deposits, up even to the 
superficial alluvium now in course of formation, so as almost to indicate that the animal 
may have outlived many, if not all, of the other quadrupeds &c. with which its remains 
are so frequently associated. 

The Reports on the Maltese Caves, read at the Meetings of the British Association in 
1865 and 1866, together with my other communications on the fossil fauna of the 
Maltese fissures and alluvial deposits, give full particulars with reference to the 
localities and mode of occurrence of this and the other members of the fossil fauna. 
One point comes out clearly in the stratigraphical distribution of the remains of 
Myoxus melitensis, viz. that the animal lived and died in the caverns of Malta ; whilst 
at the same time, from the exceedingly large numbers found strewing tlio lower portion 

VOL. V[. PART V. 2 f 



308 DR. A. LEITH ADAMS ON A F0S81L MVOXUS. 

of the fossiliferous deposit of the Muaidra-gap, there is evidence of a wholesale destruc- 
tion of this animal at all stages of its existence, from the unborn to the aged. And 
from the circumstauce that the same conditions are found to obtain with the associated 
Elephantine remains, it may be concluded that the destruction of both was due to 
something more than the ordinary process of decay. Again, the very fragmentary 
condition of the bones found in the stalagmitic deposits of the Malak cave seems to 
indicate that they had been introduced into it by carnivorous mammals or birds. 



EXPIANATION OF PLATE I.IV. 

Fig. 1. Side view of the skull of Myocnts melitensis. 

Fig. 2. Coronal view of the skull of the same individual. 

Fig. 3. Base of skull. 

Fig. 4. Lower jaw of the same individual. 

Fig. 5. Molars of upper jaw, magnified to four times the natural size. 

Fig. 6. Incisor. 

Fig. 7. Lower jaw of a young individual, the molars just appearing. 

Figs. 8, 9. Humerus. 

Fig. 10. Scapula. 

Fig. 11. Articulating surface. 



Figs. 12, 13. Femur. 
Tibia. 
N.B. Except in fig. 5 all the parts are represented of tlie natural size. 



Fig. 14. Tibia. 



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MYOXUS MELITENSI5 



[ 309 ] 



XII. On the Osteology of the Cachalot or Sperm-male (Physeter macrocephalus). 
Bi/ William Henry' Flower, F.B.S., F.R.C.S., F.L.S., F.G.S., F.Z.S., Conservator 
of the Muserim of the Royal College of Surgeons of England. 

Read November 14th, ISfiT. 

[Plates LV. to LXI.] 

Introduction. 

Our present knowledge of the osteology of the Cachalots is derived from the following 
sources : — 

1. Cuvier, in the ' Recherches sur les Ossemens Fossilcs,' has given a description, clear 
and pointed, but brief, of an imperfect skeleton, bought by him in London in 1818, and 
still preserved in the Museum of the Jardin des Plantcs. Figures on a very small scale 
are given of the cranium and lower jaw, of the scapula, humerus, radius and ulna, and of 
some of the vertebrse. The locality from which the animal was originally obtained is not 
stated. As will be presently shown, the skeleton presents certain peculiarities, especially 
in the number of the ribs and vertebrse, by which it differs from all others known. 

In the same classical work, portions of the lower jaw of three other individuals con- 
tained in the Paris Museum are figured and described. 

2. In the valuable posthumous work of Peter Camper, ' Observations Anatomiqucs 
sur la Structure interieure et le Squelette de plusieurs especes de Cetaces,' Paris, 1820, 
is a tolerably full description, and some very sketchy figures, of a mutilated cranium 
preserved in the church at Scheveningen, in Holland ; and there are also some observa- 
tions upon, and a figure of, another cranium, in the Paris Museum, taken from one of the 
individuals cast ashore near Audierne, in Brittany, in 1784. The tympanic and petrous 
bones, as well as the ossicula auditus, are figured in considerable detail ; and drawings 
are also given of the scapula and arm-bones, and of the atlas. The latter (from a speci- 
men in the British Museum) is erroneously attributed to a Balcena, while the consolidated 
mass of cervical vertebrse of a whale of this genus is described as that of a Cachalot. 

3. Lacepede (' Histoire Naturelle des Cetaces,' Paris, 1804) has given a figure of the 
skull of the Audierne Cachalot in the Paris Museum, also of one of the ribs and some 
vertebrse. 

4. Beale ('Natural History of the Sperm-Whale,' London, 1889) has given a general 
description, unaccompanied by figures, of the skeleton of a full-grown Sperm-Whale, 
mounted in the Park at Burton Constable, near Hull. Certain errors in the articulation 
of the skeleton, particularly of the hyoid bones, sternum, pelvis, and carpus, not de- 
tected by Beale, necessarily introduced confusion into his description of these parts. 

VOL. VI. — PART VI. 2 X 

ERKATUM. 

Vol. VI. 11. I<B9, li'ii-' It*, fo'' '■ I'usti-d " i-ead " i-errice." 



310 ME. \V. H. FLOWEE ON THE OSTEOLOGY OF THE SPEEM-WHALE. 

5. In a small octavo work, entitled " History and Description of the Skeleton of a 
New Sperm -Whale lately set up in the Australian Museum, by William S. Wall, 
Curator," &c., Sydney, 1851, the description, although defective in many respects, is on 
the whole the most complete yet published, as the skeleton which is the subject of it, 
although very young, was iu a tolerably perfect state. The memoir is accompanied by 
a rudely executed drawing, on a small scale, of the entire skeleton, and also of the 
sternum, hyoid, and peine bones*. 

6. In the ' Descriptive Catalogue of the Osteological Series of the Museum of the 
Royal College of Surgeons' (1853), Professor Owen has given a somewhat detailed account 
of the form and relations of the cranial bones in a vei7 instructive skull of a fcetal 
Cachalot contained in that collection. 

7. A woodcut figure of the same skull has been given by Professor Huxley in his 
'Elements of Comparative Anatomy,' 18G4. 

S. The petrotympanic bones of a Cachalot from the same Museum are figured in 
Owen's ' British Fossil jNIammals.' 

9. Dr. Gray (Proc. Zool. Soc. 1864, p. 590, and 18G5, p. 440) has given figures, taken 
from photographs, of the cervical vertebrae of two Cachalots in the Museum at Sydney, 
which he regards as belonging to distinct species. 

Numerous as the above-noticed works may appear, the information contained in them 
is but fragmentary, and very much still remains to be done before our knowledge of the 
osteological characters of this huge and strangely modified Mammal can be said to be 
placed on the footing which its interest ought to secure for it. 

In the present communication it is my intention, — 

I. To give a description, accompanied by detailed drawings, of the nearly perfect 
skeleton of an adolescent male Cachalot, which was taken in the latter part of the year 
1864 off the south-west coast of Tasmania, and the bones of which were prepared with 
great care and at considerable trouble and expense by W. L. Crowther, Esq., M.R.C.S.E., 
of Hobait Town, and by him presented to the Museum of the Royal College of Surgeons. 

II. To compare this skeleton with other skeletons or parts of skeletons which are 
available for the purpose. As materials for this portion of the work I may especially 
mention : — 

a. Various portions of the skeletons of Cachalots from the Tasmanian seas, also pre- 
sented to the Museum of the College of Surgeons by Mr. Crowther, comprising the 

* It is stated by Dr. G. Bennett (Gatherings of a N'aturalist, 1800, p. 1()2) that the real author of this work 
was the late William ftharjic Macleay. But as there is no indication of this in the work itself, as Wall's name 
alone appears on the titlepagc, and as he has been allowed by Macleay to identify himself with the author 
of the book, especially when speaking in the first person of acts connected with the preparation of the skeleton 
(see pp. 4, 5, &c.), which Dr. Bennett himself attributes to Wall, I shall always quote it under the latter name 
only. Some authors have, without any explanation, quoted this work as " Macleay's "^a practice which must 
necessarily introduce confusion into cetological literature. 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 311 

complete skull of a very young animal, four lower jaws of adults, four perfect pectoral 
limbs, some caudal vertebi-se, and several detached pelvic and petro-tympanic bones. 

h. The nearly complete skeleton of an adult male Cachalot from the north of Scotland, 
now in the British Museum. 

c. The skeleton of an adult male Cachalot preserved at Burton Constable, in 
Yorkshire. 

d. The very imperfect skeleton at the Jardin des Plantes, Paris. 

e. The foetal skull in the Museum of the Royal College of Surgeons, which was pur- 
chased in 1841 of a dealer. Unfortunately the locality whence it was obtained is not 
recorded. 

/. A disarticulated skuU, about the same size as the last, in the Museum of St. 
Bartholomew's Hospital. 

111. To compare the osteological characters of the Cachalots with those of other 
allied Cetaceans, in the hope of furnishing additional evidence as to their zoological 
position in the order. 

1\. To endeavour to ascertain whether the osteological characters furnish indications 
of more than one species of Cachalot, and, if so, to establish diagnostic characters. Not 
to prejudge this question, I shall for the present avoid using any specific scientific 
designation, but speak of the different skeletons compared, according to the localities 
from which they were obtained. 

A few notes upon the history, general characters, and condition of these skeletons vaW 
be useful before entering into details. 

1. The Tasmanian specimen. As before stated, this was captured off the coast of 
Tasmania in the year 1861. The animal was considered a full-sized male, and said to 
have measured 60 feet in length. It was towed ashore, and the skeleton prepared under 
Mr. Crowther's directions. It is perfect, with the exception of one of the pelvic bones, 
four of the chevron bones, and a few of the terminal plialanges. The condition of 
ossification shows that it is not quite adult : the epiphyses are still loose on the upper 
end of the humerus, and on all the dorsal and lumbar, as well as the anterior eleven 
caixdal vertebrae ; beyond this they are united to the bodies. The vertebral formula 
is C. 7. D. 11. L. 8. C. 24 = 50, the vertebra which bears the anterior pair of chevron 
bones at its hinder end being reckoned as the first caudal. The length of the vertebral 
column when the vertebrae were placed close together, and in a straight line, was 30' 4". 
The cranium measures 1 6' 9" in length. The entire length of the skeleton as articulated 
is 50' 1", three feet having been allowed for the intervertebral spaces*. There are eleven 

* Perhaps more should have been allowed ; for by measuring the vertebral column of an adult Porpoise, in 
the recent state, and again after maceration, I find that the length of the whole of the vertebral bodies placed 
close together is to the recent column, with the intervertebral substances, as 100 to 115. Assuming that the 
relation is the same in the Cachalot, the recent vertebral column of the animal described above would be 
34' lOi", and the entire skeleton 51' 7|". 

2x2 



312 JIE. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 

pairs of ribs, the first ten fully developed, the eleventh rudimentary. The chevron bones 
are ten in number ; but the articular surfaces on the vertebrae show that at least four 
are wanting. The teeth are complete, and very little worn. 

2. The animal from which the next specimen was prepared was washed ashore in a 
much decomposed state, in July 1863, near Thurso, in the county of Caithness. 
The skeleton was presented by Captain Macdonald, upon whose property it was 
stranded, to the British Museum ; and I am indebted to the kindness of the Keeper 
of the Zoological Department of that institution, Dr. Gray, for the opportunity of 
making a close examination of it while undergoing the process of preparation 
necessary to fit it for exhibition. This was also a male. The condition of the bones 
shows it to be quite aged : the epiphyses of all the vertebrae are firmly united to the 
bodies, and so is the head of the humerus to the shaft ; the teeth are very much worn 
down. The skeleton is unfortunately far from being perfect. The cranium has been 
much injured, most of the teeth lost ; several of the posterior caudal vertebrae, the 
hyoid bones, the pelvic bones, and many of the phalanges are missing. The vertebral 
formula is C. 7. D. 11. L. 8. C. 21-f ? The length of the column, the bones being 
placed close together, is 28' 7"; the cranium is about 17' 9" (the ends of the praemaxillae 
being broken, it cannot be measured exactly) ; so that, allowing for the terminal caudal 
vertebrae, the skeleton may be estimated at 47' without the intervertebral spaces, or 
rather more than 50' with them. The ribs, as in the former specimen, are ten well- 
developed pairs, and one rudimentary pair — that is, 13^" and 12i" long respectively, and 
nearly straight, having, apparently, been attached to the ends of the transverse pro- 
cesses of the eleventh dorsal vertebra. There are twelve chevron bones, the first anky- 
losed to its corresponding vertebra. I shall speak of this as the Caithness skeleton. 

3. The Yorkshire skeleton, as it may be conveniently termed, was prepared from an 
animal stranded in 1825, at Tunstall, in the Holderness, and which fortunately, while 
still entire, came under the observation of Dr. Alderson, then residing in Hull, who has 
given a figure and description of its external characters, with some anatomical notes, in 
the ' Cambridge Philosophical Transactions ' for the same year. No less fortunately for 
science. Sir Clifford Constable, Bart., in his capacity of " Lord Paramount of the 
Seigniories of Holderness," claimed the body of the animal, and had the skeleton 
prepared and mounted in his park at Burton Constable. With his kind permission 
I had the opportunity in June 1866 of making a careful examination, with measure- 
ments and drawings, of this specimen. Like the last, it is a perfectly adult male : the 
epiphyses of all the bodies of the vertebra? are united, only slight traces of the original 
separation still remaining in the anterior lumbar region ; and there is only a slight 
indication of tlie original epiphysial condition of the head of the humerus. The 
vertebral formula is C. 7. D. 11. L. 8. C. 23 = 49. The total length of the skeleton 
is 48' 4", the vertebral bodies being placed close together ; of this, the head occupies 
18' 11". Ten pair-s of ribs are present ; but the vertebra which I have reckoned as the 



ME. W. H. FLOWER ON THE OSTEOLOGY OP THE SPEEM- WHALE. 313 

eleventh dorsal, appears, from the condition of its transverse process, to have carried a 
rudimentary rib. The body of the hyoid bone, composed of three pieces, is present ; but 
the basihyal is articulated to the posterior end of the sternum, and the two thyro-hyals, 
joined together, form a sort of pelvis. I mention these circumstances, as a knowledge 
of them will clear up some of the diflSculties in Beale's description of this skeleton. 
The stylo-hyals are absent ; but the bone which Beale took for an os penis was evidently 
one of them. The true pelvic bones are absent. There are ten chevron bones present. 
The carpus and phalanges are nearly complete on both sides, but incorrectly articulated. 
The teeth have all been removed and are replaced by wooden models. Notwithstanding 
these defects, it is a noble-looking specimen ; and it would be a matter of great regret if 
it should become still further deteriorated by a long continuance of the exposure to 
all weathers to which it is now subjected. I cannot forbear mentioning as a curious 
incident connected with it, that at the time of my visit a Starling had formed her nest 
and was rearing her young brood in the cavity (certainly now most convenient for her 
purpose, but) which once contained the brain of this monster of the deep. 

4. In the courtyard of the Anatomicnl Museum at the Jardin des Plantes, at Paris, is 
the decayed wreck of the Cachalot's skeleton mentioned by Cuvier as having been bought 
in London in 1818 *, and which furnished what for many years was the standard and, 
indeed, only description of the osteology of the animal extant. Yet this skeleton pre- 
sents peculiarities in the number of the ribs and vertebrae, which separate it at once 
from all the preceding. While they, as well as the skeleton in the Sydney Museum 
described by "Wall, all agree in having but ten pairs of well-developed ribs, the Paris 
specimen has fourteen, besides indications of a rudimentary fifteenth ; and while in none 
of the others does the total number of vertebrae exceed fifty, this one has sixty, and 
still wants the terminal portion of the tail. Notwithstanding these differences, greater 
than are to be found in many animals generically separated, in general character 
the cranium, vertebrse, ]fibs, and other bones closely resemble those of the three 
former skeletons — so much so that one cannot avoid the suspicion that the specimen lias 
been made up of the bones of more than one individual. On account of this circum- 
stance, as well as its very imperfect condition (the hyoids, sternum, hands, pelvic bones, 
and terminal caudal vertebrae being absent), I have not made so much use of it in the 
comparisons as of the three skeletons at present in this country. It should be men- 
tioned that the animal was adolescent : the teeth are slightly worn, the epiphyses not 
united to the head of the humerus, or to the majority of the vertebrae. The entire length 
of the skeleton as it now stands is 56', the head being about 16'; but the tips of the 
premaxillaries are made of wood, and many of the epiphyses are lost from the vertebral 

* This is probably the skeleton exhibited in Rackstrow's Museum, Fleet-street, described in the Catalogue 
(1794) as " The Astonishing and Complete Skeieton of a full-grown Sperma-ceti Whale, being the real bonen 
joined together; near 70 feet in length. The Head, or Skull alone, measures IG feet." I am indebted to 
Mr. Gore, of Bath, for this reference. 



314 ME. W. H. FLOWEK ON THE OSTEOLOGY OF THE SPERM-WHALE. 

column, and insufficient spaces left between the bones ; so that even these measm-ements 
afford but an approximative comparison. 

Cranixim. 

In no known mammal does the cranium depart from the ordinary type to such an 
extent as in the Cachalot. The expansion, elongation, flattening, and distortion of 
many of the cranial and facial bones, met with in a certain degree in all Cetaceans, is 
here carried so far as to render it by no means easy, at least in the adult animal, to 
recognize their homologies. Comparison with, the skulls of young individuals and of 
less-modified cetacean forms, however, clear up most of the apparent difficulties. 

The size of the skull in the adult animal is larger in proportion to the remainder of 
the body than in any other known mammal *. As in all animals in which the great 
bulk of the skull is made up of the face and jaws and not of the brain-case, the rela- 
tive size of the entire head increases with age, at all events up to maturity ; and it is 
probable that the jaws continue to augment in size and weight after the growth of most 
other osseous structures has ceased. The relative length of the cranium to the vertebral 
column (the vertebrse being placed in contact) in the Sydney skeleton (according to 
Wall) is as 4G to 100, in the Tasmanian as 57, in the Caithness as GO, and in the York- 
shire as 67 to 100. The fii-st is scarcely more than half-grown, the second adolescent, 
the last two adult. 

As seen in the section, PI. LVI. fig. 1, the cerebral cavity is of comparatively limited 
dimensions, being of a somewhat spherical form, with an average diameter of about 10", 
and a capacity of 900 cubic inches. In front of this stretch out horizontally the 
enormously developed bones of the upper jaw, to which the great size of the entire skull 
is mainly due ; while rising above it is the high, compressed, vertical, transverse wall 
of bone forming the great occipital crest, the posterior boundary of the enormous 
supracranial basin, so remarkable a feature in this singular skull. 

The general form of the cranium may be compared to that of a huge pointed slipper, 
with a high heel-piece, and the front part trodden down. The lower surface is remark- 
ably straight and flat, though sloping upwards at the sides. The outline, seen from 
above, is long and narrow, rounded behind, maintaining a tolerably uniform breadtli for 
the posterior two-thirds of its length, and acutely pointed at the anterior extremity. 
The upper surface is, except quite in front, concave, the vast hollow in which the so- 
called " head-matter " of the whalers (composed of nearly pure spermaceti) lies being 
limited behind by the occipital crest, continued laterally into the elevated edges of 
the Vjroadly expanded maxillae, which rise from the median line towards the margin of 
the skull, instead of felling away as in most Cetaceans. The great breadth of these 
bones in front of the antorbital notch takes away the appearance of a distinct rostrum 
or beak, generally characteristic of the long-snouted dolphins. 

* The skull of Baloena mysticetus is rather longer in proportion to the vertebral column, but it is less massive. 



JIB. W. II. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 315 

The absence of bilateral symmetry so generally met with in the skulls of toothed 
whales, is carried to its extreme in the Cachalot. It is chiefly manifested in the parts 
around the nasal passages. One of these orifices, the left, is immensely developed, the 
other reduced in a corresponding degree. But the distortion thus occasioned is not 
confined to the bones immediately concerned in the formation of these apertures, it 
affects the entire wall of the great supracranial basin, as seen in the upper view of the 
young skull (PI. LVII. fig. 1). 

As the indi\idual bones of the fcetal Cachalot's skull have been described by Professor 
Owen*, I will confine my present account chiefly to the structure of the diffc'rent 
regions of the cranium of the adult Tasmanian specimen, using the young skull from 
the same locality for illustration in cases where the ankylosed state of the former renders 
it impossible to make out the nature of the parts. I will also add some comparisons 
with the Hyperoodon, as one of the most nearly related of the ordinary toothed whales, 
and finally point out such differences as I have observed in the other specimens of 
Cachalot examined. 

Commencing with the cranial cavity, as the central point around which the whole 
head is developed, and of which a view was aff'orded by a median section of the skull 
(see PI. LVI. tig. 1, and woodcuts, figs. 12 and 13, p. 372), its general form is, as said 
before, roughly speaking, spherical, although slightly flattened on its upper anterior, 
and also on its lower posterior aspect. Its greatest diameter is diagonally from below 
upwards and backwards. The extent in this direction is best seen in the woodcut, 
fig. 12, as the cavity projects upwards for 1^" on each side of a median ridge, through 
which the section is made. The cerebral hemispheres must have a remarkable deve- 
lopment in this direction, projecting considerably beyond the cerebellum, which, as its 
limits appear to be indicated by a rather obscure nearly horizontal ridge, would have a 
small proportionate development. On the other hand, the magnitude of the apertures 
for the principal nerves, as well as the canal for the medulla oblongata, shows that these 
were of great relative size. The greatest transverse diameter of the cerebral ca\T.ty is 13". 

The planes and angles formed by the different parts of the wall of the cranial cavity 
are very remarkable. The lowest part is at the junction of the basioccipital with the 
basisphenoid. Behind this the basioccipital rises upwards at an angle of 45° from a 
horizontal line drawn from one end of the cranium to the other, so that the long and 
capacious canal in the occipital bone, leading to the foramen magnum, rises abo-^e the 
level of the brain-cavity. The basisphenoid is inclined slightly upwards towards its 
junction with the presphenoid. The concave anterior wall of the cavity formed by the 
united presphenoid and ethmoid is nearly vertical in its general direction. The anterior 
half of the roof, formed by the frontal, is straight (somewhat depressed in the young 
skull), and directed upwards and backwards at an angle of 45° to the horizontal line. 
The hinder part of the roof, formed by the occipital, arches downwards and backwards. 
» Cat. Osteol. Ser. Mus. Hoy. Coll. Surg. vol. ii. p. 442, 1853. 



316 MR. W. H. FLOWER OX THE OSTEOLOGY OF THE SPERM-WHALE. 

Ill the section of the large skull, the suture between the basisphenoid and presphe- 
noid, and also that between the ethmoid and the frontal, are distinct ; all the others are 
obliterated ; but they can be traced more or less distinctly in the young skull. 

The portions of the wall formed by the basioccipital below, and by the frontal above, 
are very thin, while those formed by the occipital above, and the conjoined presphe- 
noid and ethmoid in front, are of vast thickness. The portion of the frontal divided in 
the section belongs to the bone of the right side, which extends somewhat over the 
middle line. As in the dolphins generally, the parietal forms no part of the boundary 
of the brain-cavity in the middle line, but is seen to form part of the lateral wall of 
the cavity, resting on the alisphenoid, and having the frontal in front and the occipital 
behind. Above, it has so completely coalesced with the occipital, that even in the young 
skull its limits cannot be distinguished. The bones forming the periotic capsule are 
completely excluded from the brain-cavity, and in the adult skuU removed to a distance 
of 14" from it. By the aid of the mobility of the squamosal in the young specimen, a 
minute portion of this bone, ^" by J" in dimensions, can be traced in the lateral wall of 
the interior of the cranium, between the alisphenoid and the parietal. 

The foramina at the base of the cranium are only five in number. Several of the 
nerves find their exit from the cavity by common canals, which divide in passing through 
the immense mass they have to traverse before reaching their destination. 

1. An oval foramen, ^" in its greatest diameter, situated about 1" from the middle 
line, at the junction of the frontal and ethmoid*. It leads into a canal (9" long on the 
left side) which runs forwards and outwards, traversing the last-named bone, and 
opening into a wide fissure in the posterior part of the navial passage between the 
ethmoid and the vomer. On the right side, the foramen is smaller, and, owing to the 
conformation of the bones, the canal runs a much shorter course, opening rather behind 
the upper margin of the blowhole, between the frontal and the ethmoid. This would 
probably allow the exit of a small olfactory nerve, distributed in the simplest possible 
manner on the mucous membrane of the air-passage. In several dolphins there are 
similar channels through the ethmoid bone, though unaccompanied by any increase of 
extent of olfactory surface by turbinal bones as in the Whalebone-Whales. 

2. The exit of the optic nerve is by a comparatively small foramen in the anterior 
wall of the skull, transversely oval, 1" by "6", 2" from the middle line, 3" below the last- 
mentioned foramen, and about the same distance above the junction of the presphenoid 
with the basisphenoid. The course of the long canal to which this leads, is outwards, 
and slightly forwards. The distance from the wall of the brain-cavity to the upper 
margin of the orbit, is nearly 4 feet. 

3. A large oval foramen, situated between the orbitosphenoid plate of the presphe- 
noid and the alisphenoid, 2" in greatest diameter, and leading into a canal which passes 

* In the cast taken from the interior of the cranial Ciivity, figured at p. 372, this email foramen has yielded 
no impression. The rclatire size and position of the four others are well seen. 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 317 

outwards and forwards. In the young skull the division of this canal into three 
branches takes place close to the cranial cavity. The iirst represents the sphenoidal 
fissure ; the second and third, which perforate the alisphenoid, represent the foramen 
rotundum and foramen ovale respectively. 

4. Immediately behind and rather lower than the last is a nearly circular and mucli 
smaller foramen, perforating obliquely the outer side of the basisphenoid near its union 
with the alisphenoid. This appears to be the carotid canal. 

5. The remaining nerve-openings are collected into one, large, elongated, funnel-shaped 
canal, leading outwards and downwards from the side of the lowest part of the brain- 
cavity. At the bottom of this canal, and 14" distant from its commencement, is 
placed the organ of hearing. 

Upper surface of the Shdl. — The general shape of this region has been already 
described. We may commence a more special description by taking the narial apertures 
as central points. The left, which extends close to the median line of the cranium, 
is nearly circular iu outline above, having a diameter of about a foot, and is directed 
slightly forwards and to the left side. It contracts somewhat, and is more oval in form, 
below, with the long diameter fore and aft. At its narrowest part it is 1\" by 11|". 
Its upper margin is formed by the vomer on the inner side, the premaxillary in front 
and the outer side, the maxillary behind this on the outer side, and posteriorly by a 
rough spongy mass growing out from the left side of the ethmoid, forming a kind of 
operculum projecting over the narial opening (PI. LVI. fig. 1, e). In the young specimen 
this singular mass is not fully ossified, and is therefore a much less conspicuous feature 
in the skull (PI. LVI. fig. 2). Lower down, the vomer passes all round the back and 
nearly as far as the middle of the outer side of the passage, while the pterygoid forms 
the remainder of its inferior boundary. In the anterior and outer wall, a small slip of 
the palatal appears. 

The right blow-hole is placed nearer the hinder part of the skull than the left. It 
is an ii-regularly circular canal, with an average diameter of 3", directed upwards and 
backwards. The septum which divides it from the left is 3" wide at the narrowest 
part. 

The great semicircular wall which rises up behind the narial apertures is formed by the 
extremely compressed hinder portion of the maxillaries and premaxillaries, the frontals 
and the nasals, the whole being backed up behind and on each side by the supra- 
occipital, and perhaps containing some portion of the parietals concealed within. The 
maxillaries form the greatly thickened and sloping lateral edges of the crest. They rise 
to the highest part of it, but do not meet in the middle line by the space of more than a 
foot. Their inner edge is extremely thin. They present no special asymmetry in develop- 
ment. On the other hand, the premaxillaries of the two sides difier greatly : passing 
backwards along from the upper surface of the rostrum, where they lie on each side of 
the median vomerine groove, the left, turned out of its course by the great blow-hole, 

VOL. VI. PART VI. 2 Y 



318 ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 

ends abruptly in a narrow bifid extremity on a level with tire posterior margin of this 
passage ; the right continues onwards, and, passing the blow-hole, expands into a thin 
broad plate, applied to the anterior surface of the frontal, and reaching to within 6" 
of the summit of the crest. 

On tlie left side, corresponding to this plate, overlying the frontal, and resting below 
on the top of the ethmoid, is a large, flat, very thin, loose lamina of bone («). Its lower 
edge is thicker and rounded ; but it terminates above by delicate irregular digitations. 
This measures 14" in breadth, and rather more in height; but a considerable part 
appears to be wanting from its upper edge in the adult specimen, having probably been 
lost in maceration, or perhaps never completely ossified. It is more perfect in the young 
skull, and partially united to the frontal (see PI. LVII. fig. 1). This I take to be the 
left nasal bone, as it corresponds in situation and relations with that bone in the 
Hyperoodon, although, in common with the other bones of the crest, it is excessively 
flattened out. 

In none of the skulls examined could any trace of a right nasal bone be seen. Its 
development appears to be interfered with by the ascending plate of the premaxillary ; 
or it is possible that it is concealed beneath this. In the foetal skull in the Museum of 
the College of Surgeons, described by Owen and Huxley, the left nasal is absent ; pro- 
bably it was not ossified at this early age. 

In front of the blow-holes the upper surface of the skull is comparatively flat, 
although still rising in the greater part of its extent from the centre towards tlie sides. 
This region is formed by the premaxillaries, in the shape of a pair of long narrow bands 
of varying width, with very sharp edges overhanging the median vomerine groove, but 
mainly by the greatly developed rostral portion of the maxillaries. These bones are very 
massive, and expand in width in front of the deeply marked antorbital notch (a on). 
'I'heir flat upper surface is formed of a very thin plate of bone of remarkably dense and 
Inittle texture. Their internal structure is cancellous, the large distinct cells, almost 
like those of a honeycomb, being filled in the natural state with oil. 

The infraorbital foramen (if) is represented by a fissure 10" in length, and 2" in 
breadth, placed between the blow-hole and tlie antorbital notch, but nearer the latter. 
This gives passage to the great branches of the fifth nerve destined to supply the enor- 
mous upper lip and face. 

In the right premaxillary, 15" in front of the blow-hole, is an oval fossa, 3" in 
length, leading into a canal which runs outwards, and communicates with that leading 
to the infraorbital foramen. There is no corresponding opening on the left side. 

Lateral surface of the Skull (PI. LV.). — The temporal fossa is vei-y small, though 
scarcely so much reduced as in Hyperoodon. It is especially compressed from before 
backwards, lying deeply between the great projecting masses formed by the squamosa] 
behind and the orbital process of the frontal in front. Above, it has not any distinct 
limiting ridge as in Hyperoodon and most Dolphins, but passes almost insensibly on to 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 319 

the great convex surface formed by the occipital crest. The chief peculiarity of this 
region is the apparent suppression of the parietal bone, the squamosal and the frontal 
uniting in a vertical suture for more than the lower half of the fossa, and being 
separated by a wedge-shaped piece (jj) of the supraoccipital above. A faint superficial 
groove, more strongly marked, but still only a groove, both in the young skull and in 
the foetal skull but 34" long, indicates that this wedge-shaped piece may be really the 
parietal, ankylosed at a very early period to the occipital, even before the proper elements 
of the latter have coalesced. 

The orbit is small, oval, 6^" high and 11" long, with very prominent and distinct 
boundaries, complete, except for a space of 1^" behind, where it is continuous with the 
temporal fossa. This completeness and solidity of the margins of the orbit, especially 
of the lower side, is quite peculiar among Toothed Whales [Kogia even, not excepted), 
and depends upon the remarkable conformation of the jugal bone {j). This consists of 
two parts, meeting at an acute angle at the prominent rounded antorbital process. One 
(the body of the bone) is wedged in between the under surface of the orbital process of 
the frontal and the maxillary. The other is a strong process projecting freely back- 
wards along the inferior margin of the orbit, flattened from above downwards, and 
gradually narrowing behind, where it articulates by an oblique surface with the under- 
side of the end of the zygomatic process of the squamosal. This represents the styli- 
form part of the jugal, common to the Hyperoodon and all other Toothed Whales, 
though widely differing from it in character. 

None of the skulls examined showed any trace of a separation of the body of the 
jugal bone into two parts, as in Hyperoodon and the Ziphioids, where one of the two 
di\isions has been taken for the representative of the lachrymal bone. It is probable 
that the entire bone must be considered to be composed of the malar and lachrymal 
coalesced, as in the ordinary Dolphins. 

The orbital process of the frontal is longer and naiTower than in the Dolphins 
generally, approaching somewhat to the form it assumes in the true Whales. The 
supraorbital margin is much arched, and largely uncovered by the maxillary. The 
postorbital process is strongly marked and pointed, and, as before indicated, does not 
quite come in contact with the squamosal. The antorbital process is formed by the 
maxillary and the jugal, being cut at its most prominent part by the horizontal suture 
between these bones. In front of this is the deep antorbital notch. 

The side of the rostrum commences by a broad flat surface of the maxillary, a foot 
deep, looking outwards and upwards. The borders of this gradually approximate until, 
at about one-third of the length, they are luiited into a single sharp edge, much 
uptm-ned in the middle third, but gradually flattening towards the tip. The last 
twenty-two inches of the rostrum is formed by the premaxillary alone. 

Base of the Skull. — This region is chiefly remarkable for the extent and massiveness of 
the pterygoids, although falling short of that in the Hyperoodon in this respect. They 

2 y2 



320 ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEKM-WHALE. 

unite in the middle line for a space of 11", presenting a broad flattened surface. 
Behind this they become thin and separate from each other, the usual notch on their 
posterior free margin being represented by a deep narrow fissui'e, of which the sides are 
almost in contact, except at the bottom, whei'e it widens into a triangular space. 
Tlie external surface of the pterygoid is slightly hollowed, as in Ilyperoodon and the 
Ziphioids ; but there is no reflexion of the margin to form an outer bony wall to the 
postpalatal air-sinus, as in the ordinary Dolphins. 

The palate bones are largely apparent on the surface, much more than in Hyperoodon. 
The outline of the exposed surface of each is convex in front, and concave behind, where 
it is overlapped by the pterygoid*. The vomer is widely exposed for the whole of its 
length in front of the palate bones. A narrow strip of the premaxillary becomes 
apparent between the vomer and the maxillary near the middle of the rostrum, and 
gradually widens forwards. 

The greater part of the under surface of the rostrum is formed by the broad, convex, 
triangular maxillary — generally smooth, but having a strongly marked groove {d.(j) 
running longitudinally near the middle (evidently the remains of the dental groove), 
enlarged at intervals by the foramina for the passage of the branches of the superior 
maxillary nerve and artery, which supply the thick fibrous covering of the palate. It 
is probable that the rudimentary teeth concealed in the gum of the upper jaw, the 
existence of which has been repeatedly afiirmed and denied, are situated in or near this 
groove. 

Ten such teeth were sent with the present skeleton. Some of _i„ 

these are of hard, solid, yellow ivory ; but others are white and 
friable, splitting into concentric layers, as if they had been cal- 
cined. They are all between 2" and 3" in length, and about |" 
in diameter at the thickest part. Some are straight, but most of 
them are more or less curved, one forming a complete half circle. 
All have a distinct blunt conical crown, ^" long and from ^" 
to T^" of an inch in diameter, separated by a slight constriction 
from the expanded root, which constitutes the largest part of the 
tooth. The surface of the cro\vn differs from that of the fang in 
being slightly granulated. It shows no signs of attrition ; but 
the apex in all is roughly truncated, giving the appearance of RucUmontaiT^uaxmary 
having been broken off". The pulp-cavity is completely closed at tooth, uat. size. 

the base of the tooth, which, in most of the specimens, is more or less surrounded by 
rough, irregular, spiculated outgrowths. 

* -i thin and narrow plate of the pterygoid, with rounded margin projecting forwards, and partially covering 
the palatal, both in the adult and young skull (see Pl.LVII. fig. 2), is not ossified in the foetal skull figured 
in Huxley's ' Elements,' where the ptei-ygo-palatinc suture appears straight and transverse. 




MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 321 

The bones containing the organ of hearing have been figured by Camper* and 
Owenf . They are remarkable for their small size, compared with that of the cranium. 
Not only are they much inferior to those of the true Whales, but actually less than 
those of the common Killer {Orca gladiator). In general conformation the tympanic 
and petrosal bones do not differ from those of other toothed Whales ; their principal 
peculiarity is the development of a large mass of curiously laminated bone from the 
posterior and outer end of the tympanic, close to its attachment to the petrosal J. This 
is 6" long, and thicker at its outer than at its attached extremity. It is composed 
of a large number of distinct thin plates, only held together by their common attach- 
ment to the tympanic. The whole mass partly overlaps and embraces the hinder edge 
of the squamosal, and partly fits into a groove between the latter and the exoccipital, 
and serves to attach the petrotympanic much more firmly to the cranium than is the 
case with other Toothed Whales. It evidently coiTesponds to the strong tenon-like 
process of corresponding situation and functic»n in the Whalebone-Whales. The con- 
tiguous edge of the squamosal has a laminated character, the ridges and grooves on its 
surface exactly fittmg into those of the appendage to the tympanic. 

The petrotympanic is, as in most Dolphins, further steadied in its place by a long, 
narrow, flat process, which runs out from the squamosal downwards and backwards 
immediately in front of it. The length of the tympanic is 2"'6, the greatest breadth 
of the united tympanic and petrosal is 2"- 9. As might be expected, there is scarcely 
any appreciable diflerence in the size or form of these bones in the young and the adult 
animal. 

The principal dimensions of the cranium are as follows : — 

inches. 

Extreme length 201 

Extreme breadth (across posterior part of orbital processes of frontals) 87 

Extreme height (top of occipital crest to bottom of pterygoids) ... 65 
Length of rostrum (from tip to line drawn across bottom of antorbital 

notches) 14<j 

Width of commencement of rostrum (inside antorbital notches) . . Gl 

Width of rostrum at quarter distance from base, in straight line . . 58 



•->■?, 



■J 



Eight maxillary 

Left maxillary 24 

Right premaxillary 5 

Left premaxillary 6 

Space between premaxillaries -1 

* Op. cit. plates xxiv., xxv., and xxvi. 

t Brit. Foss. Mammals, p. .526. Natural size, though stated by mistake to be half the natural size. 

t In the specimens figured by Camper and Owen this process has been broken off. 



322 ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPERM-WHALE. 

inches. 

Width of rostrum at middle 47 

Right maxillary 13 

Left maxillary 13 

Right premaxillary 9 

Left premaxillary .... 9 

Space between premaxillaries 2 

Width of rostrum at three-quarters distance from base 21 

Right maxillary 2 

Left maxillary 2 

Right premaxillary 8 

Left premaxUlary 8f 

Space between premaxillaries ^ 

Premaxillaries extending beyond maxillarics 22 

vomer 43 

Antero-posterior length of orbital process of frontal 21 

Length of jugal 29 

Height of occipital crest above the bottom of the great supracranial 

basin behind ethmoid 42 

Height of occipital crest above upper edge of foramen magnum . . 85 

Width of occipital condyles 23| 

Height of right condyle (vertical) 14^ 

AVidth of foramen magnum at upper end, between condyles .... 8 

The crania of the different skeletons presented no very marked distinguishing features, 
beyond such as might depend upon age or indi\idual peculiarity. Of the adult, or 
nearly adult specimens, those from Caithness and Tasmania are most alike, the former, 
however, being rather larger and, especially, higher in the occipital crest. The York- 
shire specimen differs from both in the greater development of the rostrum, which is 
both broader and longer than in the others : and on this chiefly depends the immense 
size attained by this gigantic skull ; for the portion of the cranium behind the antorbital 
notches is of exactly the same length as in the Caithness, and but one inch longer than 
in the Tasmanian Cachalot. 

In the following Table the individuals are arranged according to their presumed 
ages, judging by the entire length of the skull. By comparing the figures showing the 
proportions, it will be seen that, taking the length of the skull without the rostrum as 
the basis of comparison, there is a relative increase in all the other dimensions during 
growth, but that both in height and breadth, especially the former, this increase is com- 
paratively slight and irregular, compared with the steady lengthening of the rostrum 
which occurs as age advances. 



ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPEEM-WHALE. 



323 





i 


Young; Tasmania. 
Mus.Eoj. Coll. Surg. 


Young; Sydney. 
(Wall.) 


.s 

1 






1 
1 

P 


Length of skull without rostrum . . 


14" 


24" 


34' 


57" 


55" 


56" 


56" 


Proportiou 


100 


100 


100 


100 


100 


100 


100 




Length of rostrum 


20 


38 


80 


139 


146 


156 


171 




Proportion 


143 


158 


235 


244 


265 


279 


305 




Extreme breadth 


18 


36 


60 


94 
165 


87 


92 


102 




Proportion 


128 


150 


176 


158 


164 


182 






15 


26 


44 


66 


65 


70 


67 






107 


108 


129 


116 


118 


125 


120 





Many of the differences of the skull, dependent upon age, are well illustrated in 
PI. LVI., where drawings of median sections of the crania of the young and adult 
Tasmanian Cachalots are given on the same scale. Extraordinary as the disproportion 
of the facial part of the skull to the cerebral cavity appears in the older skull here 
figured, a drawing of the Yorkshire specimen would show the same character in an 
even more exaggerated degree. 

In the same Plate a figure of the section of the cranium of a Hyperoodon has been 
introduced, as that of the Whale which (except Eogia) approaches most nearly in its 
general characters to Physeter. It is easy to see, by this section, how those fantastic 
and apparently meaningless developments of the cranial bones of Hyperoodon and the 
Ziphioid Cetaceans may become, with little modification, the regular and definitely dis- 
posed walls of the huge spermaceti-basin of the Cachalot. The crest, essentially the 
same in both, is merely flattened out and expanded, as if by pressure from within ; and 
the great maxillary protuberances are reduced in size. The most essential differences 
between the cranium of Physeter and the Ziphioids are, as already pointed out. the 
absence of a distinct lachrymal bone, and the construction of the zygomatic process of 
the malar. 

Lower Jaw. 
Perhaps no part of the skeleton of the Cachalot is so well known as the lower jaw. 
as few Museums of note do not possess one or more of these tangible trophies of a 

* Owing to the imperfect condition of this skiiU, the dimensions given cannot be relied 011 as quite accurate. 



324 



ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPEEM-WHALE. 



successful whaling-campaign. It will suffice hei"e to point out that, in this part of its 
organization, Physeter conforms with the other Dolphins in the vertically expanded 
hinder part of the ramus, and immensely wide opening of the dental canal (see PI. LVI. 
fig. 1), characters which separate them from the Whalebone-Whales. It differs from 
the true Bel^iMnidce in the excessive length and narrowness of the symphysial portion, 
and consequently great and sudden lateral divergence of the rami posteriorly ; but in 
this it resembles the Platanistidce, particularly Inia *. A special peculiarity is, that 
the rami appear never to become united by osseous ankylosis at the symphysis ; at least 
this is not the case in the oldest specimen that I have had an opportunity of examining. 
In the largest jaw in the Museum of the College of Surgeons, it was observed, when the 
rami were separated, that the contiguous surfaces were not flat, but that of the left ramus 
somewhat convex in its whole extent, fitting into a corresponding concavity in the right 
ramus. In the jaw belonging to the Tasmanian skeleton, the symphysial portion is 
not perfectly straight, but has a distinct lateral curve, the concavity towards the left. 
This is, however, an individual (though not uncommon) peculiarity. Instances are fre- 
quently met with among Cachalots of excessive curvature of the lower jaw, amounting 
to serious deformity f. 

The gradual increase in the length of the symphysis, compared with that of the 
entire jaw, and the relative decrease in width behind, as age advances, are illustrated by 
the dimensions of three specimens of different sizes in the Museum of the College of 
Surgeons. 





Entire 
length}. 


Length of 
Symphysis. 


Width 
behind. 


Mandible of the young skull from Tasmania 


49" 


21" 


31" 




Proportion 


100 


43 


63 




Mandible of Tasmanian skeleton 


174 


102 


72 




Proportion 


100 


59 


41 




Largest mandible from Tasmania, presented by Mr. Crowther . . 


194 


124 


75 


Proportion 


100 


64 


38 





The mandible of the Yorkshire Cachalot is almost identical in dimensions with the 
last of these three. 

The form and structure of the mandibular teeth, their changes with age, and mode 
of implantation are fully described in Owen's ' Odontography.' They present great 
differences both in number and character in different individuals. In the Tasmanian 



* See Description of the skeleton of Inia geoffrensis. Trans. Zool. Soe. vol. vi. p. 89 et seqq. 

t See Murie, ' Proc. Zool. Soc.' 1865, p. 390. 

J The length is taken from the apes to the middle of a line drawn across the posterior ends of the rami. 



MK. W. II. FLOWER OX THE OSTEOLOGY OF THE SPERM-WHALE. 325 

skeleton the teeth are all retained in their place by the tough iibrous gum, in whicli 
they are to a large extent imbedded, and which supplies the place of close-fitting bony 
alveolar walls. Those near the middle of the series are about 5" in length, of which 
not more than 1^" projected above the gum in the living animal. The crowns are 
conical, recurved, and pointed, showing but little signs of wear. The pulp-cavity is 
widely open at the base. At the hinder end of the series they become smaller, and 
more pointed; but the last on both sides has a flat and oval crown, scarcely projecting 
above the level of the gum. They are not symmetrically placed in pairs along the jaw, 
and axe even unequal in number on the two sides, as there are twenty-eight on the left, 
and but twenty-five on the right. 

In the young cranium presented by Mr. Crowther, obtained from a sucking Cachalot, 
killed by the side of its mother, the teeth were still concealed within the gum. Although, 
unfortunately, most of them had been lost in preparing the specimen, a suflScient number 
were preserved to show their general characters. These are simple, cylindrical, nearly 
straight, obtusely pointed, 1^" long and rather less than i" in tlieir greatest diameter. 
It is interesting to observe that they show no trace of an enamel covering to the apex, 
a point which has hitherto been one of uncertainty. Judging by the alveolar depres- 
sions at the bottom of the dental groove, there appear to have been 24 teeth on each 
side in this specimen. 

The largest jaw from Tasmania, in the Museum of the College, has 25 teeth on each 
side; two others from the same locality have 2G-26 and 24-23 respectively; and a very 
old jaw, with massive and much woin teeth, locality unknown, has but 21-20 ; and a 
small, but adult specimen (female 1), has 22-22. The exact number of teeth of the 
Caithness skeleton cannot be ascertained, as the anterior portion of the mandible is 
wanting. Beale gives 24-24 as the number in the Yorkshire skeleton ; but it is doubtful 
whether this statement refers to the actual teeth, or to the wooden models now in their 
place, on which of course it is impossible to place absolute reliance. 

Hi/oid Bones. 

The bones of the hyoid arch are very remarkable, not only from their great relative 
size, but especially for the peculiar breadth and flatness of the basihyal (l/h) and thy- 
rohyais (th) (Plate LX. fig. 1). 

The stylohyals (sh) are large, subcylindrical, and slightly curved, truncated at both 
ends, 25" long, and 4" to 4|" in diameter. 

The basihyal and thyrohyals are not ankylosed; and, judging by their opposed sur- 
faces, a considerable space occupied by cartilage must have existed between them. 
These bones are also distinct in the Yorkshire skeleton. The basihyal is nearly flat, 
though the under surface is somewhat concave from side to side, and convex from before 
backwards. It is 17" long and 18" broad. A truncated process projects forward, for the 
attachment of the cartilages connecting the stylohyals : this is not bifid, as in most Ceta- 

VOL. VI. PART VI. 2 z 



326 ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 

ceans. The bone is very thick at the sides, where the thyrohyals are attached, but 
becomes gradually thinner towards the posterior (slightly emarginated) border. 

The thyrohyals are somewhat triangular, with a thick, rounded, anterior and outer 
edge, and much thinner behind. The greatest length of each of these is 21", and the 
greatest breadth 12". 

The basi- and thyrohyals of the Yorkshire Cachalot, which alone are preserved, 
only differ from those of the Tasmanian skeleton in their superior size. The basihyal 
is 18|" long by 20i" broad; each thyrohyal is 26" long by 14" broad. 

In the breadth and flatness of these bones, Kogia alone (as ascertained from a cast 
kindly sent to me by Mr. Krefft, of Sydney) resembles Physeter. In the Hyperoodon the 
thyrohyals are broader posteriorly than in Dolphins generally, and, so far, present an 
approximation to those of the Cachalot ; but, on the other hand, the stylohyals are of 
quite a different form. The absence of union between the basi- and thyrohyals in an 
animal showing all other signs of maturity,.as tlie Yorkshii-e Cachalot, is a very peculiar 
feature among Cetaceans. 

Vertebral Column. 
The vertebrae of the Cachalot, especially as contrasted with the large Whalebone- 
AVhales, present generally a rough or, rather, rugged surface, and a coarse and some- 
what spongy texture. In all the bones, in fact, there is a tendency to the development 
of rough, tuberous and spiculated outgrowths from the surface, and also to irregular epi- 
physial ossifications in the cartilaginous portions of the bone, which afterwards become 
ankylosed, as around the carpal bones, and on the articular surfaces for the chevi-on 
bones on the lower surfiices of the bodies of the caudal vertebrae. The advancing 
ossifying surfaces have generally a much more spiculated character than in other Ce- 
taceans, being covered with pointed conical eminences, which surround the channels 
for blood-vessels. This is also particularly well exemplified in the partially ossified 
carpal bones (see PI. LXI. fig. 4). The ossification of the ends of the vertebral bodies 
presents a common character by which they can be distinguished from those of all other 
Cetaceans with which I am acquainted. When the epiphysis is removed, rather above 
the centre of the surface (in the dorsal region) is a depressed circular patch, in dia- 
meter rather less than one-third of that of the vertebra ; in this part the bone has a 
nodulated appearance, — rather conspicuous pointed tubercles, projecting directly out- 
wards, being scattered over it without definite arrangement (see woodcuts, figs. 5& 10). 
Outside of this patch, the more elevated surface is roughened by furrows and inter- 
mediate ridges of various lengths and sizes, but all arranged in a tolerably regular 
manner,- radiating towards the circumference of the bone. The epiphyses of course 
correspond to this surface, being thickest in the central part. On their outer side, or 
tlrat connected with the intervertebral substance, the limits of this thickened area are 
distinctly seen in the smoother character of the surface, which towards the margin is 



MR. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPERM-WHALE. 



327 



roughened by numerous concentrically placed ridges. This arrangement is found 
throughout the whole vertebral column, as far as the epiphyses are separable, on both 
anterior and posterior surfaces. In the caudal region, the internal thickened area of 
the epiphysis is more centrally placed, and occupies a larger relative area, its diameter 
being three-sevenths of that of the whole centrum. 

In a Hyperoodon of about corresponding age, when the epiphysis is removed from 
the centrum the ridges and grooves of the exposed surface radiate from a central point 
to the circumference, without any such depressed tuberculated area. 

As mentioned before, the entire number of vertebrae is fifty, of which, according to 
the usual method of division, seven may be reckoned as cervical, eleven dorsal, eight 
lumbar, and twenty-four caudal. In this enumeration the vertebra that bears the 
rudimentary last pair of ribs is counted among the dorsal (although in many characters 
it approaches one of the lumbar series), and the caudal vertebrae commence with the 
one that bears at its hinder end the first pair of chevron bones *. Placed in series with 
their bodies in contact, the vertebrae measure 30' 4", — the seven cervical being 1', the 
eleven dorsal 6' 8", the eight lumbar 7' 6", the twelve anterior caudal 11' 8", and the 
twelve posterior caudal 3' 6". 

The following Table gives the weights of all the vertebrae, as well as the length of 
the body, the greatest width (measured between the extremities of the transverse pro- 
cesses), and the gi'eatest vertical height (from the summit of the spinous process to the 
most depending part of the body). Corresponding measurements are given of the 
vertebrae of the Caithness and the Yorkshire specimen, as far as circumstances would 
permit: — 



1st cervical 

2nd-7th cervical ^united) 1 

1st dorsal J 

2nd „ 

3rd „ 

4th „ 

5th „ 

6th , 

"th „- 

8th „ 

9th „ 

10th „ 

11th „ 

1st lumbar 

2nd „ 

3rd „ 



Tasmanian Skeleton. 



Weight. 



lbs. oz. 

51 

68 o| 

29 

29 

29 

30 

31 

31 8 

32 8 
31 
34 

36 
34 8 

37 8 

38 



Length 

of 
body. 



in. 

6 

5| 

2| 

51 

7 

6| 

6| 

64 



n 

9| 



Extreme Extreme 
breadth, height. 



37i 
32" 
26| 
25| 

21f 

21 i 

20^ 

19| 

19| 

19 

23| 

31 

31 

311 

33 



in. 
18* 
22| 
21i 
22 
24 
25 
25| 
26| 
26| 
26i 
26^ 
28 
29 
30i 
30i 
324 



Caithsess Skeleton. 



Length 

of 
body. 



Extreme 
breadth. 



6| 
9 
5 
6 

n 

H 

7 



8 

9 

91 
10 
lOi 



36 

36i 

28 

24i 

23| 

221 

221 

21 

20| 

20 

22i 

27i 

31| 

32 

32 

32 



Extreme 
height. 



21i 

24 

22i 

23| 

26 

28 

284 

28j 

29 

29 

30 

3U 

32' 

33 

341 

35i 



ToKKSHiRE Skeleton. ' 



Lengtli 

of 
body. 



9 
6 



H 

6| 

7 



8| 

9 
10 
lOi 



Extreme 
breadth. 



in. 

40 

35i 

27" 

25 

25 

24 

23 



261 

31 

29i 

29 

29i 



Extreme 
height. 



in. 
17 



36 
35i 



* The arguments in favour of including this vertebra among the caudal series are given in a former paper, 
Description of the siceleton of Inia geoffrensix, p. 100, of the present volume. 

2z2 



328 



MR. W. H. PLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 



4th lumbar 

5th „ 

6th „ 

7th „ 

8th „ 

1st caudal . 

2nd ., . 

3rd „ . 

4th „ . 

5th „ . 

6th „ . 

7th „ . 

8th „ . 

9th ., . 

10th „ , 

11th „ . 

12th „ . 

13th „ . 

14th ., . 

1.5th „ . 

16th „ . 

17th „ . 

18th „ . 

19th „ . 

20th „ . 

21st „ . 

22nd „ . 

23rd „ . 

24th „ . 



Tasmanian Skeleton. 



Weight. 



Iba. oz. 

39 8 

40 
38 8 
43 
43 
46 
49 
58 
65 
66* 
58 
55 
45 
40 
34 4 
28 
20 
10 5 

4 13 

3 0| 

2 15 

2 7 

1 11 

1 

5 

3i 

2 

H 

0| 



Length 

of 
body. 



Extreme 
breadth. 



Extreme 
height. 



in, 

lOi 
lOi 

11 

IH 

11* 

12 

12 

12i 

12i 

12i 

12| 

12 

111 

ll4^ 

lOi 
9* 

n 

6 

4i 
4 

4 
3| 
34 
3 

2* 
2i 



33| 

34 

33i 

33 

32 

31 

29i 

27i 

26i 

23 

t 

17i 

16 

141 

125f 

12 
11 



7| 
7| 
6i 

5| 
5 

3| 
3i 
2i 
2 



30| 
32 
3H 
32| 
31 
31i 
31 i 
30 
29 
28i 
25i 
24 
22 
21 
20 
17i 
14| 
12| 
9i 

7| 
7 

54 
4| 
4 
31 

2| 
2 

n 



Caithkess Skeleton. 



Length 

of 
body. 



Extreme 
breadth. 



11 

Hi 
114 

12 
12 
12 
12 
12 
12 
12 
114 

11 i 
10| 

lOi 



44 
4 

3| 

3i 

91 
^i 

3 
2| 

95 
"8 

24 



Extreme 
height. 



33 
33 
33 
32 

t 
t 
t 
t 
t 
24 

9>)1 
4 

20 

17* 

15i 

13| 
12i 
12 
10| 



•^4 

74 
64 

54 
5 



354 

36 

341 
t 
t 
t 
t 
t 

28 

26 

244 

22 

21 

19 

184 

16 

134 

94 

8 

6| 

6 

5| 

H 

4| 
4 

34 



YoKKsnraE Skeleton. 



Length 

of 
body. 



Extreme 
breadth. 



lOi 
10| 

11 

111 

124 
124 

12' 
12 
12 
111 

11? 

104 

10 
94 
74 
54 

3| 

H 
3 

-1 

2I- 

2 
14 



Extreme 
height. 



m. 
294 
30 
30 
31 
314 
314 
314 
27 
254 
20| 
184 
16 
134 
13 
114 
11 
101 



"4 
S4 



54 
44 
3| 
2| 
14 



35i 

34 

32 

311 

32 

30i 

30 

30 

29 

27 

23 

22 

21 

194 

171 

144 

12| 

9| 

8 

7 

54 
4| 

44^ 
34 
3 

?l 
iff 



The Cervical Vertel/rw exhibit in a very marked degree the antero-posterior compres- 
sion so characteristic of the Cetacean neck. The length of the bodies of the seven 
vertebrse of this region taken together, when compared with their own width, or with 
the length of the whole vertebral column, appears to be less than in any other 
Mammal. 

The atlas is distinct from the other vertebrte ; the remaining six are united by their 
bodies and spines into one consolidated mass, which, in the case of the Tasmanian 
specimen, is further united with the body of the first dorsal vertebra. This disposition 
of the cervical vertebrae (the distinctness of the atlas and union of the posterior six) 
appears to be the rule in all Cachalots' skeletons, although unknown among other Ceta- 
ceans. In the most nearly allied forms, Kogia, Ey2)eroodon, and 3Iicroj)teron, they are 
all united into one solid mass, as in BaJmna among the true Whales. In Platanista, 



* The vertebrae in this region, as is usuallj- the case in Cetacean skeletons, contain much more oil than at 
other parts, which partiaUy accounts for their great weight. Elsewhere the Tcrtebrse were free from oil, and 
dry at the time they were weighed. 

t Processes so much injured that accurate measurements cannot be made. 



MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 329 

Inia, Pontoporia, Beluga, and Monodon* all are separate, as in the Balcenopteridce. In 
the other Delphinoids the atlas, axis, and generally one or more of the other vgutebra; 
are confluent ; if any are free, it is at the hinder end of the series. 

The form of the atlas of the Cachalot is very characteristic. The great vertical depth 
of the obliquely truncated transverse processes, and the comparative straightness of the 
upper and lower border, especially the former, give it, when seen either from before 
or behind, a transversely extended quadrangular figure, quite unlike that of any other 
Cetacean atlas. It of course partakes of the regional character of great antero-posterior 
compression, though not to the same extent as the succeeding vertebrae. The whole 
bone, with all its inequalities, will lie between two planes 8" apart ; and nowhere does 
the actual thickness exceed 6". The anterior surface is hollowed out to a depth of 4|" 
for the reception of the condyles of the occiput. The posterior surface is remarkable 
for its flatness. 

Fig. 2. 




Anterior surface of atlasf. 

The form of the neural canal, where it pierces the atlas, is very nearly that of an 
equilateral triangle, with one of the angles directed downwards. The upper side is 
almost straight, the outer angles rounded : the lateral sides converge rather rapidly to 
a point rather below the middle, where their posterior margins form an angular pro- 
jection, causing a constriction of the opening. Below this the sides approach more 
gradually towards the inferior angle, which is truncated at the apex. If the anterior 
margin of the aperture alone could be seen, it would appear more perfectly triangular, 
with straight sides. Seen from behind, the opening appears di\ided by the above- 
mentioned projection on the posterior margin into two parts, an upper transversely 
elongated oval portion, and an inferior narrow vertically elongated part. The former 
alone corresponds to the neural canal of the succeeding vertebrte ; the latter fits over the 
rough surface of the axis to Avhich the odontoid ligaments are attached, and affords a 
passage for them. 

* In two skeletons of male Narwhals in the Mus. Roy. Coll. Surg, the bodies of the 2nd and 3rd cerncal 
vertebrae are firmly united. 

t The woodcuts of the vertebrae (as the figures in Tlates LVIII. and LIX.) are all reduced to yL the size of 
nature. The anterior surfaces are represented in every case. 



330 ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPERM-WHALE. 

In the Caithness Cachalot the inferior part of the opening is altogether smaller and 
especially constricted laterally, the sides being nearly parallel and 1" apart for a 
distance of 2^". 

The articular surfaces for the occipital condyles are broad and shallow, without any 
sharply defined projecting border, except at the superior internal angle. They approach 
each other below, but do not meet by a space of about 2". The bone between their 
inner margins and the edge of the neural canal is hollowed into several very rough 
irregular depressions, especially on the right side. 

The neural arch, arising on either side immediately above the condyles, is a nearly 
straight horizontal bar of bone, slightly thicker from before backwards than from above 
downwards. Its anterior is thicker than its posterior edge. It presents no appreciable 
spine, but, on the contrary, is rather hollowed than otherwise above. There is, how- 
ever, a slight rounded prominence on the middle of the anterior margin. In the 
Caithness specimen this is more developed, forming an irregular, low, tuberous spine. 
Posteriorly, on the right side, near the root of the arch, is a nearly circular flattened 
surface, 4" in diameter, with an irregular depression in the middle, and of which the 
edges are developed above and below beyond the surface of the bone from which it 
springs. This facet closely ftts a corresponding one on the commencement of the arch 
of the axis. It is evidently irregular, no trace of it being present on the other side ; 
but it is worthy of note that a similar articulating facet between the atlas and axis, in a 
corresponding situation, but on the opposite (left) side, exists in the Caithness specimen. 

The neural arch of many Cetaceans and of other Mammals is perforated laterally by 
a large foramen, through which the first cervical nerve finds a passage. In some, the 
part constituting the anterior wall of the foramen, and which joins the upper edge of 
the anterior articulating surface, is absent ; and the foramen is then represented by 
a deep groove with more or less overhanging edges. In Jlt/iJeroodon the foramen is 
complete ; in Orca the same ; in Glohiocephalus very nearly so ; also in some of the 
true jDelphini, as D. tursio. In Belwja, Platanista, and Phocwna it is a mere groove. 
In the Cachalot the last-named condition is found, though the groove is relatively 
smaller and shallower than in other genera. It is bounded in front by the sharp, 
prominent upper edge of the condylar articular surface, behind by the lateral part of 
the neural arch; internally it descends into the neural canal at its upper and outer 
angle, and externally is gradually lost in the anterior surface of the upper part of the 
transverse process. 

The inferior edge of the bone presents a tolerably regular curve, the middle part 
descending 4 inches lower than the sides. When seen from below, it appears 
slightly hollowed in the middle in front, and posteriorly presents a broad obtuse 
triangular prominence, which fits into a corresponding depression in the axis. A 
similar process of the atlas occurs in all other Cetaceans in which this bone is separate ; 
but in the Cachalot it is shorter and more massive than in Beliuja, Monodon, or 



ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPEEM-WHALE. 



331 



Platanista, and, unlike these, has no smooth articular facet on its hinder and upper 
surface. 

There remain now to be described only the lateral processes, which constitute a very 
peculiar feature in the bone. In all other Cetaceans the transverse processes of the 
atlas, whether confluent or not with those of the axis, bear but a small proportion in 
vertical height to the whole bone, but appear as more or less conical (generally 
obliquely flattened) projections arising opposite to the middle portion of the articular 
surface, or, as in Hyperoodon, from near the lower edge. In the Cachalot they form 
two short, but very deep, vertically placed crests, rising as high as the top of the 
neural arch, and extending below almost to the level of the inferior edge of the 
condylar articular surface. At the ends they are obliquely truncated, being longer at 
the lower than the upper angles. The middle of the posterior surface near the outer 
edge is hollowed. The external margin seen from the side appears thicker below than 
above ; it is rough and nodulated, especially near the inferior angle, being probably 
not completely ossified. It will be observed in the annexed table of dimensions that, 
although the atlas of the Caithness Whale is in all other respects somewhat larger 
than that of the Tasmanian, the lateral development of the transverse processes is not so 
great. This is chiefly due to their being vertically truncated, without any production of 



the inferior angle. 



Dimensions of the Atlas *. 





Tasmanian. 


Caithness. 


Extreme breadth 


in. 

37i 
23 
8i 
l-H 

9| 
10 

4 

?• 

18i 
14 

5 

2i 

f 

25 


in. 
36 
26 
10 

i? 

lOi 
2i 

1 
1 

7j 
21j 

3i 

H 


Width between outer edges of anterior articular surfaces 

Extreme width of each of these surfaces 


Extreme height of each of these surfaces 


Width of neui'al canal 


Height of neural canal 


Height of contracted lower portion of the canal 


Width of contracted lower portion of the canal at the upper end . 
Contracting below to 


Height of portion of bone below the neural canal 


Extreme height in middle line 


Vertical height of transverse processes at outer end 


Greatest antero-posterior thickness of the same 


Height of neural arch in middle 


Antero-posterior thickness of neural arch in middle 


Length of inferior surface of bone in middle line 


Width between the outer edges of posterior articular facets .... 



* My friend Mr. J. W. Clark, Superintendent of the Cambridge Anatomical Museum, has kindly given me a 
drawing of the atlas of a young Cachalot, stranded at Hartlepool more than two centuries ago, and which is now 
preserved, with other bones of the same animal, in the crypt beneath the library of the Cathedral at Durham. 
The extreme width of this atlas is 29", its height 15". A comparison of this specimen with that of the Tasma- 
nian and Caithness Cachalots, as representing three different ages, shows that the principal change which takes 
place is the gradual contraction of the lower part of the central opening, that part, below the true neural canal 
which corresponds to the odontoid surface of the axis. 



332 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 



The remaining six cervical vertebrae are completely soldered together, both by their 
bodies and their neural spines, into one mass. Their individuality can be traced 
distinctly enough at the root of the neural arches, where, for a short space, they are 
separated to admit of the passage of the cervical nerves ; but their conjoined spines 
present a solid mass in which no trace can be detected of separate vertebral elements ; 
and their bodies are almost as completely fused together, slight grooves and rows of 
foramina for blood-vessels on the lateral parts faintly indicating the limits of the several 
component vertebrae. The union of the seventh to the antecedent vertebrae is more 
complete than in any other known Cetacean ; for even in the Hyperoodon the whole 
of its neural arch is free. To this mass, in the Tasmanian specimen, the first dorsal 
vertebra is partially united by its centrum only. 

The most remarkable characteristic of these vertebrae taken together is their extreme 
antero-posterior compression, the four middle bones being most affected. The greatest 
length, the lateral part of the conjoined bodies, is 9" ; and the whole group will lie between 
two parallel planes no greater distance apart. The conjoined centrums are somewhat 
flattened from above downwards, and very broad from side to side at the anterior end, 
but less so posteriorly. 

Fig. 3. 




Anterior siu'face of second cervical vertebra. 



The anterior surface is, in the main, flat. Its median portion, 8" broad, is irregular, 
rough, and slightly raised ; in the centre of this is a ridge-like prominence, placed with 
its longest diameter vertically, and raised not more than f" above the surrounding 
bone ; this represents the odontoid process. The lower edge is smoothly hollowed 
out in the middle to receive the process on the contiguous portion of the atlas. 
On each side of this hollow and of the median rough surface are the comparatively 
smooth, slightly depressed, and nearly flat articular facets for the atlas, of an irre- 
gularly quadrilateral figure, each measuring 10" in height, and 8" in width. They 
extend quite to the lower edge of the bone. Beyond these externally are the flat 
anterior surfaces of the broad, obtusely pointed, transverse processes, projecting 5" from 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 333 

the outer margin of the articular facets. The broad flattened lateral parts of the 
neural arch rise from directly over the upper edge of the articular facets, and converge 
rapidly together to form the anterior surface of the broad rugged mass constituting tlie 
neural spine. On the right side is a raised flat surface corresponding and fitting to 
that described on the commencement of the arch of the atlas. The opening of the 
neural canal, as seen in this aspect, is a transversely elongated lozenge, with the angles 
rounded oft', 8^' broad, and 6^" high. The inferior margin is very indistinctly marked 
in the middle, — the anterior surface of the axis abo^e the rudiment of the odontoid 
process gradually passing into the flattened floor of the neural canal, which contuiues to 
rise throughout the cervical region. 

The sides of the mass formed by the conjoined bodies slope gradually downwards 
and inwards, converging towards the middle line, where they meet in a slightly elevated, 
rounded, longitudinal keel, in which all trace of the original separation of these vertebrae 
and even of the first dorsal is entirely lost. 

As in the Toothed Whales generally, the transverse process of the axis consists of a 
single, broad, imperforate plate, springing from the gi'eater part of the side of the body 
of the bone and the lower part of the neural arch, representing, in situation at least, 
the upper and lower processes found in the succeeding vertebree (and in the axis of the 
Whalebone-Whales), coalesced and with the intermediate space tilled up. In relation 
to the large size of the body of the bone, these processes may be considered short : 
the condition of the ends, in the Tasmanian specimen, shows that they have not quite 
attained their complete ossification ; but they are only very slightly longer proportionally 
in the completely adult Caithness Whale. They are much compressed from before back- 
wards, and obliquely truncated externally, the nearly straight end looking upwards and 
outwards. In the older specimen more advanced ossification of this apophysis has caused 
the end to approach nearer to a vertical line. 

In most Cetacea the inferior trans\erse process of the cervical vertebne {parapophysis, 
Owen), arising from the side of the body, increases in development from the third to 
the sixth, and suddenly becomes obsolete, or nearly so, in the seventh, where the 
articular facet for the head of the first rib appears as it were in its place, situated, 
however, not precisely at the same spot on the side of the vertebra?, but rather above 
and posterior to it. In all known genera of Delphinoids the inferior process of the 
sixth vertebra attains a considerable development, most strikingly so in those in which 
the vertebrae are free, as Beluga and, especially, Flatanista. In Hyperoodon it is very 
conspicuous, although the third, fourth, and fifth show no rudiment of the process. 
In that genus also, at least in one example (Mus. Roy. Coll. Surg. No. 2480 a), con- 
trary to the general rule, a tolerably long inferior process is developed from the body of 
the seventh vertebra, but on the right side only. In the Tasmanian Sperm- Whale there 
is no trace of an inferior transverse process on the smooth sides of the bodies of any of 
the cervical vertebrae as far as that which appears to be the sixth, inclusive. The lower 

VOL. VI. PART VI. 3 A 



334 JME. W. H. FLOWER OX THE OSTEOLOGY OP THE SPERM-WHALE. 

part of the side of the posterior end of the conjoined bodies, or that part which appears 
to be connected with the seventh neural arch, and may therefore be regarded as re- 
presenting the centrum of the seventh vertebra, occupies a space as large as the three 
antecedent vertebrae together (deciphered by the same test), and is raised into two 
rugged ridges with a groove between, of which ridges the anterior is rather the more 
prominent. Besides these, there is no indication either of process or of articular facet 
for the first rib. As the more posterior of these ridges is situated quite at the edge of 
the vertebral body, and rather higher up than the other, it may be regarded as the 
representative of this facet ; and the rib may have had a ligamentous connexion with it, 
for the form of its head would not allow it to come in contact with the bone itself. 
The other is probably a rudiment of an inferior transverse process. 

A distinct upper transverse process (dicqiopJii/sh, Owen) is present only on the 
seventh vertebra. It springs from the middle of the side of the neural arch by a base 
of about 2" in breadth, and is of the same length, irregularly triangular, and very 
much compressed. Certain small irregular projections from the edges of the deli- 
cate lamellsB of bone which constitute the lateral parts of the neural arches of the 
third, fourth, fiftli, and sixth vertebra? arc the only representatives of their transverse 
processes. 

The neural arches of these vertebrae may be considered in the two portions into which 
they naturally resolve themselves. 1. The lateral portions, springing up from the con- 
joined bodies. 2. The united mass forming the roof of the neural canal, composed of 
the conjoined spines. The lateral parts of the arches are, as before said, all distinct. 
The first (that of the axis) far surpasses the others in breadth and in stoutness. Next 
to it the last or seventh is the best developed. Between these two are placed four 
delicate brittle lamellae, scarcely thicker than cardboard, the third and sixth being 
rather stouter than the two middle ones. So brittle are these plates, that in neither of 
the three sets of cervical vertebrae examined have the whole of them escaped destruc- 
tion in the cleanmg-process ; suflScient remains, however, in every case to show that 
their general arrangement is the same. The intervals between them are of nearly equal 
width (^"), but diminish in height from before backwards, the first being 2^", and the 
last scarcely more than 1" high. The upper part of the arch is formed of a transversely 
elongated rugged mass of bone, flattened from before backwards, with two prominent 
square shoulder-like lateral projections rising from the antei'ior surface, and with a 
small pointed spine in the middle line sui-mounting the posterior edge. Between this 
spine and the two shoulders rising on each side and in front of it is a distinct groove. 
These lateral expansions, which appear to belong chiefly to the axis, are, among Tootlicd 
Whales at least, quite peculiar to I'hyseter. Even Hi/peroodon shows no trace of tliem, 
as the conjouied neural arches of the first six cervical vertebrae rise smoothly and 
gradually into a greatly elevated spinous process. The axis of the Balwnopteroe presents 
rugged lateral processes somewhat similar to those of the Sperm-\Miale. 



ME. W. H. FLOWEK ON THE OSTEOLOGY OF THE SPERM-WHALE. 



335 



As the posterior surface of the body of the cervical vertebrae is ankylosed with 
the first dorsal, little can be said of its characters. The centrum, however, is veiT 
deeply concave, recei\-ing the convex anterior surface of that of the first dorsal ; and 
there is a smooth articular facet (j'ostefwr zyg apophysis) at each side of the neural 
arch. This facet is more vertical (inclining less backwards at the upper end) than 
in other Cetacea. 



Measuiements of the conjoined six posterior Cervical Vertebrae. 





Taemanian.! Caithness. 


Extreme breadth 


in. 

32 

22| 

23 
8i 
6i 
9J 
4 

H 
9 

10| 


in. 
36i 
24 
25 
10 

m 

8i 
IH 
15 

m 


Extreme height 


Breadth between outer edges of articular facets for atlas 

Breadth of anterior opening of neural canal 


Height of anterior opening of neural canal 


Height of body of bone below this 


Antero-posterior length of superior surface (floor of neural canal). 
Antero-posterior length of inferior surface 


Antero-posterior length of lateral surface 


Breadth of posterior surface of body of seventh cervical vertebra . 
Height of posterior surface of body of seventh cervical vertebra . . 



The Dorsal Vertehrce (Pis. LVIII. and LIX.) are eleven in number. The first ten 
support well-developed ribs ; the eleventh, which in many of its characters resembles 
a lumbar vertebra, has only a rudimentary pair of ribs attached to the extremity 
of the transverse processes. The first dorsal is ankylosed by the middle part of its 
centrum (at all events at the upper and lower margins near the middle line, where no 
trace of a suture remains) to the hinder part of the conjoined six posterior cer- 
vical vertebrae. The lateral parts of the centrum are free, as also is the whole of the 
neural arch. 

The body of the first is by \ery far the shortest in the antero-posterior du-ection ; the 
second is nearly twice as long ; and they continue to increase gi-adually and progressively 
throughout the series, as will be .seen by the table of measurements. The body of the 
first is extremely concave on its posterior aspect, the middle part being 3" deeper than 
the sides. The second is convex in front, and concave behind to a less degi'ee. In the 
succeeding vertebrae the anterior and posterior surfaces are nearly flat and parallel. 
The bodies, at fii'st very broad in proportion to their height, rapidly become narrower ; 
in the fourth the breadth is already less than the lieight ; the sixth, seventh, and eighth 
are the narrowest ; after these there is a slight increase of width. 

The neural canal in the first two vertebrae is triangular, but rapidly assumes a trans- 
versely oval form, and gi-adually diminishes in both height and breadth, so much more, 
however, in the last dimension that in the tenth vertebra the long diameter of the oval 
is vertical ; in the eleventh the lateral contraction is still more marked. 

3a2 



336. 



JIR. W. H. FLOWEE ON THE OSTEOLOGY OP THE SPERM-WHALE. 



The spinous processes progressively increase in height from before backwards. That 
of the first is a small irregular tubercle broader than long, and scarcely higher than that 
of the conjoined cervical vertebrae. That of the second is still inconspicuous, but more 
compressed from side to side. The great antero-posterior width characteristic of the 
remainder begins to be seen in the third. The spines of the fifth, sixth, and seventh 

FiK. 4. 




Anterior surface of the fourth dorsal vertebra and head of right rib. 

scai'cely differ in form and dimensions, being, when seen from the side, nearly square. 
From the eighth a more rapid increase in vertical height takes place, until the last, 
by its elevation, narrowness at the base, and expansion at the extremity in the antero- 
posterior direction, as well as in thickness, resembles that of one of the lumbar vertebi-ae. 
As far as the sixth their general duection is vertical ; afterwards they have a slight but 
gradually increasing backward slope. 

Well developed rough processes, which can hardly be called articular surfaces, but 
representing the posterior zygapophyses, are formed on the hinder edge of the sides of 
the neural arch, at the root of the spine, from the fii'st to the ninth vertebra inclusive. 
Jn the tenth they have almost lost their distinctive character, and they are quite obsolete 
in the eleventh. They lie above and within the prozygapophyses of the succeeding ver- 
tebra ; but, except in the case of the first two or three, they appear very rudely coadjusted, 
compared with those of ordinary Mammalia. 

The prozygapophyses are at first represented by a fiattened stirface on the angle 
formed by the junction of the pedicle, lamina, and transverse process of the vertebra (to 
use the familiar terms of anthropotomy), or at the root of the diapophysis. Gradually 
this part increases in prominence, and forms a distinct rounded eminence, projecting 
upwards and forwards from the side of the neural arch, and bearing the smooth 
articular surface (distinct as far as the tenth vertebra), to its imier side. The sudden 
diminution of the diapophysis on the tenth vertebra, and its disappearance on the 
eleventh (see figs. 7 and 8), leaves this, though somewhat contracted in bulk, a con- 



MB. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 



33'; 



spicuous and important process on the nem-al arch, corresponding with those which in 
the lumbar region form such a characteristic feature in the Cetacean vertebral column. 
Owen has pointed out, in the description of the skeleton of Belphinus delphis (Cat. 
Osteol. Prep. Mus. Roy. Coll. Surg. vol. ii. p. 450), that these processes belong to tlie 
metapophyses of his system, as, although near their commencement they bear the 
so-called prozygapophyses, they soon become distinct from them. This is less readily 
demonstrated in the Sperm-Whale. 

Upper transverse processes for the articulation of the tubercles of the ribs (diapo- 

Fig. 5. Fig. 6. 





Eighth dorsal vertebra, epiphysis removed. 



Ninth dorsal vertebra. 



physes) exist from the first to the tenth vertebra inclusive, arising in all from the side of 
the neurapophysis, at nearly the same height tliroughout. They gradually decrease in 

Fig. 7. 




Tenth dorsal vertebra. 



length from the first to the tenth (see PI. LVIIL). The first is compressed from before 
backwards ; tlie next two subcylindrical ; the following four are very short, thick, and 



338 



MH. W. H. FLOWEE OK THE OSTEOLOGY OF THE SPERM-AVHALE. 



almost confluent with the bulky, rounded metapophyses. After this the transverse process 
oiadually resolves itself more distinctly into the ascending and forward-directed rounded 
metapophysis, and the slightly descending and backward-directed diapophysis. The 
latter rapidly diminish in bulk, the last (tenth, fig. 7) being a comparatively slender 
conical process which, on the right side, meets within a line's breadth an ascending 
tubercle of the inferior transverse process to be presently described ; on the left side 
the end of the process appears to be broken oflf. On the eleventh dorsal vertebra there 

Fig. 8. 




Elc've7ith dorsal vertebra and riglit rib. 

is no trace of diapophysis, except perhaps a slight thickening on the corresponding part 
of the right neurapophysis. The free extremities of all these transverse processes (except 
the tenth) present roughened articular facets for the tubercles of the corresponding ribs. 
In the first this facet has a compressed oval figure, with the long diameter vertical ; 
in the second it is subcircular ; in the third, fourth, and fifth triangular; in the 
remainder o\al, with the long diameter antero-posterior. 

On the upper pait of the sides of the bodies are conspicuous articular fiicets for the 
reception of the heads of the ribs. These deserve particular attention for the pecu- 
liarities they present in the hinder part of the region. They are disposed as follows : — 
The first vertebra has one facet on its hinder edge for articulation with the second rib. 
It is in the form of a low rough tuberosity. The second vertebra has a cup-shaped 
depression, with elevated margins on its hinder edge, for the third rib. The third 
vertebra has a similar facet for the fourth rib. The fourth vertebra receives in the 
same manner the fifth rib. The fifth vertebra shows on its anterior edge an indication 
of an articular surface for the hinder part of the fifth rib, and a large facet on its 
hinder edge for the sixth rib. The sixth vertebra has a more strongly marked 
articular facet on the anterior edge, while that on its hinder edge (for the seventh rib) 
is of reduced dimensions. In the seventh vertebra the articular surface near its front 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERJI-WHALE. 339 

edge, for the seventh rib, is raised upon a small tubercle ; wliile the posterior surface, 
for the eighth rib, is still further reduced and flat. In the eighth (tig. 5) the anterior 
facet is still more raised, and the posterior is nearly obsolete. In the ninth (fig. (!) the 
articular surftxce on the anterior edge of the body is developed into a distinct process, 2" 
in length, springing from the anterior half of the side of the body, obliquely truncated, 
and having on its outer and anterior surface an oval excavated articular facet, which 
bears the head of the ninth rib. There is no trace of any articular surface on tlie pos- 
terior edge for the tenth rib. In the tenth vertebra (tig. 7) the above-described tubercle, 
developed from the anterior articular facet, takes the shape of a long massive process, 
C" in length and nearly as much in antero-posterior thickness, S2)ringing from the 
middle of the side of the body, slightly ascending, compressed from above downwards, 
more expanded at the end than the middle, ending in a large, oval, conca\e articular 
facet for the tenth rib, and having at its upper surface, near the extremity, a sub- 
conical, compressed (from before backwards) process, which rises to meet the small 
subcylindrical upper transverse process, approaching so closely on the right side as to 
be separated by scarcely a line's breadth. The eleventh vertebra (tig. 8) has no articular 
facets on the side of the centrum ; but the process arising from this part on a level « ith 
that of the last-described vertebra, is still longer and more compressed, in fact precisely 
resembling in situation and general character those found on the lumbar vertebrae, but 
having at its extremity a small rough articular surface for the attachment of the 
rudimentary eleventh rib. 

In the Caithness and Yorkshire skeletons the same essential characters are found, 
differing only in details arising chiefly from more advanced age. In the Yorkshire speci- 
men, in the ninth dorsal vertebra, the inferior process is so far developed as to meet the 
upper one, forming a complete ring of bone on both sides. In the Caithness Whale this 
ring is complete only on one side in the ninth, but on both sides in the tenth vertebra. 

In all known Cetacea (with the few exceptions to be presently mentioned) the 
transverse process which arises in the fore part of the dorsal region from the side of 
the neural arch, and is evidently serially homologous with the upper transverse pro- 
cess {(liapophijsis) of the cervical vertebroe, falls gradually and almost insensibly in 
its point of origin from the vertebra, until, leaving the arch, it comes to be placed upon 
the body of the vertebra, and is perfectly continuous serially with the transverse pro- 
cesses of the lumbar vertebrae, which, from their situation, would be taken to represent 
the inferior processes [pampoplnjses) of the cervical region. We tind, moreover, after 
leaving the region of the neck, no trace of two lateral processes on the same vertebra. 

The remarkable peculiarity of the Cachalot's spinal column is, that, tracing the upper 
transverse process backwards from the neck, it never descends from its original position 
on the arch, but, after a great reduction in importance, it completely disappears in the 
eleventh vertebra ; while, on the other liand, a new process, springing from the side of the 
centrum of the eighth or nintli vertebra, and being at tirst only a development of the 



340 MR. W. H. FLOWEE ON THE OSTEOLOaT OF THE SPEKM-WHALB. 

articular facet for the head of the rib, gradually becomes more distinct, and ultimately 
forms the main transverse process, serially continuous with those of the lumbar region. 
We have thus in the hinder part of the dorsal region of the Sperm-Whale a perfect 
repetition of the characters of the cervical vertebrae of ordinary Mammals, as far as the 
transverse processes are concerned, an upper one springing from the arch, a lower one 
springing from the body, and uniting at their extremities so as to form a complete bony 
ring. 

In this important character of the spinal column Hyperoodon agrees with Physeter ; 
and this appears to me one of the most striking points of affinity between these two 
genera. In the skeleton of the Hyperoodon in the Huuterian Collection in the 
College of Surgeons, upper transverse processes are regularly developed from the arches 
of the dorsal vertebrae as far as the seventh, but cease in the eighth. The lower pro- 
cesses arising from the body begin in the seventh, which has thus two processes, which 
(in this immature specimen at least) do not meet at the ends; but the bony ring is 
completed by the attachment of the proximal end of the rib to both processes, 
recalling exactly the condition of the cervical vertebrae of the Crocodiles. In several 
adult Hyperoodons skeletons, I have seen the upper and lower transverse processes of 
the seventh dorsal \ertebra united so as to form a complete ring. 

It will be very important to ascertain whether Kotjia and the Ziphioids agree in this 
respect with Physeter and Hyperoodon, as certainly might be predicated from their 
general affinities. With regard to the first-named, the only published description* of 
the skeleton at Sydney gives no information on this point ; but the unique skeleton of 
the Micropteron sowerbyense, belonging to the last-named group, now in the Royal 
Museum at Brussels, shows the following characters : — The seventh dorsal vertebra has 
a transvei'se process springing from tlie arch ; to this the tubercle of the seventh rib is 
articulated, while its head joins the body in the usual way. The eighth vertebra has 
no process from the arch, but one projecting from tlie body at a level with the facet 
for the attachment of the head of the seventh rib ; to this the end of the eighth rib is 
fixed. The transposition of the transverse process from the arch to the body is thus 
as abrupt as in Physeter and Hyperoodon, the only difference being that the two pro- 
cesses do not coexist on any one vertebra as in those genera. Tlie principle, however, 
is the samef. 

Lumbar Vertebra. — The eight lumbar vertebrae present a remarkable similarity to 
one another, both in form and dimensions. Their bodies increase slightly but pro- 
gi-essively in length from the first to the last. One of their most characteristic features 
is the form of their infero-lateral surfaces, much hollowed out and converging to a strong 

* Wall, op. cit. 

t For the condition of the rib-attachments in Platanistti and Inia. in some respects intermediate between 
those of the Physeteridm and Delphinida, see " Description of the Skeleton of Inia geoffrensis," &c.. Trans. 
Zool. Sno. vol. vi. pp. 98-103. 



ME. AV. H. FLOAVER OX THE OSTEOLOGY OF THE SPEEM-WHALE. 341 

keel ninning along the middle line below. They are also flattened between the neural 
arch and the transverse processes, so that the general form of the end of the body of 
one of these vertebrae is pentagonal, or like a lozenge with the upper angle cut off. 
The first of the series is rather dift'erent in form, having no distinct keel. In the older 
individuals, as the Caithness specimen, the keels of the lumbar vertebrae are developed 
to a still greater extent than in the Tasmanian skeleton. 

Fij?. 9. 




Fifth lumbar vertebra. 

I'he transverse processes are comparatively short (considerably less than the dia- 
meter of the centrum) ; they have a very slight downward inclination, are of moderate 
breadth, flat, and roundly truncated at their ends. They increase slightly in length to 
the fifth, and then slightly diminish. 

The spinous, processes are high and broad, with a moderate slope backwards ; tlie 
club-like lateral expansion noted in the upper end of the spinous process of the last 
dorsal vertebra is seen also in the first lumbar, though in a less marked degree, and 
gradually subsides in the following two or three. The height of the spine of the last 
lumbar vertebra presents a considerable diminution. As in the Cetacea generally, the 
metapophyses are strong, well-marked processes, projecting upwards and forwards from 
the neural arch, embracing the hinder margin of the spine of the antecedent vertebra. 
They do not form such broad expanded plates as in the true Whales ; but, on the other 
hand, they are more distinct than in many other Delphinoids. One great peculiarity 
which distinguishes them from most others, even including Hyperoodon, is their 
gradual elevation upon the neurapophyses in passing from before backwards. In tlu- 
first lumbar vertebra they arise from the laminae of the neural arch by the sides of tlie 
canal, and their upper edge is 6|" above the top of the body ; in the last they spring 

VOL. VI. PAKT VI. 3 B 



342 ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPBEM-WHALE. 

liom the fore part of the spine, altogether above the junction of the lamina) at the roof 
of the neural arch, and their upper margin is 11" above the top of the bodj'. There 
are not any corresponding processes on the hinder edge of the spine, as in Platanista. 

The Caudal Vertebrm are twenty-four in number, of vvliich twelve or more appear, by 
the form of their inferior surfaces, to ha-se supported chevron bones, although but ten 
of these bones were sent with the skeleton. 

As is usual in this extensive region, the characters of the different vertebrae vary 
greatly. The first differs but slightly from the last lumbar vertebra. The centnrm is 
more regularly circular when seen from the end, but rather higher than broad, the keel 
is no longer distinct, the sides are less hollowed, and on the hinder edge of the under 
surface are two distinct articular facets for the first chevron bones. The transverse 
processes are short, horizontal, and rounded at the ends, as in the lumbar region ; the 
spinous process is slightly shorter, the metapophyses higher upon it, and the neural 
canal more contracted. The body of this vertebra is slightly larger than that of the 
last lumbar. In the succeeding seven there is very little change in respect to size ; after- 
wards the diminution is more rapid. The reduction in the size of the centrum of the 
thirteenth as compared with the twelfth, and of the fourteenth as compared witli the 
thirteenth, is so extraordinary that, unless there were abundant evidence to the con- 
trary, one might be tempted to suppose that several vertebrae had been lost from each 
interval. The fifteenth, again, is considerably smaller than its predecessor. Then follow 
two of almost equal size, after which a gradual and steady reduction takes place down to 
the terminal vertebra. 

Fiff. 10. 




Fourth caudal vertebra, witliout the epiphysis. 



The great irregularity in the diminution of the caudal vertebrae will be best appre. 
ciated by a reference to the table of the weights of the different members of the series 
given at p. 328. 



MK. W. H. FLOWER ON THE OSTEOLOGY OF THE SPERM-WILALE. 343 

The neural arch forms a distinct canal as far as the fifteenth caudal vertebra ; in the 
sixteenth and following vertebra; the processes which represent it do not meet across 
the middle line. In the Caithness Cachalot the canal terminates at the eleventh caudal, 
in the Yorkshire specimen at the twelfth. In the anterior vertebrae (with elongated 
bodies) the neural laminae arise from little more than the anterior half of the body, but 
posteriorly, as the bodies decrease and the laminae actually increase in the antero- 
posterior direction, they come to occupy nearly the whole length of the upper surface 
of the centrum. This is best seen in the ninth and tenth. In the first caudal vertebra 
the metapophyses form prominent flattened wing-like processes, projecting forwards 
and outwards from the front border of the neural spine, having a deep groove between 
them for the reception of the hinder edge of the spine of the last lumbar vertebra. 
Their upper and anterior edges are chickened, rough, and slightly everted. They are 
placed about halfway between the upper surface of the centrum and the highest part 
of the spine. In the five following vertebrae their height from the centrum remains 
almost precisely the same ; but the spine gradually diminishes, so that in the sixth 
its upper edge comes to be on a level with these processes; at the same time they 
alter in character, becoming shorter and thicker, and terminating in a nearly cir- 
cular, flat, but very tuberculated or rugged surfece, looking upwards and forwards ; 
after the third they are so truncated and placed so near together upon the anterior 
thickened edge of the neural spine, as to have scarcely any groove between them, 
and to have quite lost the " clasping "' character they exhibit in the lumbar region. 
In the seventh caudal vertebra they have disappeared altogether, in consequence of 
the diminution of the height of the spine, or are only slightly indicated in the laterally 
thickened anterior extremity of the spinous process. Thereafter the spinous processes 
are of a very simple nature, compressed from side to side, elongated from before back- 
wards, and truncated above. The eleventh is less compressed ; the twelfth of quite 
another form, broad and flat above, the sides meeting at a very open angle ; the thir- 
teenth a mere irregular low tuberosity, perforated by a small canal -^g" high by ■^" 
wide. In the fourteenth and fifteenth the canal is much smaller in front than behind ; 
in the last it is not so large as a goose-quill, and bridged over only for the space of 
half an inch. In the remaining vertebrae the upper surface of the bone has four 
tubercles, arranged in pairs — two near the front and two near the hinder edge, the 
anterior pair being the largest. Traces of these can be discovered as far as the penul- 
timate vertebra. 

In the anterior vertebrae of the series the transverse processes project outwards 
with an inclination slightly downwards and forwards, from rather below the middle 
of the side of the body. They soon diminish in length, and increase in width from 
before backwards. In the ninth vertebra they are reduced to roughened longitudinal 
ridges ; in the tenth they have entirely disappeared, and tlieir place is taken by a 
slight groove. In the Caithness Cachalot the transverse process is rudimentary in the 

3b 2 ' 



344 MB. W. H. FLOWER OX THE OSTEOLOGY OF THE SPEEM-WHALE. 

tenth, and absent in the eleventh caudal vertebra ; in the Yorkshire skeleton it is 
indicated by a ridge in the eighth, and is completely wanting in the ninth ; so that in 
this respect the Tasmanian specimen holds an intermediate position between the two 
northern ones. 

On the hinder edge of the transverse process of the fifth caudal vertebra, close to its 
origin from the body, is a deep notch, for the passage of a blood-vessel ; in the two fol- 
lowing vertebrse this notch increases in depth, being continuous with a vertical groove on 
the side of the body of the bone. In the eighth the notch is enclosed so as to form a 
large o^al foramen -i%" by xo" in diameter, perforating the base of the transverse process, 
rather behind the middle of the centrum. In the ninth vertebra the foramen is placed 
nearer the median line, and therefore still further enclosed in the bone. It is here and 
in the succeeding vertebra nearly midway between the anterior and posterior extremity. 
This foramen is an important feature in all the remaining caudal vertebrae, being found 
even in the penultimate, and presents several peculiarities worthy of note. As before 
said, it first appears as a distinct foramen in the eighth vertebra ; or it may rather be 
described as a broad vertical groove on the side of the centrum, bridged over for a space 
of 2i" by the base of the short transverse process. In the tenth vertebra it has become 
a canal, 7" in length ; but the lower opening is still placed on the side of the body of 
the vertebra. In the eleventh the lower opening is placed at the angle between the 
side and the inferior surface, 5" from the opening of the canal of the opposite side. In 
the twelfth they have moved toward the middle of the inferior surface, being separated 
from each other only by a harrow bony septum (less than 1" in width). In the next 
tlu'ee vertebrse the septum is still narrower ; in the sixteenth it suddenly widens to 2" ; and 
thence onward the inferior openings are gradually placed rather furtlier from each other, 
until in the last few vertebra; the canals have gained the lateral position they possessed 
on their first appearance. In the meanwhile the upper orifices of these canals, advancing 
up the sides of the bodies of the vertebra?, soon come to be placed on each side of the 
middle of the upper surface, and maintain this situation to the end, only approaching 
nearer to each other as the vertebrae diminish in size. 

The articular surfaces for the chevron bones are remarkably large and prominent, 
with elevated edges, and rough cup-like depressions in tlieir middle. In many of these 
depressions a small, loose, irregular epiphysial plate is lodged. The first caudal ver- 
tebra has a pair of subcircular facets at its hinder edge, of which the left is considerably 
the larger, though somewhat less prominent ; to these the first (ununited) hsemapophyses 
fit closely. In the Caithness Cachalot, these bones are ankylosed to the body of the 
vertebra, though not meetmg each other at their free ends. In the second vertebra, 
the liinder facets are of larger size ; but none ai'e developed on the anterior edge. In 
the third, the facets on the anterior edge are slightly indicated, the posterior ones have 
increased still more. In the following nine vertebrae, the two pairs of facets are well 
developed, the anterior ones at first comparatively small, but towards the end of the 



JIR. W. II. FLOWER ON THE OSTEOLOGY OF THE SPERM-WHALE. 345 

sei'ies almost equalling the hinder ones in importance. On the thirteenth vertebra arti- 
cular facets are faintly ind'cated on the anterior edge only. 

Having now examined the modifications of the different parts of the caudal vertebrte, 
their bodies, neural arches, transverse processes, arterial foramina, and articular facets 
for Iniemapophyses, a few words must be said on the peculiarities of the different vertebrse 
of the hinder part of the series taken as a whole. 

The first twelve, or chevron-bone-bearing vertebrae, present a gradual and steady 
change in size and general character from first to last, this change consisting of the 
gradual reduction in size of the centrum, reduction of the neurapophysis, and reduction, 
to absolute disappearance, of the transverse processes. In the thirteenth caudal vertebra, 
a great change takes place in the size of the body, though the same general form is 
maintained. The anterior and posterior surfaces are nearly subcircular ; but the absence 
of any transverse processes, and the presence of a roughened low tuberosity representing 
the neurapophysis, and of the inferior tuberosities for the attachment of the hsemapo- 
physes, make the whole bone higher than broad. The most remarkable feature about 
this vertebra is the rapid manner in which it diminishes posteriorly, the posterior sur- 
face for the attachment of the inter^-ertebral substance being 2" less in diameter than 
the anterior. 

Fis. n. 





Twc-ltth and seventeenth caudal vertebra;. 

The succeeding vertebra is of very simple character, being reduced to a centrum with 
a slight tubercle covering the neural canal to represent the spmous process ; in conse- 
quence of this, the whole bone is still slightly higher than broad. After this a sudden 
change in form takes place ; in the fifteenth and all the succeeding vertebrae the breadth 
predominates over the height. The fifteenth to the twentieth inclusive present very 
uniform characters. They may be described generally as transversely elongated oblong 
bodies, flattened before and behind, above and below, and at each side. The upper 
and under surfaces are remarkably alike, so much so that it requires close attention to 
very minute characters to be able to distinguish them. The upper surface shows on 
each side, about midway between the median line and the outer edge of the bone, the 
funnel-shaped opening of the large circular arterial canal, which perforates the vertebra 
nearly vertically. Tiie quadrangular space between these perforations bears at each 
corner four tuberosities, an anterior and a posterior pair, remnants of tlie neurapophysis, 



346 MK. AV. H. FLOWEK OX THE OSTEOLOaT OF THE SPERM-WHALE. 

divided longitudinally by the narrow groove representing the neural canal, and cut, 
as it were, in two in the opposite direction by a wider groove connecting the two 
lateral, vertical, vascular canals. Of these four tuberosities, the anterior are larger 
than the posterior pair. In the first vertebra of the series now being described, the 
longitudinal groove is bridged over by a narrow bar of bone, and the anterior and pos- 
terior tuberosities of each side are confluent, but having a canal running under them 
connecting the vascular canals. In the second there is no connexion across the middle 
line; but the anterior and posterior tuberosities unite across the groove on the right 
side. 

The under surfaces also present four tuberosities in the corresponding situation, but 
placed slightly further from the middle line, and showing less dissimilarity in size 
between the anterior and posterior pair. They very much resemble in form and situa- 
tion the processes which bear the h;emapopliyses in the anterior caudal vertebrae. 
The lower openings for the vascular canals, which in the first vertebra of this group 
are placed so near as not to allow of an interval of more than ^" between them, in the 
next are 2" apart, and then gradually become somewhat more distant proportionally 
to the size of the vertebrae. As their upper ends remain in the same position, the 
canals, from being quite vertical, gradually acquire an upward and inward direction. 
Along the middle of the lateral surface of all these vertebrse is a strongly marked longi- 
tudinal groove, bounded above and below by large rough tuberosities, of which the 
upiier one is rather the more prominent, especially at its anterior corner. This peculiar 
form of the side is one of the most characteristic features of these vertebrae, and begins 
suddenly in the first of tliis group. 

It now only remains to speak of the special peculiarities of the last four caudal ver- 
tebrae. If we take any one of these, say the third from the end (the twenty-second 
caudal), we shall find that it consists of much more than a simple centrum, the part 
which often is alone represented in this region ; but to recognize in it any of the 
ordinary elements of a typical vertebra is by no means easy. We can, however, 
trace a gradual modification from those last described to the one which terminates 
the whole series. Excluding for the present this last named, the other three dimi- 
nish rapidly in breadth, though but slightly in length. The anterior and posterior 
surfaces, to which the intervertebral substance is attached, are deeply hollowed. All 
the processes gradually subside, with the exception of the anterior part of the upper 
lateral tuberosity, which obtains an increased development, forming a distinct process, 
projecting upwards and forwards, from the anterior upper angle of the bone. The 
vertical canals continue ; but their lower openings (which were already somewhat further 
apart in the hinder vertebrae of the la<t group), have, as it were, cut their way out of 
the bone, and form deep grooves on the sides, before perforating the upper lateral pro- 
cess, much in the same manner as in those vertebne wliere these holes first made their 
appearance. The penultimate vertebra is so much elongated, and constricted in the 



MR. W. H. FLOWER ON THE OSTEOLOGY OF THE SPER:\I-WHALE. 347 

middle by these grooves on each side below, and by the one connecting the two fora- 
mina (here raucli reduced in diameter) on the upper surface, tliat at first sight it looks 
like two vertebree which have coalesced, as not unfrequently occurs in this region among 
true Whales. The last vertebra is a rough, broad, oval nodule of bone, truncated in 
front, obtusely pointed behind, flattened from above downwards. Traces of the vascular 
grooves are to be detected on its sides. The contiguous surfaces of this and the penul- 
timate are both concave ; but tlie irregularities of their prominent edges correspond 
exactly, and they have evidently been in close apposition. 

The last twelve vertebrae of the tail of another Cachalot, sent to the Museum by 
Mr. Crowther in 1866, resemble those of the skeleton in their general characters, 
and evidently correspond vertebra for vertebra, but witli considerable individual de- 
viations. They are, in the first place, all rather shorter in proportion to their breadth. 
The entire length of the twelve placed in apposition is 38", while the twelve cor- 
responding vertebrae of the skeleton measure 42", the breadth of the former slightly 
exceeding that of the latter. The neural arch is not completely closed in by bone 
even in the first of the series. The penultimate is much shorter than that of the 
specimen described above. The terminal vertebra is altogether smaller, and does not 
present the broad, depressed character of the other, but is a simple rough subconical 
nodule. The first of the set had a perfect chevron bone attached to its hinder edge, 
b^" in greatest depth, and 4" in lengtli. The second had another, consisting of two 
distinct oval plates 2^" deep by 2^" long. There were no further traces of ha>ma- 
pophyses, which, from the state of the specimen, must have been preserved if they had 
ever existed. 

The Tasmanian, Sydney *, Caithness, and Yorkshire Cachalots resemble each other in 
possessing a distinct atlas, followed by six ankylosed cervical, and ten dorsal vertebra,- 
bearing well-developed ribs. They agree, moreover, in possessing eight true lumbar 
vertebrse. The Sydney specimen, however, according to the published description, wants 
the vertebra intermediate between the dorsal and the lumbar series, found in the other 
three, and hence has one vertebra less in front of the first chevi-on bone. Of remain- 
ing or caudal vertebrae, the Tasmanian and Sydney specimens have twenty-four, the 
Yorkshire specimen (presuming it to be complete) twenty-three, while in the Caith- 
ness one the number cannot be stated with certainty. The Yorkshire and Sydney 
Cachalots, therefore, agree in the entire number of vertebrse bemg forty-nine, while 
the Tasmanian skeleton possesses fifty. These differences are so trifling that they 
may be the result of accident or individual peculiarity; they certainly by them- 
selves afford no assistance in discriminating between a northern and southern species of 
the genus. 

Nor do we find, on reverting to the special characters of the vertebrre, any m ue 

* Wall, 0^. cil. 



348 MK. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEKM-WHALE. 

certain indications of difFerence. Each of the three vertebral columns which have 
passed under more immediate observation shows certain peculiarities ; but what similar 
part of the organization of any given species of animal will not do so 1 And it must be 
remembered that, in these huge bones, differences of form and proportion are rendered 
most conspicuous which would almost escape observation in specimens of the size of 
those with which we are more accustomed to deal, and, further, tliat the ossification in ' 
the Sperm-Whale, more even than in other large Cetaceans, seems to have a special 
tendency to exuberant and irregular development, producing a great amount of indivi- 
dual character in the rugged masses composing the skeleton. 

It may be as well, however, to bring together the principal points of difFerence, such 
as they are, in the three skeletons. The first to be mentioned relates to the length of 
the column when the vertebra; are placed in close apposition. This is, in the Yorkshire 
one, 29' 5", in the Caithness one (allowing for the missing caudal vertebrcc) 29' 3", in the 
Tasmanian one 30' 4". It must be remarked that the last, though the longest, is the 
least mature of the three animals. In the first two the correspondence is exceedingly 
close. With regard to the third, something must be probably allowed for the fact 
that the loose epiphyses of the ends of the vertebrae, having become detached in 
maceration, could not be made to fit again so closely as they would if ankylosed ; and 
hence the length of each vertebra was increased to a very slight extent. The differ- 
ences in the proportions of the individual vertebrae are given in the Table at p. 327. 
They are not inconsiderable, but are as great in the case of the two British specimens 
as between either of these and the Tasmanian one. The greater height of the lumbar 
vertebrae of the two former as compared with the latter, is chiefly due to the increased 
development of the keel of the body, apparently a consequence of superior age. Slight 
differences in the atlas, chiefly relating to the form of the lower part of the central 
opening, and to the development of the transverse processes, have been already spoken 
of in the description of that bone. 

In the annexed Table I have given the vertebral formula of the principal members 
of the group of Delphinoid Cetaceans, as far as I have been able to ascertain them from 
perfectly reliable sources. 

Phy.skterid.e. 

Physeter macrocephalus australis (Mus. Roy. Coll. Surg.), C. 7, D. 11, L. 8, C. 24=50. 
P. macrocephalus australis (Wall), C. 7, I). 10, L. 8, C. 24=49. 
P. macrocephalus borealis (Yorkshire), C. 7, D. 11, L. 8, C. 23=49. 
Kogia cjrayii (Wall), C. 7, D. 14, L. C. 30 = 51. 

Hyperoodon rostratum (Mus. Oxford and Vrolik), C. 7, D. 9, L. 10, C. 19=45. 
H. rostrahim (Amsterdam and Roy. Coll. Surg.), C. 7, D. 9, L. 10, C. 18 = 44. 
Micropteron sowerbyense (Van Beneden), C. 7, D. 10, L. 10, C. 19 = 46. 
Ziphius cryptodon (Burmeister), C. 7, D. 10, L. 12, C. 20 = 49. 



MR. W. H. I'LOWEB OX THE OSTEOLOGY OF THE SPERM-WHALE. 349 

Platanistid^. 

Platanista gangetica (Eschricht), C. 7, D. 11, L. 8, C. 25=51. 
Inia geoffrensis (Brit. Mus.), C. 7, D. 13, L. 3, C. 18=41. 
Pontoporia blainvillii (Burmeister), C. 7, D. 10, L. 7, C. 18=42. 

Delphiis'id^. 

Beluga leucas (Mus. Louvain), C. 7, D. 10, L. C. 33=50. 

Monodon monocews (Mus. Roy. Coll. Surg.), C. 7, D. 11, L. G, C. 26=50. 

Phocmna communis (Mus. Leyden), C. 7, D. 13, L. 14, C. 30 = 64. 

P. communis (Mus. Roy. Coll. Surg.), C. 7, D. 13, L. 14, C. 30=64. 

P. communis ? (Mus. Roy. Coll. Surg.), C. 7, D. 13, L. 14, C. 32=66. 

P. communis S (Mus. Oxford), C. 7, D. 12, L. C. 46=65. 

Neomeris phoccenoides (Mus. Leyden), C. 7, D. 13, L. 13, C. 30=63. 

Orca gladiator (Mus. Leyden), C. 7, D. 11, L. 10, C. 23=51. 

0. gladiator (Mus. Cambridge), C. 7, D. 12, L. C. 33=52. 

Psetidorca crassidens (Reinliardt), C. 7, D. 10, L. 9, C. 24=50. 

Globiocephalus melas (Mus. Roy. Coll. Surg.), C. 7, D. 11, L. 12, C. 28=58. 

G. melas (Middlesex Hosp.), C. 7, U. 11, L. 13, C. 28=59. 

G. ] (Tasmania, Mus. Roy. Coll. Surg.), C. 7, D. 11, L. 14, C. 27=59. 

Delphinus sinensis (Mus. Roy. Coll. Surg.), C. 7, D. 12, L. 10, C. 22=51. 
I), guianensis (Van Beneden), C. 7, D. 12, L. 14, C. 22 = 55. 
D. tursio (Mus. Leyden), C. 7, D. 13, L. 17, C. 25=62. 
D. heavisidii (Mus. Leyden), C. 7, D. 13, L. 15, C. 30 = 65. 
B. superciliosus (Mus. Leyden), C. 7, D. 13, L. 23, C. 30=73. 
D. delphis (Mus. Roy. Coll. Surg.), C. 7, D. 13, L. 24, C. 31 = 75. 
Lagenorhjnchus leucopleurus (Mus. Leyden), C. 7, D. 15, L. C. 59=81. 
L. alUrostris (Mus. Cambridge), C. 7, D. 14, L. C. 67=88. 

It will be seen from this Table that the total number of vertebrse varies extremely 
in different genera, and even in species of some natural genera, the variation being 
chiefl)- in the lumbar and caudal regions. Tlie well-marked group Physeterida-, com- 
prising the five genera placed at the head of the list, are all characterized by rather a 
small number of vertebrae ; but among the other families there are many which have 
quite as few as some of the former. For instance, the vertebral formulae of Platanista, 
Beluga, Monodon, and Orca closely resemble that of Physeter. It is singular that the 
most nearly allied genus, Kogia, differs so much in the number of dorsal vertebrae. On 
the whole, little reliance can be placed on these numbers for classificatory purposes, 
as shown by the extraordinary excess to which they run in Lagenorhynchus, a form 
otherwise not possessing any special modification. 

The characters of the cervical region of the genus Physeter have already been shown 
to be quite peculiar to itself. The dorsal region presents a remarkable disposition of 

VOL. VI. — PART VI. 3 c 



350 



JIE. W. H. FLOWER ON THE OSTEOLOGY OE THE SPERM-WHALE. 



the transverse processes, which points to a close affinity with Hyperoodon and the 
Ziphioids. The lumbar and caudal regions also, in some characters, especially their 
short transverse processes, approximate to those of the same forms ; the spinous pro- 
cesses, however, fall far short of the excessive height, and the bodies of the vertebrse 
present but little of the peculiar elongation, so characteristic of those groups. 

Chevron Bones. 

Ten chevi'on bones were sent with the skeleton ; but it would appear from the distance 
to which these bones extend backwards, as ascertained with certainty on the detached 
tail described at p. 347, that fourteen is the complete number. By a careful comparison 
of the articular surfaces on the vertebrse with those on the diiFerent chevron bones, the 
missing ones were ascertained to be the fifth, sixth, and the thirteenth and fourteenth. 

The first (See PI. LIX.) is represented by a pair of small styliform bones. They are 
not quite alike, the right being rather the longest, and the left much the thickest. 
They both end in a point, and are slightly hollowed on their inner side, and flattened 
externally. They fit most closely, by an expanded, rough, subcircular base, to the corre- 
sponding facets on the hinder margin of the body of the first caudal vertebra. The 
dimensions of these bones are : — 





Right. 


Left. 


Length ,' 


inches. 
0" 
2"-2 
l"-9 
l"-3 


inches. 
5"-2 
2'--8 
2"-3 
l'-5 




Diameter at middle, aiitcro-posterior 







From the Yorkshire skeleton these bones are absent. In the Caithness specimen 
they are more massive and irregular in shape, and are completely ankylosed to the body 
of the vertebra. Their free extremities diverge from each other and project strongly 
backwards. 

In the succeeding bones (up to the fourteenth) the two lateral lamina; are united 
below, and form a prominent spine. The second is comparati^■ely small and narrow, 
and has the spine slightly developed. In the third the spine is very large, both long 
and broad, and truncated below. In the fourth it is somewliat shorter, but more mas- 
sive and rounded at the lower end. The entire length of the second bone is lOf", of 
the third 21", and of the fourth 20". The lower edge of nearly all the remaining bones 
has been cut off by a sharp instrument in preparing the skeleton ; so I am not able to 
give their correct length or configuration. The surfaces for contact with the vertebrce 
are expanded, massive, rough, and hollowed ; and many of them have imbedded within 
them separate epiphysial ossifications, more or less ankylosed either to the chevron 
hone or to the body of the vertebra. 

In both the Caithness and Yorkshire skeletons the number of the chevron bones is 



MR. W. H. FLOWER OX THE OSTEOLOGY OP THE SPERM-WHALE. 



M51 



obviously incomplete ; the former has only eleven, and the latter ten. Those that are 
present differ very notably in form and dimensions both from each other, and from 
those of the Tasmanian skeleton ; but this is the most unsatisfactory part of the whole 
exammation, owing to the imperfection of the materials and the difficulty, in incomplete 
sets, of ascertaining with precision which are the corresponding bones. 

Bibs. 

The ribs, though tolerably long, are (with the exception of the first) slender and light 
for the size of the animal, at least when compared with those of a Balmna of correspond- 
ing magnitude. This character is best illustrated by stating that in the skeleton of the 
nearly adult Balcena mysticetus, in the Mus. of Koy. Coll. of Surgeons, 46' in length, 
the twelve pairs of ribs weighed 7 cwt. 60 lbs. ; in the Cachalot, several feet longer 
and of corresponding age, the eleven pairs weigh less than 3 cwt. Nevertheless their 
tissue is dense and compact. 

It is stated by Wall that the ribs of the left side are of larger dimensions than the 
corresponding ones of the right. In order to ascertain whether there is a similar want 
of symmetry in the specimen under consideration, I weighed all the ribs; and the 
result shows considerable individual differences m corresponding ribs, and a very trifling 
general preponderance of the left side over the right — the total weight of the ribs of 
the right side being 163 lbs. 9i oz., and those of the left side 164 lbs. 5^oz. 



Weight and Measurements of the Eibs (excluding the rudimentary eleventh pair). 





Weight. 


Extreme 

length in 

straight line. 


Circumference 
at middle. 


Circumference 

at inferior 
extremity. 


Curre* 


, f R 


lbs. oz. 
27 6 
27 4 
25 13 
25 1 
25 
25 13 
21 8 
23 2 
19 8 
21 7 
19 5 
19 7 
17 7 
16 15 
15 3 
13 15 
10 6 
10 5 
9 7i 

8 ^ 


in. 

45 

47 

63i 

63 

70 

7U 

71 

74 

68 

72 

67i 

68i 

651 

66| 

63 

65 

61 

63 

56 

55| 


in. 
14 
14 
111 

H 

H 
H 

H 

8j 

82 

8 

61 

7 

6| 


in. 

29 

28 

201 

20 

m 

18 

16 

161 

14 

15 

12| 

12 

10 

8| 

^ 

f 


in. 

101 
11 
22 
21 

28 

27 

29 

29 

30 

29 

29 

28 

26 

25 

22 

22| 

19 

18 

11 

Hi 


i{l 


„ f R 


2 ■ L. 


^ R 


3 L 


, R 


Ml. .;..::;:..:: 


.JR 

5 L 


A-R, 


Ml. ..::::..;,...... 


„ f R 


''■L 


,, f R 


S • L 


„ f R 


9 L 




^^{l: ■.::;:::::::::::: 



* Taken by standing the rib with its two ends on the floor, and measuring from the latter to the interior of 
the highest part of the arch. 

3c2 



352 ME. W. H. FLOWEE ON THE OSTEOLOGY OF THE SPEEM- WHALE. 

'J'he first rib (PI. LX. fig. 2) is very diiferent from the others in form. Though 
by far the shortest, it weighs absolutely more than any of the others, this being 
(jccasioned by its great thickness, especially towards the inferior end. It is very broad 
throughout, but much compressed from before backwards. At the angle it bends very 
abruptly, the main part of the rib, or that below this point, being almost straight. The 
inner concave border is very sharp, the outer one more rounded. The tubercle is very 
largely developed, forming a great expanded rugged crest at the upper extremity of 
the bone, by which it articulates with the end of the transverse process of the first 
dorsal vertebra. The capitular process is quite rudimentary, being a slight angular 
projection marked ofi" from the tubercle by a shallow depression. It may have had a 
ligamentous union with the rough elevation near the hinder border of the body of the 
seventh cervical vertebra; but there must have been an interval of fully 6 inches 
between them. At the lower end the whole bone is twisted on itself, the anterior 
surface turning outwards, and the inner edge forwards. The size of the bone also is 
increased greatly, not only in breadth, but in thickness — the last alteration mainly 
affecting the inner or anterior part, so that the sharp edge previously mentioned 
becomes quite rounded, and the surface of the truncated inferior extremity is mucli 
wider at its inner than its outer end. 

The second rib partakes somewhat of the massiveness of the first, though it is con- 
siderably longer, and of very diff'erent general form. Its curve is tolerably regular, 
though the principal bend is, as usual, in the neighbourhood of the angle. It is 
flattened from before backwards, and has a sharp internal edge. The tubercle is 
much reduced in size, and presents a broad oval surface for articulation with the 
transverse process of the second dorsal vertebra. The neck is a well-developed com- 
pressed process, tapering towards the apex, where it is somewhat dilated, and must 
liave reached very nearly, if not quite, to the small articular surface on the side of the 
body of the first dorsal vertebra. A very prominent crest extends backwards from 
the tubercle, terminating rather abruptly at the angle. This distinguishes the second 
rib from the first, as well as from those that come after it in the series. The outer 
surface is much more regularly convex than that of the first rib. The anterior surface 
gradually becomes external towards the lower end ; and the expansion of this part (less 
marked than in the first) affects chiefly tlie middle of the bone, the truncated end pre- 
senting a regularly oval surface. 

The thu'd rib is longer, but thinner than the second. Its curve is rather wider ; it is 
less compressed and wants the prominent sharp inner concave border, and also the crest 
above the angle ; its tubercle is less rounded, but rises into a sharp angular process 
surmounting the flat inwardly turned articular surface ; its capitular process is longer 
and more dilated at the end. The inferior extremity is still considerably enlarged, but 
less than in the last. 

The fourth, fifth, and sixth ribs closely resemble the third, and each other; but the 



ME. W. H. FLOWER ON THE OSTEOLOGY OF THE SPEEM-WHALE. 353 

following changes may be obseiTed gradually occurring in them. They become more 
slender and rounded in the middle part of the shaft. The capitular process becomes 
slightly shorter, but at the same time thicker ; and the capitulum itself becomes larger 
and more irregular in shape. The tubercle becomes rather less prominent. The 
inferior end of the rib presents a less marked enlargement, and is more regularly 
cylindrical. 

Each of these ribs articulates in the usual way by the tubercle with the transverse 
process of the corresponding dorsal vertebra, and by the head vnth the facet on the 
hinder edge of the body of the preceding vertebra. The last two may also have had a 
slight connexion with the body of the corresponding vertebra. 

The seventh rib has a shorter but very thick neck, the capitular surface of which is 
produced backwards, so that it articulates distinctly with the bodies of two vertebrae, 
chiefly, however, with the one in front of that to which its transverse process is 
attached. The articular surface of the tubercle is much elongated in the fore-and-aft 
direction. 

The eighth repeats much the same condition at the upper end ; it may be distin- 
guished, however, by its greater slendemess, especially at the lower end, which presents 
scarcely any appreciable enlargement. 

The form of the upper end of the ninth rib is very characteristic. The neck is still 
shorter than in the last, and very obliquely truncated ; but its articular surface is large, 
and, contrary to what obtains in the preceding, particularly so at the hinder part, as it 
articulates chiefly with the raised facet on the body of its own vertebra. The tubercle 
is large and overhangs the posterior hollowed surface of the capitular process. Below 
the tubercle the rib is very slender, and has a slight twist backwards. The lower part 
is flattened from without inwards, and has a prominent sharp ridge on the posterior 
margin. 

The tenth rib is considerably shorter than the ninth. The capitular process is 
entirely absent; but the expanded upper end, which articulates only with the end of 
the transverse process of the tenth dorsal vertebra, is divided by a vertical groove into 
two unequal surfaces, of which the anterior larger one represents the base of the neck, 
and the smaller posterior one the tubercle. The lower end is flattened and twisted 
somewhat backwards. 

The eleventh rudimentary pair of ribs are nearly straight, the longer one having a 
slight sigmoid curve. They are largest at their attached extremity, and gradually taper, 
but are slightly enlarged again at the tip. In the greater part of their length they arc 
flattened from above downwards ; but, having a slight twist on their own axis, the longest 
one becomes towards its distal extremity flattened almost in the opposite direction. The 
proximal ends are truncated, having an oval, slightly convex articular surflice, which 
was tipped with cartilage. These bones offer few characters indicating the side of the 
body to which they respectively belong. One (probably the right) is 13f" in length, 



354 MR. W. H. TLOWEE ON TliE OSTEOLOGY OF THE SPERM-WHALE. 

and 2f" in greatest diameter at the proximal end ; the other is 11" in length, and 2|" 
in diameter at the same point. 

The ribs of the Caithness and Yorkshire skeletons agree in their number and general 
characters with those of the present specimen. In the former, the two rudimentary 
eleventh ribs are preserved, and are 13^" and 12|" long respectively, and somewhat 
stouter than in the Tasmanian Cachalot. In the skeleton at Burton Constable they are 
not present; but the condition of the ends of the transverse processes of the coiTe- 
sponding vertebrae clearly indicate then- former existence. In the Sydney skeleton, ten 
pairs of fully developed ribs are described. The small eleventh pair, if they existed, 
may easily have escaped observation, especially as the animal was scarcely more than 
half-grown. 

The skeleton at Paris has fourteen paii-s of large ribs, with indications on the trans- 
verse processes of a fifteenth. Th